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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."]}