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Type: Article
Published: 2022-08-10
Page range: 201-232
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Systematics of the Central African Spiny Reed Frog Afrixalus laevis (Anura: Hyperoliidae), with the description of two new species from the Albertine Rift

Department of Biological Sciences, University of Texas at El Paso, 500 W. University Ave., El Paso, TX 79912, USA
Herpetology Department, Institute for Biodiversity Science and Sustainability, California Academy of Sciences, San Francisco, CA 94118, USA
Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Lawrence, KS 66045, USA 4Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
Department of Biological Sciences, University of Texas at El Paso, 500 W. University Ave., El Paso, TX 79912, USA
Department of Ecology and Aquatic Biodiversity, Biodiversity Monitoring Centre, University of Kisangani, Kisangani, Democratic Republic of the Congo
6 Museum für Naturkunde Berlin—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany
Department of Environment & Social Safeguards (DESS), Uganda National Roads Authority (UNRA), P.O. Box 28487, Kampala, Uganda
Department of Biological Sciences, University of Texas at El Paso, 500 W. University Ave., El Paso, TX 79912, USA
Conservation of Biodiversity and Natural Resources and Climate Protection by Sustainable Agriculture and Forestry at Cyamudongo Forest (Rwanda), c/o Cooperation University of Koblenz-Landau/Rwanda, SH8 AVE 26, PO Box 344, Huye, Rwanda
Laboratory of Zoology, Faculty of Science, University of Yaoundé I, P.O. Box 812, Yaoundé, Cameroon
Museum für Naturkunde Berlin—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany
Department of Biology, Coe College, 1220 1st Avenue NE, Cedar Rapids, Iowa 52402, USA
Institut Congolais pour la Conservation de la Nature, P.O. Box 852, Bukavu, Democratic Republic of the Congo
Laboratoire d’Hérpétologie, Département de Biologie, Centre de Recherche en Sciences Naturelles, Lwiro, Democratic Republic of the Congo
Department of Environment & Social Safeguards (DESS), Uganda National Roads Authority (UNRA), P.O. Box 28487, Kampala, Uganda
Department of Ecology and Biodiversity of Earth Resources, Centre de Surveillance de la Biodiversité of the University of Kisangani, Kisangani, Democratic Republic of the Congo
6 Museum für Naturkunde Berlin—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany
6 Museum für Naturkunde Berlin—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany
Department of Biology, University of Texas at Arlington, Box 19498, Arlington, Texas 76017, USA
Department of Biological Sciences, University of Texas at El Paso, 500 W. University Ave., El Paso, TX 79912, USA
Institut für Integrierte Naturwissenschaften, Abteilung Biologie, AG Zoologie, Universität Koblenz-Landau, Universitätsstraße 1, 56070 Koblenz, Germany
Amphibia Afromontane Endemism Conservation Species Delimitation Phylogeny


The geographically widespread species Afrixalus laevis (Anura: Hyperoliidae) currently has a disjunct distribution in western Central Africa (Cameroon, Equatorial Guinea, Gabon, and possibly adjacent countries) and the area in and near the Albertine Rift in eastern Democratic Republic of the Congo and neighboring countries. At least two herpetologists have previously suggested that these disjunct populations represent distinct species, and herein, we utilize an integrative taxonomic approach with molecular and morphological data to reconcile the taxonomy of these spiny reed frogs. We sequenced 1554 base pairs of the 16S and RAG1 genes from 34 samples of A. laevis and one sample of A. orophilus (sympatric with eastern populations of A. laevis), and combined these data with previously sequenced GenBank Afrixalus samples via the bioinformatics toolkit SuperCRUNCH. Phylogenetic trees, dated phylogenetic analyses, and species-delimitation analyses were generated with RAxML, BEAST, and BPP, respectively. Eleven mensural characters were taken from multiple specimens of A. laevis and A. orophilus, and compared with paired t-tests and analyses of covariance. These combined results suggested populations of A. laevis in western Central Africa (Cameroon and Bioko Island, Equatorial Guinea) represent one species, whereas populations from the Albertine Rift and nearby forests represent two undescribed taxa that are sister to A. dorsimaculatus. The two new species (A. lacustris sp. nov. and A. phantasma sp. nov.) are distinguished by our phylogenetic and species-delimitation analyses, significant differences in several mensural characters, qualitative morphological differences, and by their non-overlapping elevational distribution.



  1. Ahl, E. (1930) Ueber die afrikanischen Arten der Baumfroschgattung Megalixalus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1930, 89–102.
    Amiet, J.-L. (1991) Images d’Amphibiens camerounais. IV. Les constructeurs de nids. Alytes, 9 (3), 71–77.
    Amiet, J-L. (2009) Observations sur les Afrixalus du Cameroun (Amphibia, Anura, Hyperoliidae). Revue Suisse de Zoologie, 116, 53–92.
    Amiet, J-L. (2012) Les Rainettes du Cameroun (Amphibiens Anoures). La Nef des Livres, Saint-Nazaire, 591 pp.
    Amiet, J.-L. & Goutte, S. (2017) Chants d’Amphibiens du Cameroun. J.-L. Amiet, Nyons, France & Editions Petit Génie, Saint-Nazaire, 280 pp., 4 audio CDs.
    Ayebare, S., Plumptre, A.J., Kujirakwinja, D. & Segan, D. (2018) Conservation of the endemic species of the Albertine Rift under future climate change. Biological Conservation, 220, 67–75.
    Bachman, S., Moat, J., Hill, A., de la Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial Conservation Assessment Tool. ZooKeys, 150, 117−126.
    Badjedjea, G., Masudi, F.M., Dudu Akaibe, B. & Gvoždik, V. (2022) Amphibians of Kokolopori: An introduction to the amphibian fauna of the Central Congolian Lowland Forests, Democratic Republic of the Congo. Amphibian & Reptile Conservation, 16 (1), 35–70.
    Barej, M.F., Schmitz, A., Menegon, M., Hillers, A., Hinkel, H., Böhme, W. & Rödel, M.-O. (2011) Dusted off—the African Amietophrynus superciliaris-species complex of giant toads. Zootaxa, 2772 (1), 1–32.
    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. & Zamudio, K.R. (2017) Idiosyncratic responses to climate-driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands. Molecular Ecology, 26 (19), 5223–5244.
    Channing, A. & Howell, K.M. (2006) Amphibians of East Africa. Cornell University Press, Ithaca, New York, New York, xi + 418 pp.
    Channing, A. & Rödel, M.-O. (2019) Field Guide to the Frogs & Other Amphibians of Africa. Struik Nature, Cape Town, 408 pp.
    Channing, A., Rödel, M.-O. & Channing, J. (2012) Tadpoles of Africa: The Biology and Identification of All Known Tadpoles in sub-Saharan Africa. Edition Chimaira, Frankfurt am Main, 402 pp.
    Charles, K.L., Bell, R.C., Blackburn, D.C., Burger, M., Fujita, M.K., Gvoždík, V., Jongsma, G.F.M., Kouete, M.T., Leaché, A.D. & Portik, D.M. (2018) Sky, sea, and forest islands: Diversification in the African leaf-folding frog Afrixalus paradorsalis (Anura: Hyperoliidae) of the Lower Guineo-Congolian rain forest. Journal of Biogeography, 45 (8), 1781–1794.
    Chiari, Y., Vences, M., Vieites, D.R., Rabemananjara, F., Bora, P., Ramilijaona Ravoahangimalala, O. & Meyer, A. (2004) New evidence for parallel evolution of colour patterns in Malagasy poison frogs (Mantella). Molecular Ecology, 13 (12), 3763–3774.
    de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, New York, New York, pp. 57–75.
    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879–886.
    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 (Anura: Hyperoliidae). Herpetozoa, 28 (1/2), 39–47.
    Dewynter, M. & Frétey, T. (2019) Liste taxonomique commentée et catalogue illustré des Amphibiens du Gabon. Les cahiers de la fondation Biotope, 27, 1–84.
    Doumenge, C. (1998) Forest diversity, distribution, and dynamique in the Itombwe Mountains, South-Kivu, Congo Democratic Republic. Mountain Research and Development, 18 (3), 249–264.
    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, i–x + 1–70.
    Drewes, R.C. & Vindum, J.V. (1994) Amphibians of the Impenetrable Forest, southwest Uganda. Journal of African Zoology, 108, 55–70.
    Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29 (8), 1969–1973.
    Dubois, A. (1986 “1985”) Miscellanea nomenclatorica batrachologica (IX). Alytes, 4, 97–100.
    Flouri, T., Jiao, X., Rannala, B. & Yang, Z. (2018) Species tree inference with BPP using genomic sequences and the multispecies coalescent. Molecular Biology and Evolution, 35 (10), 2585–2593.
    Frost, D.R. (2022) Amphibian Species of the World: An Online Reference. Version 6.1 American Museum of Natural History, New York, New York. Electronic Database accessible. Available from: (accessed 27 April 2022)
    Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P.E., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370.[0001:TATOL]2.0.CO;2
    Greenbaum, E. (2017) Emerald Labyrinth: A Scientist’s Adventures in the Jungles of the Congo. ForeEdge, Lebanon, New Hampshire, xv + 329 pp.
    Greenbaum, E., Meece, J., Reed, K. & Kusamba, C. (2015) Extensive occurrence of the amphibian chytrid fungus in the Albertine Rift, a Central African amphibian hotspot. The Herpetological Journal, 25 (2), 91–100.
    Hirschfeld, M, Blackburn, D.C., Burger, M., Greenbaum, E., Zassi-Boulou, A.-G. & Rödel, M.-O. (2015) Two new species of Cardioglossa (Anura: Arthroleptidae) from the Central African rainforests. African Journal of Herpetology, 64 (2), 81–102.
    Hughes, D.F., Tolley, K.A., Behangana, M., Lukwago, W., Menegon, M., Dehling, J.M., Stipala, J., Tilbury, C.R., Khan, A.M., Kusamba, C. & Greenbaum, E. (2018) Cryptic diversity in Rhampholeon boulengeri (Sauria: Chamaeleonidae), a pygmy chameleon from the Albertine Rift biodiversity hotspot. Molecular Phylogenetics and Evolution, 122, 125–141.
    Hydeman, M.E., Longo, A.V., Velo-Antón, G., Rodríguez, D., Zamudio, K.R. & Bell, R.C. (2017) Prevalence and genetic diversity of Batrachochytrium dendrobatidis in Central African islands and continental amphibian communities. Ecology and Evolution, 7 (19), 7729–7738.
    IUCN (2021) IUCN Red List Categories and Criteria: Version 2021-3. Available from: (accessed 25 May 2022)
    Jaynes, K.E., Myers, E.A., Gvoždík, V., Blackburn, D.C., Portik, D.M., Greenbaum, E., Jongsma, G.F.M., Rödel, M.-O., Badjedjea, G., Bamba-Kaya, A., Baptista, N.L., Akuboy, J.B., Ernst, R., Kouete, M.T., Kusamba, C., Masudi, F.M., McLaughlin, P.J., Nneji, L.M., Onadeko, A.B., Penner, J., Vaz Pinto, P., Stuart, B.L., Tobi, E., Zassi-Boulou, A.-G., Leaché, A.D., Fujita, M.K. & Bell, R.C. (2022) Giant tree frog diversification in West and Central Africa: Isolation by physical barriers, climate, and reproductive traits. Molecular Ecology 31: 3979–3998.
    Jongsma, G.F.M., Barej, M.F., Barratt, C.D., Burger, M., Conradie, W., Ernst, R., Greenbaum, E., Hirschfeld, M., Leaché, A.D., Penner, J., Portik, D.M., Zassi-Boulou, A.-G., Rödel, M.-O. & Blackburn, D.C. (2018) Diversity and biogeography of frogs in the genus Amnirana (Family Ranidae) across sub-Saharan Africa. Molecular Phylogenetics and Evolution, 120, 274–285.
    Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution, 30 (4), 722–780.
    Katoh, K., Kuma, K., Toh, H. & Miyata, T. (2005) MAFFT version 5: Improvement in accuracy of multiple sequence alignment. Nucleic Acids Research, 33 (2), 511–518.
    Katoh, K., Misawa, K., Kuma, K. & Miyata, T. (2002) MAFFT: A novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research, 30 (14), 3059–3066.
    Laurent, R. (1950) Genres Afrixalus et Hyperolius (Amphibia Salientia). Exploration du Parc National Albert, Mission G.F. De Witte (1933–1935), 64, 5–120, 7 pls.
    Laurent, R. (1952) Batraciens et reptiles récemment acquis par le Musée du Congo belge. Revue de zoologie et de botanique africaines, 45, 198–203.
    Laurent, R.F. (1955) Aperçu de la biogéographie des batraciens et des reptiles de la région des grand lacs. Bulletin de la Société Zoologique de France, 79, 290–310.
    Laurent, R.F. (1964) Adaptive modifications in frogs of an isolated highland fauna in Central Africa. Evolution, 18 (3), 458–467.
    Laurent, R.-F. (1965) Contribution à l’histoire de l’herpétologie congolaise et bibliographie générale. Académie royal des sciences d’outre-mer. Classe des Sciences naturelles et médicales, Nouvelle Série, 16 (3), 1–55.
    Laurent, R.F. (1972) Amphibiens. Exploration du Parc National des Virunga, Deuxième Série, 22, 1–125, 11 pls.
    Laurent, R.F. (1982) Le genre Afrixalus Laurent (Hyperoliidae) en Afrique Centrale. Annales du Musée Royal de l’Afrique Centrale, Série in Octavo, Science Zoologique, 235, 1–93.
    Laurent, R.F. (1983) About the herpetofauna of Central African montane forest. In: Rhodin, A.G.J. & Miyata, K. (Eds.), Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Cambridge, Massachusetts, pp. 350–358.
    Leaché, A.D. & Fujita, M.K. (2010) Bayesian species delimitation in West African forest geckos (Hemidactylus fasciatus). Proceedings of the Royal Society B: Biological Sciences, 277 (1697), 3071–3077.
    Li, W. & Godzik, A. (2006) Cd-hit: A fast program for clustering and comparing large sets of protein or nucleotide sequences. Bioinformatics, 22 (13), 1658–1659.
    Liedtke, H.C., Soler-Navarro, D.J., Gomez-Mestre, I., Loader, S.P. & Rödel, M.-O. (2021) Parallel diversification of the African tree toad genus Nectophryne (Bufonidae). Molecular Phylogenetics and Evolution, 162, 107184.
    Liem, S.S. (1970) The morphology, systematics, and evolution of the Old World treefrogs (Rhacophoridae and Hyperoliidae). Fieldiana Zoology, 57, i–vii + 1–145.
    Mertens, R. (1938) Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 442, 1–52.
    Mertens, R. (1940) Amphibien aus Kamerun. Senckengergiana, 22, 103–135.
    Meyers, J.B., Rosendahl, B.R., Harrison, C.G. & Ding, Z.-D. (1998) Deep-imaging seismic and gravity results from the offshore Cameroon Volcanic Line, and speculation of African hotlines. Tectonophysics, 284 (1–2), 31–63.
    Nečas, T., Badjedjea, G., Vopálenský, M. & Gvoždík, V. (2021) Congolius, a new genus of African reed frog endemic to the central Congo: A potential case of convergent evolution. Scientific Reports, 11, 8338.
    Packard, G.C. & Boardman, T.J. (1999) The use of percentages and size-specific indices to normalize physiological data for variation in body size: Wasted time, wasted effort? Comparative Biochemistry and Physiology Part A, 122, 37–44.
    Palumbi, S., Martin, A., Roman, S., McMillan, W.O., Stice, L. & Grabowski, G. (2002) The Simple Fool’s Guide to PCR. Version 2. University of Hawaii, Honolulu, 45 pp.
    Parker, H.W. (1936) The amphibians of the Mamfe Division, Cameroons—I. Zoogeography and systematics. Proceedings of the Zoological Society of London, 106 (1), 135–163.
    Passmore, N.I. & Carruthers, V.C. (1978) Further comment on English common names for South African frogs. Journal of the Herpetological Association of Africa, 19, 2–7.
    Perret, J.-L. (1960) Batraciens du genre Afrixalus du Cameroun. Revue de zoologie et de botanique africaines, 61, 366–374.
    Perret, J.-L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie, 93, 289–464.
    Pickersgill, M. (1984) Three new Afrixalus (Anura: Hyperoliidae) from south-eastern Africa. Durban Museum Novitates, 13, 203–220.
    Pickersgill, M. (1992) A new species of Afrixalus (Amphibia, Anura, Hyperoliidae) from eastern Africa. Steenstrupia, 18, 145–148.
    Pickersgill, M. (1995) Afrixalus dabagae Pickersgill, 1992, relegated to synonymy. Journal of the Herpetological Association of Africa, 44, 58.
    Pickersgill, M. (2007) Frog Search: Results of Expeditions to Southern and Eastern Africa. Edition Chimaira, Frankfurt am Main, 574 pp.
    Plumptre, A.J., Davenport, T.R.B., Behangana, M., Kityo, R., Eilu, G., Ssegawa, P., Ewango, C., Meirte, D., Kahindo, C., Herremans, M., Kerbis Peterhans, J., Pilgrim, J.D., Wilson, M., Languy, M. & Moyer, D. (2007) The biodiversity of the Albertine Rift. Biological Conservation, 134 (2), 178–194.
    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., Leaché, A.D., Loader, S.P., Lötters, S., van der Meijden, A., Menegon, M., Müller, S., Nagy, Z.T., Ofori-Boateng, C., Ohler, A., Papenfuss, T.J., Rößler, D., Sinsch, U., Rödel, M.-O., Veith, M., Vindum, J.V., Zassi-Boulou, A.-G. & McGuire, J.A. (2019) Sexual dichromatism drives diversification within a major radiation of African amphibians. Systematic Biology, 68 (6), 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.
    Portik, D.M., Leaché, A.D., Rivera, D., Blackburn, D.C., Rödel, M.-O., Barej, M.F., Hirschfeld, M., Burger, M. & Fujita, M.K. (2017) Evaluating mechanisms of diversification in a Guineo-Congolian forest frog using demographic model selection. Molecular Ecology, 26 (19), 5245–5263.
    Portik, D.M. & Wiens, J.J. (2020) SuperCRUNCH: A bioinformatics toolkit for creating and manipulating supermatrices and other large phylogenetic datasets. Methods in Ecology and Evolution, 11 (6), 763–772.
    Portillo, F. & Greenbaum, E. (2014) A new species of the Leptopelis modestus complex (Anura: Arthroleptidae) from the Albertine Rift of Central Africa. Journal of Herpetology, 48 (3), 394–406.
    Pramuk, J.B., Robertson, T., Sites, J.W. Jr. & Noonan, B.P. (2008) Around the world in 10 million years: Biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae). Global Ecology and Biogeography, 17 (1), 72–83.
    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 (2), 543–583.
    Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904.
    Ranwez, V., Douzery, E.J.P., Cambon, C., Chantret, N. & Delsuc, F. (2018) MACSE v2: Toolkit for the alignment of coding sequences accounting for frameshifts and stop codons. Molecular Biology and Evolution, 35 (10), 2582–2584.
    Richards, C.M. & Moore, W.S. (1996) A phylogeny for the African treefrog family Hyperoliidae based on mitochondrial rDNA. Molecular Phylogenetics and Evolution, 5 (3), 522–532.
    Rödel, M.-O. (2000) Herpetofauna of West Africa. Vol. I. Amphibians of the West African Savanna. Edition Chimaira, Frankfurt am Main, 332 pp., audio CD.
    Sabaj, M.H. (2019) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: An online reference. Version 7.1. American Society of Ichthyologists and Herpetologists, Washington, D.C. Electronically accessible. Available from: (accessed 22 December 2021)
    Sánchez-Vialas, A., Calvo-Revuelta, M., Castroviejo-Fisher, S. & De la Riva, I. (2020) Synopsis of the amphibians of Equatorial Guinea based upon the authors’ field work and Spanish natural history collections. Proceedings of the California Academy of Sciences, 66, 137–230.
    Sanderson, I.T. (1936) The amphibians of the Mamfe Division. Cameroons—II. Ecology of the frogs. Proceedings of the Zoological Society of London, 106 (1), 165–208.
    Schiøtz, A. (1964) A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjøbenhavn, 127, 1–17.
    Schiøtz, A. (1967) The treefrogs (Rhacophoridae) of West Africa. Spolia Zoologica Musei Hauniensis, 25, 1–346.
    Schiøtz, A. (1974) Revision of the genus Afrixalus (Anura) in eastern Africa. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjøbenhavn, 137, 9–18.
    Schiøtz, A. (1975) The treefrogs of Eastern Africa. Steenstrupia, Copenhagen, 232 pp.
    Schiøtz, A. (1999) Treefrogs of Africa. Edition Chimaira, Frankfurt am Main, 350 pp.
    Schiøtz, A. (2007) Zoogeography of the treefrogs in Africa’s tropical forests. Alytes, 25 (1–2), 1–37.
    Spawls, S., Howell, K.M. & Drewes, R.C. (2006) Pocket Guide to the Reptiles and Amphibians of East Africa. A & C Black Publishers, London, 240 pp.
    Stamatakis, A. (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30 (9), 1312–1313.
    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 (2), 407–450.
    Vaughan, E.R., Teshera, M.S., Kusamba, C., Edmonston, T.R. & Greenbaum, E. (2019) A remarkable example of suspected Batesian mimicry of Gaboon Vipers (Reptilia: Viperidae: Bitis gabonica) by Congolese Giant Toads (Amphibia: Bufonidae: Sclerophrys channingi). Journal of Natural History, 53 (29–30), 1853–1871.
    Vences, M., Kosuch, J., Glaw, F., Böhme, W. & Veith, M. (2002) Molecular phylogeny of hyperoliid treefrogs: Biogeographic origin of Malagasy and Seychellean taxa and re-analysis of familial paraphyly. Journal of Zoological Systematics and Evolutionary Research, 41 (3), 202–215.
    Vonesh, J. (2001) Natural history and biogeography of the amphibians and reptiles of Kibale National Park, Uganda. Contemporary Herpetology, 2001. [published online]
    Wiens, J.J. & Penkrot, T.A. (2002) Delimiting species using DNA and morphological variation and discordant species limits in spiny lizards (Sceloporus). Systematic Biology, 51 (1), 69–91.
    Yang, Z. (2015) The BPP program for species tree estimation and species delimitation. Current Zoology, 61 (5), 854–865.
    Yang, Z. & Flouri, T. (2020) BP&P version 4.2 documentation. Accessible at: