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Published: 2017-08-02

Uncovering a hidden diversity: two new species of Breviceps (Anura: Brevicipitidae) from northern KwaZulu-Natal, South Africa

Unit for Environmental Sciences and Management, North-West University, Potchefstroom 2520, South Africa.
Unit for Environmental Sciences and Management, North-West University, Potchefstroom 2520, South Africa. Laboratory of Aquatic Ecology, Evolution and Conservation, University of Leuven, Charles Debériotstraat 32, B-3000 Leuven, Belgium.
Unit for Environmental Sciences and Management, North-West University, Potchefstroom 2520, South Africa. South African Institute for Aquatic Biodiversity, Somerset Street, Grahamstown 6140, South Africa.
Amphibia Breviceps carruthersi sp. nov. Breviceps passmorei sp. nov. advertisement calls bioacoustics cryptic species molecular genetics morphology Zululand


Breviceps carruthersi sp. nov. and Breviceps passmorei sp. nov. are described from northern KwaZulu-Natal. Both new species are distinguished from B. adspersus, B. mossambicus, B. poweri and B. cf. sopranus by substantial interspecific genetic divergence (> 6.8 uncorrected p-distance) in the mitochondrial 16S rRNA gene, differences in advertisement call structure and limited morphological variation. Breviceps carruthersi sp. nov. has a short, pulsatile call with minimal amplitude modulation and groups its calls. Breviceps passmorei sp. nov. differs significantly from B. adspersus and B. mossambicus in call duration and pulse number, but not in pulse rate or dominant frequency. For both new species morphological characters are of limited diagnostic value, necessitating a re-evaluation of the taxonomic status of populations currently assigned to B. adspersus and B. mossambicus in KwaZulu-Natal.



  1. Altschul, S.F., Gish, W., Miller, W., Myers, E.W. & Lipman, D.J. (1990) Basic local alignment search tool. Journal of Molecular Biology, 215, 403–410.

    Angulo, A. & Icochea, J. (2010) Cryptic species complexes, widespread species and conservation: lessons from Amazonian frogs of the Leptodactylus marmoratus group (Anura: Leptodactylidae). Systematics and Biodiversity, 8, 357–370.

    Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K.L., Meier, R., Winkler, K., Ingram, K.K. & Das, I. (2007) Cryptic species as a window on diversity and conservation. Trends in Ecology and Evolution, 22, 148–155.

    Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences, 97, 6585–6590.

    Boulenger, G.A. (1907) Description of a new Engystomatid frog of the genus Breviceps from Namaqualand. The Annals & Magazine of Natural History, Series 7, 20, 46–47.

    Broadley, D.G. (1971) The reptiles and amphibians of Zambia. Puku, Chilange, 143 pp.

    Channing, A. & Wahlberg, K. (2011) Distribution and conservation status of the desert rain frog Breviceps macrops. African Journal of Herpetology, 60, 101–112.

    Channing, A. (2012) A new species of Rain Frog from Namaqualand, South Africa (Anura: Brevicipitidae: Breviceps). Zootaxa, 3381, 62–68.

    Channing, A. & Baptista, N. (2013) Amietia angolensis and A. fuscigula (Anura: Pyxicephalidae) in southern Africa: A cold case reheated. Zootaxa, 3640 (4), 501–520.

    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 (5), 518–550.

    Channing, A., Measey, G.J., De Villiers, A.L., Turner, A.A. & Tolley, K.A. (2017) Taxonomy of the Capensibufo rosei group (Anura: Bufonidae) from South Africa. Zootaxa, 4232 (2), 282–292.

    Conradie, W. (2014) The King of the Dwarves: a new cryptic species of Dainty Frog (Anura: Pyxicephalidae: Cacosternum) from the eastern Great Escarpment of South Africa. Zootaxa, 3785 (3), 438–452.

    Cuvier, P.M.L.B. (1829) Le Règne Animal Distribué d'Après son Organisation, pour Servir de Base à l'Histoire Naturelle des Animaux et d'Introduction à l'Anatomie Comparée. Vol. 2. Nouvelle Edition, Revue et Augmentée par P.A. Latreille. Deterville, Paris, 406 pp.

    Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature methods, 9, 772–772.

    Dawood, A., Channing, A. & Bogart, J.P. (2002) A molecular phylogeny of the frog genus Tomopterna in southern Africa: examining species boundaries with mitochondrial 12S rRNA sequence data. Molecular Phylogenetics & Evolution, 22, 407–413.

    Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and 546 high throughputs. Nucleic Acids Research, 32, 4673–4680.

    Elmer, K.R. & Cannatella, D.C. (2008) Three new species of leaflitter frogs from the upper Amazon forests: cryptic diversity within Pristimantis “ockendeni” (Anura: Strabomantidae) in Ecuador. Zootaxa, 1784, 11–38.

    FitzSimons, V. (1930) Descriptions of new South African Reptilia and Batrachia, with distribution records of allied species in the Transvaal Museum collection. Annals of the Transvaal Museum, 14, 20–48.

    Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N.J. (2007) Underestimation of species richness in Neotropical frogs revealed by mtDNA analysis. PLoS ONE, 2, e1109.

    Frost, D.R. (2017) Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History, New York, USA. Electronic Database. Available from: (accessed 12 June 2017)

    Funk, W.C., Caminer, M. & Ron, S.R. (2012) High levels of cryptic species diversity uncovered in Amazonian frogs. Proceedings of the Royal Society B: Biological Sciences, 279, 1806–1814.

    Guindon, S. & Gascuel, O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum likelihood. Systematic Biology, 52, 696–704.

    Hewitt, J. (1925) Descriptions of three new toads belonging to the genus Breviceps Merrem. Annals of the Natal Museum, 5, 189–194.

    Hewitt, J. (1932) Some new species and subspecies of South African batrachians and lizards. Annals of the Natal Museum, 7, 105–128.

    Hoffman, A.C. (1944) Frogs from Chitiala, Nyasaland, together with description of new species. Indago, 1, 173–182.

    Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Mentjies, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.

    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, 86, 6196–6200.

    Köhler, J., Glaw, F., Pabijan, M. & Vences, M. (2015) Integrative taxonomic revision of mantellid frogs of the genus Aglyptodactylus (Anura: Mantellidae). Zootaxa, 4006 (3), 401–438.

    Köhler, J., Jansen, M., Rodriguez, A., Kok, P.J., Toledo, L.F., 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), 1–124.

    Kurabayashi, A. & Sumida, M. (2013) Afrobatrachian mitochondrial genomes: genome reorganization, gene rearrangement mechanisms, and evolutionary trends of duplicated and rearranged genes. BioMed Central Genomics, 14, 633.

    Lambiris, A.J.L. (1989) A review of the amphibians of Natal. Lammergeyer, 1989, 391–212.

    Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. 10th Edition. L. Salvii. Stockholm, 824 pp.

    Merrem, B. (1820) Tentamen Systematis Amphibiorum. J.C. Krieger, Marburg, Hesse, 191 pp.

    Minter, L.R. (1995) The advertisement call and other aspects of the reproductive biology of Breviceps adspersus Peters (Anura, Microhylidae). Madoqua, 19, 37–44.

    Minter, L.R. (1997) Advertisement call structure and morphology of Breviceps mossambicus Peters and B. poweri Parker (Anura: Microhylidae) from northern Mozambique. Annals of the Natal Museum, 38, 5–19.

    Minter, LR. (1998) Aspects of the reproductive biology of Breviceps. Unpublished PhD thesis. University of the Witwatersrand, Johannesburg, 480 pp.

    Minter, L.R. (2003) Two new cryptic species of Breviceps (Anura: Microhylidae) from southern Africa. African Journal of Herpetology, 49, 3–8.

    Minter, L.R. (2004a) Breviceps adspersus Peters, 1882. 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. SI/MAB Series #9. Smithsonian Institution, Washington, D.C., pp 172–173.

    Minter, L.R. (2004b) Breviceps mossambicus Peters, 1854. 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. SI/MAB Series #9. Smithsonian Institution, Washington, D.C., pp 184–186.

    Minter, L.R. (2004c) Breviceps sopranus Minter, 2003. 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. SI/MAB Series #9. Smithsonian Institution, Washington, D.C., pp 189–191.

    Mucina, L. & Rutherford, M.C. (2006) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia. Vol. 19. South African National Biodiversity Institute, Pretoria, 805 pp.

    Parker, H.W. (1934) A monograph of the frogs of the family Microhylidae. Trustees of the British Museum, London, 208 pp.

    Parker, W.K. (1868) A monograph on the structure and development of the shoulder girdle and sternum in the Vertebrata. No. 42. Ray Society Series. Robert Hardwicke, London, 237 pp.

    Passmore, N.I. (1985) Sibling species, the acoustic environment and the anuran specific-mate recognition system. In: Vrba, E.S. (Eds.), Species and speciation. Transvaal Museum Monograph No. 4. Transvaal Museum, Pretoria, pp. 125–127.

    Paterson, H.E.H. (1985) The recognition concept of species. In: Vrba, E.S., (Eds.), Species and speciation. Transvaal Museum Monograph No. 4. Transvaal Museum, Pretoria, pp. 21–29.

    Paterson, H.E.H. (1991) The recognition of cryptic species among economically important insects. In: Zalucki, M.P., (Eds.), Heliothis: Research Methods and Prospects, 9, pp. 1–10.

    Peloso, P.L., Frost, D.R., Richards, S.J., Rodrigues, M.T., Donnellan, S., Matsui, M., Raxworthy, C.J., Biju, S.D., Lemmon, E.M., Lemmon, A.R. & Wheeler, W.C. (2015) The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in narrow-mouthed frogs (Anura, Microhylidae). Cladistics, 32, 113–140.

    Peters, W.C.H. (1854) Diagnosen neuer Batrachier, welche zusammen mit der früher (24. Juli und 18. August) gegebenen Übersicht der Schlangen und Eidechsen mitgetheilt werden. Bericht über die zur Bekanntmachung geeigneten Verhandlungen der Königlich Preußischen Akademie der Wissenschaften zu Berlin, Berlin, 15 pp. [pp. 614–628]

    Peters, W.C.H. (1882) Naturwissenschaftlich Reise nach Mossambique auf befehl Seiner Mäjestat des Königs Friedrich Wilhelm IV in den Jahren 1842 bis 1848 ausgeführt. Zoologie 3 (Amphibien). Berlin: G. Reimer.

    Pienaar, U.d.V. (1963) The zoogeography and systematic list of Amphibia in the Kruger National Park. Koedoe, 6, 76–82.

    Power, J.H. (1926) A monographic revision of the genus Breviceps, with distribution records and descriptions of new species. Annals of the South African Museum, 20, 451–471.

    Poynton, J.C. (1963) Descriptions of southern African amphibians. Annals of the Natal Museum, 15, 319–332.

    Poynton, J.C. (1964) The amphibians of southern Africa: a faunal study. Annals of the Natal Museum, 17, 1–334.

    Poynton, J.C. (1982) On species pairs among southern African amphibians. South African Journal of Zoology, 17, 67–74.

    Poynton, J.C. & Broadley, D.G. (1985) Amphibia Zambeziaca I. Scolecomorphidae, Pipidae, Microhylidae, Hemisidae, Arthroleptidae. Annals of the Natal Museum, 26, 503–553.

    Ronquist, F. & Huelsenbeck, J.P. (2003) Mr Bayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.

    Swofford, D. L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods), Ver. 4. Sinauer Associates, Sunderland, Massachusetts.

    Vences, M., Thomas, M., Van der Meijden, A., Chiari, Y. & Vieites, D.R. (2005) Comparative performance of the 16S rRNA gene in DNA barcoding of amphibians. Frontiers in Zoology, 2, 5.

    Werner, F. (1899) Beschreibung einiger neuer Schlangen und Batrachier. Zoologischer Anzeiger, 22, 114–117.

    Werner, F. (1903) Über Reptilien und Batrachier aus Guatemala und China in der zoologischen Staats-Sammlung in München nebst einem Abhang über seltene Formen aus Anderer gebieten. Abhandlungen der Mathematisch-Physikalischen Classe der Königlich Bayerischen Akademie der Wissenschaften, München, 43 pp. [pp. 342–384]