Abstract
Eight new specimens are reported for the caecilian amphibian Boulengerula fischeri, a species known in detail previously only from its holotype collected in 1987. The new material was collected at the type locality in Rwanda in 2009, and is used to expand and refine the morphological characterization of the species. Mitochondrial and nuclear gene sequences were used to assess the phylogenetic position of B. fischeri in the context of a recent molecular phylogeny of six of the seven other species of Boulengerula (from Kenya and Tanzania). Among nominal species, only B. denhardti remains to be included in molecular phylogenetic studies of Boulengerula. Boulengerula is recovered as monophyletic, with either B. fischeri or (more probably) B. boulengeri sister to all other sampled species. There are at least three deep lineages within Boulengerula: (1) B. boulengeri, (2) B. fischeri, and (3) all other Eastern Arc Mountain and Coastal Forest species from Kenya and Tanzania. The status of Afrocaecilia, a genus erected by Taylor in 1968 to contain all Boulengerula except B. boulengeri, is not yet resolved.References
Behangana, M., Kasoma, P.M.B. & Luiselli, L. (2009) Ecological correlates of species richness and population abundance patterns in the amphibian communities from the Albertine Rift, East Africa. Biodiversity and Conservation, 18, 2855–2873.
Castresana, J. (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution, 17, 540–552.
Fischer, E. & Hinkel, H. (1992) Natur und Umwelt Ruandas. Einführing in die Flore und Fauna Ruandas und ihre wichtigsten Biotopsysteme. Ministerium des Innern und für Sport, Rheinland-Pfalz, Mainz, Germany. 452 pp.
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sà. R.O., Channing, A., Wilkinson, M., Donnellan, S.C. Raxworthy, C.J., Campbell, J.A., Blotto, B., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J., Green, D.M. & Wheeler, W.C. (2006) The amphiban tree of life. Bulletin of the American Museum of Natural History, 297, 1–370.
Gower, D.J., Loader, S.P., Moncrieff, C.B. & Wilkinson, M. (2004) Niche separation and comparative abundance of Boulengerula boulengeri and Scolecomorphus vittatus (Amphibia: Gymnophiona) in an East Usambara forest, Tanzania. African Journal of Herpetology, 53, 183–190.
Gower, D.J., Wilkinson, M., Sherratt, E. & Kok, P.J.R. (2010) A new species of Rhinatrema Duméril & Bibron (Amphibia: Gymnophiona: Rhinatrematidae) from Guyana. Zootaxa, 2391, 47–60.
Herrel, A. & Measey, G.J. (2010) The kinematics of locomotion in caecilians: effects of substrate and body shape. Journal of Experimental Zoology, 313A, 303–309.
Jones, D.T., Loader, S.P. & Gower, D.J. (2006) Trophic ecology of East African caecilians (Amphibia: Gymnophiona), and their impact on forest soil invertebrates. Journal of Zoology, London, 269, 117–126.
Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P.A., McWilliam, H., Valentin, F., Wallace, I.M., Wilm, A., Lopez, R., Thompson, J.D., Gibson, T.J. & Higgins, D.G. (2007) Clustal W and Clustal X version 2.0. Bioinformatics, 23, 2947–2948.
Loader, S.P., Wilkinson, M., Cotton, J.A., Müller, H., Menegon, M., Howell, K.M. & Gower, D.J. (in press) Molecular phylogenetics of Boulengerula (Amphibia: Gymnophiona: caeciliidae) and implications for taxonomy, biogeography and conservation. Herpetological Journal.
Malonza, P.K. & Measey, G.J. (2005) Life history of an African caecilian: Boulengerula taitanus Loveridge 1935 (Amphibia Gymnophiona Caeciliidae). Tropical Zoology, 18, 49–66.
Malonza, P.K. & Müller, H. (2004) A rediscovery after two decades: The Changamwe lowland caecilian Boulengerula changamwensis Loveridge, 1932 (Amphibia: Gymnophiona: Caeciliidae). Journal of East African Natural History, 93, 57–61.
Measey, G.J. (2004) Are caecilians rare? An East African perspective. Journal of East African Natural History, 93, 1–21.
Measey, G.J. & Barot, S. (2006) Evidence of seasonal migration in a tropical subterranean vertebrate. Journal of Zoology, 269, 29–37.
Measey, G.J., Mejissa, J. & Müller, H. (2006) Notes on the distribution and abundance of the caecilian Boulengerula uluguruensis (Amphibia: Gymnophiona: Caeciliidae) in the Uluguru Mountains, Tanzania. African Journal of Ecology, 44, 6–13.
Nussbaum, R.A. & Hinkel, H. (1994) Revision of East African caecilians of the genera Afrocaecilia Taylor and Boulengerula Tornier (Amphibia: Gymnophiona: Caeciliaidae). Copeia, 1994, 750–760.
Posada, D. (2008) jModelTest: Phylogenetic Model Averaging. Molecular Biology and Evolution, 25, 1253–1256.
Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K. & Bossuyt, F. (2007) Patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences of the United States of America, 104, 887–892.
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
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.
San Mauro, D., Gower, D.J., Massingham, T., Wilkinson, M., Zardoya, R. & Cotton, J.A. (2009) Experimental design in caecilian systematics: phylogenetic information of mitochondrial genomes and nuclear rag1. Systematic Biology, 58, 425–438.
Stamatakis, A. (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis using Parsimony. Version 4.0b. Sunderland, MA: Sinauer Associates.
Taylor, E.H. (1968) The caecilians of the World: a taxonomic review. University of Kansas Press, Lawrence, KA, USA. 848 pp.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
Wilkinson, M. & Loader, S. (2004) Boulengerula fischeri. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.2. <www.iucnredlist.org.apps/redlist/details/59497/0>. Downloaded on 25 August 2010.
Wilkinson, M. & Nussbaum, R.A. (2006) Caecilian phylogeny and classification. In: Exbrayat, J.-M. (Ed.), Reproductive Biology and Phylogeny of Amphibia Volume 3. Gymnophiona. Science Publishers Inc., Enfield, NH, USA, pp. 39–78.
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.
Wilkinson, M., Loader, S.P., Müller, H. & Gower, D.J. (2004) Taxonomic status and phylogenetic relationships of Boulengerula denhardti Nieden, 1912 (Amphibia, Gymnophiona, Caeciliidae). Mitteilungen aus dem Museum für Naturkunde Berlin, Zoologische Reihe, 80, 41–51.
Wilkinson, M., Nussbaum, R.A. & Hoogmoed, M. (2009) A new species of Microcaecilia (Amphibia: Gymnophiona: Caeciliidae) from Suriname. Herpetologica, 65, 413–418.
Zhang, P. & Wake, M.H. (2009) A mitogenomic perspective on the phylogeny and biogeography of living caecilians (Amphibia: Gymnophiona). Molecular Phylogenetics and Evolution, 53, 479–491.