Abstract
The taxonomic status of Amietia vertebralis and Strongylopus hymenopus, two frogs restricted to the Maluti-Drakensberg highlands in southern Africa, is unclear. Here, morphological examination and phylogenetic analyses elucidate the systematic position of these two species. Type specimens of both species were examined and compared with more recent collections to clarify their identity. These comparisons revealed discrepancies between the original application of these names and their current usage. The holotype and original description of A. vertebralis match specimens from an extant population at that species’ type locality that are currently assigned to S. hymenopus. Furthermore, the type specimen of S. hymenopus is of uncertain provenance and does not match well with either of the forms currently associated with these names. We assessed both intraspecific and interspecific variability using DNA sequence data. Broad sampling of the form currently assigned to A. vertebralis revealed very little genetic variation thereby dispelling the hypothesis that this is a compound taxon. The generic placement of both species within the family Pyxicephalidae was resolved using a combined dataset of mitochondrial 16S rDNA and nuclear RAG1 gene sequences. The form currently recognised as S. hymenopus was excluded from the otherwise monophyletic genus Strongylopus, and was placed instead within Amietia. Based on these data a revised application of both species names is recommended here. Populations currently assigned to Amietia vertebralis are referred to Amietia umbraculata and those attributed to Strongylopus hymenopus are referred instead to Amietia vertebralis.References
Bates, M.F. (2002) The distribution of Amietia vertebralis (Hewitt, 1927) (Anura: Ranidae), with comments on its taxonomic and conservation status. Annals of the National Museum of Bloemfontein, 18, 78–94.
Bates, M.F. & Haacke, W.D (2003) The frogs of Lesotho: diversity and distribution. Annals of the National Museum of Bloemfontein, 19, 101–157.
Bates, M.F. (2004) Amietia vertebralis (Hewitt, 1927) species account. 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, DC, USA, pp. 277–280.
Boulenger, G.A. (1920) Descriptions of three new frogs in the collection of the British Museum. Annals and Magazine of Natural History, 9, 106.
Channing, A. (1979) Ecological and systematic relationships of Rana and Strongylopus in southern Natal (Amphibia: Anura). Annals of the Natal Museum, 23, 797–831.
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, 3763–3774.
Cunningham, M.J. & Cherry, M.I. (2004) Molecular systematics of African 20-chromosome toads (Anura: Bufonidae). Molecular Phylogenetics and Evolution, 32, 671–685.
Bush, S.F. (1952) On Rana umbraculata, a new frog from South Africa. Annals of the Natal Museum, 2, 153–164.
Dubois, A. (1987) Miscellanea taxinomica batrachologica (I). Alytes, 5, 7–95.
Dubois, A. (1992) Notes sur la classification des Ranidae (amphibians anoures). Bulletin Mensuel de la Société Linnéenne de Lyon, 61, 305–352.
Ewer, R.F. (1952) Some observations on the eye of Rana umbraculata Bush. Annals of the Natal Museum, 2, 165–174.
Fitzsimons, V. (1948) Notes on some reptiles and amphibians from the Drakensberg, together with a description of a new Platysaurus from Northern Natal. Annals of the Transvaal Museum, 21, 73–80.
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., Da Sá, R.O., Channing, A., Wilkonson, M., Donnellan, S.C. 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, New York, 297, 240–241.
Hewitt, J. (1927) Further descriptions of reptiles and batrachians from South Africa. Records of the Albany Museum, 3, 371–415.
Huelsenbeck, J.P. & Ronquist, F. (2001) MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Huelsenbeck, J.P. & Ronquist, F. (2004) Bayesian analysis of molecular evolution using MrBayes. Available from: http://workshop.molecularevolution.org/software/mrbayes/files/mrbayes_2004.pdf (accessed October 2007).
Kumar, S., Tamura, K. & Nei, M. (2004) MEGA version 3.1. Integrated software for Molecular Evolutionary Genetics Analysis and sequence alignment. Briefings in Bioinformatics, 5, 150–163.
Lambiris, A.J.L. (1987) Description of the tadpole of Strongylopus hymenopus (Boulenger, 1920) (Amphibia: Anura: Ranidae) and a key to described southern African tadpoles of the genus. Annals of the Natal Museum, 28, 455–462.
Lambiris, A.J.L. (1988) A review of the amphibians of Natal. Lammergeyer, 39, 1–209.
Lambiris, A.J.L. (1991) The use of laryngeal and buccopharyngeal morphology in anuran taxonomy. Doctoral thesis. Department of Biology, University of Natal.
Palumbi, S., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool’s guide to PCR. Version 2. Privately published document compiled by S Palumbi, Department of Zoology, University Hawaii, Honolulu.
Pearse, R.O. (1973) Barrier of Spears. Drama of the Drakensberg. Howard Timmins, Durban.
Poynton, J. C. (1964) The Amphibia of southern Africa: a faunal study. Annals of the Natal Museum, 17, 1–334.
Scott, E. (2005) A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis of morphological and molecular data. Cladistics, 21, 507–574.
Shimodaira, H. & Hasegawa, M. (1999) Multiple comparison of log-likelihoods with applications to phylogenetic inference. Molecular Biology and Evolution, 16, 1114–1116.
Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (* and other methods). Version 4.0b.10. Sinauer Associates, Sunderland, MA.
Van der Meijden, A., Vences, M., Hoegg, S. & Meyer, A. (2005) A previously unrecognised radiation of ranid frogs in southern Africa revealed by nuclear and mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 37, 674–685.
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.