Skip to main content Skip to main navigation menu Skip to site footer
Published: 2021-12-13

A new species of red toad, Schismaderma Smith, 1849 (Anura: Bufonidae), from central Angola

CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Universidade do Porto, Campus Agrário de Vairão, Rua Padre Armando Quintas, 4485-661 Vairão, Portugal. Departamento de Biologia, Faculdade de Ciências, Universidade do Porto, 4169-007 Porto, Portugal
CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Universidade do Porto, Campus Agrário de Vairão, Rua Padre Armando Quintas, 4485-661 Vairão, Portugal. Fundação Kissama, Luanda, Angola
Zoology and Entomology Molecular Lab, Department of Zoology and Entomology, Rhodes University, Makhanda, South Africa 8South African Institute for Aquatic Biodiversity (SAIAB), Makhanda, South Africa
Zoology and Entomology Molecular Lab, Department of Zoology and Entomology, Rhodes University, Makhanda, South Africa
Museum für Naturkunde—Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, Germany. Berlin-Brandenburg Institute of Advanced Biodiversity Research—BBIB, Altensteinstr. 34, 14195 Berlin, Germany
South African Institute for Aquatic Biodiversity (SAIAB), Makhanda, South Africa
Amphibia allopatric speciation cryptic diversity endemism integrative taxonomy Angolan Miombo woodlands


A new species of red toad, from the previously monotypic genus Schismaderma, is described. The new species was found in Malanje Province, and seems endemic to central Angola, occurring approximately 500 km west of the closest known records of Schismaderma carens. Unusual adult colouration and geographical distance to remaining S. carens populations suggested specific differentiation. In an integrative approach, we compared the red toads from central Angola with S. carens from across the entire range, including molecular data, morphology of adults and tadpoles, and male advertisement calls. Phylogenetic analysis based on mitochondrial (12S, 16S, COI) and nuclear (CXCR4, RAG1) markers, retrieved the Angolan clade as monophyletic, and revealed intra-specific substructuring among the remaining Schismaderma. Genetic distances supported specific differentiation of the central Angolan material compared with other S. carens. Adults from the new Angolan species have bolder ventral patterning and smaller body size than S. carens. No obvious differences were detected between the tadpoles and the advertisement calls of the two Schismaderma species. This discovery adds to the knowledge of the herpetofauna of the Angolan Miombo woodlands, a poorly understood ecoregion, and likely more biodiverse than previously assumed. The result of past river basin dynamics in central Angola likely led to the evolution of this new species of Schismaderma.



  1. Altig, R. & McDiarmid, R.W. (1999) Body plan: development and morphology. In: McDiarmid, R.W & Altig, R. (Eds.), Tadpoles: the biology of anuran larvae. University of Chicago Press, Chicago, Illinois, pp. 24–51.
    Altig, R. (2007) A primer for the morphology of anuran tadpoles. Herpetological Conservation and Biology, 2 (1), 71–74.
    AmphibiaWeb. (2008) Amphibiaweb, University of California, Berkeley, California. Schismaderma carens: Red Toad. Available from: (accessed 30 July 2020)
    American Society of Ichthyologists and Herpetologists, The Herpetologists’ League & Society for the Study of Amphibians and Reptiles. (1987) Guidelines for use of live amphibians and reptiles in field research. Available from: (accessed 20 February 2021)
    American Veterinary Medical Association. (2020) AVMA Guidelines for the Euthanasia of Animals: 2020 Edition. Available from: (accessed 20 February 2021)
    Balinsky, B.I. (1969) The reproductive ecology of amphibians of the Transvaal highveld. African Zoology, 4 (1), 37–93.
    Baptista, N., Conradie, W., Vaz Pinto, P. & Branch, W.R. (2019) The amphibians of Angola: early studies and the current state of knowledge. In: Huntley, B.J., Russo, V., Lages, F., & Ferrand, N. (Eds.), Biodiversity of Angola. Science & conservation: a modern synthesis. Springer Nature, Cham, pp. 243–281.
    Baptista, N.L, Tolley, K.A., Bluhm, M., Finckh, M. & Branch, W.R. (2020) Rediscovery, range extension, habitat and phylogenetic relation of the endemic scaled sandveld lizard Nucras scalaris Laurent, 1964 (Sauria: Lacertidae) in the central Angolan plateau. African Journal of Herpetology, 69 (1), 12–28.
    Bittencourt-Silva, G.B. (2019) Herpetological survey of western Zambia. Amphibian & Reptile Conservation, 13 (2), 1–28.
    Bruford, M.W., Hanotte, M., Brookfield, J.F.Y. & Burke, T. (1992) Single locus and multilocus DNA fingerprint. In: Hoelzel, A.R. (Ed.), Molecular genetic analysis of populations: a practical approach. IRL Press, Oxford, pp. 225–270.
    Burgess, N., Hales, J.A., Underwood, E., Dinerstein, E., Olson, D., Itoua, I., Schipper, J., Ricketts, T. & Newman, K. (2004) Terrestrial ecoregions of Africa and Madagascar: a conservation assessment. Island Press, Washington, 497 pp.
    Center for Conservation Bioacoustics. (2019) Raven Pro: interactive sound analysis software. Version 1.6.1. The Cornell Lab of Ornithology, Ithaca, New York. Available from: (accessed 30 August 2020)
    Ceríaco, L.M., Marques, M.P., André, I., Afonso, E., Blackburn, D.C. & Bauer, A.M. (2020) Illustrated type catalogue of the “lost” herpetological collections of Museu do Dundo, Angola. Bulletin of the Museum of Comparative Zoology, 162 (7), 379–440.
    Channing, A. (2001) Amphibians of central and southern Africa. Cornell University Press, New York, New York, 470 pp.
    Channing, A. & Rödel, M.-O. (2019) Field guide to the frogs and 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.
    Charter, R.R. & MacMurray, J.B.C. (1939) On the “frilled” tadpole of Bufo carens Smith. South African Journal of Science, 36, 386–389.
    Ceríaco, L.M., Parrinha, D. & Marques, M.P. (2021) Saving collections: taxonomic revision of the herpetological collection of the Instituto de Investigação Científica Tropical, Lisbon (Portugal) with a protocol to rescue abandoned collections. ZooKeys, 1052, 85–156.
    Conradie, W., Keates, C., Lobón-Rovira, J., Vaz Pinto, P., Baptista, N., Harvey, J. & Júlio, T. (2020) New insights into the taxonomic status, distribution and natural history of De Witte’s clicking frog (Kassinula wittei Laurent, 1940). African Zoology, 55 (4), 311–322.
    de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: A conceptual unification and terminological recommendations. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless forms: species and speciation. Oxford University Press, Oxford, pp. 57–75.
    de Vienne, D.M., Giraud, T. & Martin, O.C. (2007) A congruence index for testing topological similarity between trees. Bioinformatics, 23, 3119–3124.
    Du Preez, L. & Carruthers, V. (2017) Frogs of Southern Africa, a complete guide. Struik Nature, Cape Town, 519 pp.
    Estes, R.D. (2013) Hippotragus niger, sable antelope. In: Kingdon, J. & Hoffmann, M. (Eds.), Mammals of Africa. Vol. IV Pigs, hippopotamuses, chevrotain, giraffes, deer and bovids. Bloomsbury, London, pp. 556–565.
    Flügel, T.J., Eckardt F.D. & Cotterill F.P.D. (2015) The present day drainage patterns of the Congo River system and their Neogene evolution. In: de Wit, M.J., Guillocheau, F. & de Wit, M.C.J. (Eds.), Geology and resource potential of the Congo Basin. Regional geology reviews. Springer, Berlin and Heidelberg, pp. 315–337.
    FrogMap. (2020) Virtual Museum of the FitzPatrick Institute of African Ornithology. Available from: (accessed 5 October 2020)
    Frost, D.R. (2021) Amphibian Species of the World: an Online Reference. Version 6.1 Electronic Database. American Museum of Natural History, New York. Available from: (accessed 20 January 2021)
    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
    Gosner, K.L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16 (3), 183–190.
    Günther, A.C.L.G. (1858) Catalogue of the batrachia salientia in the collection of the British Museum. Taylor and Francis, London, 160 pp.
    Hall, T.A. (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
    Hallermann, J., Ceríaco, L.M., Schmitz, A., Ernst, R., Conradie, W., Verburgt, L., Marques, M.P. & Bauer, A.M. (2020) A review of the Angolan house snakes, genus Boaedon Duméril, Bibron and Duméril (1854) (Serpentes: Lamprophiidae), with description of three new species in the Boaedon fuliginosus (Boie, 1827) species complex. African Journal of Herpetology, 69 (1), 29–78.
    Harper, E.B., Measey, G.J., Patrick, D.A., Menegon, M., Vonesh, J.R. & Swilla, I. (2010) Field guide to the amphibians of the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya. Camerapix Publishers International, Nairobi, 114 pp.
    Huntley, B.J. (2019) Angola in outline: physiography, climate and patterns of biodiversity. In: Huntley, B.J., Russo, V., Lages, F. & Ferrand, N. (Eds.), Biodiversity of Angola. Science & conservation: a modern synthesis. Springer, Cham, pp. 15–42.
    iNaturalist (2020) iNaturalist. Available from: (accessed 5 October 2020)
    IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. 2nd Edition. IUCN, Gland and Cambridge, 32 pp.
    IUCN SSC Amphibian Specialist Group (2013) Schismaderma carens. The IUCN Red List of Threatened Species, 2013, e.T54885A3021025.
    Köhler, J., Jansen, M., Rodriguez, A., Kok, P.J., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F., 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.
    Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences USA, 86, 6196–6200.
    Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34, 772–773.
    Laurent, R.F. (1964) Reptiles et amphibiens de l’Angola (Troisème contribution). Publicações Culturais da Companhia de Diamantes de Angola, 67, 1–165.
    Liedtke, H.C., Muller, H., Hafner, J., Penner, J., Gower, D.J., Mazuch, T., Rödel, M.-O. & Loader, S.P. (2017) Terrestrial reproduction as an adaptation to steep terrain in African toads. Proceedings of the Royal Society B: Biological Sciences, 284, 20162598.
    Liedtke, H.C., Müller, H., Rödel, M.-O., Menegon, M., Gonwouo, L.N., Barej, M.F., Gvozdik, V, Schmitz, A., Channing, A., Nagel, P. & Loader, S.P. (2016) No ecological opportunity signal on a continental scale? Diversification and life-history evolution of African true toads (Anura: Bufonidae). Evolution, 70, 1717–1733.
    Marques, M.M. (1973) Contribuição para o estudo geológico-geomorfológico do troço superior do vale do Cuanza (Distrito do Bié– Angola). Boletim dos Serviços de Geologia e Minas de Angola, 24, 15–22.
    Marques, M.M. (1992) La cuirasse ferrugineuse conglomératique de Cabatuquila (Malanje-RP d’Angola) et sa signification paléogéographique. Ciências da Terra/Earth Sciences Journal, 11, 142–148.
    Marques, M.P., Ceríaco, L.M., Bandeira, S., Pauwels, O.S. & Bauer, A.M. (2019) Description of a new long-tailed skink (Scincidae: Trachylepis) from Angola and the Democratic Republic of the Congo. Zootaxa, 4568 (1), 51–68.
    Marques, M.P., Ceríaco, L.M., Buehler, M.D., Bandeira, S.A., Janota, J.M. & Bauer, A.M. (2020) A revision of the dwarf geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with the description of three new species. Zootaxa, 4853 (3), 301–352.
    MCZBASE. (2020a) The Database of the Zoological Collections. Harvard University: Museum of Comparative Zoology. Available from: (accessed 5 September 2020)
    MCZBASE. (2020b) The Database of the Zoological Collections. Harvard University: Museum of Comparative Zoology. Available from: (accessed 5 September 2020)
    Mendelsohn, J. & Weber, B. (2015) Atlas e perfil do Moxico, Angola. An atlas and profile of Moxico, Angola. RAISON (Research & Information Services of Namibia), Windhoek, 44 pp.
    National Geographic Okavango Wilderness Project (2017) Initial findings from exploration of the upper catchments of the Cuito, Cuanavale, and Cuando Rivers, May 2015 to December 2016. Unpublished report, 368 pp.
    Nielsen, S.V., Conradie, W., Ceríaco, L.M.P., Bauer, A.M., Heinicke, M.P., Stanley, E.L. & Blackburn, D.C. (2020) A new species of rain frog (Brevicipitidae, Breviceps) endemic to Angola. ZooKeys, 979, 133–160.
    Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7 (1), 1–14.
    Páez-Moscoso, D.J. & Guayasamin, J.M. (2012) Species limits in the Andean toad genus Osornophryne (Bufonidae). Molecular Phylogenetics and Evolution, 65 (3), 805–822.
    Palumbi, S.R. (1996) Nucleic acids II: the polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular systematics. Sinauer Associates, Sunderland, Massachusetts, pp. 205–247.
    Pauly, G.B., Hillis, D.M. & Cannatella, D.C. (2004) The history of a Nearctic colonization: molecular phylogenetics and biogeography of the Nearctic toads (Bufo). Evolution, 58 (11), 2517–2535.
    Plug, C. (2014) Wahlberg, Mr Johan August (zoological collection, plant collection). In: S2A3 Biographical database of southern African science. S2A3, South Africa. Available from: Biograph_final.php?serial=3018 (accessed 18 March 2021)
    Power, H.J. (1925) Notes on the habits and life-histories of certain little-known Anura, with descriptions of the tadpoles. Transactions of the Royal Society of South Africa, 13 (2), 107–117.
    Poynton, J.C. (1964) The Amphibia of southern Africa: a faunal study. Annals of the Natal Museum, 17, 1–334.
    Poynton, J.C. (1996) Diversity and conservation of African bufonids (Anura): some preliminary findings. African Journal of Herpetology, 45, 1–7.
    Poynton, J.C., Broadley, D.G. (1988) Amphibia Zambesiaca 4. Bufonidae. Annals of the Natal Museum, 29 (2), 447–490.
    Pramuk, J.B. (2006) Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence. Zoological Journal of the Linnean Society, 146 (3), 407–452.
    Pramuk, J.B., Robertson, T., Sites, J.W. & 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, 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.
    R Core Team (2020) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: (accessed 19 March 2021)
    RStudio Team (2019) RStudio: Integrated Development for R. RStudio, Inc., Boston, Massachusetts. Available from: (accessed 19 March 2021)
    Rambaut, A. & Drummond, A.J. (2007) Tracer. Version 1.4. Available from: (accessed 12 June 2020)
    Rambaut, A. (2014) FigTree. Version 1.4.2. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh.
    Rödel, M.-O. (2000) Herpetofauna of West Africa, Vol. I: amphibians of the West African savanna. Edition Chimaira, Frankfurt am Main, 322 pp.
    Ruas, C. (1996) Contribuição para o conhecimento da fauna de batráquios de Angola. Garcia de Orta, Série de Zoologia, 21 (1), 19–41.
    Ruas, C. (2002) Batráquios de Angola em colecção no Centro de Zoologia. Garcia de Orta, Série de Zoologia, 24 (1–2), 139–154.
    Ronquist, F., Teslenko, M., Van Der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3) 539–542.
    San Mauro, D., Gower, D.J., Oommen, O.V., Wilkinson, M. & Zardoya, R. (2004) Phylogeny of caecilian amphibians (Gymnophiona) based on complete mitochondrial genomes and nuclear RAG1. Molecular Phylogenetics and Evolution, 33, 413–427.
    Smith, A. (1848) Illustrations of the zoology of South Africa; consisting chiefly of figures and descriptions of the objects of natural history collected during an expedition into the interior of South Africa, in the years 1834, 1835, and 1836; fitted out by the Cape of Good Hope Association for Exploring Central Africa Vol. III. Reptilia. Part 27. Smith, Elder, and Co. Cornill, London, 606 pp.
    Smith, A. (1849) Illustrations of the zoology of South Africa; consisting chiefly of figures and descriptions of the objects of natural history collected during an expedition into the interior of South Africa, in the years 1834, 1835, and 1836; fitted out by the Cape of Good Hope Association for Exploring Central Africa. Vol. III. Reptilia. Part 28. Errata sheet. Smith, Elder, and Co. Cornill, London, 768 pp.
    Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30 (9), 1312–1313.
    Sueur, J., Aubin, T. & Simonis, C. (2008) Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics, 18, 213–226.
    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
    Tandy, M. & Keith, R. (1972) Bufo of Africa. In: Blair, W.F. (Ed.), Evolution in the genus Bufo. University of Texas Press, Austin, Texas and London, pp. 119–170.
    Tandy, M. & Tandy, J. (1976) Evolution of acoustic behaviour in African Bufo (Anura: Bufonidae). African Zoology, 11 (2), 349–368.
    Van Bocxlaer, I., Loader, S.P., Roelants, K., Biju, S.D., Menegon, M. & Bossuyt, F. (2010) Gradual adaptation toward a range-expansion phenotype initiated the global radiation of toads. Science, 327 (5966), 679–682.
    Van Dijk, D.E. (1966) Systematic and field keys to the families, genera and described species of southern African anuran tadpoles. Annals of the Natal Museum, 18, 231–286.
    Vaz Pinto, P. (2018) Evolutionary history of the critically endangered giant sable antelope (Hippotragus niger variani). Insights into its phylogeography, population genetics, demography and conservation. PhD Thesis, University of Porto, Porto, 287 pp.
    Viertel, B. & Channing, A. (2017) The larva of Schismaderma carens (Smith, 1849) (Anura: Bufonidae) - a redescription. Alytes, 33, 38–46.
    Watters, J.L., Cummings, S.T., Flanagan, R.L. & Siler, C.D. (2016) Review of morphometric measurements used in anuran species descriptions and recommendations for a standardized approach. Zootaxa, 4072 (4), 477–495.
    Wickham, H. (2016) ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. Available from: (accessed 20 August 2021)
    Xia, X. (2013) DAMBE5: A comprehensive software package for data analysis in molecular biology and evolution. Molecular Biology and Evolution, 30, 1720–1728.