Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2019-04-29
Page range: 342–366
Abstract views: 185
PDF downloaded: 3

Rediscovery, taxonomic status, and phylogenetic relationships of two rare and endemic snakes (Serpentes: Psammophiinae) from the southwestern Angolan plateau

Port Elizabeth Museum, Beach Road, Humewood, Port Elizabeth, South Africa Research Associate, Department of Zoology, Nelson Mandela University, Port Elizabeth, South Africa
Instituto Superior de Ciências da Educação da Huíla (ISCED-Huíla), Rua Sarmento Rodrigues, Lubango, Huíla, Angola CIBIO-InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Laboratório Associado, Universidade do Porto, Campus Agrário de Vairão, Rua Padre Armando Quintas, 4485-661 Vairão, Portugal
Zoology & Entomology Molecular Laboratory, Department of Zoology & Entomology, Rhodes University, Grahamstown, South Africa
Zoology & Entomology Molecular Laboratory, Department of Zoology & Entomology, Rhodes University, Grahamstown, South Africa
Psammophis ansorgii Psammophylax ocellatus new distribution phylogenetic relationships Angolan escarpment montane grassland Reptilia

Abstract

Two rare and endemic psammophines (Serpentes: Psammophiinae) occur in Angola. The taxonomic status of Psammophylax rhombeatus ocellatus Bocage, 1873 and Psammophis ansorgii Boulenger, 1905 have long remained problematic, with both having varied past and present taxonomic assignments, and whose distributions may therefore present zoogeographic anomalies. Little was known of their biology, habitat associations, or phylogenetic relationships. New material was collected during biodiversity surveys of the Humpata Plateau, near Lubango, Angola. It allowed fuller descriptions of scalation and live coloration for both species, and resolution of their taxonomic status. Genetic analysis confirms that both are distinct at the specific level. In addition, within Psammophis, Jalla’s Sand Snake (Psammophis jallae Peracca, 1896), of which P. rohani Angel, 1925, remains a synonym, is sister to P. ansorgii, and Boulenger’s comment on similarities with P. crucifer are not supported. The status of an unusual skaapsteker from Calueque, Cunene Province, Angola, is discussed and its assignment to Ps. ocellatus is provisional and requires additional material for taxonomic resolution. The new P. ansorgii records from Tundavala represent a range (+400 km southwest) and altitude (1800 m to 2286 m a.s.l) extension from the previous only known precise locality of Bela Vista (= Catchiungo), Huambo Province, whilst that for Ps. ocellatus doubles the known altitude from 1108 m to 2286 m a.s.l and extends the range about 122 km to the northwest from historical material from the plateau of Huíla and Cunene provinces.

References

  1. Akani, G.C., Barieenee, I.F., Capizzi, D. & Luiselli, L. (1999) Snake communities of moist rainforest and derived savanna sites of Nigeria: biodiversity patterns and conservation priorities. Biodiversity Conservation, 8, 629–642.

    https://doi.org/10.1023/A:1008849702810

    Arévalo, E., Davis, S.K, & Sites, J.J.W. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387–418.

    https://doi.org/10.1093/sysbio/43.3.387

    Baptista, N., António, T., Branch, W.R. (2018) Amphibians and reptiles of the Tundavala region of the Angolan Escarpment. Biodiversity & Ecology, 6, 397–403.

    https://doi.org/10.7809/b-e.00351

    Barbosa, L.A.G. (1970) Carta fitogeográfica de Angola. Instituto de Investigação Científica de Angola, Luanda, 323 pp.

    Bocage, J.V.B. (1873) Melanges erpetologiques. II. Sur quelques reptiles et batraciens nouveaux, rares ou peu connus d‘Afrique occidentale. Jornal de Sciências, Mathemáticas, Physicas e Naturaes, Lisboa, 4, 209–227.

    Bocage, J.V.B. (1895) Herpétologie d'Angola et du Congo. Imprimerie Nationale, Lisbonne, 203 pp., 20 pls.

    Boettger, O. (1883) Liste von Reptilien von Smithfield, Transvaal. Bericht Offenbacher Vereins für Naturkunde (Offen-bach am Main), 1883, 155–156.

    Bogert, C.M. (1940) Herpetological results of the Vernay-Angola Expedition, with notes on African reptiles in other collections. I. Snakes, including an arrangement of African Colubridae. Bulletin of the American Museum of Natural History, 77, 1–107.

    Boulenger, G.A. (1887) A synopsis of the snakes of South Africa. Zoologist, Series 3, 11, 171–182.

    Boulenger, G.A. (1905) A list of the batrachians and reptiles collected by Dr. W. J. Ansorge in Angola with descriptions of new species. Annals and Magazine of Natural History, Series 7, 16, 105–115.

    https://doi.org/10.1080/03745480509443656

    Bourgeois, M. (1968) Contribution à la morphologie comparée du crâne des Ophidiens de l’Afrique Centrale. Publications de l’Université Officielle du Congo, 18, 1–293.

    Branch, W.R. (2018) Snakes of Angola—an annotated checklist. Amphibian & Reptile Conservation, 12 (2), 41–82.

    Broadley, D.G. (1966) A review of the African Stripebellied Sand snakes of the genus Psammophis. Arnoldia, Rhodesia, 2 (36), 1–9.

    Broadley, D.G. (1977a) A review of the genus Psammophis in southern Africa (Serpentes: Colubridae). Arnoldia, Rhodesia, 8 (12), 1–29.

    Broadley, D.G. (1977b) A revision of the African snakes of the genus Psammophylax Fitzinger (Colubridae). Occasional Papers of the National Museum of Rhodesia, Series B, 6 (1), 1–44.

    Broadley, D.G. (1990) FitzSimons' Snakes of Southern Africa. Revised Edition): Jonathan Ball & Ad. DonkerPublishers, Parklands. 387pp.

    Broadley, D.G. (1997) A review of Hemirhagerrhis viperina (Bocage) (Serpentes: Colubridae), a rupicolous Psammophine snake. Madoqua, 19 (2), 161–169.

    Broadley, D.G. (2002) A review of the species of Psammophis Boie found south of Latitude 12° S (Serpentes: Psammophiinae). African Journal of Herpetology, 51 (2), 83–119.

    https://doi.org/10.1080/21564574.2002.9635466

    Bruford, M.W., Hanotte, O., Brookfield, J.F.Y. & Burke, T. (1992) Singlelocus and multilocus DNA fingerprint. In: Rus Hoelzel, A. (Ed.), Molecular genetic analysis of populations: a practical approach. IRL Press, Oxford, pp. 225–269.

    Cadle, J.E. (1994) The colubrid radiation in Africa (Serpentes: Colubridae): phylogenetic relationships and evolutionary patterns based on immunological data. Zoological Journal of the Linnaean Society, 110, 103–140.

    https://doi.org/10.1111/j.1096-3642.1994.tb01473.x

    Chapin, J.P. (2018) Manuscript: Ansorge, William J., summary of itineraries in Angola, 1903–1910, American Geographical Society of New York Records, 1723–2010, bulk 1854–2000. [undated]

    de Vienne, D.M., Giraud, T. & Martin, O.C. (2007) A Congruence Index for Testing Topological Similarity between Trees. Bioinformatics, 23 (23), 3119–3124.

    https://doi.org/10.1093/bioinformatics/btm500

    Dowling, H.G. (1951) A proposed standard system for counting ventral scales in snakes. British Journal of Herpetology, 1 (5), 97–99.

    Feio, M. (1981) O relevo do sudoeste de Angola, estudo de geomorfologia. Junta de Investigações Científicas do Ultramar, Lisboa, 326 pp.

    Figueroa, A., McKelvy, A.D., Grismer, L.L., Bell, C.D. & Lailvaux, S.P. (2016) A Species-Level Phylogeny of Extant Snakes with Description of a New Colubrid Subfamily and Genus. PLoS One, 11b (9), e0161070.

    https://doi.org/10.1371/journal.pone.0161070.

    Fischer, J.G. (1881) Herpetologische Bemerkungen vorzugsweise über Stücke des Naturhistorischen Museums in Bremen. Abhandlungen des Naturwissenschaftlichen Vereins in Bremen, 7, 225–238.

    Forstner, M.R.J., Davis, S.K. & Arevalo, E. (1995) Support for the hypothesis of Anguimorph ancestry for the suborder Serpentes from phylogenetic analysis of mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 4, 93–102.

    https://doi.org/10.1006/mpev.1995.1010

    Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.

    Hellmich, W. (1957) Die Reptilienausbeute der Hamburgischen Angola-Expedition. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 55, 39–80.

    Hoffmann, L.A.C. (1989) An annotated list of amphibian and reptiIe observations from Etosha National Park. Madoqua, 16 (2), 87–92.

    Hughes, B. (1999) Critical review of a revision of Psammophis (Linnaeus 1758) (Serpentes, Reptilia) by Frank Brandstätter. African Journal of Herpetology, 48, 63–70.

    https://doi.org/10.1080/21564574.1999.9651073

    Hughes, B. (2004) Misidentification of Dromophis lineatus (Dumèril & Bibron, 1854) as Psammophis sibilans (Linné 1758) and the perpetuation of error. African Journal of Herpetology, 53, 63–76.

    https://doi.org/10.1080/21564574.2004.9635498

    Hughes, B. & Wade, E.O.Z. (2002) On the African leopard whip snake Psammophis leopardinus Bocage 1887 (Serpentes: Colubridae), with the description of a new species from Zambia. Bulletin of the natural History Museum, London, Zoology, 68, 75–81.

    Hughes, B. & Wade, E.O.Z. (2004) Is Psammophis sibilans occidentalis Werner, 1919 a junior synonym of P. phillipsi (Hallowell, 1844)? (Squamata: Serpentes: Colubridae). Herpetozoa, 16 (3/4), 127–132.

    Huntley, B. (2009) SANBI/ISCED/UAN Angolan Biodiversity Assessment and Capacity Building Project, Report on Pilot Project. Unpublished Report, South African National Biodiversity Institute, Pretoria, 111 pp.

    Huntley, B.J. & Matos, E.M. (1994) Botanical diversity and its conservation in Angola. Strelitzia, 1, 53–74.

    Jürgens, N., Strohbach, B., Lages, F., Schmiedel, U., Finckh, M., Sichone, P., Hahn, L.-M., Zigelski, P. (2018) Biodiversity observation—an overview of the current state and first results of biodiversity monitoring studies. Biodiversity & Ecology, 6, 382–396.

    https://doi.org/10.7809/b-e.00350

    Kelly, C.M.R., Barker, N.P., Villet, M.H., Broadley, D.G. & Branch, W.R. (2008) The snake family Psammophiidae (Reptilia: Serpentes): phylogenetics and species delimitation in the African Sand Snakes (Psammophis Boie, 1825) and allied genera. Molecular Phylogenetics and Evolution, 47, 1045–1060.

    https://doi.org/10.1016/j.ympev.2008.03.025

    Kelly, C.M.R., Branch, W.R., Broadley, D.G., Barker, N.P. & Villet, M.H. (2011) Molecular systematics of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903. Molecular Phylogenetics and Evolution, 58 (2), 415–426.

    https://doi.org/10.1016/j.ympev.2010.11.010

    Largan, M.J. & Rasmussen, J.B. (1993) Catalogue of the snakes of Ethiopia (Reptilia Serpentes), including identification keys. Tropical Zoology, 6, 313–434.

    https://doi.org/10.1080/03946975.1993.10539231

    Laurent, R.F. (1964) Reptiles et Amphibiens de l’Angola (Troisième contribution). Publicações culturais da Companhia de Diamantes de Angola, 67, 11–165.

    Loveridge, A. (1940) Revision of the African snakes of the genera Dromophis and Psammophis. Bulletin of the Museum of comparative Zoology, 87 (1), 1–69.

    Luiselli, L., Akani, G.C., Angelici, F.M., Eniang, E.A., Ude, L. & Politano, E. (2004) Local distribution, habitat use, and diet of two supposed ‘species’ of the Psammophis “phillipsi’ complex (Serpentes: Colubridae), sympatric in southern Nigeria. Amphibia-Reptilia, 25, 415–423.

    https://doi.org/10.1163/1568538042788915

    Maddison, W.P. & Maddison, D.R. (2017) Mesquite: a modular system for evolutionary analysis. Version 3.3. Available from: http://mesquiteproject.org (accessed 27 November 2018)

    Mendelsohn, J., Jarvis, A. & Robertson, J. (2013) A profile and atlas of the Cuvelai-Etosha Basin. Publ. Raison & Gondwana Collection, Windhoek, 170 pp.

    Nagy, Z.T., Joger, U., Wink, M., Glaw, F. & Vences, M. (2003) Multiple colonization of Madagascar and Socotra by colubrid snakes: evidence from nuclear and mitochondrial gene phylogenies. Proceedings of the Royal Society of London, B, 270, 2613–2621.

    https://doi.org/10.1098/rspb.2003.2547

    Posada, D. (2008) jModelTest: Phylogenetic Model Averaging. Molecular Biology and Evolution, 25, 1253–1256.

    https://doi.org/10.1093/molbev/msn083

    Pyron, R.A., Burbrink, F.T. & Wiens, J.J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13, 93.

    https://doi.org/10.1186/1471-2148-13-93

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2013) Tracer v1.5. University of Oxford, Rambaut Research Group. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 14 April 2018)

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539–542.

    https://doi.org/10.1093/sysbio/sys029

    Ruane, S., Myers, E.A., Lo, K., Yuen, S., Welt, R.S., Juman, M., Futterman, I., Nussbaum, R.A., Schneider, G., Burbrink, F.T. & Raxworthy, C.J. (2017) Unrecognized species diversity and new insights into colour pattern polymorphism within the widespread Malagasy snake Mimophis (Serpentes: Lamprophiidae), Systematics and Biodiversity, 16 (3), 229–244.

    https://doi.org/10.1080/14772000.2017.1375046

    SASSCAL ObservationNet (2018) http://www.sasscalobservationnet.org/ (accessed 13 April 2018)

    Shine, R. (1994) Sexual dimorphism in snakes revisited. Copeia, 1994 (2), 326–346.

    https://doi.org/10.2307/1446982

    Slowinski, J.B. & Lawson, R. (2002) Snake phylogeny: evidence from nuclear and mitochondrial genes. Molecular Phylogenetics Evolution, 24, 194–202.

    https://doi.org/10.1016/S1055-7903(02)00239-7

    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.

    https://doi.org/10.1093/molbev/mst197

    Vaz Pinto, P., Veríssimo, L. & Branch, W.R. (2018) Hiding in the bushes for 107 years: rediscovery of a lost Angolan gecko (Phyllodactylus ansorgii Boulenger, 1907, Sauria: Gekkonidae). [in preparation]

    Vidal, N. & Hedges, S.B. (2002) Higher-level relationships of caenophidian snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus de l’Académie des Sciences, 325, 987–995.

    https://doi.org/10.1016/S1631-0691(02)01509-3

    Volynchik, S. (2012) Morphological variability in Vipera palaestinae along an environmental gradient. Asian Herpetological Research, 3 (3), 227–239.

    https://doi.org/10.3724/SP.J.1245.2012.00227

    Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. Taylor and Francis, CRC Press, Florida, 1237 pp.

    https://doi.org/10.1201/b16901

    Whiting, A.S., Bauer, A.M. & Sites, J.W. (2003) Phylogenetic relationships and limb loss in sub-Saharan African scincine lizards (Squamata: Scincidae). Molecular Phylogenetics and Evolution, 29, 582–598.

    https://doi.org/10.1016/S1055-7903(03)00142-8

    Xia, X. (2013) DAMBE5: a comprehensive software package for data analysis in molecular biology and evolution. Molecular Biology and Evolution, 30 (7), 1720–1728.

    https://doi.org/10.1093/molbev/mst064

    Zaher, H. (1999) Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History, 240, 1–168. Zwickl, D.J. (2006) Genetic algorithm approaches for the phylogenetic analysis of large biological sequence datasets under the maximum likelihood criterion. Ph.D. dissertation, The University of Texas at Austin, Texas, 115 pp.