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Recovering the evolutionary history of Africa’s most diverse viper genus: morphological and molecular phylogeny of Bitis (Reptilia: Squamata: Viperidae)

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Abstract

Assessing evolutionary relationships among wide-ranging species can be particularly beneficial to our understanding of speciation patterns and biogeography of taxa, with broad implications for conservation and applications for human health. Integrative phylogenetic analyses that incorporate multiple independent datasets (e.g., DNA, protein, phenotype) can resolve many problematic issues in systematics such as cryptic diversity and incongruence between datasets. Vipers in the genus Bitis are widely distributed throughout much of sub-Saharan Africa, filling a variety of ecological niches and presenting an important public health problem. However, evolutionary relationships among this medically and ecologically important genus have not been fully resolved due to inadequate taxon sampling and lack of informative characters. Here, we conduct the first phylogenetic study incorporating complete sampling of known species within the genus Bitis. Using morphological, molecular, and combined approaches under multiple criteria, we recovered many of the species groups detected by previous investigators, further validating four currently recognized subgenera. Bitis arietans and Bitis worthingtoni appear to be early-diverging, monotypic lineages, while the “big Bitis” group and the small southern African species form distinct clades. Although our study provides additional information regarding the interspecific relationships within Bitis, the placement of Bitis albanica, Bitis heraldica, and Bitis inornata remains problematic. This study enhances our understanding of the evolutionary history of species within the genus Bitis incorporating a combined evidence approach to phylogenetics.

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Acknowledgments

We thank the following curators who kindly made specimens or other material under their care available to us: L.S. Ford (AMNH); N. Gilmore (ANSP); S. Rogers and J.J. Wiens (CM); A. Resetar (FMNH); J.E. Simmons, E. Greenbaum, and L. Trueb (KU); A.D. Leaché, F.T. Burbrink, and J.A. McGuire (LSUMZ); J. Rosado and J. Hanken (MCZ); C. Cicero and D.B. Wake (MVZ); W.R. Branch (PEM); G. Schneider and A.G. Kluge (UMMZ); R. Reynolds, J. Poindexter III, S.W. Gotte, and R.W. McDiarmid (USNM); R. Ackley and J.A. Campbell (UTA); and J.B. Rasmussen (ZMUC). We thank A. Saunders and B. Shipley of the Denver Zoo for graciously providing us photographs of live Bitis specimens for our figures. We thank D. Killebrew, D. Pogue, N.B. Ford, and S.A. Orlofske for reviewing an earlier version of this manuscript. Stephen Spawls was an invaluable field companion as RDW and RLG conducted research in the southwestern forests of Ethiopia. Bill Branch graciously contributed correspondence and advice to RDW during this study. Funding for a trip to Ethiopia by RDW and RLG in search of Bitis parviocula was provided through a University of Texas at Tyler Fund for Excellence grant and an East Texas Herpetological Society grant. RCJ received funding through an NSF GK-12 Fellowship (DGE 0742544).

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Authors and Affiliations

  1. Department of Biology, University of Texas at Tyler, Tyler, TX, 75799, USA

    Rod D. Wittenberg, Robert C. Jadin, Allyson M. Fenwick & Ronald L. Gutberlet Jr.

  2. U.S. Fish and Wildlife Service, Crescent Lake National Wildlife Refuge, 10630 Road 181, Ellsworth, NE, 69340, USA

    Rod D. Wittenberg

  3. Department of Biology, Northeastern Illinois University, Chicago, IL, 60625, USA

    Robert C. Jadin

  4. Department of Biology, University of Central Oklahoma, Edmond, OK, 73034, USA

    Allyson M. Fenwick

  5. Department of Biological Sciences, Salisbury University, Salisbury, MD, 21801, USA

    Ronald L. Gutberlet Jr.

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  1. Rod D. Wittenberg
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  4. Ronald L. Gutberlet Jr.
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Correspondence to Robert C. Jadin.

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Rod D. Wittenberg and Robert C. Jadin are joint first authors.

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Appendix S1

Specimens examined. Museum acronyms follow Leviton et al. (1985). (DOC 258 kb)

Appendix S2

Morphological character descriptions used in the morphological phylogenetic analyses. (DOC 256 kb)

Table S1

Taxa and data used in DNA analysis. Abbreviations are as follows: ND4 = NADH dehydrogenase subunit 4, Cyt b = cytochrome b, 16S and 12S = small ribosomal RNA fragments, ND2 = NADH dehydrogenase subunit 2, PRLR = prolactin receptor, Ba34 = anonymous nuclear locus from Barlow et al. (2012). (DOC 256 kb)

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Wittenberg, R.D., Jadin, R.C., Fenwick, A.M. et al. Recovering the evolutionary history of Africa’s most diverse viper genus: morphological and molecular phylogeny of Bitis (Reptilia: Squamata: Viperidae). Org Divers Evol 15, 115–125 (2015). https://doi.org/10.1007/s13127-014-0185-3

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