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Hyperolius nitidulus Peters, 1875 Hyperolius nitidulus Peters, 1875. ZMB 7729, Lagos, Nigeria. Colour in alcohol: Above violet-grey, a dark brown stripe from the nostrils, over the eye, to the flank, where it becomes many flecks against a pale background. Below this band, white with many small black flecks. Hyperolius pallidus Mertens, 1940. NHMW 16862, “Poli bei Garua, Nordkamerun” (Poli near Garua, Cameroon). Synonymy by Rödel (2000). Above pale grey with fine black speckling on the head and sides of females. Juveniles uniform pale, or with two to four pale longitudinal stripes. Hyperolius viridiflavus angeli Laurent, 1951. MNHNP 1921.34b, “Gribingui, Oubang-Chari” Central African Republic. The H. viridiflavus clade does not extend far west into the Central African Republic, and the type is here assigned to H. nitidulus. Dorsal colour uniform, with fine black speckling on side of head, tibia and lower leg, which may not be visible in some specimens. Hyperolius nitidulus aureus Perret, 1966. MHNG 1038.39 “Ngaoundér”, Cameroon. Golden-yellow above, with a dark brown stripe from the side of the snout over the eye to the vent with a black border, and a similar dark line along the tibia. Table 3 lists the localities of the synonyms. Sequences were available from Côte d’Ivoire, Benin, Ghana, and Cameroon (Fig. 8A). The minimum uncorrected p- distances from other clades were 3.3–7.2% (Table 2). Hyperolius nitidulus bangwae Perret, 1966 was removed from the synonymy of H. nitidulus and elevated to a full species. Rödel et al. (2010) showed that H. nitidulus was genetically different from H. spatzi, and confirmed its status, following earlier proposals (Wieczorek et al. 2001, Schiøtz 1967, Drewes 1984, and Rödel 2000). The 16S rRNA sequences from Rödel et al. (2010) were used to identify this clade. The localities of taxa regarded as junior synonyms of H. nitidulus are listed in Table 8. The type locality of Hyperolius nitidulus Peters, 1875 is Lagos, Nigeria. Although there were no sequences available from Lagos or anywhere else in Nigeria, this clade consists of sequences that were identified in the field as H. nitidulus or H. viridiflavus nitidulus, and that occurs west and east of Nigeria. Only two clades (H. nitidulus and H. spatzi) are known from West Africa. Portik et al. (2019) recognised two sub-groups, one in Cameroon and the other in Ghana. Although the Cameroon sample is monophyletic, and has about a 2% difference from the Ghana samples, it is not reciprocally monophyletic from the samples from Benin, Ghana, Nigeria and Côte d’Ivoire. The most common pattern elements in the descriptions of the types included in H. nitidulus are a dark lateral band (50%), and a uniform back (50%). The percentages are not additive, as the pattern elements may be shared. The advertisement call (no voucher) is described as ‘a single, pure, metallic and very loud tone’. Mean call length is 0.08 s, at a mean frequency of 2638 Hz (Rödel et al. 2010)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 334-335, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Rodel, M. - O. (2000) Herpetofauna of West Africa. Vol 1 Amphibians of the West African Savanna. Edition Chimaira, Frankfurt am Main, 332 pp.","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie, 93, 289 - 464.","Rodel, M. - O., Sandberger, L., Penner, J., Man, Y. & Hillers, A. (2010) The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae). Bonn zoological Bulletin, 57, 177 - 188.","Schiotz, A. (1967) The treefrogs (Rhocophoridae) of West Africa. Spolia zoologica Musei hauniensis, 25, 1 - 346.","Drewes, R. C. (1984) A phylogenetic analysis of the Hyperoliidae (Anura): Treefrogs of Africa, Madagascar, and the Seychelles Islands. Occasional Papers of the California Academy of Sciences, 139, 1 - 70.","Portik D. M., Bell, R. C., Blackburn, D. C., Bauer, A. M., Barratt, C. D., Branch, W. R., Burger, M., Channing, A., Colston, T. J., Conradie, W., Dehling, J. M., Drewes, R. C., Ernst, R. Greenbaum, E., Gvozdik, V., Harvey, J., Hillers, A., Hirschfeld, M., Jongsma, G. F. M., Kielgast, J., Kouete, M. T., Lawson, L. P., Leach, A. D., Loadcer, S. P., Lotters, S., Van der Meijden, A., Menegon, M., Muller, S., Nagy, Z. T., Ofori-Boateng, C., Ohler, A., Papenfuss, T. J., Rossler, D., Sinsch, U., Rodel, M. - O., Veith, M., Vindum, J., Zassi-Boilou, A. - G. & McQuire, J. A. (2019) Sexual dichromatism drives diversification within a major radiation of African amphibians. Systematic Biology, 68, 859 - 875. https: // doi. org / 10.1093 / sysbio / syz 023"]}

Hyperolius mariae Barbour & Loveridge, 1928 Rappia vermiculata Pfeffer, 1893. Syntype ZMB 10988 (other ZMB syntypes destroyed) “Sansibar (= Zanzibar, Tanzania) The name is preoccupied by Hyperolius vermiculatus Peters, 1882, and so is not available, although it appears to be the oldest name for this taxon. Above brown with many small yellow flecks. A dark band runs from eye to snout tip. Eyelid very dark with thin yellow margin. Hyperolius mariae Barbour & Loveridge, 1928. MCZ 13267, “near the mill dam at Derema, Usambara Mtns., Tanganyika Territory ” Tanzania. Colour in life: Above uniform greyish-white, with a black speck on snout, nostrils ringed with black, edge of upper eyelid black. A broad cream-coloured stripe laterally with a broad black one below it. Lower lip with red marks, belly blood-red. Hyperolius callichromus paratypes from Dar es Salaam ZMB 85869–85872 (Tillack et al. 2021), are assigned as synonyms of H. mariae as this is the only clade known from Dar es Salaam. Hyperolius vermicularis Ahl, 1931. ZMB 10988, Zanzibar, Tanzania. (Replacement name for Rappia vermiculata Pfeffer, 1893). Only the H. mariae clade is known from Zanzibar. Sequences from the adjacent mainland form part of the clade. Above brown with very many small yellow flecks, forming vermiculation. A dark band from snout tip to eye, eyelid with a thin yellow margin, tibia with small dark speckles. Hyperolius melanopthalmus Ahl, 1931. ZMB 85670 –85672, “ Sansibar ”, (Zanzibar, Tanzania). Synonymy by Loveridge (1936). Colour in alcohol: Above pale grey, nostrils ringed with black, upper eyelid with dark fleck. A broad lateral dark or violet band between the limbs, sometimes marbled. Hyperolius rubripes Ahl, 1931. ZMB 36110 and 57530, “Kililana”, Kenya. Synonymy by Wieczorek et al. (2000). Colour in alcohol: Above uniform pale yellow, pink or brownish. Hyperolius renschi Ahl, 1931. ZMB type lost (Tillack et al. 2021) Zanzibar, Tanzania. Synonymy by Loveridge (1936). Colour in alcohol: Above pale yellow, with a lateral violet band. Hyperolius marmoratus mafianus Laurent, 1961. BMNH 1937.7.1.42–44, This is a new name for specimens of H. albifrons Ahl, 1931, from Mafia Island, reported on by Parker (1937). Only one clade is known from northern coastal Tanzania. Back and sides yellowish-brown. Upper surface of snout silvery-white or yellowish-green, extending back to the eyes, continuing as lateral stripes to the level of the shoulder. The lateral stripes may branch to form a dorsal marbling, or may unite over the coccyx.A dark subdermal lateral band is present. Limbs the same colour as the back. Hyperolius udjidjiensis Ahl, 1931 was removed from the synonymy of H. mariae and placed in the synonymy of Hyperolius parallelus Günther, 1858 by Wieczorek et al. (2000). Sequences were available from Mafia Island (as H. mariae and H. viridiflavus mariae) and Mtai Forest, Tanzania; Arabuko Sokoke, Kenya (as H. viridiflavus mariae and H. v. rubripes). The minimum uncorrected p- distances from other clades were 4.0–6.5% (Table 2). The sample occurs in coastal Tanzania and Kenya (Fig. 9C). The localities of taxa now regarded as junior synonyms of H. mariae are listed in Table 3. The most common pattern elements in the descriptions of the types included in H. mariae are various markings on the head such as black-ringed nostrils (50%), followed by a dark lateral band (33%). The percentages are not additive, as the pattern elements may be shared. Advertisement calls from Mikumi, Tanzania (no vouchers), show a series of brief notes, with an emphasised frequency of about 3200 Hz, and a note rate of 5 s-1 (Schiøtz 1975, 1999 as H. v. mariae). This is similar to another call (no voucher) from the same locality (Pickersgill 2007, as H. mariae). A call from Kingupira, Tanzania (no voucher) has an emphasised frequency of 2750 Hz (Schiøtz 1975, as H. marginatus), similar to calls from Karonga, Malawi (no vouchers) (Pickersgill, 2007, as H. nyassae) and Bububu, Zanzibar (no vouchers) (Pickersgill 2007, as H. renschi)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 339-340, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Loveridge, A. (1936) Scientific results of an expedition to rain forest regions in eastern Africa. VII. Amphibians. Bulletin of the Museum of Comparative Zoology, 79, 369 - 430.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Parker, H. W. (1937) Reptiles and amphibians from the Mafia Archipelago. Annals and Magazine of Natural History, Series 10, 20, 629 - 632. https: // doi. org / 10.1080 / 00222933708655396","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp."]}

Hyperolius dintelmanni Lötters & Schmitz, 2004 Hyperolius dintelmanni Lötters & Schmitz, 2004. ZFMK 67871 (Table 3), “ From the Edib Hills (ca. 1,200 m above sea level), Bakossi Mountains, South-West Cameroon (4° 57′ N, 9° 39′ E).” Colour in life, phase J: back olive to translucent dark green extending to limbs, with a pale green hourglass pattern. Phase F: back dark brown to black with greenish spots and bright red areas on upper legs, feet and hands. The dorsal surface has a goldenyellow dense marbling. The females may be brown with reddish speckles (Portik et al. 2016). Sequences were available from Edib Village, the type locality, Manjo, Mt Kupe, Mantem, Southwest, Littoral, West, all in Cameroon. The minimum uncorrected p- distances from other clades were 1.9–6.0% (Table 2). The clade was identified following Bell et al. (2017). It is known from eastern Cameroon (Fig. 10D). The most common pattern element in the description of the type of H. dintelmanni is dorsal marbling (100%), followed by green spots (50%) or a red speckle (50%). The percentages are not additive, as the pattern elements may be shared. The advertisement call is unknown., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 343, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Lotters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west- Cameroon. Bonner zoologische Beitrage, 52, 149 - 154.","Portik, D. M., Jongsma, G. F. M., Kouete, M. T., Scheinberg, L. A., Freiermuth, B., Tapondjou, W. P. & Blackburn, D. C. (2016) A survey of amphibians and reptiles in the foothills of Mount Kupe, Cameroon. Amphibian & Reptile Conservation, 10 [Special Section], e 131, 37 - 67.","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., Zassi-Boulou, A. G. & 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, 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260"]}

Hyperolius spatzi Ahl, 1931 Hyperolius spatzi Ahl, 1931. Lectotype ZMB 32602, “Bakel-Kidira (oberes Senegalgebiet)” Senegal (Table 3). Colour in alcohol: Above chalk-white or sometimes with fine dark brown stippling. Three sequences were available, all identified as H. spatzi. The minimum uncorrected p- distances from other clades were 3.4–6.5% (Table 2). The sequences all come from Senegal (Fig. 10C). Rödel et al. (2010) showed that H. spatzi was distinct from H. nitidulus using genetics, morphology and advertisement calls. The clade is identified as H. spatzi as the sequences derive from material collected by Rödel et al. (2010). This species is known from Gambia and Senegal, and its distribution was discussed by Rödel et al. (2010). The pattern elements in the descriptions of the type of H. spatzi are a uniform back (100%), or fine brown stipple (100%). The percentages are not additive, as the pattern elements may be shared. The advertisement call (no voucher) is described as ‘a single, pure, metallic and very loud tone’. Mean call length is 0.08 s, at a mean emphasised frequency of 2638 Hz (Rödel et al. 2010)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 342-343, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Rodel, M. - O., Sandberger, L., Penner, J., Man, Y. & Hillers, A. (2010) The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hyperolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoliidae). Bonn zoological Bulletin, 57, 177 - 188."]}

Hyperolius bangwae Perret, 1966 Hyperolius nitidulus bangwae Perret, 1966. MHNG 1039.2 “Bangwa”, Cameroon (Table 3). Back beige or brown sometimes with a dark lateral line, or with dark speckles. This clade consists of 50 sequences from Gabon, Equatorial Guinea, Republic of Congo and Cameroon (Fig. 9A), all reported in GenBank as Hyperolus tuberculatus. The sequence reported in Veith et al. (2009) as Hyperolius tuberculatus Complex B forms part of this clade. The uncorrected p- distances from other clades were 1.9–6.3% (Table 2). Two species occur at Bangoua in western Cameroon, H. nitidulus and a taxon previously identified as H. tuberculatus. Perret (1966) described Hyperolius nitidulus bangwae from Bangoua, and I remove it from the synonomy of H. nitidulus and regard it as a full species. The most common pattern elements in the descriptions of the type of H. bangwae are a dark lateral line (100%), and dark dorsal speckling (100%). An advertisement call from Monts Cristal (ZFMK 73143) consists of a series of brief notes at a mean note rate of 9.5 s- 1 with a mean emphasised frequency of 2271 Hz (Lötters et al. 2004, as H. tuberculatus). A specimen from the same locality was sequenced (AY329321) and identified as H. bangwae. The advertisement call attributed to H. tuberculatus from Binguéla, Cameroon (Schiøtz 1999) is within the range of H. bangwae as here envisaged, and is regarded as a call of H. bangwae. The call consists of a series of brief notes, about 13 s-1, with an emphasised frequency of 1900 Hz. This is quite different from the call of H. tuberculatus from Río Muni, Equatorial Guinea (no vouchers), as shown by Bosch et al. (2000). A call from Olemb, Cameroon (as H. tuberculatus) (no voucher) that consists of a series of simple notes with a note rate of 14 s-1, and an emphasised frequency of 3000 Hz (Amiet & Goutte 2017) is provisionally regarded as that of H. bangwae, although the call of H. sp. 1 is unknown., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 338, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Perret, J. - L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbucher. Abteilung fur Systematik, Okologie und Geographie, 93, 289 - 464.","Veith, M., Kosuch, J., Rodel, M. - O., Hillers, A., Schmitz, A., Burger, M. & Lotters, S. (2009) Multiple evolution of sexual dichromatism in African reed frogs. Molecular Phylogenetics and Evolution, 51, 388 - 393. https: // doi. org / 10.1016 / j. ympev. 2008.12.022","Lotters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west- Cameroon. Bonner zoologische Beitrage, 52, 149 - 154.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Bosch, J., De la Riva, I. & Marquez, R. (2000) Advertisement calls of seven species of hyperoliid frogs, from Equatorial Guinea. Amphibia-Reptilia, 21, 246 - 255.","Amiet, J. - L. & Goutte, S. (2017) Chants d'Amphibiens du Cameroun. J. - L. Amiet & Editions Petit Genies, Nyons, 280 pp."]}

Hyperolius noblei Ahl, 1931 Hyperolius noblei Ahl, 1931. ZMB 85765, “Kilwa, Deutsch-Ostafrika ” Tanzania (Table 3). Colour in alcohol: Uniform yellow-brown. A silver-white stripe runs from the side of the snout, over the upper eyelid to form a larger mark behind the eye. Sequences were available from Nhica and Palma, Mozambique (as Hyperolius cf. marmoratus), and Nhica do Rovuma (as Hyperolius marmoratus taeniatus), Mozambique. The two sequences from Nhica (MG829620 and MK509512) were derived from the same specimen (MNHNP 2010.185). The minimum uncorrected p- distances from other clades were 4.2–6.9% (Table 2). Presently this species is only known from southern coastal Tanzania and northern coastal Mozambique (Fig. 10A). The nearest described taxon to this clade is Hyperolius noblei Ahl, 1931. The description matches the specimens collected and sequenced from northern Mozambique (Table 1). The pattern elements in the description of the type of H. noblei are a uniform back, with a transverse silver stripe forming a large mark behind the eye. The advertisement call (no voucher) was recorded on the Afungi Peninsula at Palma, Mozambique (10.8456°S, 40.4732°E). The call is a long (0.08 s) whistle, frequency modulated, with the emphasised frequency at 3500 Hz. A typical call starts at 3000 Hz, and rises steeply in pitch to 3595 Hz in 0.03 s, then maintains that pitch for 0.05 s. The emphasised frequency varies in the population from 3500 Hz to 4565 Hz., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 340, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132."]}

Hyperolius marmoratus Rapp, 1842. Hyperolius marmoratus Rapp, 1842. Types unknown. “Natal”, KwaZulu-Natal Province, South Africa. Locality obscure. Back dark brown with irregular pale grey markings. Lower limbs flecked. Hyperolius verrucosus Smith, 1849. Types unknown. “Kaffirland to the eastward of the Cape Colony” (see comment above), South Africa. Locality obscure. Synonymy by Hewitt (1937). Back greyish brown variegated with spots, stripes or broken bars of pale sienna-yellow or cream-yellow. Extremities pale buff-orange, thighs speckled with dull olive-green. Inner surfaces of thighs reddish-orange. Hyperolius taeniatus Peters, 1854. ZMB 4531, Boror, Mozambique. Synonymy by Boulenger (1882). Yellow lateral stripes join at the snout. Legs silver with dark flecks, with a dark lateral band from snout to leg. Hyperolius sugillatus Cope, 1862. ANSP 11319, Umvoti, South Africa. Synonymy by Loveridge (1941). Back straw colour, with a yellow band from snout to sacral region, which has a dark purple line forming a lower border to the yellow band on the snout. A purple spot on each eyelid. Hyperolius citrinus Günther, 1864. BMNH 1947.2.9.61 and 1947.2.9.63. “Zambesi Expedition” Luabo River, Zambezi Delta, Mozambique. Synonymy by Laurent (1952). Lemon-coloured above and below. Hyperolius granulosus Peters, 1867. Syntypes ZMB 4811, ZMB 75652, “ Capanga am Flüsschen Mutizi östlich von Tette” Capanga, Mozambique. Synonymy by Laurent (1961). Back, sides and upper surfaces of limbs signal red with yellow vermiculation that shows occasional black pigmentation. Hyperolius variegatus Peters, 1882. ZMB 10249, 75602, “Cabaceia, Quelimane, Inhambane ”, Mozambique. Synonymy by Laurent (1952). ZMB 75603 –75604 were listed as types, but these are Afrixalus sp. (Tillack et al. 2021). Above dark brown with golden flecks and lines. Lips and ventral surfaces of limbs black with golden dots with red borders. Limbs blood-red, throat marbled in yellow. Hyperolius marmoratus pondoensis FitzSimons, 1930. TMP (now DNMNH) 13689. “Port St Johns, Pondoland Coast”, South Africa. Synonymy by Laurent (1952). Olive to chocolate-brown above, with irregular sized grass-green spots, darkly edged. Upper sides irregular and black edged. Ventral surfaces of limbs pink. Hyperolius breviceps Ahl, 1931. ZMB 86026, “Chemba”, Mozambique. Synonymy by Laurent (1952). Colour in alcohol: Back, sides and limbs chalk-white to pale grey-brown, with irregular small black or dark brown markings, some of which are rounded. An interrupted dark stripe on the side of the head and snout. Small dark dots on a white throat. Hyperolius albifrons Ahl, 1931. ZMB 36095. “Afrika (ohne genauern Fundort) [without precise locality]. Colour in alcohol: Back and sides yellowish-brown. Upper surface of snout silvery-white, extending back to the eyes, continuing as lateral stripes to the level of the shoulder. Limbs the same colour as the back. I follow Tillack et al. (2021) in retaining this taxon in H. marmoratus. Hyperolius dermatus Ahl, 1931. ZMB 85999, “Cabayra” (=Cabaceia?), Mozambique. The locality is obscure (Tillack et al. 2021). Synonymy by Laurent (1985). Colour in alcohol: Above chalk white with a yellow-brown tint that is more pronounced behind the head. Upper lip pale yellow. Yellow lateral band between the limbs. Hyperolius fuelleborni Ahl, 1931. ZMB 71184 –71186, “Neu-Helgoland” Papya Island, Malawi. Illustrated in Tillack et al. (2021, fig. 7). Other specimens included in the description by Ahl (1931) are placed as synonyms of H. parallelus because the H. marmoratus clade does not occur in the northern Lake Malawi area. Back pale grey with dark brown to black marbling. Sides of head without flecks. Hyperolius swynnertoni FizSimons, 1941. TMP 18911 and 18916, “Chirinda Forest”, Zimbabwe. Presently regarded as a good species. I follow the synonymy by Poynton & Broadley (1987). Dark vermiculation on a pale background, sometimes with pale stippling on a dark background. Hyperolius albofasciatus Hoffman, 1944. NMB A2622, “Umbe, Nyasaland ” (Malawi). Implied synonymy by Poynton & Broadley (1987). Dark brown back with three white bands running the length of the body. The middle band starts between the nostrils and joins with the lateral bands just above the vent. Femur pink, tibia above brown with narrow white bands. Arms pale with scattered brown spots. Hyperolius marmoratus broadleyi Poynton, 1963. NMZB 5370, “Fern Valley, Umtali” Zimbabwe. Back black with a regular white band from the snout tip to above the vent, and a lateral band from behind the eye to the groin. A thin red line present in the middle of each white band. Limbs with black and red markings. Hyperolius fumosus Pickersgill, 2007. LIV 1998.46.67 “Beira, Mozambique ”. It is assigned as a synonym as the colour pattern is part of the known variation. The phase F is silver-white with black speckling with the typical red or pink on the hidden parts of the limbs. The advertisement call (no voucher) has a duration of 64–90 ms, with an emphasised frequency of 2.6–3.8 kHz (Pickersgill 2007) which matches those along the South African east coast. Only a single clade, H. marmoratus, occurs along the coast including Beira. Nearly 40 taxa were previously placed in the synonomy of H. marmoratus (Frost 2021). The type locality is Kwa- Zulu-Natal, which agrees with the clade here assigned to H. marmoratus. Sequences were available from South Africa, Zimbabwe, eSwatini, and Mozambique. The minimum uncorrected p- distances from other clades were 4.0–6.1% (Table 2). Names applied to the sequences in this clade include H. viridiflavus, H. v. broadleyi, H. v. swynnertoni, H. v. marmoratus, H. v. taeniatus, H. v. verrucosus, H. v. albofasciatus, H. marmoratus, H. m. marmoratus, H. m. taeniatus, and H. swynnertoni. This clade occurs from the south coast of South Africa, to eSwatini, Zimbabwe, Malawi and Mozambique (Fig. 5). See Table 3 for synonym localities. The following extralimital taxa were removed from the synonymy of H. marmoratus and placed as synonyms of H. parallelus: Hyperolius vermiculatus Peters, 1882 (from Angola); Hyperolius plicifera Bocage, 1893 (after Perret, 1976); Rappia rhodoscelis Boulenger, 1901 (from the DRC); Hyperolius nossibeensis Ahl, 1931 (from Angola); Hyperolius decoratus Ahl, 1931 (from Angola); Hyperolius microstictus Ahl, 1931 (from Angola); Hyperolius marungaensis Ahl, 1931 (from Angola); Hyperolius latirostris Laurent, 1943 (from the DRC); Hyperolius melanoleucus kandoensis Laurent, 1943 (from the DRC); Hyperolius fuliginosus Laurent, 1943 (from the DRC); Hyperolius marmoratus epheboides Laurent, 1957 (from the DRC); Hyperolius marmoratus alborufus Laurent, 1964 (from Angola); Hyperolius fuelleborni Ahl, 1931 types from Songwe, Miramba, Rungwe, and Lumbira in Tanzania (types of Hyperolius fuelleborni Ahl, 1931 from Papya Island, Malawi are retained in H. marmoratus); Hyperolius nyassae Ahl, 1931 (from Tanzania). The following taxa were removed from the synonymy of H. marmoratus and placed as synonyms of H. glandicolor: Hyperolius asper Ahl, 1931 (after Wieczorek et al. 2000); Hyperolius marmoratus campylogrammus Laurent, 1951, which occurs north of the range of the H. marmoratus clade. Hyperolius lestagei Laurent, 1943 (from the DRC) was removed from the synonymy of H. marmoratus, which does not occur in the DRC, and placed as a synonym of H. marginatus. Hyperolius marmoratus mafianus Laurent, 1961 was removed from the synonymy of H. marmoratus and placed as a synonym of H. mariae, which occurs north of the H. marmoratus clade. Hyperolius marmoratus Rapp, 1842, was described from KwaZulu-Natal Province, South Africa. Only one clade extends along the east coast of South Africa, and this clade is assigned to Hyperolius marmoratus (Fig. 5). The localities of types now regarded as junior synonyms of H. marmoratus are listed in Table 3. The range of this species has expanded southwards to Cape Town and its environs due to multiple introductions (Bishop 2004, Tolley et al. 2008). The most common pattern elements in the descriptions of the types included in H. marmoratus are dorsal marbling (25%), and a pale lateral band (25%), followed by small black speckle or spots (19%), and a pale vertebral band (19%). The percentages are not additive, as patterns may be shared. Pickersgill showed how the colour patterns intergrade along the South African coast (2007: 384–385). An advertisement call from Chimanimani (no voucher) is a brief whistle (0.06 s) with an emphasised frequency of 2608 Hz; a call from Gqeberha (no voucher) is a whistle (0.1 s) showing a slight rise in frequency, at an emphasised frequency of 2608 Hz. Calls from Howick (no vouchers) are long, rising whistles, 0.13 s, at an emphasised frequency of 2583–2928 Hz., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 325-328, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Hewitt, J. (1937) A Guide to the Vertebrate Fauna of the Eastern Cape Province, South Africa. Part II. Reptiles, Amphibians, and Freshwater Fishes. Albany Museum, Grahamstown, 141 pp., 34 pls.","Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London, 256 pp., 16 plates.","Loveridge, A. (1941) South African frogs of the genus Hyperolius in the Museum of Comparative Zoology, Cambridge, Massachusetts. Annals of the Transvaal Museum, 20, 283 - 291.","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. F. (1985) Hyperoliidae. In: Frost, D. R. (Ed.), Amphibian Species of the World: A Taxonomic and Geographical Reference. Association of Systematics Collections and Allen Press, Lawrence, Kansas, pp. 203 - 233.","Poynton, J. C. & Broadley, D. G. (1987) Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae, Annals of the Natal Museum, 28, 161 - 229.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp.","Perret, J. - L. (1976) Revision des amphibiens africains et principalement des types, conserves au Musee Bocage de Lisbonne. Arquivos do Museu Bocage, Lisboa, Serie 2 6, 15 - 34.","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Laurent, R. F. (1957) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National de l'Upemba. Mission G. F. de Witte (1946 - 1949), 42, 3 - 47.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Bishop, P. J. (2004) Hyperolius marmoratus Rapp, 1842. In: Minter, L. R., Burger, M., Harrison, J. A., Braack, H. H., Bishop, P. J. & Kloepfer, D. (Eds.), Atlas and Red Data Book of the Frogs of South Africa, Lesotho and Swaziland. Smithsonian Institution - Man and Biosphere Biodiversity Program, Washington, D. C., pp. 141 - 143.","Tolley, K. A., Davies, S. J. & Chown, S. L. (2008) Deconstructing a controversial local range expansion: conservation biogeography of the painted reed frog (Hyperolius marmoratus) in South Africa. Diversity and Distributions, 14, 400 - 411. https: // doi. org / 10.1111 / j. 1472 - 4642.2007.00428. x"]}

Hyperolius goetzei Ahl, 1931 Hyperolius goetzei Ahl, 1931. ZMB 53181, “ Uhehe, Massai-Steppe” (= Iringa), Tanzania. Colour in alcohol: Above bright grey-red, with large dark brown marbling on the side of the head and body which has darker margins. Hyperolius viridiflavus reesi Schiøtz, 1982. ZMUC R77205, Magombero Forest, Tanzania. The type was collected close to the localities of the sequenced material, and this form is known to be widely distributed on the central Tanzania highlands (Schiøtz, 1999). Phase J colour in life: back silvery-brown with a darker pattern of spots and an irregular hourglass pattern. Sides of head and body dark, without a subdermal dark lateral streak. Phase F: Back yellow to light green, flanks with a conspicuous subdermal lateral streak, overlain by 2–3 yellow vertical stripes. Sequences were available from Kibasira Swamp and Mang’ula, (close to Magombero Forest) Tanzania (Fig. 10B). The minimum uncorrected p- distances from other clades were 4.0–7.0% (Table 2). Poynton (1977) recognised Hyperolius sp. from central Tanzania with lateral black markings that show through the skin, which Schiøtz (1982) described as H. v. reesi. Poynton later (1991) recognised Hyperolius reesi as a good species. The dark lateral band is common in other species, as are yellow transverse stripes in some animals. However, Ahl (1931) described Hyperolius goetzei from Iringa in central Tanzania, which Schiøtz (1999) illustrated in colour from Dabaga near Iringa. The back is uniform with mottled sides. This species is known from central Tanzania (Schiøtz 1999). The localities of the type and taxon regarded as a junior synonym are listed in Table 3. The most common pattern elements in the descriptions of the types included in H. goetzei are dorsal marbling (67%), followed by lateral marbling (33%), small dark spots (33%), or a dark lateral band (33%). The percentages are not additive, as the pattern elements may be shared. The advertisement call (no voucher) is a series of frequency modulated whistles or clicks with a mean length of 0.04 s and an emphasised frequency of 2239 Hz (Iringa, Tanzania)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 340-341, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Schiotz, A. (1982) Two new Hyperolius (Anura) from Tanzania. Steenstrupia, 8, 269 - 276.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Poynton, J. C. (1977) A new Bufo and associated Amphibia from southern Tanzania. Annals of the Natal Museum, 23, 37 - 41."]}

Hyperolius glandicolor Peters, 1878 Hyperolius glandicolor Peters, 1878. ZMB 9299, ZMB 77768, Taita, Kenya. Brown to yellow background, covered densely with very small dark dots. Hyperolius striolatus Peters, 1882. ZMB 9300, Taita, Kenya. Synonymy by Laurent (1961). Above rust coloured with black dots and stripes Megalixalus pantherinus Steindachner, 1891. NHMW 3734, “Leikipia” (Laikipia), Kenya. Synonymy by Wieczorek et al. (2000). Yellow with panther-like black markings on the back and behind the head. Rappia Ferniquei Mocquard, 1902. MNHNP 1902.184, “de la rivière Atchi, l’altitude de 1,450 mètres” Athi River, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Beige-grey above with a slight brown tint from the nostril to the arm, many dark speckles over the dorsal surface and limbs. Rappia symetrica Mocquard, 1902. MNHNP 1924.30–34. “de la rivière Atchi, l’altitude de 1,450 mètres” Athi River, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Above a very pale brown, with a black T shaped mark between the eyes. Many speckles along the midline. A dark band from the nostril, over the eye to the arm. A second mark, sometimes paired, extends obliquely towards the vent. These marks become gradually pink towards the ventral surface. A third irregular mark over the vent, edged with white. Rappia platyrhinus Procter, 1920. BMNH 1947.2.9.46, “Nairobi”, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). “Uniformly brownish-grey above, finely speckled with black pigment cells which are slightly concentrated along canthus rostralis” (Proctor 1920). Hyperolius albolabris Ahl, 1931. ZMB 58748, “Kwa Buosch in Süd Kavirondo”, south-western Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Above pale pink, with numerous dark pink or violet-brown flecks and stripes, usually arranged longitudinally, and with indistinct margins. A wide white stripe above the upper lip, extending back to the flank, and separating the colour of the upper and lower surfaces. Hyperolius asper Ahl, 1931. ZMB 4807, Nairobi, Kenya. Placed in the synonymy of H. glandicolor by Wieczorek et al. (2000). Colour in alcohol: Pale grey above and on the sides of the head, covered with many very small spots resembling a spray of ink. Hyperolius bergeri Ahl, 1931. ZMB but type not found (Tillack et al. 2021). “Guaso Narok (Englisch-Ostafrika)” Uaso Narok, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above pale yellow-green, with distinct blue marbling on the back and tibia. Upper thigh dark brown (red in life?) with a thin yellow-green longitudinal stripe. Hyperolius coeruleopunctatus Ahl, 1931. ZMB 36115, “Nairobi”, Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above pale grey, with many small, round blue-black dots (bright blue in life), resembling sprayed ink. Hyperolius pulchromarmoratus Ahl, 1931. ZMB 77751, “ Britisch-Ostafrika ” (Kenya). Locality obscure. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above pale yellow on back, head, and upper surfaces of limbs, with thin sulfur-yellow and blue marbling. In between irregular large, dark brown markings of different sizes. Hyperolius scheffleri Ahl, 1931. ZMB 85759, “Kibwezi, Britisch-Ostafrika ” Kenya. Synonymy with H. marmoratus striolatus by Laurent (1952). Colour in alcohol: Above uniform pale yellow. Thigh dark, probably red in life. Hyperolius marmoratus ommatostictus Laurent, 1951. MNHNP 1912.383 “Kibonoto, Kilimandjaro ” (Kibongoto), Tanzania. Synonymy by Wieczorek et al. (2000). Colour in alcohol: The back pattern consists of round dark brown eye-shaped markings, with a red-brown centre surrounded by a pale zone. Hyperolius marmoratus campylogrammus Laurent, 1951. MNHNP 1924.28, “Bura, alt 1050 m ” (Taita Hills). The H. marmoratus clade does not occur as far north as Kenya. The type occurs close to sequenced material of H. glandicolor from Taita. A transverse band between the eyes, with three longitudinal bands, the lateral bands curved inwards. Often lumbar spots present. Males with uniform backs. Hyperolius viridiflavus pitmani Laurent, 1952. Type unknown, “Lac Bunyoni”, Uganda. Synonymy by Wieczorek et al. (2000). Back with many small yellow spots. Hyperolius sheldricki Duff-MacKay and Schiøtz, 1971. NMK A/366/3, “north of Aruba Dam, Tsavo National Park (East) 3° 19’ S, 30° 54’ E ”, Kenya. It is a typical member of the H. viridiflavus group (Duff-MacKay and Schiøtz 1971). Sequences from material collected in the field as H. sheldricki are embedded in the clade identified as H. glandicolor. The back is light brown, with two dark brown stripes along the midline, joined over the snout. Other thin dark brown lines run from the eye back to the flank, and a pair of interrupted dorso-lateral lines, in places with a pale centre. Hyperolius viridiflavus ngorongoriensis Schiøtz, 1975. ZMUC R079324, Ngorongoro Crater Rim, Tanzania. Synonymy by Wieczorek et al. (2000). Dark brown stripes on a light brown background. Dark spots scattered evenly on the back and upper surfaces of the limbs. Ventral surface of femur and tibia maroon in life, pink in alcohol. Hyperolius orkarkarri Drewes, 1997. CAS 186087, “ Tanzania: Mara Region: Serengeti Dist: Serengeti National Park: Moru Kopjes, on Gong Rock. 02° 43.05 S; 34° 47.6 E at an elevation of 1585 m.” Synonymy by Channing & Howell (2006). Back grey, with a broad white vertebral band. This is interrupted by a dark triangle between the eyes, pointing backwards, and three pairs of lateral extensions, one from the scapular region to the eye, a second medio-dorsal saddle, and a posterior pair from the pelvic region. Hyperolius goetzei Ahl, 1931 was removed from the synonymy of H. glandicolor and elevated to a full species below. This clade consists of 20 sequences from Kenya, Tanzania, Uganda and Rwanda. The minimum uncorrected pdistances from other clades were 2.8–5.9% (Table 2). The sequences were reported in GenBank as Hyperolius glandicolor, H. goetzei, H. viridiflavus, H. v. ferniquei, H. v. glandicolor, H. v. ngorongoriensis, H. v. pantherinus, and H. v. pitmani. This clade occurs in Rwanda, Uganda, northern Tanzania and southern Kenya (Fig. 8B). The clade distribution closely matches the distribution of types that are regarded as junior synonyms, and sequences occur at the type locality. The localities of the type and taxa now regarded as junior synonyms of H. glandicolor are listed in Table 3. The most common pattern element in the descriptions of the types included in H. glandicolor is dorsal marbling (39%), followed by speckle or small spots that may be black or blue (33%). The percentages are not additive, as the pattern elements may be shared. An advertisement call from Kakoneni, Kenya (no voucher) shows individual notes with a length of about 0.1 s, and an emphasised frequency of 2800–3150 Hz (Schiøtz 1975, 1999, as H. sheldricki). A call from Arusha National Park in Tanzania (no voucher) shows brief whistles with frequency modulation, with a mean emphasised frequency of 2200 Hz (Schiøtz 1975). This is similar to calls from Kisessa, Tanzania (no vouchers), as H. goetzei; Arusha, Tanzania, as H. ommatostictus and Lake Naivasha, Kenya, as H. ferniquei; Nyahuru, Kenya, as H. pantherinus (Pickersgill, 2007). An advertisement call consisting of a brief note with an emphasised frequency of 1950 Hz was recorded at Lake Bunyoni, Uganda (no voucher), as H. v. pitmani and at Kabale, Uganda (no voucher) as H. pitmani (Pickersgill 2007). This is similar to calls from Mwanza (no vouchers) in Tanzania that have an emphasised frequency of 2800 Hz (Schiøtz 1975, 1999, Pickersgill 2007, as H. mwanzae) and Kisiizi, Uganda (no vouchers) (Pickersgill 2007, as H. variabilis)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 336-337, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Duff-Mackay, A. & Schiotz, A. (1971) A new Hyperolius (Amphibia Anura) from Kenya. Journal of the East Africa Natural History Society and National Museum, 29, 1 - 3.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Channing, A. & Howell, K. M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, 418 pp.","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp."]}

Hyperolius tuberculatus (Mocquard, 1897) Rappia tuberculata Mocquard, 1897. MNHNP 96.570 “Lambaréné”, Gabon (Table 3). Back and exposed parts of limbs light reddish-brown back with many fine tubercles. A brown stripe is sometimes present from the eye to the snout tip. Another mark of the same colour reaches the eye from the top of the head, while another starts at the lower edge of the eye, running obliquely to the angle of the jaw. This clade consists of 30 sequences from Gabon, Bioko Island (Equatorial Guinea) and Cameroon (Fig. 9B). All sequences were reported in GenBank as Hyperolius tuberculatus. The sequence reported in Veith et al. (2009) as Hyperolius tuberculatus Complex A forms part of this species. The uncorrected p- distances from other clades were 2.1–6.2% (Table 2). The pattern element in the description of the type of H. tuberculatus consists of various stripes on the head. The advertisement call (no vouchers) is a high-pitched long series of notes (Bosch et al. 2000, from Río Muni, Equatorial Guinea), with an emphasised frequency of 3228 Hz, and a mean note rate of 11.6 s- 1. A call from Adjap, Cameroon (no voucher) is a series of double pulsed notes with a note rate of 6 s-1, and an emphasised frequency of 3000 Hz (Amiet & Goutte 2017)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on page 338, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Veith, M., Kosuch, J., Rodel, M. - O., Hillers, A., Schmitz, A., Burger, M. & Lotters, S. (2009) Multiple evolution of sexual dichromatism in African reed frogs. Molecular Phylogenetics and Evolution, 51, 388 - 393. https: // doi. org / 10.1016 / j. ympev. 2008.12.022","Bosch, J., De la Riva, I. & Marquez, R. (2000) Advertisement calls of seven species of hyperoliid frogs, from Equatorial Guinea. Amphibia-Reptilia, 21, 246 - 255.","Amiet, J. - L. & Goutte, S. (2017) Chants d'Amphibiens du Cameroun. J. - L. Amiet & Editions Petit Genies, Nyons, 280 pp."]}

Hyperolius parallelus Günther, 1858. Hyperolius parallelus Günther, 1858. Syntypes BMNH 1947.2.9.41–43 “Süd Afrika”. Boulenger (1882) corrected the locality for the northern specimens to Ambriz, Angola. Above dark brown with three white parallel bands. Upper lip yellowish. Hyperolius marmoratus var. angolensis Steindachner, 1867 type in NHMW, “ Angola ”. Synonymy by Broadley 1971. Back and upper surfaces of limbs brown. Back with irregular yellow flecks or reticulation, or with a pair of yellow lateral bands and irregular yellow flecks in the middle of the back. Limbs with yellow spots and lines. Hyperolius insignis Bocage, 1868. Syntypes MBL (now MUHNAC) T. 21–164, “Benguella”, type localities corrected to São Salvador and Novo Redondo, and ZMB 6462 to Benguela, Angola by Perret 1976. Synonymy by Boettger (1888). Colour in alcohol: Overall yellow tinted with red. Two wide blue-black bands with bright red edges run from the head to the vent. A similar colour band runs from behind the eye to the shoulder, continuing interrupted to the vent. The upper lip has small blue spots with red borders, with similar markings on the upper limbs. Hyperolius toulsonii Bocage, 1868. MBL (now MUHNAC) T.27–275, “Loanda” (Luanda, Angola). Type destroyed in 1978 fire. Synonymy by Perret, 1976. Colour in alcohol: back overall grey, with three wide longitudinal white bands. Thighs uniform grey, with forearms and thighs with large rounded white markings. Hyperolius huillensis Bocage, 1873. MBL (now MUHNAC) 2-166 destroyed in 1978 fire, “Huilla”, Angola. Placed in the synonomy of H. parallelus as sequences from Huila are embedded in the H. parallelus clade. Back and upper surfaces of limbs, excluding the thighs are red with yellow spots, most with black borders. Ventral surfaces and thighs a uniform pale yellow. Hyperolius vermiculatus Peters, 1882. Syntypes ZMB 10050, “ Malanje ”, Angola. It was initially placed as a synonym of H. marmoratus by Laurent (1952), but H. marmoratus only occurs on the east coast of southern Africa. Back and upper surfaces of limbs signal red with yellow vermiculation, with black pigment sometimes visible. Rappia plicifera Bocage, 1893. Syntypes MBL (now MUHNAC) T.22-209 destroyed in 1978 fire. Syntypes NHMW 22895, Caconda, Duque de Bragança, Angola. Synonymy by Perret (1976). Colour in alcohol: Above marbled and spotted in brown-violet or red on a grey base. Lower surfaces and thighs a uniform bright yellow. Rappia marmorata var. taeniolata Bocage, 1895. Types in MBL (now MUHNAC) considered destroyed. “ Duque de Bragança ”, “Huilla”, “Caconda” and “Cahata”, Angola. Synonymy by Laurent (1952). Back and flanks burgundy with various irregular yellow lines and spots, with black borders. Rappia marmorata var insignis Bocage, 1869. Syntypes MBL (now MUHNAC) T 21–164 destroyed by fire, surviving syntype ZMB 6462 (Tillack et al. 2021). St. Salvador du Congo, Angola north of Quanza, Novo Redondo, Dombe, Angola. Synonymy by Perret (1976). Back black with three longitudinal white or yellow bands with red edges. Rappia burgeoni De Witte, 1921. MRAC 585, “Madyu (Uelé)” DRC. See the section on obscure localities, above. Watsa, Upper Uelé, DRC. This taxon from the northern DRC was regarded as a synonym of H. viridiflavus pachydermus by Laurent (1951a). Dehling (2015) demonstrated that Rappia pachyderma Werner, 1908 was a junior synonym of Hyperolius papyri (Werner, 1908), a species of the H. nasutus group (Channing et al. 2013) and that the synonymization of several species of the H. viridiflavus group with this species by Laurent (1951) were thus unwarranted. However, the clade identified as a subspecies of H. viridiflavus by Laurent (1951) does not occur in the northern DRC, while the clade identified as H. parallelus is well represented in the northern DRC. Pale yellow above, with a dark brown lateral band from the snout tip through the eye to the vent. A band of similar colour on the tibia. A darker vertebral band present. Rappia rossii Calabresi, 1925. Syntypes MSNM 584 “Regione dell’Alto Uellè (Congo Belga)” Watsa, DRC. The H. viridiflavus clade is not known as far west as Watsa. See also Dehling (2015) and image of syntype in Blackburn & Scali (2014). Above blue-grey with white tubercles with red centres. Thighs and fingers with pink speckle. Hyperolius bitaeniatus Ahl, 1931. ZMB 39004, “Konde-Nyika, Deutsch-Ost-Afrika” (northern tip of Lake Malawi), Tanzania. Placed as a synonym of H. marmoratus mariae by Laurent (1952) but here placed in H. parallelus as the H. mariae clade does not occur in the northern Lake Malawi area. Colour in alcohol: Above light brown, with two yellow or white longitudinal stripes. A dark lateral band runs between the arms and leg. Hyperolius nyassae Ahl, 1931. ZMB 39006, “Langenburg” Lumbira, Tanzania. Placed as a synonym of H. marmoratus nyassae (Laurent 1952), but here placed in H. parallelus as the H. marmoratus clade does not occur in the northern Lake Malawi area. Colour in alcohol: Back and upper surfaces of limbs white to yellow or brown. Some individuals have a large triangular mark between the eyes, pointing backwards. A thin dark line runs from the nostril to the eye, where it widens to become a lateral band that ends in the middle of the body. A pair of large markings between the flank and the sacrum, with another pair over the vent. The slightly darker markings are thinly edged in white or yellow. The tibia has a similar transverse marking. Hyperolius fuelleborni Ahl, 1931. ZMB 86128 –86131, “S’ongwe” Songwe, Tanzania, ZMB 85972 –85973, “Miramba bei Langenburg”, Miramba near Lumbira, Tanzania, ZMB 77465 –77468, “Langenburg” Lumbira, Tanzania, ZMB 85929 –85963, 85974–85988, 86017, 86132–86137, 90929–90948, “Rugwe” (Rungwe village, Mbeya Region, Tanzania). Placed as a synonym of H. marmoratus nyassae by Laurent (1952), but here moved to H. parallelus as the H. marmoratus clade does not occur in the northern Lake Malawi area. The specimens ZMB 71184 –71186, “Neu-Helgoland” Papya Island, Malawi, are retained in H. marmoratus. Colour in alcohol: Pale above with more or less distinct dark brown to black markings that overlap to form a marbled pattern. Hyperolius marungaensis Ahl, 1931. ZMB 10736, “Marunga, Angola ”, corrected to northern Marunga, DRC by Tillack et al. (2021). It was considered a synonym of H. marmoratus by Laurent (1961). The clade identified as H. marmoratus is restricted to the east coast of southern Africa. The colour pattern (Ahl 1931: fig. 226) consists of a dark back with three large round markings on the head, five large circular markings arranged in a vertebral line, and another five longer white markings behind each eye running to the flank. Upper lip with a broad white band. Hyperolius decoratus Ahl, 1931. Lectotype ZMB 36112, “Longa” (=Cuito Cuanavale), Angola. Regarded as a synonym of H. marmoratus (Perret 1962), but that clade is restricted to the east coast. Colour in alcohol: above wine-red, with many more or less large round, white markings with black borders. Three broad longitudinal stripes, the first running from snout tip to vent, the other two run from the side of the snout to the flanks. Similar patterns are found on the visible surfaces of the limbs. Hyperolius microstictus Ahl, 1931. ZMB 36100, ZMB 77762, “Longa, oberhalb Minnesera” (=Cuito Cuanavale), Angola. Synonymised with H. marmoratus decoratus by Perret (1962) but that clade is restricted to the east coast. Colour in alcohol: Above and on the visible parts of the limbs when sitting, bright wine-red with many irregular, pointed white markings. Hyperolius decipiens Ahl, 1931. ZMB 39003, “Westliches Russisi-Ufer und Nordwestufer des Tanganyika-see’s” (Rusizi River west bank and NW Lake Tanganyika, Tanzania). Synonymy with H. marmoratus argentovittis by Laurent (1952) but here moved to H. parallelus as the H. marmoratus clade does not occur in the northern Lake Malawi area. Colour in alcohol: Above brown to blue-grey without markings. Upper lip with white border. Hyperolius nossibeensis Ahl, 1930. ZMB 50098 –50100, “Nossi-Bé” Madagascar. Corrected by Tillack et al. (2021) to Lunda, Angola. Originally placed as a synonym of H. marmoratus, which is restricted to the east coast. The Angolan coastal locality places it within the distribution of the H. parallelus clade. Colour in alcohol: Above red-brown with a number of silver-white flecks and lines, that flow together to form three longitudinal bands, that may be interrupted and indistinct. The side of the head and body similarly marbled. Hyperolius callichromus Ahl, 1931. ZMB 78576, “Westliches Russisi-Ufer und nordwestufer des Tanganyika-see’s” (Rusizi River west bank and NW Lake Tanganyika, Tanzania). Synonymy with H. marginatus by Wieczorek et al. (2000) but here moved to H. parallelus as the H. marginatus clade does not occur in the northern Lake Malawi area. Colour in alcohol: Above deep red-brown, with a wide white vertebral band with black border, that runs from level of nostrils to vent. A similar band runs laterally from upper eyelid to vent. The longitudinal bands may be broken into irregular shapes. Upper lip white, sides with a wide red-brown stripe. Illustrated by Tillack et al. (2021, fig. 6). Paratypes from Dar es Salaam ZMB 85869 –85872 (Tillack et al. 2021), are assigned as synonyms of H. mariae. Hyperolius erythromelanus Monard, 1937. Syntypes in LCFM. Sangev, Angola. Synonymy with H. marmoratus huillensis by Laurent (1952). The H. marmoratus clade does not occur in Angola. Upper surfaces dark red, almost black with many small white spots, arranged in lines edged with black. Upper surfaces of limbs the same colour as the back. Hyperolius angolensis quarrei Laurent, 1943. MRAC 37366, “Lusuku (Lomami)”, DRC. Synonymy with H. marmoratus by Laurent (1952), but that clade is restricted to the east coast. Colour in alcohol: Dorsal surfaces yellow to grey, with rust or red-brown irregular marbled markings. Upper surfaces of limbs with similar markings. Hyperolius latirostris Laurent, 1943. MRAC 1394, “Kasai”, DRC. Considered a subspecies of H. marmoratus by Laurent (1952), but that clade does not extend as far west as the DRC. Back and parts of the limbs visible at rest, red-brown, dappled with pale brown and grey, with a violet tinge to the head. Tibia with a trace of brown. Hyperolius brieni Laurent, 1943. MRAC 5445, “Nyonga (s/Lac Upemba)” DRC. Schiøtz (1975) placed it as a synonym of H. marginatus, but the type locality is close to sequenced material of the H. parallelus clade, with which it is regarded as a synonym. Hyperolius fuliginosus Laurent, 1943. MRAC 35166, “Kasenga (Katanga)” DRC. Synonymy with H. marmoratus by Schiøtz (1975). Hyperolius marmoratus does not occur in the DRC, but the locality is close to sequenced material of H. parallelus. Colour in alcohol: Grey-brown above, on the forearm and tibia, with dark brown marbling with indistinct edges. A pale vertebral band and two indistinct lateral bands are present. Limbs finely dotted or with small black spots. Hyperolius hutsebauti Laurent, 1956. MRAC 52495, “Ibembo, Uelé” DRC. Sequences of specimens identified as H. hutsebauti are embedded in a clade identified as H. parallelus. Earlier studies that recognised H. hutsebauti (e.g. Bell et al. 2017) did not compare material with H. parallelus. Female colouration consists of a beige background with dark marbling edged with red. Hyperolius marmoratus epheboides Laurent, 1957. IRSNB 1.158, “Mukana, marais près de Lusinga 1.810 m ” (Mukana swamp near Lusinga) DRC. The H. marmoratus clade is restricted to the east coast, while the type locality is close to sequenced material in the H. parallelus clade. Colour in alcohol: A beige background, with black markings on the flanks, the side of the head, and on the upper surfaces of the limbs. These are intertwined with longitudinal bright red markings. In life the ventral surface has brilliant red markings. Hyperolius marmoratus alborufus Laurent, 1964. MD 5769, “Cazombo, Alto Zambeze, Moxico, Angola ”. The holotype is figured in Ceríaco et al. (2020). The H. marmoratus clade is restricted to the east coast, while the type locality is close to sequenced material in the H. parallelus clade. Back with red marbling, covering also the upper arm and tibia. The sides of the tibia and the toes with small black spots. A total of 57 sequences were available from Angola, Burundi, Rwanda, DRC, northern Zambia, and northern Namibia. The minimum uncorrected p- distances from other clades were 2.2–6.5% (Table 2). The sequences were reported in GenBank as H. angolensis, H. parallelus, H. huillensis, H. viridiflavus and H. v. argentovittis. The sample is distributed from northern Namibia, through Angola, across to southern Lake Tanganyika, Burundi, Rwanda and northern DRC (Fig. 7). The type localities of taxa now regarded as junior synonyms of H. parallelus are listed in Table 3. Hyperolius parallelus Günther, 1858 was described from “Süd-Afrika”. Boulenger (1882) corrected the locality for the northern specimens to Ambriz, on the northern coast of Angola. Only one clade has a distribution that includes coastal Angola, and it is assigned to Hyperolius parallelus. The most common pattern elements in the descriptions of the types included in H. parallelus are dorsal marbling (39%), or a pale lateral band (30%), or a pale vertebral stripe (30%), followed by small yellow spots sometimes with red centres or black edges (17%). The percentages are not additive, as the pattern elements may be shared. The underside may be red, sometimes with red speckles on the throat. Calls from Humpata, Angola, are brief (0.03 s) at an emphasised frequency of 1189–2153 Hz., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 330-334, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second Edition. Taylor and Francis, London, 256 pp., 16 plates.","Broadley, D. G. (1971) The reptiles and amphibians of Zambia. Puku, 6, 1 - 143.","Perret, J. - L. (1976) Revision des amphibiens africains et principalement des types, conserves au Musee Bocage de Lisbonne. Arquivos do Museu Bocage, Lisboa, Serie 2 6, 15 - 34.","Boettger, O. (1888) Materialien zur Fauna des unteren Congo II. Reptilien und Batrachier. Bericht uber die Senckenbergische Naturforschende Gesellschaft in Frankfurt am Main, 1888, 3 - 108.","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Dehling, J. M. (2015) On the identities of two enigmatic reed frog taxa from South Sudan, Rappia papyri Werner, 1908 and Rappia pachyderma Werner, 1908. Herpetozoa, 28, 39 - 47.","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 (3), 301 - 350. https: // doi. org / 10.11646 / zootaxa. 3620.3.1","Blackburn, D. C. & Scali, S. (2014) An annotated catalog of the type specimens of Amphibia in the collection of the Museo Civico di Storia naturale, Milan, Italy. Herpetological Monographs, 28, 24 - 45. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00008","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","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., Zassi-Boulou, A. G. & 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, 5223 - 5244. https: // doi. org / 10.1111 / mec. 14260","Laurent, R. F. (1957) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National de l'Upemba. Mission G. F. de Witte (1946 - 1949), 42, 3 - 47.","Ceriaco, L. M. P., Marques, M. P., Andr, I., Afonso, E., Blackburn, D. C. & Bauer, A. M. (2020) Illustrated type catalogue of the \" lost \" herpetological collections of Museum do Dundo, Angola. Bulletin of the Museum of Comparative Zoology, 167, 379 - 440. https: // doi. org / 10.3099 / 0027 - 4100 - 162.7.379"]}

Hyperolius viridiflavus (Duméril & Bibron, 1841) Eucnemis viridi-flavus Duméril & Bibron, 1841. Syntypes MNHNP 412, 4599, NHMW 22896, “Abyssinie”. Adwa, Ethiopia. Back green with yellow spots, limbs dark pink, ventral surfaces yellow with small red spots. Rappia bayoni Boulenger, 1911. Syntypes MSNG and BMNH 1947.2.9.32–36, Entebbe, Busso, Bululu, Jinja and Kabulmuliro, Uganda. Synonymy by Laurent (1950). Pale brown back, with a dark lateral band edged with white, sometimes with a similar colour vertebral band, and a similar band along the outer limb surfaces, which are usually brown or dotted with brown. Hyperolius kwidjwiensis Ahl, 1931. ZMB 52449, “ Insel Kwidjwi im Kivu-See” Idjwi Island, Lake Kivu, DRC. Synonymy by Laurent (1952). Colour in alcohol: Back pale yellow with a thin brown band running from the nostrils to the groin. This band becomes the colour of the back along the body, leaving two thin dark margins running backwards from the level of the eye. Tibia the same colour as the body, with a darker brown stripe. Hyperolius variabilis Ahl, 1931. ZMB 36122, “Deutsch-Ost-Afrika”. Bukoba, Tanzania. Synonymy by Schiøtz (1975). Colour in alcohol: Back silvery grey to white, with highly variable red-brown or black-brown markings. These are reproduced here as part of Fig. 3, after Ahl (1931, fig. 173), drawn from Tornier (1896, pl. 4) (Tillack et al. 2021). Ventral surface pale with reddish spots. Hyperolius punctatissimus Ahl, 1931. ZMB 39013, “Bukoba”, Tanzania. Synonymy by Laurent (1952). Colour in alcohol: Back pale grey with many small dark spots or flecks, that may cover the back to produce a uniform dark grey colour. A pale line runs from the upper lip along the side of the body to the upper leg, demarcating the dark back. Limbs similarly patterned. Hyperolius kandti Ahl, 1931. ZMB 46526, “Kivu-See”. Lake Kivu, DRC. Synonymy by Laurent (1947). Colour in alcohol: Uniform beige back and sides. a thin dark line from nostril to eye that broadens into a lateral brown band to the flank. Tibia with dark flecks. Hyperolius schubotzi Ahl, 1931. Type unknown. “Kissenji”, Rwanda. Synonymy by Laurent (1947). Colour in alcohol: Back grey-blue, covered densely with small dark spots or vermiculations. Upper surfaces of limbs as for the back. Throat pale with small round spots, belly yellow, hidden surfaces of legs red. Hyperolius wettsteini Ahl, 1931. ZMB 36103, “Bukoba”, Tanzania. Synonymy by Laurent (1952). Colour in alcohol: Back and sides violet-grey, with large dark violet flecks arranged in longitudinal rows. Ventral surfaces pale yellow, with brown stipples on the throat. Hyperolius phrynoderma Ahl, 1931. Syntypes ZMB 39000 and ZMB 77734 –36, “Zentrales Deutsch-Ostafrika ”, corrected to north-eastern Tanzania (Tillack et al. 2021). Provisionally placed as a synonym of H. viridiflavus (Tillack et al. 2021). Colour in alcohol: Pale yellow all over. Hyperolius karissimbiensis Ahl, 1931. ZMB 46525, “Bambusurwald und Waldwiesen ca. 2400 m hoch, beim Dorf des Mhcabu Gahama am Karissimbi”, Mhcabu Gahama village on Mt Karissimbi, Rwanda. Synonymy by Laurent (1952). Colour in alcohol: Back pale grey with fine blue stipple. A thin dark line runs from snout tip, below the canthus rostralis to under the eye. It starts again on the flank and runs to the vent. The dark line is bordered by white. A dark vertebral stripe. Dark longitudinal line run from between the eyes, interrupted near the armpit, continuing to the vent. These lines have pale borders. Upper surfaces of limbs yellowish with a dark longitudinal line. Hyperolius mohasicus Ahl, 1931. ZMB 36094, “Mohasi-See, Ruanda ” Muhasi Lake, Rwanda. Synonymy by Schiøtz (1975). Colour in alcohol: Back, sides and upper surfaces of limbs white with many moderately large, rounded or long reddish marks, arranged in longitudinal rows. Small bright red-brown warts present. Hyperolius multicolor Ahl, 1931. ZMB 39002, 39005, 74953–74956. “Bambusurwald und Waldwiesen ca. 2400 m hoch, beim Dorf des Mhcabu Gahama am Karissimbi” Mhcabu Gahama village on Mt Karisimbi, Rwanda. Synonymy with Hyperolius karissimbiensis by Laurent (1950). Colour in alcohol: Head and shoulder yellowish to reddish, back red-brown to brown. A dark brown quadratic mark between the eyes, a thin blue vertebral stripe from the shoulder blades to the sacrum. A large dark brown lateral marking between the limbs reaches up to the level of the sacral vertebrae, with a wide black border. A white to yellow regular stripe runs from the level of the sacrum to the vent. Hyperolius macrodactylus Ahl, 1931. ZMB 39100, “Kivu-See” Lake Kivu, DRC. Synonymy by Laurent (1947). Colour in alcohol: Back and sides bright yellow, with many irregular longitudinal dark brown markings forming a marbling. The upper surfaces of the limbs similarly coloured. Hyperolius flavoguttatus Ahl, 1931. ZMB 39011, “Bukoba”, Tanzania. Synonymy by Laurent (1952). Colour in alcohol: Dark brown back with large numbers of round spots more or less in lines. These were bright yellow in life. Upper surfaces of limbs similarly coloured. Hyperolius monticola Ahl, 1931. ZMB 39010, “Niansa, Ruanda, 1500 m hoch” Nyabisundu, Nyanza District, Rwanda. Synonymy with Hyperolius variabilis by Laurent (1943). Colour in alcohol: White above, with four longitudinal lines of small brown markings. Upper surfaces of limbs similar. Hyperolius oculatus Ahl, 1931. ZMB 58570, Balaibo, DRC. Synonymy with H. viridiflavus pachydermus by Laurent (1952). Colour in alcohol: Grey-brown above, with many irregularly spaced pale brown spots with red-brown centres. Hidden surfaces of limbs bright red. In life the back is green with red spots with yellow borders. Hyperolius stuhlmanni Ahl, 1931. ZMB 13008, “Vitschumbi, Südende des Albert-Eduard-See’s” Witschumbi, Lake Edward, DRC. Synonymy by Schiøtz (1975). Colour in alcohol: Pale above, on the sides and upper surfaces of limbs, with many irregular short dark wormlike lines that divide and intertwine. There are also some paler lines in between the darker markings. Hyperolius irregularis Ahl, 1931. ZMB 36105, ZMB 75606, “Mohasi-See, Ruanda ” Muhasi Lake, Rwanda. Synonymy by Schiøtz (1975). Colour in alcohol: Silver-grey above, on the sides and on upper surfaces of limbs, with more or less large, rounded, or elongated dark to red brown markings. Hyperolius koehli Ahl, 1931. ZMB 26089, “Kissenji, Deutsch-Ost-Afrika”, Kissenji, Rwanda. Synonymy by Laurent (1950). Colour in alcohol: Uniform dark brown above. Hyperolius taeniolatus Laurent, 1940. MRAC 39471, “Lacs Mokoto” Mokoto Lake, DRC. Preoccupied by Hyperolius taeniolatus Bocage, 1895. Synonymy with H. viridiflavus bayoni by Laurent (1950). Colour in alcohol: Upper surfaces dark, with hidden surfaces pink. Head and tibia with pink vermiculation, oriented longitudinally. Tarsus and toes pink with black flecks. Ventral surfaces pale with black stipple. Hyperolius ornatus Laurent, 1940. MRAC 3836, Rutshuru, DRC. Synonymy by Laurent (1972). Colour in alcohol: Back dark with pale reticulation and red dots along the flanks. Upper limbs with large dark spots. Hyperolius schubotzi ruandensis Laurent, 1943. MRAC 38371, “Nyakatare, Ruanda ” (Rwanda). Synonymy by Laurent (1983). Colour patterns very diverse, with dark irregular marbling arranged longitudinally. Hyperolius destefanii Scortecci, 1943. Type lost (Blackburn & Scali 2014). “Nargi”, Ethiopia. Synonymy by Schiøtz (1999). Colour description from Schiøtz (1999): “Dorsum green to bluish green, unmarked or with small dark blue or black dots and freckles; posterior face of thigh iridescent pale blue to purple with blackish speckling”. Hyperolius karissimbiensis xanthogrammus Laurent, 1950. MRAC types unknown. Paratypes in LACM. “Cyumbe, Marais de la Mulindi (alt. 1.950 m.), Territoir de Biumba, Ruanda.” Mulindi Swamp, Rwanda. Synonymy by Laurent (1952). Back pale beige to red-brown, with fine yellow to orange dots and vermiculation arranged in three longitudinal lines. Hyperolius karissimbiensis françoisi Laurent, 1950. Type in MRAC, paratypes in LACM, “ Bururi alt 1900–1950m, arrondissement de Bururi ”, Bururi, Burundi. Synonymy by Laurent (1952). Back with a pale vertebral stripe, two lateral-dorsal parallel lines and another between the eyes. An oblique line on the tibia. Hyperolius karissimbiensis intermedius Laurent, 1950. MRAC B 80021.3242 “Marais Rugushu, Kayanza, alt 1900m, terr. de Ngozi ”, Rugushu Swamp, Burundi. Synonymy by Laurent (1952). The pattern is intermediate between H. karissimbiensis xanthogrammus and H. k. françoisi. A pale-yellow interorbital bar and a transverse yellow bar over the scapula, in addition to the patterns described above. Hyperolius viridiflavus hybridus Laurent, 1951. MRAC B 80021.3865 – 3883, “Etang de la Mukura, alt. 1700 m. près d’Astrida” Mukura pond (= Huye), Rwanda. Colouration variable, with a red thigh. Hyperolius viridiflavus mwanzae Schiøtz, 1975. Swamp north of Mwanza, Tanzania. Phase F translucent green with or without bluish spots. No red pigmentation. Hyperolius burgeoni De Witte, 1921 and Rappia rossii Calabresi, 1925 were provisionally removed from the synonymy of H. viridiflavus as both occur in northern DRC where the H. viridiflavus clade has not been recorded, and placed in the synonymy of H. parallelus. Hyperolius viridiflavus angeli Laurent, 1951 from the Central African Republic was removed from the synonymy of H. viridiflavus which does not occur north of the Congo Basin and placed in the synonymy of Hyperolius nitidulus. Altogether 28 sequences were available from Uganda, northern Tanzania, the Kenyan highlands, Burundi, Rwanda, and Ethiopia. The minimum uncorrected p- distances from other clades were 4.2–7.6% (Table 2). The sequences were reported in GenBank as Hyperolius viridiflavus, H. v. viridiflavus, H. v. variabilis, H. v. destefanii, and H. v. bayoni. The earliest species description in this complex is Hyperolius viridiflavus Duméril & Bibron, 1841, from Adwa in Ethiopia. Only one clade has records from Ethiopia, and this clade is assigned to H. viridiflavus (Fig. 4). See Table 3 for localities of synonyms. The map reflects the distribution in Ethiopia reported in Largen & Spawls (2010). The most common pattern elements in the descriptions of the types included in H. viridiflavus are small blue, black or red dark spots (26%), with the next most common elements being marbling on the back (22%), a dark lateral band (19%), and various markings arranged in longitudinal rows (19%). The percentages are not additive, as some individuals share pattern elements. Many of the descriptions were based on preserved material, and a large number of colour patterns is expected when fresh material is examined. Figure 3 illustrates some of the patterns recognised by Laurent (1983) and Ahl (1931) for material they described, here recognised as H. viridiflavus. This sample indicates the problems of identification based on colour patterns. Lötters et al. (2004) illustrate a call (as H. viridiflavus) from Kakamega Forest, Kenya with a distinct pulsed structure and an emphasised frequency of 2971 Hz. A specimen from the same locality was sequenced (AY323920) (Lötters et al. 2004) and found within the H. viridiflavus clade. An advertisement call from Kitale, Kenya consists of a brief whistle with an emphasised frequency of 2600–3000 Hz (Schiøtz 1975 as H. v. viridiflavus). The advertisement call of a specimen from Sango Bay, Uganda, consists of a very brief note with an emphasised frequency of 2400 Hz (Schiøtz 1999)., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 320-325, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Laurent, R. (1950) Exploration du Parc National Albert. Mission G. F. de Witte (1933 - 1935). Genres Afrixalus et Hyperolius. Institut des Parcs Nationaux du Congo Belge, Brussels, pp. 14 - 116, 7 pls.","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Tillack, F., De Ruiter, R. & Rodel, M. - O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum fur Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407 - 450. https: // doi. org / 10.3897 / zse. 97.68000","Laurent, R. F. (1947) Sur quelques rhacophorides nouveaux de Parc National Albert appartenant aux genres Megalixalus et Hyperolius. Bulletin du Musee Royal d'Histoire Naturelle de Belgique, 23, 1 - 8.","Laurent, R. F. (1983) La superespece Hyperolius viridiflavus (Dumeril & Bibron, 1841) (Anura Hyperoliidae) en Afrique Centrale. Monitore Zoologico Italiano, New Series, Supplement 18, 1 - 93.","Tornier, G. (1897) Die Kriechthiere Deutsch-Ost-Afrikas. Dietrich Reimer, Berlin, 206 pp.","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Laurent, R. F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxieme Serie, 22, 1 - 125.","Blackburn, D. C. & Scali, S. (2014) An annotated catalog of the type specimens of Amphibia in the collection of the Museo Civico di Storia naturale, Milan, Italy. Herpetological Monographs, 28, 24 - 45. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00008","Schiotz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Largen, M. J. & Spawls, S. (2010) The Amphibians and Reptiles of Ethiopia and Eritrea. Edition Chimaria, Frankfurt am Main, 693 pp.","Lotters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west- Cameroon. Bonner zoologische Beitrage, 52, 149 - 154."]}

Hyperolius marginatus Peters, 1854. Hyperolius marginatus Peters, 1854. ZMB 4806, “Macanga”, Mozambique. Back yellow with dark red spots, lower femur pink. A dark lateral line runs from the top of the snout to the sacrum, with similar dorso-lateral bands starting behind each eye, with short bands running from the sacrum to the vent. Rappia undulata Boulenger, 1901. MRAC and BMNH 1947.2.9.64, “Pweto et de Lofoi”, DRC. Synonymy by Laurent (1983). Brown above, with a symmetrical dark marking with a pale border. An irregular pale line on the side of the head and body forming an hourglass pattern on the back. Inside of limbs pink. Rappia rhodoscelis Boulenger, 1901. MRAC and BMNH 1947.2.9.44–45, “Pweto”. It was previously placed as a synonym of H. marmoratus (see Frost 2021). The H. marmoratus clade does not occur in southern DRC. Greybrown above, with a pale lateral band from the shoulder. Very fine dark speckle on the limbs. Hyperolius argentovittis Ahl, 1931. ZMB 85718, “Ujiji”, (Udjidji) Tanzania. Synonymy by Laurent (1961). Colour in alcohol: Above yellow-brown with three silver-white longitudinal bands. The lateral bands originate behind the eyes. Illustrated in Tillack et al. fig. 5 (2021). Hyperolius udjidjiensis Ahl, 1931. ZMB 36101, “Udjidji”, Tanzania. Synonymy by Wieczorek et al. (2000). Colour in alcohol: Above pale grey with many small rounded or rectangular markings, often overlapping. Visible parts of limbs the same colour. The markings are a distinct pale brown. Hyperolius graueri Ahl, 1931 ZMB 85758, “Westliches Russisi-Ufer und Nordwestufer des Tanganyika-see’s” (Rusizi River west bank and NW Lake Tanganyika, Tanzania). Considered a synonym of Hyperolius marmoratus argentovittis by Laurent (1952). Hyperolius marmoratus pyrrhodictyon Laurent, 1965. MCZ A-46182 “Mazabuka, in the water-grass fringe of the Kafue River” Zambia. Previously placed as a full species H. pyrrhodictyon Laurent, 1965, it falls within the distribution of the H. marginatus clade, and there is no molecular evidence to maintain its species status. The back has no marbling or any spots, but there is a dark red network on the throat and belly. Hyperolius melanoleucus Laurent, 1941. MRAC 35962–67 Lukafu, DRC. Synonymy by Wieczorek et al. (2000). Colour in alcohol: Back black or brown spotted, with a dark mark from between the eyes to the mid back, and two dark lateral markings, sometimes pale, or with a pattern in white consisting of a medio-dorsal stripe, sometimes interrupted, from snout tip to vent. Two lateral lines starting from the eyelid. Sometimes a dark back with a broad pale vertebral stripe, and an irregular pale lateral line. Hyperolius melanoleucus kandoensis Laurent, 1943. MRAC 1394, “Kando (près de Tenke, Katanga Province)” DRC. Implied synonymy by Wieczorek et al. (2000). Hyperolius marmoratus does not occur in the DRC. Colouration very variable. Three irregular longitudinal white bands, confluent with other markings, producing a marbling effect. Hyperolius lestagei Laurent, 1943. MRAC 39679, “Luapula, (près du Lac Bengwelo)” Luapula Province, DRC. Placed as a synonym of H. marmoratus by Schiøtz (1975). However, the clade identified as H. marmoratus does not occur in the DRC. Colour in alcohol: back and visible parts of the limbs uniform grey or blackish, without markings. Hyperolius angolensis straeleni Laurent, 1943. IRSNB 1.040, “Bukama (Lualaba)”, DRC. Placed as a synonym of H. marginatus by Wieczorek et al. (2000). Two patterns were described. The first consists of a symmetrical, or nearly symmetrical broad vertebral stripe, that may be interrupted, with large irregular markings along the flanks. Tibia marbled. The second pattern consists of a red-brown background with pale vermiculation. Hyperolius marmoratus rhodesianus Laurent, 1948. BMNH 1937.1.5.1, Matetsi, Zimbabwe. Previously regarded as a good species H. rhodesianus Laurent, 1948, it is placed as a synonym of H. marginatus as it occurs close to the localities of sequences (Fig. 6) forming the H. marginatus clade. Dark lateral area without a pale stripe, and greyish dots on the back, all weakly pigmented. Hyperolius marmoratus aposematicus Laurent, 1951. MNHNP 33.295, “Lealui s/Zambèze, Rhodesie” (Zambia). The H. marmoratus clade does not extend this far west. The type locality is between sequenced material in the H. marginatus clade. Adult colouration variable, with three wide pale bands and two pale lumbar marks, or black spots, rounded or elongated, separated from each other. Hyperolius decipiens and H. callichromus (only specimens from north-west Lake Tanganyika), were removed from the synonymy of H. marginatus which does not occur there, and placed in H. parallelus. Hyperolius albofasciatus Hoffman, 1944 was removed from the synonymy of H. marginatus and placed as a synonym of H. marmoratus based on its type locality (Umbe, Malawi) within the distribution of the H. marmoratus clade. It was previously placed as a subspecies of H. marmoratus by Loveridge (1953). Hyperolius brieni Laurent, 1943. Schiøtz (1975) placed it as a synonym of H. marginatus, but the type locality is close to sequenced material of the H. parallelus clade, with which it is regarded as a synonym. Sequences were available from Angola, Zambia, Zimbabwe, Malawi and western Tanzania (Fig. 6). The minimum uncorrected p- distances from other clades were 2.4–5.3% (Table 2). The sequences were reported in GenBank as Hyperolius marginatus, H. viridiflavus aposematicus, H. v. marginatus, H. v. melanoleucus, H. v. pyrrhodictyon, H. v. rhodesianus, H. pyrrhodictyon, and H. parallelus alborufus. Although there are no sequences available from the type locality of Macanga in northern Mozambique, the clade occupies the area where Schitz (1999) mapped H. marginatus. The conservative approach of assigning this clade to H. marginatus thus seems prudent. The type localities of taxa considered junior synonyms of H. marginatus are listed in Table 3. The most common pattern element in the descriptions of the types included in H. marginatus is a pale lateral band (58%), followed by a pale vertebral stripe (42%) and dorsal marbling (33%). The percentages are not additive, as the elements may be shared. The advertisement call of a specimen from Lusaka, Zambia consists of a series of clicks (Schiøtz 1975, (no voucher) as H. paralellus melanoleucus) with an emphasised frequency of 2150 Hz. A call from Livingstone, Zambia is similar, with an emphasised frequency of 2000 Hz (Schiøtz 1975, (no voucher) as H. p. aposematicus). A call from Kachalola, Zambia has an emphasised frequency of 2100 Hz (Schiøtz 1975, (no voucher) as H. marginatus). The call is identical to those from Kitwe, Zambia (as H. melanoleucus), Solwezi, Zambia (as H. alborufus) and Nkotakota, Malawi (as H. marginatus) (Pickersgill 2007). Calls in Kasanka National Park (no vouchers) are brief (0.07 s) at an emphasised frequency of 2497 Hz., Published as part of Channing, Alan, 2022, Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA, pp. 301-354 in Zootaxa 5134 (3) on pages 328-330, DOI: 10.11646/zootaxa.5134.3.1, http://zenodo.org/record/6538259, {"references":["Laurent, R. F. (1983) La superespece Hyperolius viridiflavus (Dumeril & Bibron, 1841) (Anura Hyperoliidae) en Afrique Centrale. Monitore Zoologico Italiano, New Series, Supplement 18, 1 - 93.","Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1 - 132.","Laurent, R. F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musee de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65 - 96.","Wieczorek, A., Drewes, R. C. & Channing, A. (2000) Phylogenetic relationships within the Hyperolius viridiflavus complex (Anura: Hyperoliidae), and comments on taxonomic status. Amphibia-Reptilia, 22, 155 - 166. https: // doi. org / 10.1163 / 15685380152030382","Laurent, R. (1952) Apercu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Societ Royale Zoologique de Belgique, 82, 379 - 397.","Laurent, R. F. (1965) The geographical variation of the frog Hyperolius marmoratus (family Hyperoliidae) in Rhodesia, Nyasaland and Tanganyika. Breviora, 219, 1 - 9.","Laurent, R. (1943 a) Les Hyperolius (Batraciens) du Musee du Congo. Annales du Musee du Congo Belge, 4, 61 - 140.","Schiotz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.","Laurent, R. (1948 \" 1947 \") Two new forms of the genus Hyperolius. Annals and Magazine of Natural History, Series 2, 14, 294 - 296.","Laurent, R. (1951 a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d'Histoire Naturelle de Paris. Annales Societ Royale Zoologique de Belgique, 82, 23 - 50.","Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp."]}

Alain Dubois has made important contributions to our understanding of African amphibians. His 1981 interpretations based only on earlier literature include the recognition of a series of subfamiles in the Ranidae. Later he added data derived from his work on the morphology of museum specimens, defining new subdivisions within the Raninae. By 2005 he was using molecular evidence to provide a testable working taxonomy, and proposed an arrangement including the epifamily, superfamily, family and subfamily divisions

Ptychadena mutinondoensis sp. nov. Channing and Willems Mutinondo Grass Frog (Figs. 3, 4, 6) Holotype. A male, ZMB 88379, (Fig. 3) collected 7 January 2017, from Mutinondo Wilderness, northern Zambia (12.456�� S; 31.293�� E, 1430 m) Paratypes. Two females, ZMB 88377 and ZMB 88380, and five males, ZMB 88381 ��� 88385, collected at the same locality. Other material. Two tadpoles ZMB 88374 and 88378, and two metamorphs ZMB 88375 ��� 76 all collected from the same locality. Sequences. 16S rRNA sequences were obtained from ZMB 88375���77 and ZMB 88379���80. Diagnosis. We assign the new species to the genus Ptychadena on the basis of the vomerine teeth reaching the anterior corners of the choanae, the outer metatarsals separated by webbing from the rest of the foot, and males with external lateral vocal sacs. Comparative data were drawn from Poynton and Broadley (1985) and Dehling and Sinsch (2013). Ptychadena mutinondoensis was compared and distinguished from all other East, Central and Southern African Ptychadena species; the following diagnosis omits the West African species. The new species has no contrasting longitudinal bands on the posterior thigh, distinguishing it from P. subpunctata, P. taenioscelis, P. keilingi, P. upembae, P. guibei, P. nilotica, and P. anchietae. The foot is longer than half SVL, distinguishing it from P. schillukorum and P. mossambica. It has no light triangle on the snout, distinguishing it from P. subpunctata, P. obscura, P. anchietae and P. oxyrhynchus. It has three phalanges free of web on the fourth toe, distinguishing it from those with more webbing (P. keilingi and P. bunoderma). There is less than one phalanx of the fifth toe free of web, distinguishing from those with less webbing (P. mossambica, P. keilingi, P. upembae, P. guibei, P. taenisocelis, P. bunoderma, P. ansorgii, P. porosissima and P. obscura). The median skin folds are all similarly developed, distinguishing it from P. bunoderma, which has enlarged median folds. The snout is without skin folds, distinguishing it from P. uzungwensis and P. broadleyi. The similar Ptychadena broadleyi was characterised by a pair of short skin folds on the snout, and another between the eyes (Stevens 1972). This feature is not present in all specimens. Stevens explains the smooth snout of some specimens as a note: "Snout ridges are very indistinct in some of (the) type specimens owing to the author's habit of injecting preservative into material during fixation, resulting in the skin being stretched and lifted from underlying muscles". Stevens was incorrect, as the skin over the top of the snout has not lifted in the type series. It would be better to state that the short skin folds on the snout are not always present in adults. There is no light longitudinal line on the tibia, distinguishing it from those with such a line (P. subpunctata, P. keilingi and P. porosissima). A light line is sometimes present in P. mossambica, P. guibei, P. taenioscelis and P. nilotica. There is no row of tubercles under the fourth metatarsal, although a row of flat white marks is present, distinguishing it from species with a row of tubercles under the fourth metatarsus (P. mossambica, P. keilingi, P. upembae, P. guibei, P. bunoderma, P. ansorgii, P. uzungwensis, P. grandisonae, P. porosissima and P. obscura). The snout is long, with the snout���nostril distance greater than the internarial distance, which distinguishes it from those without a long snout (P. schillukorum, P. mossambica, P. keilingi, P. upembae, P. guibei, P. ansorgii, P. broadleyi, P. grandisonae, P. porosissima, P. nilotica, P. obscura and P. anchietae). The foot is shorter than the tibia, distinguishing it from those with the foot equal to or longer than the tibia (P. subpinctata, P. keilingi, P. guibei, P. taenioscelis, P. ansorgii and P. nilotica). The gular pouch slits of males end below the arm insertion, similar to P. grandisonae. This character distinguishes the new species from those species where the slit ends in line with the lower arm insertion, to above the arm (P. subpunctata, P. schillukorum, P. mossambica, P. keilingi, P. upembae, P. guibei, P. taeniscelis, P. bunoderma, P. ansorgii, P. uzungwensis, P. broadleyi, P. porosissima, P. nilotica, P. obscura, P. anchietae and P. oxyrhynchus). The snout is longer and sharper in dorsal view compared to Ptychadena broadleyi (Fig. 2). Description of holotype. A male (Fig. 3), SVL 38.5, SUL 37.9; the body is gracile; head short (HL/SVL 0.34, HW/SVL 0.35), not wider than trunk, about as long as wide (HL/HW 0.97); snout long (SL/HL 0.58), sharply rounded in dorsal view, rounded in profile, projecting well beyond lower jaw, narrow (SL/ IND 2.00); canthus rostralis rounded; loreal region slightly concave; nostrils situated on slight projections, closer to the eye than snout tip (END/SL 0.28); eyes directed anterolaterally, slightly protruding, small (ED/HL 0.29); eye diameter less than half snout length (ED/SL 0.50); interorbital distance is less than upper eyelid (IOS /EL 0.63), and nearly equal to internarial distance (IOS / IND 0.76); internarial distance just less than eye diameter (IOS /ED 0.76); vomerine teeth inconspicuous; tympanum visible, smaller than eye diameter (TYM/ED 0.63), the pale upper lip becoming a thickened gland below the tympanum, extending to above the arm insertion; upper jaw with small needle-like dentition; choanae small, rounded; tongue slightly notched posteriorly, 5.2 at widest part; median lingual process absent; vocal sac double, darkly pigmented through slit; vocal sac slit starts below arm insertion; skin folds present on back; median fold complete, runs from eye to thigh; post eyelid fold complete; dorsolateral fold interrupted anteriorly and posteriorly; lateral fold interrupted; skin rough laterally and between skin folds; ventral surface of limbs, gular and abdomen smooth. Arms slender; hand moderately large (HND/SVL 0.20); tips of fingers not enlarged into discs; relative length of fingers: II I 2 ���2.2 II 2��� 3 III 2���3.2 IV 3��� 1 V; thin margin of webbing extending to tips; inner metatarsal tubercle prominent, oval smaller than eye diameter (IMT /ED 0.45); thin tarsal ridge present; outer metatarsal tubercle present as a small white bump. Measurements of the holotype and paratypes are presented in Appendix 2. Colour in life. The dorsal pattern of the holotype consists of small dark brown patches arranged along the skin folds on a beige background (Fig. 3). Reddish-brown patches are present on the upper arm and laterally. There is a thin white vertebral line, enclosed in a broad beige vertebral band. A dark brown line runs from the tip of the snout, over the nostril, to the base of the eye. The iris is dark brown. The tympanum is transparent with a beige triangular central marking. The upper lip is lightly speckled with brown, and the lower lip is black posteriorly, with the pigment extending into the vocal pouch slit. The lateral skin fold is white, as is the gland extending from the upper lip to the arm insertion. The flanks have irregular brown and pale marbling. The upper surfaces of the arm are beige, with darker brown transverse bars on the lower arm. The leg has eight transverse irregular darker bars over the upper leg, thigh and foot. The back of the thigh is unmarked, apart from some faint speckling. The venter is white with a few black speckles on the yellow throat. Colour in preservative. The pattern is clearly visible as a darker brown on a paler brown background. Paratype variation. The paratypes are remarkably similar to the holotype in proportions, with the females larger, reaching a SVL of 43.5 mm. Between 3���3.5 phalanges of the fourth toe may be free of webbing on either side. Females had bright red infusions on the upper arm, above the eye, and in spots on the flanks. Advertisement call. The calls were recorded on 14 January 2016 between 19h00 and 20h00, air temperature 24��C. Males were heard calling during daytime, linked to rainy conditions, and after dark. The description is based on the call of males visually confirmed to be of this species, near the locality where the holotype was collected. The call consists of an irregular fast series of clicks, and brief trills of five to seven pulses (Fig. 4). The pulse rate is approximately 35 s -1. The fundamental frequency is 1.1 kHz, with additional harmonics. The duration of the trills varies from 0.1��� 0.3 s. Analysis of 93 calls from five males showed that the number of pulses varied from 1 to 8 (mean 3.7), call duration varied from 13���288 ms (mean 100), and pulse rate varied from 10.5��� 58.1 s -1 (mean 29.2 s - 1, n=79). The emphasised frequency varied from 1033���3531 Hz, (mean 1738 Hz). In contrast, the advertisement call of P. broadleyi (n=3) (Fig. 4) consists of longer loud trills of 9���24 (mean 17) pulses, with a call duration of 0.84��� 2.25 s (mean 1.6 s) and pulse rate 9.6��� 10.2 s -1 (mean 10 s -1). The emphasised frequency varies from 1205���3531 Hz (mean 2670 Hz). The calls of other species of grass frogs consist of trills and chuckle calls. Breeding and tadpoles. Some notes on timing of calling and oviposition were taken during the 2016���2017 wet season. Rainfall occurred on multiple days in November and December. A few frogs were first noticed calling 22 December after dusk, the first warm evening after 48 hours of heavy rainfall and cold overcast weather. Freshly deposited eggs were found at two out of 10 known oviposition sites checked the following morning. No calling was noticed during consecutive days with mostly dry weather conditions, during which the seepage flow at oviposition sites dried up. Heavy rains set in again on 2 January leading to mass calling activity and oviposition at all known sites (checked from 4 January onwards). Calling activity continued throughout the wet season, with activity peaking during the first hours after dusk. Calling was also frequently noted at other times of day or night during warm and wet conditions. There was little if any calling after multiple days without precipitation. Eggs are deposited at the top of rock faces, where there is a film of water, under overhanging vegetation (Figs. 6, 7). The eggs are not covered by the film of water. Eggs were found until March, always during or shortly after a period of good rains. Four clutches contained 140, 180, 190 and 280 eggs (7 January 2016). Mean egg diameter was 9.3 mm, N=35. The following description is based on ZMB 88378, a stage 40 tadpole collected at Mutinondo Wilderness (Fig. 5). Two others collected at the same time and place were found to have identical 16S rRNA sequences to the adults, confirming their identity. The tadpole has a total length of 32.9 mm. The head is bluntly rounded from above, and in lateral view. The eyes are large, positioned dorsolaterally. Extra-ocular proportion: width of head minus distance between lateral limits of eyes/distance between lateral limits of eyes = 0.9. The oral disc is anteroventral, with a single row of rounded marginal papillae below, with extra papillae laterally. The rostral gap in the papillae extends across the whole labium. The labial tooth row formula (Altig 1970) is 2/1. The upper and lower jaw sheaths are deeply pigmented for more than half of their height. Both jaw sheaths are finely serrated. The external nares are closer to the eye than the tip of the snout. Rostronasal distance/ orbitonasal distance = 1.5. The narial aperture is circular without a distinct margin. No narial valve is apparent. Internarial ratio: width of nostril/internarial distance = 0.15. There is no orbitonasal line. No pineal spot is present. The spiracle opening is small, elliptical, slightly away from the body, and is visible dorsally. The opening is dorsolateral. Tail height is less than head height. Height of tail muscles at base of tail/height of trunk = 0.50. Tails are low with little fin, dorsal and ventral fins equal; the dorsal fin originates at the junction of the body and the tail muscle. The fins are nearly parallel. Tail fins taper to the tip, with nearly straight margins. The tail tip is acute. The axis of the tail, extrapolated anteriorly, passes below the eye. Tail length/length of head and trunk = 2.7. The ratio exceeds 3.2 in other specimens. The vent tube is dextral, continuous with the ventral fin margin. The major measurements for the tadpole at Gosner stage 40, are: total length 32.9; tail length 24; maximum fin height 2.8; distance from highest point on fin to tail tip 7.0; distance from snout tip to spiracle 5.3; distance from snout tip to nostril 1.6; distance from snout tip to eye 2.8; distance from eye to nostril 1.1; oral disc width 1.7. In life the tadpoles are mottled grey and yellow-brown, with a lateral row of small white dots along the tail muscle, and additional small white dots along the base of the dorsal fin. The tadpoles can be darker, a mottled black and grey. The pale spots disappear once preserved. In preservative the specimen is pigmented dorsally (Fig. 6). Dark areas are present around the nostrils, and between the eyes. A dark line runs from behind the eyes to the base of the tail. Fine black stipples are gathered in groups forming mottles along the caudal muscles and dorsal fin. The underside of the trunk, and the anterior half of the base of the tail, are unpigmented. The gut coils are clearly visible through the ventral skin. Tadpole development. Although no detailed data were systematically collected, development from hatching to metamorphosis takes 1���2 months. In both the 2015���2016 and 2016���2017 wet seasons, clear size cohorts could be distinguished within and between oviposition sites, suggesting multiple egg-laying periods within each season, presumably linked to periods of wet and warm conditions. Stage 33 tadpoles were found on 10 May 2017, the latest in the season that tadpoles were found. Habitat. The frogs have only been found in association with granite slopes where there is a 1���2 mm film of slow-flowing water, seeping from vegetation higher up the slope. These conditions at Mutinondo are on or near large granite inselbergs, visible on the skyline in fig. 7. Both the eggs and tadpoles are found in the film of water, which does not cover them. Adults have been found in and under the surrounding vegetation. Calling males were typically observed at an oviposition site. A single adult female was found inside climax evergreen riverine forest 130 meters from a known oviposition site, during the hot dry season (8 September 2016). All other frog observations were in grassy vegetation on or very close to wet granite slopes during the wet November to April months. Plant species typical of oviposition sites and surrounding vegetation include Xerophyta equisetoides, Myrothamnus flabillifolia and Pentas nobilis at sub-shrub level, and Coleochloa setifera, Aeollanthus subacaulis, Polygala africana, Actiniopteris sp., Oldenlandia herbacea, Scleria sp. and Pycreus sp. at herb level. Larger shrubs and emergent trees are typically absent. Amphibian community. Amphibian species recorded breeding at any of the 24 surveyed P. mutinondoensis breeding sites, include Tomopterna tuberculosa (1 site; calling males and tadpoles) and Mertensophryne taitana (2 sites; amplexus and tadpoles). P. mutinondoensis was the only breeding species at 22 out of 24 breeding sites surveyed. Other frogs recorded at the same general sites, but not observed breeding in association with the same seepage patches, are Arthroleptis xenodactyloides, A. xenochirus, A. stenodactylus, Sclerophrys gutturalis, S. pusilla, Kassina senegalensis, Leptopelis bocagii, Breviceps mossambicus, Amietia delalandii, Phrynobatrachus parvulus and Ptychadena oxyrhynchus. Tadpole predation. An all-black, 10���15 mm long species of terrestrial carabid beetle was noted predating on P. mutinondoensis tadpoles on multiple occasions. The tadpoles also share the seepage waters with a range of carnivorous water insects such as small dragonfly (Anisoptera) larvae, which constitute likely predators. Population size. Lodge Rock and Campers Rock were systematically searched for tadpoles on 7 March 2015. A total of 1,050 tadpoles was counted, spread over 20 seepage patches, 19 of which were located on Lodge Rock. Estimated numbers of calling males never exceeded 50 on a single evening during the 2015���2016 and 2016���2017 seasons. Repeated visits to all inselbergs in the reserve produced the following maximum numbers: Klipspringer Rock: remotely estimated at 20���50 calling males on the western flank alone (observers position Lodge Rock); Kankonde Rock: 50 tadpoles; Charlie���s Rock / Chipundu Stream: single juvenile; Mulinso Falls: 50 juveniles; Mafonie: 100 tadpoles, single adult female; all other sites: none. The Mutinondo grass frog appears to be restricted to a small area, with a relatively small population. Etymology. The species is named for the Mutinondo Wilderness area., Published as part of Channing, Alan & Willems, Frank, 2018, A new grass frog with rupicolous tadpoles from northern Zambia (Anura: Ptychadenidae), pp. 349-366 in Zootaxa 4462 (3) on pages 355-361, DOI: 10.11646/zootaxa.4462.3.3, http://zenodo.org/record/1441683, {"references":["Poynton, J. C. & Broadley, D. G. (1985) Amphibia Zambesiaca 2. Ranidae Annals of the Natal Museum, 27, 115 - 181.","Dehling, J. M. & Sinsch, U. (2013) Diversity of ridged frogs (Anura: Ptychadenidae: Ptychadena spp.) in wetlands of the upper Nile in Rwanda: Morphological, bioacoustic, and molecular evidence. Zoologische Anzeiger, 253, 143 - 157. https: // doi. org / 10.1016 / j. jcz. 2013.08.005","Stevens, R. A. (1972) A new species of Ptychadena (Amphibia: Ranidae) from southern Malawi. Arnoldia, Rhodesia, 5, 1 - 14.","Altig, R. (1970) A primer for the morphology of anuran tadpoles. Herpetological Conservation and Biology, 2, 71 - 74."]}

Amietia hymenopus (Boulenger, 1920) Phofung River Frog Rana hymenopus Boulenger, 1920. Poynton 1964 Phrynobatrachus lawrencei FitzSimons, 1947 Rana draconensis FitzSimons, 1948 Strongylopus hymenopus (Boulenger, 1920). Van Dijk 1966 Amietia vertebralis (Hewitt, 1927). Tarrant, Cunningham & Du Preez 2008; Kruger, Weldon & Du Preez 2011 Strongylopus vertebralis (Hewitt, 1927). Clarke & Poynton 2012, by implication, Published as part of Channing, Alan, 2015, The Maluti Mystery revisited: Taxonomy of African River Frogs (Pyxicephalidae, Amietia) on the Drakensberg Mountains in southern Africa, pp. 271-280 in Zootaxa 3925 (2) on page 278, DOI: 10.11646/zootaxa.3925.2.8, http://zenodo.org/record/236789, {"references":["Boulenger, G. A. (1920) Descriptions of three new frogs in the collection of the British Museum. The Annals and Magazine of Natural History, 6, 106. http: // dx. doi. org / 10.1080 / 00222932008632416","Poynton, J. C. (1964) The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum, 17, 1 - 334.","FitzSimons, V. (1947) Descriptions of new species and subspecies of reptiles and amphibians from Natal, together with notes on some other little known species. Annals of the Natal Museum, 11, 111 - 137, plates I - III.","Van Dijk, D. E. (1966) Systematic and field keys to the families, genera and described species of southern African anuran tadpoles. Annals of the Natal Museum, 18, 231 - 286.","Hewitt, J. (1927) Further descriptions of reptiles and batrachians from South Africa. Records of the Albany Museum, 3, 371 - 415, plates XX - XXIV.","Tarrant, J., Cunningham, M. J. & Du Preez, L. H. (2008) Maluti Mystery: A systematic review of Amietia vertebralis (Hewitt, 1927) and Strongylopus hymenopus (Boulenger, 1920) (Anura: Pyxicephalidae). Zootaxa, 1962, 33 - 48.","Kruger, D. J. D., Weldon, C. & Du Preez, L. H. (2011) Resolving the confusion: Amietia vertebralis and A. umbraculata tadpole morphology. African Zoology, 46, 309 - 319. http: // dx. doi. org / 10.3377 / 004.046.0210","Clarke, B. T. & Poynton, J. C. (2012) On the generic and specific identity of the holotype of Rana hymenopus Boulenger 1920 (Amphibia: Anura: Pyxicephalidae). Zootaxa, 3195, 51 - 56."]}

Identification of Rana hymenopus Boulenger, 1920 I examined the type NHMUK 1947.2. 28.65, and compared the description with representative specimens of the Maluti River Frog AACRG 1105 and the Phofung River Frog AACRG 0 648, and other material listed in the appendix. Four statements confirm that the description refers to the Phofung River Frog, while the rest of the description is not diagnostic���Snout rounded, scarcely projecting, a little longer than the eye; interorbital width three-fourths that of the upper eyelid; three phalanges of fourth toe free of web (toes half webbed); two phalanges of third and fifth toes free of web. The type (Fig. 2) is in good condition, and can confidently be recognised as a Phofung River Frog., Published as part of Channing, Alan, 2015, The Maluti Mystery revisited: Taxonomy of African River Frogs (Pyxicephalidae, Amietia) on the Drakensberg Mountains in southern Africa, pp. 271-280 in Zootaxa 3925 (2) on page 276, DOI: 10.11646/zootaxa.3925.2.8, http://zenodo.org/record/236789, {"references":["Boulenger, G. A. (1920) Descriptions of three new frogs in the collection of the British Museum. The Annals and Magazine of Natural History, 6, 106. http: // dx. doi. org / 10.1080 / 00222932008632416"]}

Amietia vertebralis (Hewitt, 1927) Maluti River Frog Rana vertebralis Hewitt, 1927. Poynton 1964 Amietia vertebralis (Hewitt, 1927). Dubois 1987 Rana umbraculata Bush, 1952 Amietia umbraculata (Bush, 1952). Tarrant, Cunningham & Du Preez 2008; Kruger, Weldon & Du Preez 2011, Published as part of Channing, Alan, 2015, The Maluti Mystery revisited: Taxonomy of African River Frogs (Pyxicephalidae, Amietia) on the Drakensberg Mountains in southern Africa, pp. 271-280 in Zootaxa 3925 (2) on pages 278-279, DOI: 10.11646/zootaxa.3925.2.8, http://zenodo.org/record/236789, {"references":["Hewitt, J. (1927) Further descriptions of reptiles and batrachians from South Africa. Records of the Albany Museum, 3, 371 - 415, plates XX - XXIV.","Poynton, J. C. (1964) The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum, 17, 1 - 334.","Bush, S. F. (1952) On Rana umbraculata, a new frog from South Africa. Annals of the Natal Museum, 12, 153 - 164, plates III - VI.","Tarrant, J., Cunningham, M. J. & Du Preez, L. H. (2008) Maluti Mystery: A systematic review of Amietia vertebralis (Hewitt, 1927) and Strongylopus hymenopus (Boulenger, 1920) (Anura: Pyxicephalidae). Zootaxa, 1962, 33 - 48.","Kruger, D. J. D., Weldon, C. & Du Preez, L. H. (2011) Resolving the confusion: Amietia vertebralis and A. umbraculata tadpole morphology. African Zoology, 46, 309 - 319. http: // dx. doi. org / 10.3377 / 004.046.0210"]}

Identification of Rana vertebralis Hewitt, 1927 The two river frogs in question can be easily distinguished on the extensive webbing reaching the last phalanx of the fourth toe in the Maluti River Frog, but with up to three phalanges free in the Phofung River Frog, and small tympanum, less than the eye-tympanum distance in the Maluti River Frog, larger in the Phofung River Frog. The description (Hewitt 1927) is brief and incomplete, with an important error. Only four statements are diagnostic, and all four identify the Maluti River Frog���Tympanum width less than eye-tympanum distance; toes completely webbed in five types (the description refers to half-webbed toes, but examination of the holotype PEM A 1550, and the paratypes PEM A 1551, A 1552, A 1562 and A 10562 shows this to be in error, only PEM A 1555 has webbing that does not reach the tip of the fourth toe). Hewitt (1927) illustrates the type in Plate 24. The quality of the reproduction is too poor to make out useful features. The left foot of the type appears to show that three phalanges of the fourth toe are free of webbing (Fig. 3 A). This would identify the type as a Phofung River Frog, in contradiction to the small tympanum. Unfortunately the type is now in very poor condition, but it clearly shows the fourth toe to be completely webbed. It appears that Fig. 2 of Plate 24 of Hewitt (1927) was retouched, as the dark line along the margin of the webbing is not present in the type (Fig. 3 B). The webbing might have been invisible in the photograph if this retouching had not been done. Whoever did it, however, got it wrong. It also appears that Hewitt's photograph was flipped horizontally, as the right foot and right hand of the type show the same orientation as the left foot and left arm in the photograph (Fig. 3). The same specimen was illustrated by Tarrant et al. (2008: Fig. 2 A) to show the poor condition. Note that the Tarrant et al. Fig. 2 A is flipped horizontally., Published as part of Channing, Alan, 2015, The Maluti Mystery revisited: Taxonomy of African River Frogs (Pyxicephalidae, Amietia) on the Drakensberg Mountains in southern Africa, pp. 271-280 in Zootaxa 3925 (2) on page 276, DOI: 10.11646/zootaxa.3925.2.8, http://zenodo.org/record/236789, {"references":["Hewitt, J. (1927) Further descriptions of reptiles and batrachians from South Africa. Records of the Albany Museum, 3, 371 - 415, plates XX - XXIV.","Holotype PEM A 1550 Mont-aux-Sources","Tarrant, J., Cunningham, M. J. & Du Preez, L. H. (2008) Maluti Mystery: A systematic review of Amietia vertebralis (Hewitt, 1927) and Strongylopus hymenopus (Boulenger, 1920) (Anura: Pyxicephalidae). Zootaxa, 1962, 33 - 48."]}

Identification of Rana umbraculata Bush, 1952 Bush (1952) distinguished R. umbraculata from R. vertebralis on the basis of its large size compared to the immature type of Rana vertebralis, and small differences in the pectoral girdle. These were later shown by Poynton (1964) to not be valid. The description, excellent photographs, and examination of the topotype NMSA 2608 shows that while most statements are not diagnostic, five refer to the Maluti River Frog���Interorbital space 1 to 1.25 the distance between nostrils; eye-tympanum slightly less than eye; fingers with small rounded discs or tubercles; web reaches terminal tubercle of toes; tympanum smaller than eye-tympanum distance. A single statement does not refer to all Maluti River Frogs���Heels meet but do not overlap when the limbs are folded at right angles to the body axis. The short legs of the type are illustrated in Tarrant et al. (2008 Fig 2 B). Note that the Tarrant et al. Figure 2 B is flipped horizontally. Examination of the paratypes NMSA 712, and NMSA 2608, a large 120 mm SUL topotype collected from the Umzimkulu River by Bob Crass in 1951 show that the heels overlap considerably, demonstrating that leg length is variable in the population. Identification of Rana draconensis FitzSimons, 1948 and Phrynobatrachus lawrencei FitzSimons, 1947 Poynton (1964) assigned both these names as junior synonyms of Rana hymenopus Boulenger, 1920. This action is not in contention. I examined the types of Rana draconensis NMSA 734 and Phrynobatrachus lawrencei NMSA 669 confirming that these are typical Phofung River Frogs (Fig. 2). DNA sequences. On the basis of a molecular study, Tarrant et al. (2008) placed the Phofung River Frog in the genus Amietia as it formed a clade with Amietia dracomontana (Channing, 1978) now a junior synonym of Amietia quecketti (Boulenger, 1894), plus two forms identified as Amietia angolensis (Bocage, 1866) and the Maluti River Frog. The genus Strongylopus formed a separate clade, represented by S. bonaespei (Dubois, 1980), S. fasciatus (Smith, 1849) and S. wageri (Wager, 1961). A new phylogeny of all the species of Amietia, using four genes, confirms that the Phofung River Frog is firmly embedded within this genus (A. Channing unpubl. data)., Published as part of Channing, Alan, 2015, The Maluti Mystery revisited: Taxonomy of African River Frogs (Pyxicephalidae, Amietia) on the Drakensberg Mountains in southern Africa, pp. 271-280 in Zootaxa 3925 (2) on page 278, DOI: 10.11646/zootaxa.3925.2.8, http://zenodo.org/record/236789, {"references":["Bush, S. F. (1952) On Rana umbraculata, a new frog from South Africa. Annals of the Natal Museum, 12, 153 - 164, plates III - VI.","Poynton, J. C. (1964) The Amphibia of Southern Africa: a faunal study. Annals of the Natal Museum, 17, 1 - 334.","Tarrant, J., Cunningham, M. J. & Du Preez, L. H. (2008) Maluti Mystery: A systematic review of Amietia vertebralis (Hewitt, 1927) and Strongylopus hymenopus (Boulenger, 1920) (Anura: Pyxicephalidae). Zootaxa, 1962, 33 - 48.","FitzSimons, V. (1947) Descriptions of new species and subspecies of reptiles and amphibians from Natal, together with notes on some other little known species. Annals of the Natal Museum, 11, 111 - 137, plates I - III.","Boulenger, G. A. (1920) Descriptions of three new frogs in the collection of the British Museum. The Annals and Magazine of Natural History, 6, 106. http: // dx. doi. org / 10.1080 / 00222932008632416"]}

Tomopterna gallmanni sp. nov. Wasonga & Channing Gallmann���s Sand Frog (Fig. 6) Holotype. NMK A/ 5039 / 1, an adult male from an artificial water pond, Dam Ya Colin, in Ol Ari Nyiro, Laikipia Nature Conservancy, Laikipia County, Kenya (0�� 34 ��� N; 36 �� 24 ��� E) collected 3 August 2009 by Domnick V. Wasonga and Michael Roberts. Paratypes. NMK A/ 5039 / 2, an adult male and NMK A/ 5039 / 3 an adult female collected together with the holotype; NMK A/ 5154 / 1, adult female and NMK A/ 5154 / 2 adult male collected from a pond at Mogwooni Ranch, Laikipia County, Kenya (0�� 14.5 ��� N; 36 �� 58.6 ��� E) on 10 and 11 May, 2010 by Michael Roberts; NMK A/ 5045 adult male collected from wooded grassland near Ngare Ndare Forest, Lewa Wildlife Conservancy, Lakipia County, Kenya (0�� 09��� 10 ��� N; 37 �� 26 ��� 16 ��� E); NMK A/ 5151 / 1-3, three sub-adults dug from the soil on the banks of a seasonal stream in Baawa village near Maralal, Isiolo County, Kenya (0�� 02��� 45 ��� N; 36 �� 48 ��� 55 ��� E) collected 5 June, 2010 by Domnick V. Wasonga, Michael Roberts and Victoria Zero; NMK A/ 5159, subadult from the roadside, between Baragoi and South Horr, Samburu County, Kenya (01�� 20.76 ��� N; 36 �� 53.1 ��� E) collected 20 June, 2010 by Michael Roberts. Material used for genetic analysis. NMK A/ 5154 / 1, NMK A/ 5045 (one male from Mogwooni Ranch), NMK A/ 5039 / 2 (one male from Laikipia Nature Conservancy), NMK A/ 5151 / 1 (one sub-adult from Baawa Village, Maralal) and NMK A/ 5159 (sub-adult from Baragoi-South Horr Road). Diagnosis. Based on both morphological similarities and phylogenetic affinities that place T. gallmanni sp. nov. within the clade that includes all other Tomopterna, we assign the new taxon to the genus Tomopterna. Appendices 3 and 4 give a summary of morphological features as well as interspecific comparisons. This species can be distinguished from other Tomopterna spp. by a combination of morphological and advertisement characters. It has divided subarticular tubercles, which are found also in T. krugerensis and T. wambensis. The nostrils are midway between the eye and snout tip, a condition only shared with T. cryptotis. The advertisement calls of T. krugerensis, T. wambensis and T. cryptotis are quite different. The body is stout, morphologically similar to T. cryptotis. It is about the same size as T. monticola (SVL 41.2, n= 1) with standard size ranging from 40.0���45.0 (n= 6); but mainly differs from T. monticola which has a deeply notched tongue, finely serrated upper jaw, smaller head width, tympanum partially hidden (although this may be due to prolonged preservation of this material), shorter snout length, smaller tympanum diameter, nostril slightly closer to snout tip than eye and smaller inner metatarsal tubercle. The advertisement call consists of a series of notes, each consisting of one or two pulses, which resemble a rapid knocking, different from the slow repetitive ���knock��� of T. krugerensis. The typical note rate of two individuals is 5.5��� 6.5 s - 1 (Fig. 5). The individual note rates in a chorus vary from 2.7��� 3.3 s - 1. This differs from T. tandyi, T. delalandii and T. cryptotis where the repetition rates are as high as 7���9 s - 1. Each note is brief, about 0.012 s long, with two harmonics (others being slightly longer e.g. T. tandyi and T. cryptotis 0.039��� 0.048 s). The mean frequency of the harmonics is 980 Hz and 1960 Hz with a fainter third harmonic at 2940 Hz. The second harmonic is emphasized (differing from T. krugerensis with multiple and higher frequency harmonics 1900���3100 Hz; T. tandyi 2500���2700 Hz; T. delalandii 1800���2200 Hz; T. cryptotis 3100���3800 Hz). Subarticular tubercles single and prominent. Webbing on feet is scanty; up to 2 �� phalanges free of web on the forth digit. Glandular ridge below the tympanum asymmetric; continuous on the left side and interrupted on the right side (vs. T. wambensis sp. nov. where it is interrupted on both sides). Outer metatarsal tubercle absent; a weak whitish bump present instead. Dorsal pattern consists of irregular grey patches on a lighter background (Figs. 6 A and D) contrasting with greenish brown patches on a lighter background in T. wambensis sp. nov.; irregular dark markings and dark spots on a lighter ground colour in T. krugerensis and also varying from the scantily marked orange-brown to grey-brown of T. marmorata. Scanty, slightly enlarged brown warts present on the dorsum spreading to dorsolateral surface, bordered with black edges (different from more denser, small reddish-brown warts present in T. wambensis sp. nov.; smooth in T. krugerensis, distinctive and prominent tubercles in T. tuberculosa). Ventral surface smooth but coarsely granular towards the vent and inner thighs (Figs. 6 G and H). Description of Holotype. Body is stout and toad like; head comparatively small (HL/SVL 0.28, HW/SVL 0.40), narrow compared to trunk, not longer than wide (HL/HW 0.70); snout short (SL/HL 0.55), rounded in dorsal view, blunt in profile (Fig. 6 A), slightly projecting beyond moderately cusped lower jaw, narrower than long (SL/ INS 1.56); canthus rostralis is moderately angled; loreal region almost straight; nostrils situated on slight projections of the canthus, approximately midway between the snout tip and the eye (ENL/SNL 1.06); eyes directed anterolaterally, protrusion less distinct, relatively small (ED/HL 0.47); snout longer than eye diameter (ED/SL 0.86); interorbital distance is shorter than upper eyelid (IOS/ED 1.25), and longer than internarial distance (INS/ED 0.75); tympanum distinct externally, vertically oval, discontinuous glandular ridge present below, smaller than eye diameter (TD/ED 0.70); upper jaw without dentition; choanae large, round, located anteriorly on the roof of the mouth and easily visible, vomerine teeth in two oblique rows between the choanae; tongue short and strongly notched behind, divided in fairly equal parts (6.0 at widest part); median lingual processes present; vocal sac single, darkly pigmented; gular flap absent, with three distinct longitudinal folds clearly visible (Fig. 6 G). Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs, gular and abdomen smooth, but turning coarsely granular in the ventral area and thighs. Fore limbs fairly thick; hand moderately large (HND/SUL 0.21); tips of fingers not enlarged into discs; relative length of fingers: IVI 1 ��� 1 II 1.5 ��� 1.5 III 1.5 +��� 1.5 IV 1.5 +��� 2 V; inner metatarsal tubercle prominent and shovel-shaped; outer absent. Colouration in life. The dorsal pattern consists of irregular grey patches on a lighter background. Scanty, brown warts present on the dorsum spreading to dorsolateral surface, bordered with black edges; darker bar present between the eyes, anteriorly bordered by a white patch extending to the snout; dorsal stripe absent, ventrum creamy white, turning pinkish on the ventral surface and palms, soles of feet black; extended dark patch on the throat; hands and thighs show two and three transverse dark markings respectively. Colouration in preservative. Reddish dorsal wart and creamy colours on the ventral surface are lost; the other patterns are visible. Paratype variation. The paratypes are similar to the holotype in measurements (Table 4). Female type specimen (SVL 55.9) is larger than males (SVL 40.0���45.0, mean 43.8, n= 6). Thin, continuous white dorsal line from the snout to the vent in the female specimen, dorsolateral white steaks originating just behind the tympanum running towards the thigh but not reaching it; in other male specimens dorsal white line present but interrupted near the head. Female throat almost clear, with only scant dark speckles (Fig. 6 H). See measurements (in mm) of the type series of Tomopterna gallmanni in Table 3. Eggs and tadpoles. Unknown. Habitat. T. gallmanni specimens were found in habitats similar to those of T. wambensis. Large breeding congregations were observed in Laikipia Nature Conservancy. Solitary males were captured crossing the road during day time following a heavy downpour (at Lewa and near Baragoi). Calling males took solitary positions, in some cases in shallow edges of a water body. The following species were found sympatrically with T. gallmanni: Ptychadena anchietae and Ptychadena mascareniensis. Distribution. The collecting records of Tomopterna gallmanni are mostly situated on the Laikipia Plateau of Kenya, with two northern populations at Maralal and Baragoi areas. Current data suggest a limited range, but with some populations known within privately protected property. The elevation of collecting localities ranged from 1314 m (Baragoi-South Horr Road) to 1952 m (Lewa Wildlife Conservancy). Etymology. The species is named for the Gallmann Memorial Foundation for contributing funding for this research. Remarks. Based on its wide distribution, and abundance at breeding sites, we propose a Least Concern status according to the current IUCN Redlist criteria., Published as part of Wasonga, Domnick V. & Channing, Alan, 2013, Identification of sand frogs (Anura: Pyxicephalidae: Tomopterna) from Kenya with the description of two new species, pp. 221-240 in Zootaxa 3734 (2) on pages 229-232, DOI: 10.11646/zootaxa.3734.2.7, http://zenodo.org/record/220044

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

Tomopterna wambensis sp. nov. Wasonga & Channing Wamba Sand Frog (Fig. 4) Holotype. NMK A/ 5152 / 1, adult male, from Sodor water pan in farmlands near Wamba township, Isiolo County, Kenya (0�� 59 ��� N; 37 �� 20 ��� E), collected 5 May, 2010 by Domnick V. Wasonga, Michael Roberts and Victoria Zero. Paratypes. NMK A/ 5152 / 2���3, adult males and NMK A/ 5152 / 4��� 5 adult females collected together with the holotype; NMK A/ 5057 / 1��� 6 adult males collected from temporary pond at Ikave Secondary School, Kitui County, Kenya (1 �� 35 ' 20 " S; 37 �� 54 ' 0" E, 948 m) collected 31 October, 2009 by P.K. Malonza; NMK A/ 5149 / 1���3 and NMK A/ 5150 / 1���2, adult males collected from a murrum pit near the Air Strip at Mpala Research Center, Laikipia County, Kenya (0�� 21 ��� 12 ��� N; 36 �� 52 ��� 20 ��� E) collected 3 May 2010 by Domnick V. Wasonga, Michael Roberts and Victoria Zero; MCNPV-CA 334 adult (not sexed) from open fields 17 km south of Allia Bay, Marsabit County, Kenya (03�� 42 ��� N; 36 �� 20 ��� E), collected 7 July, 2005 by U. Ziliani, R. Sindaco and E. Razzetti; MCNPV-CA 250 adult (not sexed) from open fields near Illeret township on the eastern shores of Lake Turkana (04�� 18 ��� N; 36 �� 13 ��� E) collected 10 January, 2005 by U. Ziliani; MCNPV-CA 252 adult (not sexed) from Karsa Well on the eastern shores of Lake Turkana, Marsabit County, Kenya (03�� 36 ��� N; 36 �� 18 ��� E) collected 11 January, 2005 by E. Baucompagni; MCNPV-CA 337 adult (not sexed) from Lugga Daudi sand river in Sibiloi National Park, eastern Lake Turkana, Marsabit County, Kenya (03�� 49 ��� N; 36 �� 21 ��� E) collected in June 2005 by U. Ziliani, R. Sindaco and E. Razzetti; NMK A/ 4323 adult (not sexed) from Bura Hasuma sand river, Marsabit County Kenya (03�� 48 ��� N; 36 �� 17 ��� E) collected 5 July 2005 by U. Ziliani, R. Sindaco and E. Razzetti; NMK A/ 5145 / 1��� 2 adult males from roadside water puddles near Kitobo Forest, Taita-Taveta County, Kenya (03�� 26 ��� 29 ��� S; 37 �� 37 ��� 13 ��� E) collected 11 April, 2010 by P.K. Malonza, B. Bwong��� and J. Nyamache. Material used for genetic analysis. NMK A/ 5057 / 1 adult male from Ikave, Kitui, NMK A/ 5149 / 1��� 3 adult males from Mpala Research Center, NMK A/ 5152 / 1 adult male from Wamba, NMK A/ 5145 / 1 adult male from Kitobo Forest, MCNPV-CA 334, MCNPV-CA 250, MCNPV-CA 252, MCNPV-CA 337, NMK A/ 4323 five unsexed adults from eastern Lake Turkana. Diagnosis. The phylogenetic analysis in this study has placed T. wambensis sp. nov. within the clade that includes all other Tomopterna, and we have no hesitation in assigning this taxon to the genus Tomopterna. The diagnostic characters and comparison with other members of the genus are presented in Appendices 3 and 4. The new species differs from all other Tomopterna species by a combination of morphological and advertisement call characteristics. These include divided subarticular tubercles, which are also found in T. gallmanni and T. krugerensis. The latter two species both have a harsh knocking-like advertisement call, unlike T. wambensis. The standard length (34.6���40.8, n= 12) is much smaller compared to the recently re-described T. monticola (SVL 41.2, n= 1) from Olengarua Village in Massai. Subarticular tubercles single and prominent. Webbing on hind toe is moderate; 2 ��� 2 �� phalanges free of web on the forth digit. Glandular ridge below the tympanum broken into two unequal parts. Outer metatarsal tubercle absent; a weak whitish bump present instead (vs. prominent and strongly protruding in some members of the genus). Dorsum consists of greenish brown patches on a lighter background (Figs. 4 A and D) contrasting with irregular dark markings and dark spots on a lighter ground colour in T. krugerensis and also varying from the scantily marked orange-brown to grey-brown of T. marmorata. Small reddish-brown warts present on dorsum (smooth in T. krugerensis, distinctive and prominent tubercles in T. tuberculosa). The advertisement call consists of a series of high-pitched rapidly repeated notes. The note rate is 7.2��� 10.9 s - 1 (T. krugerensis covers entire frequency range with a click; T. tandyi 7���8 notes s - 1; T. delalandii 5���7 notes s - 1; T. cryptotis 9 notes s - 1). The call shows two harmonics with mean frequencies of 1135 Hz and 2253 Hz, with a fainter third harmonic at 3372 Hz. The second harmonic is emphasized (differing from T. krugerensis with multiple emphasized harmonics with dominant harmonic ranging from 1900���3100 Hz; T. tandyi 2500���2700 Hz; T. delalandii 1800���2200 Hz; T. cryptotis 3100���3800 Hz, T. luganga has the first harmonic emphasized 1050���1170 Hz). Each note has a duration of 0.028 s (Fig. 5), varying from T. krugerensis 0.008��� 0.011 s; T. tandyi 0.04 s; T. delalandii 0.039 s; T. cryptotis 0.048 s. Description of Holotype. The body is stout and toad like; head comparatively small (HL/SVL 0.32, HW/SVL 0.41), not wider than trunk, not longer than wide (HL/HW 0.78); snout short (SL/HL 0.54), rounded in dorsal view, blunt in profile (Fig. 4 A), slightly projecting beyond rounded lower jaw, narrower than long (SL/INS 1.47); canthus rostralis is moderately angled; loreal region concave; nostrils situated on slight projections of the canthus, much closer to the snout tip than the eye (ENL/SNL 1.19); eyes directed anterolaterally, slightly protruding, relatively small (ED/HL 0.44); eye diameter shorter than snout (ED/SL 0.81); interorbital distance is longer than upper eyelid (IOS/ED 1.33), and longer than internarial distance (INS/ED 0.84); tympanum visible externally, discontinuous glandular ridge present below, smaller than eye diameter (TD/ED 0.73); upper jaw without dentition; choanae small, round, partly hidden by upper jaw, located anteriorially on the roof of the mouth, vomerine teeth in two oblique rows between the choanae; tongue short and strongly notched behind, one side slightly larger (5.7 at widest part); median lingual processes present; vocal sac single, darkly pigmented; gular pouch flattened anteriorly into a U-shape on the lower lip margin. Dorsal surfaces of head, trunk and limbs generally smooth; ventral surface of limbs, gular and abdomen smooth (Figs. 4 B and E). Fore limbs slender; hand moderately large (HND/SVL 0.24); tips of fingers not enlarged into discs; relative length of fingers: IV I 1 ���1.5 II 1.5 +��� 2 III 2 ��� 2 IV 1.5 -��� 1 V; inner metatarsal tubercle prominent and shovel-shaped; outer absent. Colouration in life. The dorsal pattern consists of greenish brown patches on a lighter background. Reddishbrown dorsal warts spread to surface of head, often bordered with black, larger and more prominent towards the mid dorsum; lateral sides of head and scapular region grey, anterior border of eye black; a darker bar present between the eyes; dorsal stripe absent, white flecks present; ventrum cream, throat dark; hands and thighs show two and three transverse dark markings respectively. Colouration in preservative. Dorsal wart colouration is lost; the other patterns are visible. Paratype variation. Females (SVL 42���43, mean 42.5, n= 2) are larger than males (SVL 34.6���40.8, mean 37.6, n= 12). Colouration of male paratypes is similar to that of the holotype. In some specimens, however, a white thin dorsal line reaching the snout is present in both sexes. Females have a speckled throat (Fig. 4 F). The dorsal glands are relatively larger and scattered in male individuals compared to females with small dense dorsal glands. Measurements are given in Table 3. Eggs and tadpoles. Unknown. Habitat. The specimens were found mostly in open habitats including man-made (e.g. Wamba) and natural water points (e.g. Turkana and Kitobo Forest). Most of the breeding congregations were found in seasonal wetlands. Specimens were observed from within (ca. 10 cm deep) and outside the water bodies (ca. 2 m). Males called from similar positions, but maintained some distance from each other (> 2 m). The following species were found sympatrically or syntopically with the new species: Ptychadena anchietae, Ptychadena mascareniensis, Hildebrandtia ornata and Kassina senegalensis. Distribution. The collection locations of Tomopterna wambensis sp. nov. range from northern to south-eastern Kenya as indicated in Fig. 2. Occurrence in Tanzania is confirmed by a single specimen from Arusha, Tanzania as revealed by the molecular data. However, a possible range extension into Ethiopia is implied by the proximity of the northernmost record of the series at Illeret to the Kenya-Ethiopian border (Etymology. The specific name refers to the type locality, the village of Wamba in central Kenya. Remarks. Based on the records of the type series across Kenya with at least one population in Tanzania, the species is considered widespread. We have recorded the species in both protected and unprotected areas. We presume that it is under no immediate threat and propose a Least Concern status under the current IUCN Redlist criteria., Published as part of Wasonga, Domnick V. & Channing, Alan, 2013, Identification of sand frogs (Anura: Pyxicephalidae: Tomopterna) from Kenya with the description of two new species, pp. 221-240 in Zootaxa 3734 (2) on pages 226-229, DOI: 10.11646/zootaxa.3734.2.7, http://zenodo.org/record/220044