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Published: 2011-02-23
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Dusted off—the African Amietophrynus superciliaris-species complex of giant toads

Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, 10115 Berlin, Germany Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany
Muséum d’histoire naturelle, Department of Herpetology and Ichthyology, C.P. 6434, 1211 Geneva 6, Switzerland
Museo Tridentino di Scienze Naturali, Via Calepina 14, 38100 Trento, Italy
Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, 10115 Berlin, Germany Muséum National d’Histoire Naturelle, Département Evolution et Systématique, Reptiles et Amphibiens, 25 rue Cuvier, 75005 Paris, France
c/o The World Bank, Country Office, PO Box 609, Kigali, Rwanda
Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany
Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, 10115 Berlin, Germany
Amphibia Amietophrynus channingi sp. nov. A. superciliaris chevalieri A. s. superciliaris West & Central Africa

Abstract

Amietophrynus superciliaris is known to occur in rain forests from West Africa to eastern Democratic Republic of Congo and Gabon. We herein present morphological and molecular data indicating the existence of three distinct taxa. The name A. superciliaris superciliaris is restricted to toads from the western Lower Guinean Forest (eastern Nigeria, Cameroon, Equatorial Guinea, Gabon). We resurrect A. s. chevalieri for the Upper Guinean forest (Sierra Leone, Guinea, Liberia, Ivory Coast, Ghana) and describe a new species occurring in the eastern part of the Lower Guinean Forest (eastern Democratic Republic of Congo). Amietophrynus channingi sp. nov. from eastern Lower Guinean Forest differs from both other taxa by its brownish lateral coloration (reddish-purple in the other taxa). The new species differs morphologically from western Lower Guinean A. s. superciliaris by a less pointed eyelid process, a dark coloured posterior abdominal region and a dark coloured vertebral line (both absent in A. s. superciliaris), the shape of the parotid glands (bulged and rounded at the posterior tip in the new species, slender drop shaped and pointed at the posterior tip in A. s. superciliaris), and juvenile interorbital markings (V-shaped in the new species, usually interrupted and broken in A. s. superciliaris). Amietophrynus channingi sp. nov. differs from the Upper Guinean A. s. chevalieri by the presence of an eyelid process (absent in A. s. chevalieri), presence of a dark vertebral line and a pair of dark spots on the posterior part of the back (both absent in A. s. chevalieri). The Upper Guinean A. s. chevalieri differs from western Lower Guinean A. s. superciliaris by the absence of an eyelid process, a dark coloured posterior abdominal region (absent in A. s. superciliaris) and lacking a pair of dark spots in the posterior part of the back (present in A. s. superciliaris). The new species differs from both other taxa by 2.2–2.8% in the investigated 16S rRNA gene. West African and western Central African populations differ by only 0.9–1.1% in 16S rRNA, lacking any intra-taxon variation within each clade, and are cautiously regarded as subspecies although the genetic distinction is mirrored by strong morphological differences and distinct geographic distribution which may support its elevation to species status once that more comprehensive data become available. A key to the taxa of the A. superciliaris-species complex is provided.

References

  1. Affa'a, F.-M. & Amiet, J.-L. (1990) Les modes d'association avec les espèces-hôtes chez les nyctothères d'amphibiens du sud-Cameroun. Revue d'écologie (la terre et la vie), 45, 345–355.

    Akani, G.C., Politano, E. & Luiselli, L. (2004) Amphibians recorded in forest swamp areas of the river Niger delta (southeastern Nigeria), and the effects of habitat alteration from oil industry development on species richness and diversity. Applied Herpetology, 2, 1–22.

    Amiet, J.-L. (1976a) Voix d'Amphibiens camerounais V. Bufonidae; genres Bufo, Werneria et Nectophryne. Annales de la Faculté des Sciences du Cameroun, 21–22, 139–157.

    Amiet, J.-L. (1976b) Observations anatomique et biologique sur le genre Werneria Poche, 1903. Revue Zoologique Africaine, 90, 33–45.

    Amiet, J.-L. (1986) La batrachofaune sylvicole d'un secteur forestier du Cameroun: la région de Yaoundé. Mémoires du Muséum National d'Histoire Naturelle, 132, 29–42.

    Amiet, J.-L. (1989) Quelques aspects de la biologie des Amphibiens Anoures du Cameroun. L'année biologique, 28, 73–136.

    Amiet, J.-L. & Perret, J.-L. (1969) Contributions à la faune de la région de Yaoundé (Cameroun) II. – Amphibiens anoures. Annales de la Faculté des Sciences du Cameroun, 3, 117–137.

    Angel, F. (1931) Note sur des exemplaires vivants de Bufo superciliaris Boulenger, de l'Afrique Equatoriale. Bulletin du Muséum National d'Histoire Naturelle, 2e Série, 3, 606–607.

    Anthony, N.M., Johnson-Bawe, M., Jeffery, K., Clifford, S.L., Abernethy, K.A., Tutin, C.E., Lahm, S.A., White, L.J.T., Utley, J.F., Wickings, E.J. & Bruford, M.W. (2007) The role of Pleistocene refugia and rivers in shaping gorilla genetic diversity in Central Africa. Proceedings of the National Academy of Sciences, USA, 104, 20432–20436.

    Babault, G. (1935) Note sur les Boocercus du Kivu. Bulletin du Muséum National d'Histoire Naturelle, 2e Série, 7, 224–225.

    Bailo, D., Alphonse, N. & Gu, Y. (2009) An inventory of biodiversity in the Badiar National Park, Guinea Conakry: Implication for conservation. Research Journal of Biological Sciences, 4, 948–951.

    Barbour, T. (1911) Some West African amphibians. Bulletin of the Museum of Comparative Zoölogy, 54, 129–136.

    Barej, M.F., Rödel, M.-O., Gonwouo, N.L., Pauwels, O.S.G., Böhme, W. & Schmitz, A. (2010) Review of the genus Petropedetes Reichenow, 1874 in Central Africa with the description of three new species (Amphibia: Anura: Petropedetidae). Zootaxa, 2340, 1–49.

    Bequaert, J. (1922) Chpt. III. The predaceous enemies of ants In: Wheeler, W.M. (Ed.) Ants of the American Museum Congo Expedition. A contribution to the myrmecology of Africa. Bulletin of the American Museum of Natural History, 45, 271–331.

    Berlioz, J. (1935) Étude d'une collection d'oiseaux du Congo Belge. Bulletin du Muséum National d'Histoire Naturelle, 2e Série, 7, 159–164.

    Blackburn, D.C. (2008) Biogeography and evolution of body size and life history of African frogs: Phylogeny of squeakers (Arthroleptis) and long-fingered frogs (Cardioglossa) estimated from mitochondrial data. Molecular Phylogenetics and Evolution, 49, 806–826.

    Blackburn, D.C., Gonwouo, N.L., Ernst, R. & Rödel, M.-O. (2009) A new squeaker frog (Arthroleptidae: Arthroleptis) from the Cameroon Volcanic Line with redescriptions of Arthroleptis adolfifriederici Nieden, 1911 "1910" and A. variabilis Matschie, 1893. Breviora, 515, 1–22.

    Blackburn, D.C., Kosuch, J., Schmitz, A., Burger, M., Wagner, P., Gonwouo, N.L., Hillers, A. & Rödel, M.-O. (2008) A new species of Cardioglossa (Anura: Arthroleptidae) from the Upper Guinean Forests of Africa. Copeia, 2008, 603–612.

    Blanc, C.P. & Frétey, T. (2004) Répartition écologique des amphibiens dans la Réserve de Faune de la Lopé et la station biologique de la Makandé (Gabon). Bulletin de la Société Zoologique de France, 129, 297–315.

    Böhme, W. (1994) Frösche und Skinke aus dem Regenwaldgebiet Südost-Guineas, Westafrika I. Einleitung; Pipidae, Arthroleptidae, Bufonidae. herpetofauna, 16(92), 11–19.

    Boulenger, G.A. (1888) A list of the reptiles and batrachians collected by Mr. H. H. Johnston on the Rio del Rey, Cameroons District, W. Africa. Proceedings of the Royal Society, London, 1887, 564–565.

    Boulenger, G.A. (1900) A list of the batrachians and reptiles of the Gaboon (French Congo), with descriptions of new genera and species. Proceedings of the Zoological Society of London, 1900, 433–456.

    Boulenger, G.A. (1919) Batraciens et reptiles recueillis par le Dr. C. Christy au Congo Belge dans les districts de Stanleyville, Haut-Uelé et Ituri en 1912-1914. Revue Zoologique Africaine (avec supplément botanique), 7, 1–29.

    Channing, A. (2001) Amphibians of Central and Southern Africa. Cornell University Press, New York, 470 pp.

    Chatelain, C., Dao, H., Gautier, L. & Spichiger, R. (2004) Forest cover changes in West Africa Côte d’Ivoire and Upper Guinea. In: Poorter, L., Bongers, F., Kouamé , F.N. & Hawthorne, W. (Eds.) Biodiversity of West African Forests: an Ecological Atlas of Woody Plant Species. CABI Publishing, Wallingford, 15–32.

    Chatelain, C., Gautier, L. & Spichiger, R. (1996) A recent history of forest fragmentation in southwestern Ivory Coast. Biodiversity and Conservation, 5, 37–53.

    Clifford, S.L., Anthony, N.M., Bawe-Johnson, M., Abernethy, K.A., Tutin, C.E.G., White, L.J.T., Bermejo, M., Goldsmith, M.L., McFarland, K., Jeffery, K.J., Bruford, M.W. & Wickings, E.J. (2004) Mitochondrial DNA phylogeography of western lowland gorillas (Gorilla gorilla gorilla). Molecular Ecology, 13, 1551–1565.

    Cunningham, M. & Cherry, M.I. (2000) Mitochondrial DNA divergence in southern African bufonids: are species equivalent entities? African Journal of Ecology, 49, 9–22.

    Cunningham, M. & Cherry, M.I. (2004) Molecular systematics of anuran 20-chromosome toads (Anura: Bufonidae). Molecular Phylogenetics and Evolution, 32, 671–685.

    Dayrat, B. (2010) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85, 407–415.

    de Salle, R. Egan, M.G. & Siddall, M. (2005) The unholy trinity: taxonomy, species delimitation and DNA barcoding. Philosophical Transactions of the Royal Society of London, B, 360, 1905–1916.

    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.

    Drewes, R.C. & Vindum, J.V (1994) Amphibians of the Impenetrable Forest, southwest Uganda. Journal of African Zoology, 108, 55–70.

    Dubois, A. (1995) Keratodont formula in anuran tadpoles: proposals for a standardization. Journal of Zoological Systematics and Evolutionary Research, 33, I–XV.

    Ernst, R., Linsenmair, K.E. & Rödel, M.-O. (2006) Diversity erosion beyond the species level: Dramatic loss of functional diversity after selective logging in two tropical amphibian communities. Biological Conservation, 133, 143–155.

    Falk, T.M., Teugels, G.G., Abban, E.K., Villwock, W. & Renwrantz, L. (2003) Phylogeographic patterns in populations of the black-chinned Tilapia complex (Teleostei, Cichlidae) from coastal areas in West Africa: support for the refuge zone theory. Molecular Phylogenetics and Evolution, 27, 81–92.

    Fjeldså, J. & Lovett, J.C. (1997) Geographical patterns of old and young species in African forest biota: the significance of specific montane areas as evolutionary centres. Biodiversity and Conservation, 6, 325–346.

    Frétey, T. & Blanc, C.P. (2000) Liste des Amphibiens d’Afrique Centrale. Cameroun, Congo, Gabon, Guinée-Equatoriale, République Centrafricaine, République Démocratique du Congo, São Tomé et Príncipe. Les dossiers de l’ADIE, Série Biodiversité, N°2, 1–39.

    Frétey, T. & Blanc, C.P. (2001) Inventaire systématique des amphibiens anoures du centre du Gabon. Bulletin de la Société Zoologique de France, 126, 375–390.

    Frétey, T. & Dewynter, M. (1998) Amphibiens anoures de la Forêt des Abeilles (Gabon). Journal of African Zoology, 112, 171–184.

    Frost, D. (2010) Amphibian Species of the World: an Online Reference. Version 5.4 (8 April, 2010). Electronic Database accessible at http://research.amnh.org/vz/herpetology/amphibia/ American Museum of Natural History, New York, USA. (Accessed on: 09.05.2010).

    Frost, D., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sa, R.O., Channing, A., Wilkinson, M., Donnellan, S., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., 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, 1–291.

    Germain, L. (1936) Contributions à la faune malacologique de l'Afrique Équatoriale. Bulletin du Muséum National d'Histoire Naturelle, 2e Série, 8, 146–157.

    Glaw, F. & Franzen, M. (2006) Type catalogue of amphibians in the Zoologische Staatssammlung München. Spixiana, 29, 153–192.

    Gosner, K.L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16, 183–190.

    Gossmann, V., Lötters, S., Obame, F. & Böhme, W. (2002) Zur Herpetofauna Gabuns – Teil II: Kommentierte Artenliste der gefundenen Reptilien, Bemerkungen zur Artenvielfalt. herpetofauna, 24(136), 19–33.

    Guibé, J. & Lamotte, M. (1958) La réserve naturelle intégrale du Mont Nimba. XII. Batraciens (sauf Arthroleptis, Phrynobatrachus et Hyperolius). Mémoires de l'Institut Français d'Afrique Noire, 53, 241–273.

    Haffer, J (1969) Speciation in Amazonian forest birds. Science, 165, 131–137.

    Hammer, Ø., Harper, D.A.T. & Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4, 1–9.

    Herrmann, H.-W., Böhme, W., Herrmann, P.A., Plath, M., Schmitz, A. & Solbach, M. (2005) African biodiversity hotspots: The amphibians of Mt. Nlonako, Cameroon. Salamandra, 41, 61–81.

    Hillers, A. & Rödel, M.-O. (2007) The amphibians of three national forests in Liberia, West Africa. Salamandra, 43, 1–10.

    Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, 25, 29–38.

    IUCN (2010) The IUCN red list of threatened species. Version 2010.1. Available from http://www.iucnredlist.org (accessed 16 April 2010).

    Jackson, K. & Blackburn, D.C. (2007) The amphibians and reptiles of Nouabale-Ndoki National Park, Republic of Congo (Brazzaville). Salamandra, 43, 149–164.

    Jackson, K., Zassi-Boulou, A.-G., Mavoungou, L.-B. & Pangou, S. (2007) Amphibians and reptiles of the Lac Télé Community Reserve, Linkouala Region, Republic of Congo (Brazzaville). Herpetological Conservation and Biology, 2, 75–86.

    Joger, U. (1990) The herpetofauna of the Central African Republic, with description of a new species of Rhinotyphlops (Serpentes: Typhlopidae). In: Peters, G. & Hutterer, R. (Eds.) Vertebrates in the Tropics. Alexander Koenig Zoological Research Institute and Zoological Research Museum, Bonn: 85–102.

    Köhler, J., Bwong, B.A., Schick, S., Veith, M. & Lötters, S. (2006) A new species of arboreal Leptopelis (Anura: Arthroleptidae) from the forests of western Kenya. Herpetological Journal, 16, 183–189.

    Laporte, N.T., Stabach, J.A., Grosch, R., Lin, T.S. & Goetz, S.J. (2007) Expansion of industrial logging in Central Africa. Science, 316, 1451.

    Largen, M.J. & Dowsett-Lemaire, F. (1991) Amphibians (Anura) from the Kouilou River basin, République du Congo. Tauraco Research Report, 4, 145–168.

    Lasso, C.A., Rial, A.I., Castroviejo, J. & de la Riva, I. (2002) Herpetofauna del Parque Nacionale de Monte Alén (Río Muni, Guinea Equatorial). Graellsia, 58, 21–34.

    Lawson, D.P. (1993) The reptiles and amphibians of the Korup National Park Project, Cameroon. Herpetological Natural History, 1, 27–90.

    Lötters, S., Gossmann, V., Obame, F. & Böhme, W. (2001) Zur Herpetofauna Gabuns Teil I: Einleitung, Untersuchungsgebiet und Methodik, kommentierte Artenliste der gefundenen Froschlurche. herpetofauna, 23(133), 19–34.

    Loveridge, A. (1936) African reptiles and amphibians in Field Museum of Natural History. Zoological Series of Field Museum of Natural History, 22, 1–111.

    Low, R.S. (1972) Evidence from parotid–gland secretions. In: Blair, W.F. (Ed.) Evolution in the genus Bufo. University of Texas Press, Dallas, 244–264.

    Maley, J. (1987) Fragmentation de la forêt dense humide africaine et extension des biotopes montagnards au quaternaire récent: nouvelles données polliniques et chronologiques. Implications paleoclimatique et biogeographique. Paleoecology of Africa and the surrounding islands, 18, 307–334.

    Maley, J. (1991) The African rain forest vegetation and palaeoenvironments during late Quaternary. Climatic change, 19, 79–98.

    Matthee, C.A., Tilbury, C.R. & Townsend, T. (2004) A phylogenetic review of the African leaf chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation. Proceedings of the Royal Society, B, 271, 1967–1975.

    Mausfeld-Lafdhiya, P., Schmitz, A., Ineich, I. & Chirio, L. (2004) Genetic variation in two African Euprepis species (Reptilia, Scincidae) based on Maximum-Likelihood and Bayesian Analyses: taxonomic and biogeographic conclusions. Bonner zoologische Beiträge, 52, 159–177.

    Maxson, L.R. (1981) Albumin evolution and its phylogenetic implications in African toads of the genus Bufo. Herpetologica, 37, 96–104.

    Mayr, E. & O'Hara, J. (1986) The biogeographic evidence supporting the pleistocene forest refuge hypothesis. Evolution, 40, 55–67.

    McDiarmid, R.W. & Altig, R. (1999) Tadpoles - The Biology of Anuran Larvae. University of Chicago Press, 458 pp.

    Mertens, R. (1940) Amphibien aus Kamerun, gesammelt von M. Köhler und Dr. H. Graf. Senckenbergiana, 22, 103–135.

    Mocquard, M.F. (1908) Description de quelques reptiles et d’un batracien nouveaux de la collection du muséum. Bulletin du Muséum national d’Histoire naturelle, 14, 259–262.

    Moritz, C. (1996) Defining 'Evolutionary Significant Units' for conservation. Trends in Ecology and Evolution, 9, 373–375.

    Müller, L. (1910) Beiträge zur Herpetologie Kameruns. Abhandlungen der Mathematisch-Physikalischen Klasse der königlich bayerischen Akademie der Wissenschaften, 24, 544–627.

    Nicolas, V., Mboumba, J.-F., Verheyen, E., Denys, C., Lecompte, E., Olayem, A., Missoup, A.D., Katuala, P. & Colyn, M. (2008) Phylogeographic structure and regional history of Lemniscomys striatus (Rodentia: Muridae) in tropical Africa. Journal of Biogeography, 35, 2074–2089.

    Nieden, F. (1910) Die Fauna der deutschen Kolonien, Reihe I Kamerun, Heft 2 Die Reptilien (außer den Schlangen) und Amphibien. Friedländer & Sohn, Berlin, 74 pp.

    Orts, S.G. (1970) Description et écologie des formes larvaires de Bufo superciliaris Blgr (Amphibia, Bufonidae). Revue de zoologie et de botanique africaines, 81, 207–219.

    Padial, J.M., Miralles, A., de la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, Art. 16.

    Parker, H.W. (1936) The amphibians of the Mamfe Division, Cameroons. – I. Zoogeography and systematics. Proceedings of the Zoological Society of London, 44, 135–163.

    Pauwels, O.S.G. & Rödel, M.-O. (2007) Amphibians and National Parks in Gabon, western Central Africa. Herpetozoa, 19, 135–148.

    Pellegrin, J. (1933) Poissons de la région du Kivu adressés par M. Guy Babault. Bulletin de la Société Zoologique de France, 58, 169–175.

    Pellegrin, J. (1935) Description de deux poissons nouveaux de la région du Kivu appartenant au genre Varicorhinus. Revue de zoologie et de botanique africaines, 26, 401–403.

    Penner, J. & Rödel, M.-O. (2007) Amphibians and reptiles of the Lokutu Region, Democratic Republic of Congo. In: Butynski, T.M. & McCullough, J. (Eds.) A rapid biological assessment of Lokutu, Democratic Republic of Congo. RAP Bulletin of Biological Assessment, 46, Conservation International, Washington D.C., 37–41.

    Perret, J.-L. (1966) Les amphibiens du Cameroun. Zoologische Jahrbücher (Abteilung für Systematik, Ökologie und Geographie der Tiere), 8, 289–464.

    Perret, J.-L. & Amiet, J.-L. (1971) Remarques sur les Bufo (Amphibiens Anoures) du Cameroun. Annales de la Faculté des Sciences du Cameroun, 5, 47–55.

    Perret, J.-L. & Mertens, R. (1957) Étude d'une collection herpétologique faite au Cameroun de 1952 à 1955. Bulletin de l'Institut Français d'Afrique Noire. Série A, 19, 548–601.

    Poynton, J.C. (1996) Diversity and conservation of African bufonids (Anura): Some preliminary findings. African Journal of Herpetology, 45, 1–7.

    Rasmussen, J.B., Chirio, L. & Ineich, I. (2000) The herald snakes (Crotaphopeltis) of the Central African Republic, including a systematic review of C. hippocrepis. Zoosystema, 22, 585–600.

    Rödel, M.-O. (2000) Herpetofauna of West Africa, Vol. I: Amphibians of the West African savanna. Edition Chimaira, Frankfurt am Main, 335 pp.

    Rödel, M.-O. & Agyei, A.C. (2003) Amphibians of the Togo-Volta highlands, eastern Ghana. Salamandra, 39, 207–234.

    Rödel, M.-O. & Bangoura, M.A. (2004a) A conservation assessment of amphibians in the Fôret Cassée du Pic de Fon, Simandou Range, southeastern Republic of Guinea, with description of a new Amnirana species (Amphibia Anura Ranidae). Tropical Zoology, 17, 201–232.

    Rödel, M.-O. & Bangoura, M. (2004b) Rapid survey of amphibians and reptiles in the Forêt Classée du Pic de Fon, Guinea. In: McCullough, J. (Ed.): A Rapid Biological Assessment of the Forêt Classée du Pic de Fon, Simandou Range, South-eastern Republic of Guinea / Une Évaluation Biologique Rapide de la Forêt Classée du Pic de Fon,Chaîne du Simandou, Guinée. RAP Bulletin of Biological Assessment 35, Conservation International, Washington D.C., 51–60 + appendices 3–5, 232–237.

    Rödel, M.-O., Bangoura, M.A. & Böhme, W. (2004) The amphibians of south–eastern Republic of Guinea. Herpetozoa, 17, 99–118.

    Rödel, M.-O., Boateng, C.O., Penner, J. & Hillers, A. (2009) A new cryptic Phrynobatrachus species (Amphibia: Anura: Phrynobatrachidae) from Ghana, West Africa. Zootaxa, 1970, 52–62.

    Rödel, M.-O. & Ernst, R. (2000) Bufo taiensis n. sp., eine neue Kröte aus dem Taï-Nationalpark, Elfenbeinküste. herpetofauna, 22(125), 9–16.

    Rödel, M.-O., Kosuch, J., Veith, M. & Ernst, R. (2003) First record of the genus Acanthixalus Laurent, 1944 from the Upper Guinean Rain Forest, West Africa, with the description of a new species. Journal of Herpetology, 37, 43–52.

    Roelants, K., Gower, D.J., Wilkinson, J.A., Loader, S.P., Biju, S.D., Guillaume, K., Moriau, L. & Bossuyt, F. (2007) Global patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences, USA, 104, 887–892.

    Sanderson, I.T. (1936a) The giant frog problem. The Nigerian Field, 5, 161–170.

    Sanderson, I.T. (1936b) The amphibians of the Mamfe Division, Cameroon. – II. Ecology of the frogs. Proceedings of the Zoological Society of London, 44, 165–208.

    Schiøtz, A. (1963) The amphibians of Nigeria. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I København, 125, 1–92.

    Schiøtz, A. (1964) A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening I København, 127, 1–17.

    Schmitz, A. (1998) Systematik und Zoogeographie der Herpetofauna einer montanen Regenwaldregion in Kamerun. unpublished diploma thesis, University of Bonn, 276 pp.

    Schmitz, A., Ineich, I. & Chirio, L. (2005) Molecular review of the genus Panaspis sensu lato (Reptilia: Scincidae) in Cameroon, with special reference to the status of the proposed subgenera. Zootaxa, 863, 1–28.

    Segniagbeto, G.H., Bowessidjaou, J.E., Dubois, A. & Ohler, A. (2007) Les amphibiens du Togo: état actuel des connaissances. Alytes, 24, 72–90.

    Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts, USA.

    Tandy, M. & Keith, R. (1972) Evidence from osteology. In: Blair, W.F. (Ed.) Evolution in the genus Bufo. University of Texas Press, Dallas, 119–170.

    Van Bocxlaer, I., Biju, S.D., Loader, S.P. & Bossuyt, F. (2009) Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot. BMC Evolutionary Biology, 9, 131.

    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, 679–682.

    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, 1–12.

    Vonesh, J.R. (2001) Natural history and biogeography of the amphibians and reptiles of Kibale National Park, Uganda. Contemporary Herpetology, 4, http://www.contemporaryherpetology.org/ch/2001/CH_2001.htm.

    Wagner, P., Barej, M.F. & Schmitz, A. (2009a) Studies on African Agama VII. A new species of the Agama agama-group (Linnaeus, 1758) (Sauria: Agamidae) from Cameroon & Gabon, with comments on Agama mehelyi Tornier, 1902. Bonner Zoologische Beiträge, 56, 285-297.

    Wagner, P., Böhme, W., Pauwels, O.S.G. & Schmitz, A. (2009b) A review of the African red-flanked skinks of the Lygosoma fernandi (Burton, 1836) species group (Squamata: Scincidae) and the role of climate change in their speciation. Zootaxa, 2050, 1–30.

    Werner, F. (1897) Ueber einige neue oder seltene Reptilien und Frösche der zoologischen Sammlung des Staates München. Sitzungsberichte der mathematisch-physikalischen Classe der königlich bayerischen Akademie der Wissenschaften zu München, 27, 203–220.

    Werner, F. (1898) Ueber Reptilien und Batrachier aus Togoland, Kamerun und Tunis aus dem kgl. Museum für Naturkunde in Berlin (Teil II.). Verhandlungen der kaiserlich–königlichen zoologisch–botanischen Gesellschaft in Wien, 48, 191–230.

    Wheeler, W.M. (1922) Chpt. II. The ants collection by the American Museum Congo Expedition. In: Wheeler, W.M. (Ed.) Ants of the American museum Congo Expedition. A contribution to the myrmecology of Africa. Bulletin of the American Museum of Natural History, 45, 39–269.

    Wilkinson, M., Loader, S.P., Gower, D.J., Sheps, J.A. & Cohen, B.L. (2003) Phylogenetic relationships of African caecilians (Amphibia: Gymnophiona): insights from mitochondrial rRNA gene sequences. African Journal of Herpetology, 52, 83–92.