Skip to main content Skip to main navigation menu Skip to site footer
Published: 2022-05-11

Colour patterns to sequences: a perspective on the systematics of the Hyperolius viridiflavus group (Anura: Hyperoliidae) using mitochondrial DNA

Unit for Environmental Sciences and Management, North-West University, Potchefstroom 2520, South Africa
Amphibia Africa reed frogs Hyperolius viridiflavus group taxonomy 16S rRNA distribution


African reed frogs are widespread and highly polymorphic for colour pattern. The Hyperolius viridiflavus complex includes 121 described taxa, including subspecies, which are presently synonymised as 16 species. However, the taxonomy is confused as colour patterns have been used to identify species, despite an inadequate understanding of pattern variation and overlap. This study examined 267 sequences of the 16S rRNA fragment from individuals across the range of the group, from Senegal to Ethiopia, and to the southern tip of Africa. Maximum likelihood analysis showed that there were 14 species that only partially agreed with existing taxonomy. This study recognised the following species: Hyperolius viridiflavus, H. marmoratus, H. marginatus, H. parallelus, H. nitidulus, H. glandicolor, H. tuberculatus, H. mariae, H. noblei, H. goetzei, H. spatzi, H. bangwae, H. dintelmanni and an undescribed species from southern Cameroon. Hyperolius swynnertoni was found to be embedded within H. marmoratus, H. hutsebauti was embedded within H. parallelus, H. rhodesianus and H. pyrrhodictyon were embedded within H. marginatus and H. sheldricki was embedded within H. glandicolor. There was no sequence information to determine the status of Hyperolius nimbae. The ranges of the species are presented, although these are expected to change as more sequences become available. The use of DNA sequences to identify animals is recommended in this group, as the variation in advertisement calls and morphology is not well understood.



  1. Ahl, E. (1931) Zur Systematik der afrikanischen Arten der Baumfroschgattung Hyperolius (Amph. Anur.). Mitteilungen aus dem Zoologischen Museum in Berlin, 17, 1–132.
    Amiet, J.-L. & Goutte, S. (2017) Chants d’Amphibiens du Cameroun. J.-L. Amiet & Éditions Petit Génies, Nyons, 280 pp.
    Barej, M.F., Rödel, M.-O., Loader, S.P., Menegon, M., Gonwouo, N.L., Penner, J., Gvozdík, V., Günther, R., Bell, R.C., Nagel, P. & Schmitz, A. (2014) Light shines through the spindrift – Phylogeny of African torrent frogs (Amphibia, Anura, Petropedetidae). Molecular Phylogenetics and Evolution, 71, 261–273.
    Barratt, C.D., Bwong, B.A, Onstein, R.E., Rosauer, D.F., Menegon, M., Doggart, N., Nagel, P., Kissling, W.D. & Loader, S.P. (2017) Environmental correlates of phylogenetic endemism in amphibians and the conservation of refugia in the coastal forests of Eastern Africa. Diversity and Distributions, 23, 875–887.
    Bell, R.C. (2016) A new species of Hyperolius (Amphibia: Hyperoliidae) from Príncipe Island, Democratic Republic of São Tom and Príncipe. Herpetologica, 72, 343–351.
    Bell, R.C., Parra, J.L., Badjedjea, G., Barej, M.F., Blackburn, D.C., Burger, M., Channing, A., Dehling, J.M., Greenbaum, E., Gvoždík, V., Kielgast, J., Kusamba, C., Lötters, S., McLaughlin, P.J., Nagy, Z.T., Rödel, 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.
    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.
    Bittencourt-Silva, G.B. (2019) A herpetological survey of western Zambia. Amphibian & Reptile Conservation, Special Section, 13 (2), e181, 1–28.
    Bittencourt-Silva, G.B., Bayliss, J. & Conradie, W. (2020) First herpetological surveys of Mount Lico and Mount Socone, Mozambique. Amphibian & Reptile Conservation, 14, 198–217.
    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.
    Boettger, O. (1888) Materialien zur Fauna des unteren Congo II. Reptilien und Batrachier. Bericht über die Senckenbergische Naturforschende Gesellschaft in Frankfurt am Main, 1888, 3–108.
    Bosch, J., De la Riva, I. & Márquez, R. (2000) Advertisement calls of seven species of hyperoliid frogs, from Equatorial Guinea. Amphibia-Reptilia, 21, 246–255.
    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.
    Center for Conservation Bioacoustics (2019) Raven Pro: Interactive Sound Analysis Software. Version 1.6.1. Computer software. The Cornell Lab of Ornithology, Ithaca, New York. Available from: (accessed 26 April 2022)
    Ceríaco, 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.
    Channing, A., Dehling, J.M., Lötters, S., Ernst, R. (2016) Species boundaries and taxonomy of the African river frogs (Amphibia: Pyxicephalidae: Amietia). Zootaxa, 4155 (1), 1–76.
    Channing, A., Hillers, A., Lötters, S., Rödel, M.-O., Schick, S., Conradie, W., Rödder, 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.
    Channing, A. & Howell, K.M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, 418 pp.
    Channing, A. & Rödel, M.-O. (2019) Frogs and other amphibians of Africa. Struik Nature, Cape Town, 408 pp.
    Channing, A., Schmitz, A., Burger, M., Kielgast, J. (2013) A molecular phylogeny of African dainty frogs, with the description of four new species (Anura: Pyxicephalidae: Cacosternum). Zootaxa, 3701 (50, 518–550.
    Conradie, W., Branch, W.R., Measey, G.J. & Tolley, K.A. (2012) A new species of Hyperolius Rapp, 1842 (Anura: Hyperoliidae) from the Serra da Chela Mountains, south-western Angola. Zootaxa, 3269 (1), 1–17.
    Conradie, W., Branch, W.R. & Tolley, K.A. (2013) Fifty shades of grey: giving colour to the poorly known Angolan ashy reed frog (Hyperoliidae: Hyperolius cinereus), with the description of a new species. Zootaxa, 3635 (3), 201–223.
    Conradie, W., Bittencourt-Silva, G.B., Farooq, H., Loader, S.P., Menegon, M. & Tolley, K.A. (2018a) New species of Mongrel Frogs (Pyxicephalidae: Nothophryne) for northern Mozambique inselbergs. African Journal of Herpetology, 67, 61–85.
    Conradie, W., Verburgt, L., Portik, D.M., Ohler, A., Bwong, B.A. & Lawson, L.P. (2018b). A new reed frog (Hyperoliidae: Hyperolius) from coastal northeastern Mozambique. Zootaxa, 4379 (2), 177–198.
    Dehling, J.M. (2012) An African glass frog: A new Hyperolius species (Anura: Hyperoliidae) from Nyungwe National Park, southern Rwanda. Zootaxa, 3391 (1), 52–64.
    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.
    Dehling, J.M. & Sinsch, U. (2019) Partitioning of morphospace in larval and adult reed frogs (Anura: Hyperoliidae: Hyperolius) of the Central African Albertine Rift. Zoologische Anzeiger, 280, 65–77.
    Deichmann, J.L., Mulcahy, D.G., Vanthomme, H., Tobi, E., Wynn, A.H., Zimkus, B.M. & McDiarmid, R.W. (2017) How many species and under what names? Using DNA barcoding and GenBank data for west Central African amphibian conservation. PLoS ONE, 12 (11), e0187283.
    De Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.
    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.
    Drewes, R.C. & Wilkinson, J.A. (2004) The California Academy of Sciences Gulf of Guinea Expedition (2001). I. The taxonomic status of the genus Nesionixalus Perret, 1976 (Anura: Hyperoliidae), treefrogs of São Tom and Príncipe, with comments on the genus Hyperolius. Proceedings of the California Academy of Sciences, 55, 395–407.
    Duff-Mackay, A. & Schiøtz, A. (1971) A new Hyperolius (Amphibia Anura) from Kenya. Journal of the East Africa Natural History Society and National Museum, 29, 1–3.
    Du Preez, L. & Carruthers, V. (2017) Frogs of southern Africa. Struik Nature, Cape Town, 519 pp.
    Foskett, R. (1965) The Zambezi journal and letters of Dr John Kirk 1858–63. Vol. 1. Oliver & Boyd, Edinburgh and London, xxi + 317 pp.
    Friis, I. & Edwards, S. (2001) By whom and when was the flora of Ethiopia and Eritrea named? Biologiske Skrifter, 54, 103–136.
    Frost, D.R. (2022) Amphibian Species of the World: an online reference. Version 6.1. American Museum of Natural History, New York. Electronic database accessible. Available from: (accessed 4 January 2022)
    Guichenot, A. (1851) Reptiles in: des Murs, O., Prévost, F., Guichenot, A. & Guérin-Menneville, F.E. (Eds.), Voyage en Abyssinie. Quatrième Partie. Histoire Naturelle. Zoologie. Tome Sixième. Arthus Bertrand, Paris, pp. 185–222.
    Hayes, D.T., Ceriaco, L.M.P., Jongsma, G.F.M., Bandeira, S., Hedley, R.W., Stanley, E.L., Bauer, A.M. & Blackburn, D.C. (undated) Amphibian surveys at two important type localities in Malanje Province, Angola. [unpublished]
    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.
    Hoang, D.T., Chernomor, O., Von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2017) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35, 518–522.
    Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., Von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589.
    Kishino, H. & Hasegawa, M. (1989) Evaluation of the maximum-likelihood estimate of the evolutionary tree topologies from DNA sequence data, and the branching order in Hominoidea. Journal of Molecular Evolution, 29, 170–179.
    Kurabayashi, A., Matsui, M., Belabut, D.M., Yong, H.S., Ahmed, N., Sudin, A., Kuramoto, M., Hamidy, A. & Sumida, M. (2011) From Antarctica or Asia? New colonization scenario for Australian-New Guinea narrow mouth toads suggested from the findings on a mysterious genus Gastrophrynoides. BMC Evolutionary Biology, 11, 175.
    Köhler, J., Scheelke, K., Schick, S., Veith, M. & Lötters, S. (2005) Contribution to the taxonomy of hyperoliid frogs (Amphibia: Anura: Hyperoliidae): advertisement calls of twelve species from East and Central Africa. African Zoology, 40, 127–142.
    Köhler, J., Jansen, M., Rodríquez, A., Kok, P.J.R., Toledo, L.P., Emmrich, M., Glaw, F., Haddad, C.F.B., Rödel, M.-O. & Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa, 4251, 1–124.
    Kouamé, N.G., Assemian, N.E., Tohé, B., Adeba, P.J., Gourène & Rödel, M.-O. (2016) Rediscovery of the Mount Nimba Reedfrog, Hyperolius nimbae Laurent, 1958, in western Ivory Coast. Herpetozoa, 29, 3–13.
    Lanza, B. (1990) Amphibians and reptiles of the Somali Democratic Republic: check list and biogeography. Biogeographia, 14, 407–465.
    Largen, M.J. (2001) Catalogue of the amphibians of Ethiopia, including a key for their identification. Tropical Zoology, 14, 307–402.
    Largen, M.J. & Spawls, S. (2010) The Amphibians and Reptiles of Ethiopia and Eritrea. Edition Chimaria, Frankfurt am Main, 693 pp.
    Larson, J.G. & Zimkus, B.M. (2018) Preliminary assessment of the frog assemblages from sites adjacent to three national parks in Gabon. Herpetological Conservation and Biology, 13, 240–256.
    Laurent, R. (1943a) Les Hyperolius (Batraciens) du Musée du Congo. Annales du Musée du Congo Belge, 4, 61–140.
    Laurent, R. (1943b) Contribution a l’étude des genres Megalixalus et Hyperolius. Bulletin du Musée royal d’Histoire naturelle de Belgique, 19 (30), 1–20.
    Laurent, R.F. (1947) Sur quelques rhacophorides nouveaux de Parc National Albert appartenant aux genres Megalixalus et Hyperolius. Bulletin du Musée Royal d’Histoire Naturelle de Belgique, 23, 1–8.
    Laurent, R. (1948 “1947”) Two new forms of the genus Hyperolius. Annals and Magazine of Natural History, Series 2, 14, 294–296.
    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.
    Laurent, R. (1951a) Catalogue des rainettes africaines (genres Afrixalus et Hyperolius) de la collection du Museum National d’Histoire Naturelle de Paris. Annales Sociét Royale Zoologique de Belgique, 82, 23–50.
    Laurent, R. (1951b) Quelques données nouvelles sur la systématique et l’ecologie du genre Hyperolius. Annales Sociét Royale Zoologique de Belgique, 82, 329–339.
    Laurent, R. (1952) Aperçu des formes actuellement reconnaissables dans la superespece Hyperolius marmoratus. Annales Sociét Royale Zoologique de Belgique, 82, 379–397.
    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.
    Laurent, R.F. (1961) Note sur les Hyperolius et quelques Afrixalus (Salientia) du Musée de Berlin. Revue de Zoologie et de Botanique Africaines, 64, 65–96.
    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.F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxième Série, 22, 1–125.
    Laurent, R.F. (1976) Nouveaux commentaires sur la superespece Hyperolius viridiflavus (Anura). Annales de Musee Royal de l’Afrique Centrale, Series 8, 213, 69–114.
    Laurent, R.F. (1983) La superespèce Hyperolius viridiflavus (Duméril & Bibron, 1841) (Anura Hyperoliidae) en Afrique Centrale. Monitore Zoologico Italiano, New Series, Supplement 18, 1–93.
    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.
    Lefebvre, T., Petit, A, Quartin-Dillon, R. & Vignaud, J. (1845) s.n. In: Bertrand, A. (Ed.), Voyage en Abyssinie. Libraire de la Sociét de Geographie, Paris, pp. 1–386.
    Li, J., Che, J., Bain, R.H., Zhao, E. & Zhang, Y. (2008) Molecular phylogeny of Rhacophoridae (Anura): A framework of taxonomic reassignment of species within the genera Aquixalus, Chiromantis, Rhacophorus, and Philautus. Molecular Phylogenetics and Evolution, 48, 302–312.
    Lötters, S. & Schmitz, A. (2004) A new species of tree frog (Amphibia; Hyperolius) from the Bakossi Mountains, south-west-Cameroon. Bonner zoologische Beiträge, 52, 149–154.
    Lötters, S., Schick, S., Scheelke, K., Teege, P., Kosuch, J., Rotich, D. & Veith, M. (2004) Bio-sketches and partitioning of sympatric reed frogs, genus Hyperolius (Amphibia; Hyperoliidae), in two humid tropical African forest regions. Journal of Natural History, 38, 1969–1997.
    Loveridge, A. (1933) Reports on the scientific results of an expedition to the southwestern highlands of Tanganyika Territory. VII. Herpetology. Bulletin of the Museum of Comparative Zoology, 74, 197–416.
    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.
    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.
    Marques, M.O., Ceríaco, L.M.P., Blackburn, D.C. & Bauer, A.M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840–2017). Proceedings of the California Academy of Sciences, Series 4, 65, 1–501.
    Matsui, M., Hamidy, A., Belabut, D.M., Ahmad, N., Panha, S., Suding, A., Khonsue, W., Oh, H.-S., Yong, H.-S., Jiangi, J.-P. & Nishikawa, K. (2011) Systematic relationships of Oriental tiny frogs of the family Microhylidae (Amphibia, Anura) as revealed by mtDNA genealogy. Molecular Phylogenetics and Evolution, 61, 167–176.
    Maya-Soriano, M.J., Holt, W.V. & Lloyd, R.E. (2012) Biobanked amphibian samples confirmed to species level using 16S rRNA DNA barcodes. Biopreservation and Biobanking, 10, 22–28.
    Mercurio, V. (2011) Amphibians of Malawi. Edition Chimaira, Frankfurt am Main, 393 pp.
    Nguyen, L.-T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum likelihood phylogenies. Molecular Biology and Evolution, 32, 268–274.
    Nneji, L.M., Adeola, A.C., Okeyoyin, A.O., Oladipo, O.C., Saidu, J.Y., Ugwumba, A.A. & Adeola, A.C. (2019) Assessing the effectiveness of molecular data for species identification and diversity studies of Nigerian herpetofauna. Mitochondrial DNA B Resource, 4, 589–593.
    Parker, H.W. (1937) Reptiles and amphibians from the Mafia Archipelago. Annals and Magazine of Natural History, Series 10, 20, 629–632.
    Perret, J.-L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie, 93, 289–464.
    Perret, J.-L. (1976) Révision des amphibiens africains et principalement des types, conservés au Musée Bocage de Lisbonne. Arquivos do Museu Bocage, Lisboa, Série 2 6, 15–34.
    Pickersgill, M. (2007) Frog Search. Edition Chimaira, Frankfurt am Main, 574 pp.
    Pilsbry, H.A. (1919) A review of the land Molluscs of the Belgian Congo chiefly based on the collections of the American Museum Congo Expedition, 1900–1915. Bulletin of the American Museum of Natural History, 40, 1–370.
    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], e131, 37–67.
    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., Gvoždík, 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., Lötters, S., Van der Meijden, A., Menegon, M., Müller, S., Nagy, Z.T., Ofori-Boateng, C., Ohler, A., Papenfuss, T.J., Rossler, D., Sinsch, U., Rödel, 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.
    Portik, D.M. & Blackburn, D.C. (2016) The evolution of reproductive diversity in Afrobatrachia: A phylogenetic comparative analysis of an extensive radiation of African frogs. Evolution, 70-9, 2017–2032.
    Poynton, J.C. (1977) A new Bufo and associated Amphibia from southern Tanzania. Annals of the Natal Museum, 23, 37–41.
    Poynton, J.C. (1991) Amphibians of Southeastern Tanzania, with special reference to Stephopaedes and Mertensophryne (Bufonidae). Bulletin of the Museum of Comparative Zoology, 152, 451–473.
    Poynton, J.C. & Broadley, D.G. (1987) Amphibia Zambesiaca 3. Rhacophoridae and Hyperoliidae, Annals of the Natal Museum, 28, 161–229.
    Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–583.
    Rambaut, A. (2018) FigTree Tree Figure Drawing Tool. Version 1.4.4. Available from: (accessed 26 April 2022)
    Reeder, J.E. (2019) Amphibians of northern KwaZulu-Natal: A phylogenetic study. Master of Science dissertation, North-West University, Potchefstroom, 114 pp.
    Rödel, M.-O. (2000) Herpetofauna of West Africa. Vol 1 Amphibians of the West African Savanna. Edition Chimaira, Frankfurt am Main, 332 pp.
    Rödel, 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.
    Sabaj, M.H. (2019) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: An Online Reference. Version 7.1. 21 March 2019. American Society of Ichthyologists and Herpetologists, Washington, D.C. Electronically accessible. Available from: (accessed 26 April 2022)
    Schick, S., Kielgast, J., Rödder, D., Muchai, V., Burger, M. & Lötters, S. (2010) New species of reed frog from the Congo basin with discussion of paraphyly in cinnamon-belly reed frogs. Zootaxa, 2501 (1), 23–36.
    Schiøtz, A. (1967) The treefrogs (Rhocophoridae) of West Africa. Spolia zoologica Musei hauniensis, 25, 1–346.
    Schiøtz, A. (1971) The superspecies Hyperolius viridiflavus (Anura). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 134, 21–76.
    Schiøtz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.
    Schiøtz, A. (1982) Two new Hyperolius (Anura) from Tanzania. Steenstrupia, 8, 269–276.
    Schiøtz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.
    Shimodaira, H. (2002) An approximately unbiased test of phylogenetic tree selection. Systematic Biology, 51, 492–508.
    Shimodaira, H. & Hasegawa, M. (1999) Multiple comparisons of log-likelihoods with applications to phylogenetic inference. Molecular Biology and Evolution, 16, 1114–1116.
    Sinsch, U., Lümkemann, K., Rosar, K., Schwarz, C. & Dehling, J.M. (2012) Acoustic niche partitioning in an anuran community inhabiting an Afromontane wetland (Butare, Rwanda). African Zoology, 47, 60–73.
    Starnberger, I., Poth, D., Peram, P.S., Schulz, S., Vences, M., Knudsen, J., Barej, M.F., Rödel, M.-O., Walzl, M. & Hödl, W. (2013) Take time to smell the frogs: vocal sac glands of reed frogs (Anura: Hyperoliidae) contain species-specific chemical cocktails. Biological Journal of the Linnean Society, 110, 828–838.
    Sweeney, B.A., Hoksza, D., Nawrocki, E.P., Ribas, A.E., Madeira, F., Cannone, J.J., Gutell, R., Maddala, A., Meade, C.D., Williams, L.D., Petrov, A.S., Chan, P.P., Lowe, T.M., Finn, R.D. & Petrov, A.I. (2021) R2DT is a framework for predicting and visualising RNA secondary structure using templates. Nature Communications, 12, 3494.
    Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [software]
    Tillack, F., De Ruiter, R. & Rödel, M.-O. (2021) A type catalogue of the reed frogs (Amphibia, Anura, Hyperoliidae) in the collection of the Museum für Naturkunde Berlin (ZMB) with comments on historical collectors and expeditions. Zoosystematics and Evolution, 97, 407–450.
    Tornier, G. (1897) Die Kriechthiere Deutsch-Ost-Afrikas. Dietrich Reimer, Berlin, 206 pp.
    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.
    Trifinopoulos, J., Nguyen, L.-T., Von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44, W1, W232–W235.
    Turner, A.A. & Channing, A. (2008) A new species of Arthroleptella Hewitt, 1926 (Anura: Pyxicephalidae) from the Klein Swartberg Mountain, Caledon, South Africa. African Journal of Herpetology, 57, 1–12.
    Van den Abeele, M. (1968) s.n. In: Burgeon (Louis-Jules-Léon). Biographie belge d'Outre-mer. Academie royale des Sciences d'Outre-mer, Brussels, pp. 147–149.
    Veith, M., Kosuch, J., Rödel, M.-O., Hillers, A., Schmitz, A., Burger, M. & Lötters, S. (2009) Multiple evolution of sexual dichromatism in African reed frogs. Molecular Phylogenetics and Evolution, 51, 388–393.
    Vences, M. (2000) Phylogenetic studies on ranoid frogs (Amphibia: Anura) with a discussion of the origin and evolution of the vertebrate clades of Madagascar. PhD thesis, Rheinische Friedrich-Wilhelms-Universität Bonn, Bonn, 294 pp.
    Vences, M, Thomas, M., Van der Meijden, A., Chiari, Y. & Vietes, D.R. (2005) Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology, 2, 5.
    Vences, M., Köhler, J., Crottini, A. & Glaw, F. (2010) High mitochondrial divergence meets morphological and bioacoustic conservatism: Boophis quasiboehmei sp. n., a new cryptic treefrog species from south-eastern Madagascar. Bonn zoological Bulletin, 57, 241–255.
    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.
    Wilson, L. & Channing, A. (2019) A new sand frog from Namaqualand, South Africa (Pyxicephalidae: Tomopterna). Zootaxa, 4609 (2), 225–246.
    Wollenberg, K.C., Glaw, F., Meyer, A. & Vences, M. (2007) Molecular phylogeny of Malagasy reed frogs, Heterixalus, and the relative performance of bioacoustics and color-patterns for resolving their systematics. Molecular Phylogenetics and Evolution, 45, 14–22.