Skip to main content Skip to main navigation menu Skip to site footer
Article
Published: 2021-04-27

Night stalkers from above: A monograph of Toxicodryas tree snakes (Squamata: Colubridae) with descriptions of two new cryptic species from Central Africa

University of Texas at El Paso, Department of Biological Sciences, 500 W. University Avenue, El Paso, Texas 79912, USA
Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Lawrence, Kansas 66045, USA Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
University of Texas at El Paso, Department of Biological Sciences, 500 W. University Avenue, El Paso, Texas 79912, USA
Royal Belgian Institute of Natural Sciences, Rue Vautier 29, 1000 Brussels, Belgium
4 Potter Park, Cambridge, Massachusetts 02138, USA.
Laboratoire d’Hérpétologie, Département de Biologie, Centre de Recherche en Sciences Naturelles, Lwiro, Democratic Republic of the Congo
Laboratoire d’Hérpétologie, Département de Biologie, Centre de Recherche en Sciences Naturelles, Lwiro, Democratic Republic of the Congo
Institut Supérieur d’écologie pour la Conservation de la Nature, Katana Campus, Democratic Republic of the Congo
Department of Ecology and Biodiversity of Earth Resources, Centre de Surveillance de la Biodiversité of the University of Kisangani, Kisangani, Democratic Republic of the Congo
Department of Ecology and Aquatic Biodiversity, Biodiversity Monitoring Centre, University of Kisangani, Kisangani, Democratic Republic of the Congo
Museum für Naturkunde—Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany
Museum für Naturkunde—Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany
University of Texas at El Paso, Department of Biological Sciences, 500 W. University Avenue, El Paso, Texas 79912, USA
Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Lawrence, Kansas 66045, USA Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Lawrence, Kansas 66045, USA Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Lawrence, Kansas 66045, USA Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
Department of Ecology and Evolutionary Biology, University of Kansas, 1200 Sunnyside Avenue, Lawrence, Kansas 66045, USA Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045, USA
Reptilia Congo Basin arboreal cryptic species rivers taxonomy

Abstract

The genus Toxicodryas, historically included with the renowned Australasian cat-eyed snakes of the colubrid genus Boiga, currently includes two widespread species (T. blandingii and T. pulverulenta) in western, central, and eastern Africa. We leverage findings from a recent phylogenomic and historical demographic analysis of this genus (based on 2848–4471 Rad-seq loci from across the genome), with robust sampling from throughout the ranges of both species, to define two additional taxonomic units, with species boundaries corresponding to river barriers. Additional morphometric data from scores of examined museum specimens and literature records bolster the recognition of these two new cryptic species. We hypothesize that T. blandingii occurs west of the confluence of the Congo and Ubangi rivers, whereas a cryptic new species that is found east of this biogeographic barrier has significantly higher numbers of ventral scale counts in both sexes, additional significant differences in several scale counts, and lower venom toxicity. Toxicodryas pulverulenta occurs west of the Niger Delta in West Africa, whereas a cryptic new species that is found east of this biogeographic barrier has significantly higher numbers of subcaudal scale counts in both sexes. A review of published information regarding morphological variation, ecology, natural history, habitat, and venom is summarized for these four Toxicodryas species.

 

References

  1. Akaffou, M.H., Chippaux, J.-P., Kouadio Allali, B., Coulibaly, Z. & Dosso, M. (2017) Peuplement ophidien des plantations d’Hevea brasiliensis d’Anguédédou (sud-est de la Côte d’Ivoire). Bulletin de la Société Herpétologique de France, 162, 31–38.

    Akani, G.C., Amadi, N., Dendi, D. & Luiselli, L. (2018) Do community metrics vary in reptile communities from Niger Delta forests subjected to slash-and-burn agricultural practices? African Journal of Ecology, 56 (4), 1044–1048.

    https://doi.org/10.1111/aje.12547

    Akani, G.C., Ebere, N., Luiselli, L. & Eniang, E.A. (2007) Community structure and ecology of snakes in fields of oil palm trees (Elaeis guineensis) in the Niger Delta, southern Nigeria. African Journal of Ecology, 46 (4), 500–506.

    https://doi.org/10.1111/j.1365-2028.2007.00885.x

    Allen, K.E., Greenbaum, E., Hime, P.M., Tapondjou Nkonmeneck, W.P., Sterkhova, V.T., Kusamba, C., Rödel, M.-O., Penner, J., Towsend Peterson, A. & Brown, R.M. (2021) Rivers, not refugia, drove diversification in arboreal, sub-Saharan African snakes. Ecology and Evolution. [in press]

    Amri, K. & Chippaux, J.-P. (2012) Envenimation bénigne par morsure de Toxicodryas blandingii (Hallowell, 1844) en Suisse. Bulletin de la Société Herpétologique de France, 142–143, 145–148.

    Angel, F. (1933) Les Serpents d’Afrique Occidentale Française. Larose Éditeur, Paris, 246 pp.

    Angel, F., Guibé, J., Lamotte, J. & Roy, R. (1954) La réserve naturelle intégrale du Mont Nimba. XXXII. Serpents. Mémoires de l’Institut Française d’Afrique Noire, 40, 381–402.

    Auliya, M., Wagner, P. & Böhme, W. (2012) The herpetofauna of the Bijagós archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist. Bonn zoological Bulletin, 61 (2), 255–281.

    Ayanlade, A. & Proske, U. (2015) Assessing wetland degradation and loss of ecosystem services in the Niger Delta, Nigeria. Marine and Freshwater Research, 67 (6), 828–836.

    https://doi.org/10.1071/MF15066

    Barry, D.H. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 42–43.

    Blackwell, K. (1967) A preliminary survey of the reptiles of the Ibadan area. British Journal of Herpetology, 3, 307–311.

    Bocage, J.V.B. (1895) Herpétologie d’Angola et du Congo. Ministério da Marinha e das Colónias, Lisbonne, XX + 203 pp., pls. I–XIX.

    Bogert, C.M. (1940) Herpetological results of the Vernay Angola expedition. Bulletin of the American Museum of Natural History, 77, 1–107, pl. I.

    Böhme, W. (2000) Diversity of a snake community in a Guinean rain forest (Reptilia, Serpentes). In: Rheinwald, G. (Ed.), Isolated Vertebrate Communities in the Tropics. Proceedings of the 4th International Symposium in Bonn. Bonner zoologische Monographien, 46, 69–78.

    Booth, A.H. (1958) The Niger, the Volta and the Dahomey Gap as geographic barriers. Evolution, 12 (1), 48–62.

    https://doi.org/10.1111/j.1558-5646.1958.tb02927.x

    Boulenger, G.A. (1896) Catalogue of the Snakes in the British Museum (Natural History). Vol. III. Containing the Colubridæ (Opisthoglyphæ and Proteroglyphæ), Amblycephalidæ, and Viperidæ. Trustees of the British Museum (Natural History), London, xiv + 727 pp.

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

    https://doi.org/10.5962/bhl.part.15108

    Bourgeois, M. (1968) Contribution a la morphologie comparée du crane des ophidiens de l’Afrique centrale. Publications de l’Université Officielle du Congo à Lubumbashi, 18, 1–293 + tables I–IV.

    Branch, W.R. (2005) A Photographic Guide: Snakes Other Reptiles and Amphibians of East Africa. Ralph Curtis Publishing, Inc., Sanibel Island, Florida, 132 pp.

    Branch, W.R. & Rödel, M.-O. (2003) Herpetological survey of the Haute Dodo and Cavally forests, western Ivory Coast, Part II: Trapping results and reptiles. Salamandra, 39 (1), 21–38.

    Broaders, M. & Ryan, M.F. (1997) Enzymatic properties of the Duvernoy’s secretion of Blanding’s tree snake (Boiga blandingi) and of the mangrove snake (Boiga dendrophila). Toxicon, 35 (7), 1143–1148.

    https://doi.org/10.1016/S0041-0101(96)00207-3

    Broaders, M., Carlos, F. & Ryan, M.F. (1999) Partial purification of acetylcholine receptor binding components from the Duvernoy’s secretions of Blanding’s tree snake (Boiga blandingi) and the mangrove snake (Boiga dendrophila). Journal of Natural Toxins, 8 (2), 155–166.

    Broadley, D.G. & Cotterill, F.P.D. (2004) The reptiles of southeast Katanga, an overlooked ‘hot spot.’ African Journal of Herpetology, 53 (1), 35–61.

    https://doi.org/10.1080/21564574.2004.9635497

    Broadley, D.G., Doria, C.T. & Wigge, J. (2003) Snakes of Zambia: An Atlas and Field Guide. Edition Chimaira, Frankfurt am Main, 280 pp.

    Butler, J.A. & Reid, J.C. (1986) Habitat preferences of snakes in the southern Cross River State, Nigeria. In: Roček, Z. (Ed.), Studies in Herpetology. Charles University, Prague, pp. 483–488.

    Butler, J.A. & Reid, J.C. (1990) Records of snakes from Nigeria. The Nigerian Field, 55 (1), 19–40.

    Butynski, T.M., Kingdon, J. & Kalina, J. (Eds.) (2013) Mammals of Africa. Volume II. Primates. Bloomsbury, London, New Delhi, New York, and Sydney, 556 pp.

    Cansdale, G. (1954) Gold Coast snakes—a complete list. The Nigerian Field, 19, 118–132.

    Cansdale, G. (1955) Reptiles of West Africa. Penguin Books, London, United Kingdom, 104 pp.

    Cansdale, G.S. (1965) West African Snakes. Longmans, Green and Co. Ltd., London, vi + 74 pp.

    Carlino, P. & Pauwels, O.S.G. (2015) An updated reptile list of Ivindo National Park, the herpetofaunal hotspot of Gabon. Bulletin of the Chicago Herpetological Society, 50 (3), 25–39.

    Ceríaco, L.M.P., Marques, M.P. & Bauer, A.M. (2018) Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the terrestrial herpetofauna of São Tomé, Príncipe and Annobon islands. Zootaxa, 4387 (1), 091–108.

    https://doi.org/10.11646/zootaxa.4387.1.4

    Ceríaco, L.M.P., Marques, M.P., Schmitz, A. & Bauer, A.M. (2017) The “Cobra-preta” of São Tomé Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae). Zootaxa, 4324 (1), 121–141.

    https://doi.org/10.11646/zootaxa.4324.1.7

    Chabanaud, P. (1917a) énumération des ophidiens non encore étudiés de l’Afrique occidentale, appartenant aux collections du Muséum avec la description des espèces et des variétés nouvelles. Bulletin du Muséum d’Histoire Naturelle, Paris, 22, 362–382.

    https://doi.org/10.5962/bhl.part.17132

    Chabanaud, P. (1917b) Note complémentaire sur les ophidiens de l’Afrique occidentale, avec la description d’une espèce nouvelle. Bulletin du Muséum d’Histoire Naturelle, Paris, 23, 7–14.

    Chabanaud, P. (1917c) Révision de quelques reptiles d’Afrique et description de trois espèces nouvelles. Bulletin du Muséum d’Histoire Naturelle, Paris, 23, 442–454.

    Chan, K.O., Hutter, C.R., Wood, Jr., P.L., Grismer, L.L., Das, I. & Brown, R.M. (2020) Gene flow creates a mirage of cryptic species in a Southeast Asian spotted stream frog complex. Molecular Ecology, 29 (20), 3970–3987.

    https://doi.org/10.1111/mec.15603

    Chippaux, J-P. (2006) Les Serpents d’Afrique Occidentale et Centrale. 3rd Edition. IRD Éditions, Paris, 311 pp.

    Chippaux, J.-P. & Jackson, K. (2019) Snakes of Central and Western Africa. Johns Hopkins University Press, Baltimore, Maryland, x + 429 pp.

    Chirio, L. & Ineich, I. (2006) Biogeography of the reptiles of the Central African Republic. African Journal of Herpetology, 55 (1), 23–59.

    https://doi.org/10.1080/21564574.2006.9635538

    Chirio, L. & LeBreton, M. (2007) Atlas des Reptiles du Cameroun. Publications Scientifiques du MNHN, IRD Éditions, Paris, 686 pp.

    Condamin, M. (1958) La collection de serpents de l’I.F.A.N. (acquisitions 1956). Bulletin de l’Institut Français d’Afrique Noire, 20A (1), 243–262.

    Condamin, M. (1959) Serpents récoltés à Sérédou (Guinée) par R. Pujol. Bulletin de l’Institut Français d’Afrique Noire, 21A (4), 1351–1366.

    Couvreur, T.L.P., Dauby, G., Blach-Overgaard, A., Deblauwe, V., Dessein, S., Droissart, V., Hardy, O.J., Harris, D.J., Janssens, S.B., Ley, A.C., Mackinder, B.A., Sonké, B., Sosef, M.S.M., Stévart, T., Svenning, J.-C., Wieringa, J.J., Faye, A., Missoup, A.D., Tolley, K.A., Nicolas, V., Ntie, S., Fluteau, F., Robin, C., Guillocheau, F., Barboni, D. & Sepulchre, P. (2021) Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna. Biological Reviews, 96 (1), 16–51.

    https://doi.org/10.1111/brv.12644

    Cozens, B. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 44.

    Dashevsky, D., Debono, J., Rokyta, D., Nouwens, A., Josh, P. & Fry, B.G. (2018) Three-finger toxin diversification in the venoms of cat-eye snakes (Colubridae: Boiga). Journal of Molecular Evolution, 86, 531–545.

    https://doi.org/10.1007/s00239-018-9864-6

    de Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation. In: Howard, D.J. & Berlocher, S.H. (Eds.), Endless Forms: Species and Speciation. Oxford University Press, New York, New York, pp. 57–75.

    de Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879–886.

    https://doi.org/10.1080/10635150701701083

    de Witte, G.-F. (1933) Reptiles récoltés au Congo Belge par le Dr H. Schouteden et par M. G.-F. de Witte. Annales du Musée du Congo Belge, C.—Zoologie, 3 (1), 55–98.

    de Witte, G.-F. (1962) Genera des serpents du Congo et du Ruanda-Urundi. Annales du Musée Royal de l’Afrique Centrale, Sciences Zoologique, Série in Octavo, 104, i–viii + 1–203.

    de Witte, G.-F. (1966) Exploration du Parc National de la Garamba. Reptiles. Institut du Parcs Nationaux du Congo, 48, 1–108, pls. I–V.

    de Witte, G.-F. (1975) Serpents, Boidae, Colubridae, Elapidae et Viperidae. Fondacion pour Favoriser les Recherches Scientifiques en Afrique, Série 2, 24, 5–121.

    Dekeyser, P.L. (1955) Les Mammifères de l’Afrique Noire Française. 2nd Edition. Institut Français d’Afrique Noire, Initiations Africaines, Dakar, Senegal, 426 pp.

    Demenou, B.B., Piñeiro, R. & Hardy, O.J. (2016) Origin and history of the Dahomey Gap separating West and Central African rain forests: Insights from the phylogeography of the legume tree Distemonanthus benthamianus. Journal of Biogeography, 43 (5), 1020–1031.

    https://doi.org/10.1111/jbi.12688

    Doucet, J. (1963) Les serpents de la République de Côte d’Ivoire. Serpents venimeux. Acta Tropica, 20 (3-4), 297–340.

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

    Dowling, H.G. & Savage, J.M. (1960) A guide to the snake hemipenis: A survey of basic structure and systematic characteristics. Zoologica, 45 (Part 1), 17–28, pls. I–III.

    Droissart, V., Dauby, G., Hardy, O.J., Deblauwe, V., Harris, D.J., Janssens, S., Mackinder, B.A., Blach-Overgaard, A., Sonké, B., Sosef, M.S.M., Stévart, T. & Svenning, J.-C. (2018). Beyond trees: Biogeographical regionalization of tropical Africa. Journal of Biogeography, 45 (5), 1153–1167.

    https://doi.org/10.1111/jbi.13190

    Duméril, A.H.A. (1859) Reptiles et poissons de l’Afrique occidentale. étude précédée de considérations générales sur leur distribution géographique. Archives du Muséum National d’Histoire Naturelle, 10, 137–240.

    Dunger, G.T. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 43.

    Eriksson, J., Hohmann, G., Boesch, C. & Vigilant, L. (2004) Rivers influence the population genetic structure of bonobos (Pan paniscus). Molecular Ecology, 13 (11), 3425–3435.

    https://doi.org/10.1111/j.1365-294X.2004.02332.x

    Farooq, H., Azevedo, J.A.R., Soares, A., Antonelli, A. & Faurby, S. (2021) Mapping Africa’s biodiversity: More of the same is just not good enough. Systematic Biology. [in press]

    https://doi.org/10.1093/sysbio/syaa090

    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, 11 (9), e0161070.

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

    Fischer, J.G. (1856) Neue Schlangen des Hamburgischen Naturhistorischen Museums. Abhandlungen aus dem Gebiete der Naturwissenschaften herausgegeben von Naturwissenschaftlichen Verein in Hamburg, 3, 79–116 + 3 pl.

    Fuchs, J. & Bowie, R.C.K. (2015) Concordant genetic structure in two species of woodpecker distributed across the primary West African biogeographic barriers. Molecular Phylogenetics and Evolution, 88, 64–74.

    https://doi.org/10.1016/j.ympev.2015.03.011

    Ganesh, S.R., Achyuthan, N.S., Chandramouli, S.R. & Vogel, G. (2020) Taxonomic revision of the Boiga ceylonensis group (Serpentes: Colubridae): reexamination of type specimens, redefinition of nominate taxa and an updated key. Zootaxa, 4779 (3), 301–332.

    https://doi.org/10.11646/zootaxa.4779.3.1

    Gans, C. (1961) Mimicry in procryptically colored snakes of the genus Dasypeltis. Evolution, 15 (1), 72–91.

    https://doi.org/10.2307/2405844

    Gauduin, P. (1970) Contributions à la faune de la région de Yaoundé. Annales de la Faculté des Sciences du Cameroun, 4, 73–99.

    Goodman, J.D. (1985) Two record size Blanding’s tree snakes from Uganda. East African Natural History Society Bulletin, 1985, 56–57.

    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.

    Greenbaum, E. (2017) Emerald Labyrinth: A Scientist’s Adventures in the Jungles of the Congo. ForeEdge, Lebanon, New Hampshire, xv + 329 pp.

    Greenbaum, E. & Carr, J.L. (2005) The herpetofauna of Upper Niger National Park, Guinea, West Africa. Scientific Papers, Natural History Museum, The University of Kansas, 37, 1–21.

    https://doi.org/10.5962/bhl.title.8470

    Greenbaum, E., Portillo, F., Jackson, K. & Kusamba, C. (2015) A phylogeny of Central African Boaedon (Serpentes: Lamprophiidae), with the description of a new cryptic species from the Albertine Rift. African Journal of Herpetology, 64 (1), 18–38.

    https://doi.org/10.1080/21564574.2014.996189

    Greene, H.W. (1989) Ecological, evolutionary, and conservation implications of feeding biology in Old World cat snakes, Genus Boiga (Colubridae). Proceedings of the California Academy of Sciences, 46 (8), 193–207.

    Greene, H.W. (1997) Snakes: The Evolution of Mystery in Nature. University of California Press, Berkeley, Los Angeles and London, xiii + 351 pp.

    Groves, F. (1973) Reproduction and venom in Blanding’s tree snake. International Zoo Yearbook, 13, 106–108.

    https://doi.org/10.1111/j.1748-1090.1973.tb02114.x

    Günther, A. (1858) Catalogue of Colubrine Snakes in the Collection of the British Museum. Trustees of the British Museum, London, xvi + 281 pp.

    https://doi.org/10.5962/bhl.title.13272

    Hallermann, J. (1998) Annotated catalog of the type specimens of the herpetological collection in the Zoological Museum of the University of Hamburg. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 95, 197–223.

    Hallermann, J., Ceríaco, L.M.P., Schmitz, A., Ernst, R., Conradie, W., Verburgt, L., Marques, M.P. & Bauer, A.M. (2020) A review of the Angolan house snakes, genus Boaedon Duméril, Bibron and Duméril (1854) (Serpentes: Lamprophiidae), with the description of three new species in the Boaedon fuliginosus (Boie, 1827) species complex. African Journal of Herpetology, 69 (1), 29–78.

    https://doi.org/10.1080/21564574.2020.1777470

    Hallowell, E. (“1844” 1845) Description of new species of African reptiles. Proceedings of the Academy of Natural Sciences of Philadelphia, 2 (6), 169–172.

    Hallowell, E. (1854) Remarks on the geographical distribution of reptiles, with descriptions of several species supposed to be new, and corrections of former papers. Proceedings of the Academy of Natural Sciences of Philadelphia, 7 (1854–1855), 98–105.

    Hallowell, E. (1857) Notice of a collection of reptiles from the Gaboon country, West Africa, recently presented to the Academy of Natural Sciences of Philadelphia, by Dr. Henry A. Ford. Proceedings of the Academy of Natural Sciences of Philadelphia, 9 (1857–1858), 48–72.

    Happold, D.C.D. (1996) Mammals of the Guinea–Congo rain forest. Proceedings of the Royal Society of Edinburgh, 104B, 243–284.

    https://doi.org/10.1017/S0269727000006151

    Happold, M. & Happold, D. (Eds.) (2013) Mammals of Africa. Vol. IV. Hedgehogs, Shrews and Bats. Bloomsbury, London, New Delhi, New York, and Sydney, 800 pp.

    Hedges, N.G. (1983) Reptiles and Amphibians of East Africa. Kenya Literature Bureau, Nairobi, Kenya, xii + 139 pp.

    Hellmich, W. (1957a) Herpetologische Ergebnisse einer Forschungreise in Angola. Veröffentlichungen der Zoologischen Staatssammlung München, 5, 1–92.

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

    Hirschfeld, M., Blackburn, D.C., Burger, M., Greenbaum, E., Zassi-Boulou, A.-G. & Rödel, M.-O. (2015) Two new species of long-fingered frogs of the genus Cardioglossa (Anura: Arthroleptidae) from Central African rainforests. African Journal of Herpetology, 64 (2), 81–102.

    https://doi.org/10.1080/21564574.2015.1052102

    Hughes, B. (1983) African snake faunas. Bonner zoologische Beiträge, 34 (1–3), 311–356.

    Hughes, B. (1988) Herpetology in Ghana (West Africa). British Herpetological Society Bulletin, (25), 29–37.

    Hughes, B. (2000) Polymorphism, sex, size and other aspects of the African snake, Boiga blandingi (Hallowell, 1844). Herpetological Bulletin, 74, 8–12.

    Hughes, B. & Barry, D.H. (1969) The snakes of Ghana: a checklist and key. Bulletin de l’Institut fondamental d’Afrique noire. Série A, Sciences naturelles, 31 (3), 1004–1041.

    Isemonger, R.M. (1962) Snakes of Africa: Southern, Central and East. Thomas Nelson and Sons (Africa) (Party) Ltd., Johannesburg, viii + 236 pp.

    IUCN (2020) The IUCN Red List of Threatened Species. Version 2020-2. Available from: https://www.iucnredlist.org (accessed 30 November 2020)

    Jan, G. & Sordelli, F. (1870–1881) Iconographie Générale des Ophidiens. Tome Troisième (Livrais. 35 à 50), Contenant Quatre Vingt Seize Planches. Georges Jan and Ferdinand Sordelli, Milan, 96 pls.

    Johnsen, P. (1962) Notes on African snakes, mainly from northern Rhodesia and Liberia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjobenhavn, 124, 115–130.

    Jones, T. (1961) Notes on bat-eating snakes. The Nigerian Field, 26, 69–70.

    Jongsma, G.F.M., Barej, M.F., Barratt, C.D., Burger, M., Conradie, W., Ernst, R., Greenbaum, E., Hirschfeld, M., Leaché, A.D., Penner, J., Portik, D.M., Zassi-Boulou, A.-G., Rödel, M.-O. & Blackburn, D.C. (2018) Diversity and biogeography of frogs in the genus Amnirana (Anura: Ranidae) across sub-Saharan Africa. Molecular Phylogenetics and Evolution, 120, 274–285.

    https://doi.org/10.1016/j.ympev.2017.12.006

    Kennis, J., Nicolas, V., Hulselmans, J., Katuala, P.G.B., Wendelen, W., Verheyen, E., Dudu, A.M. & Leirs, H. (2011) The impact of the Congo River and its tributaries on the rodent genus Praomys: speciation origin or range expansion limit? Zoological Journal of the Linnean Society, 163 (3), 983–1002.

    https://doi.org/10.1111/j.1096-3642.2011.00733.x

    Kingdon, J. (1989) Island Africa: The Evolution of Africa’s Rare Animals and Plants. Princeton University Press, Princeton, New Jersey, 287 pp.

    Knoepffler, L.-P. (1966) Faune du Gabon (amphibiens et reptiles). I. Ophidiens de l’Ogooué-Ivindo et du Woleu N’tem. Biologia Gabonica, 2 (1), 3–23.

    Kroon, C. (1973) A second specimen of the colubrid snake Boiga philippina. Herpetologica, 29 (1), 51–53.

    https://doi.org/10.2307/1443124

    Laurent, R.F. (1954) Reptiles et batraciens de la région de Dundo (Angola). Deuxième note. Publicaçŏes Culturais da Companhia de Diamantes de Angola, 23, 35–84.

    Laurent, R.F. (1956) Contribution à l’herpétologie de la région des Grands Lacs de l’Afrique centrale. I. Généralités—II. Chéloniens—III. Ophidiens. Annales du Musée Royal du Congo Belge, Série in-8°, 48, 1–390, pls. I–XXXI.

    Laurent, R.F. (1960) Notes complémentaires sur les chéloniens et les ophidiens du Congo Oriental. Annales du Musée Royal du Congo Belge, Série in-8°, 84, 1–86.

    Laurent, R.F. (1964) Subsídos para o estudo da biologia na Lunda, Museo do Dundo. Reptiles et amphibiens de l’Angola (Troisième contribution). Publicaçŏes Culturais da Companhia de Diamantes de Angola, 67, 17–165.

    Lawson, D. (1993) The reptiles and amphibians of the Korup National Park Project, Cameroon. Herpetological Natural History, 1 (2), 27–90.

    Leaché, A.D. (2005) Results of a herpetological survey in Ghana and a new country record. Herpetological Review, 36 (1), 16–19.

    Leaché, A.D., Grummer, J.A., Miller, M., Krishnan, S., Fujita, M.K., Böhme, W., Schmitz, A., Lebreton, M., Ineich, I., Chirio, L., Ofori-Boateng, C., Eniang, E.A., Greenbaum, E., Rödel, M.-O. & Wagner, P. (2017) Bayesian inference of species diffusion in the West African Agama agama species group (Reptilia, Agamidae). Systematics and Biodiversity, 15 (3), 192–203.

    https://doi.org/10.1080/14772000.2016.1238018

    Leaché, A.D., Oaks, J.R., Ofori-Boateng, C. & Fujita, M.K. (2020) Comparative phylogeography of West African amphibians and reptiles. Evolution, 74 (4), 716–724.

    https://doi.org/10.1111/evo.13941

    Leeson, F. (1950) Identification of Snakes of the Gold Coast. The Crown Agents for the Colonies, London, x + 142 pp.

    Leston, D. (1970) Some snakes from the forest zone of Ghana. British Journal of Herpetology, 4 (6), 141–144.

    Leston, D. & Hughes, B. (1968) The snakes of Tafo, a forest cocoa-farm locality in Ghana. Bulletin de l’Institut Française d’Afrique Noire, 30A, 737–770.

    Levinson, S.R., Evans, M.H. & Groves, F. (1976) A neurotoxic component of the venom from Blanding’s tree snake (Boiga blandingi). Toxicon, 14 (4), 307–312.

    https://doi.org/10.1016/0041-0101(76)90027-1

    Loveridge, A. (1937) Zoological results of the George Vanderbilt African expedition of 1934. Part VII—Reptiles and amphibians. Proceedings of the Academy of Natural Sciences of Philadelphia, 89, 265–296.

    Loveridge, A. (1938) On a collection of reptiles and amphibians from Liberia. Proceedings of the New England Zoölogical Club, 17, 49–74.

    Loveridge, A. (1941) Report on the Smithsonian-Firestone expedition’s collection of reptiles and amphibians from Liberia. Proceedings of the United States National Museum, 91 (3128), 113–140.

    https://doi.org/10.5479/si.00963801.91-3128.113

    Loveridge, A. (1942) Scientific results of a fourth expedition to forested areas in east and central Africa. IV. Reptiles. Bulletin of the Museum of Comparative Zoology, Harvard, 91 (4), 237–373, pls. 1–6.

    Loveridge, A. (1955) On snakes collected in the Anglo-Egyptian Sudan by J.S. Owen, Esq. Sudan Notes and Records, 36 (1), 37–56.

    Loveridge, A. (1957) Check list of the reptiles and amphibians of East Africa (Uganda; Kenya, Tanganyika; Zanzibar). Bulletin of the Museum of Comparative Zoology, Harvard, 117 (2), 151–362 + i–xxxvi (index).

    Luiselli, L., Akani, G.C. & Barieenee, I.F. (1998a) Observations on habitat, reproduction and feeding of Boiga blandingi (Colubridae) in south-eastern Nigeria. Amphibia-Reptilia, 19 (4), 430–436.

    https://doi.org/10.1163/156853898X00098

    Luiselli, L., Akani, G.C. & Capizzi, D. (1998b) Food resource partitioning of a community of snakes in a swamp rainforest of south-eastern Nigeria. Journal of Zoology, London, 246 (2), 125–133.

    https://doi.org/10.1111/j.1469-7998.1998.tb00141.x

    Luiselli, L., Amori, G., Akani, G.C. & Eniang, E.A. (2015) Ecological diversity, community structure and conservation of Niger Delta mammals. Biodiversity and Conservation, 24, 2809–2830.

    https://doi.org/10.1007/s10531-015-0975-8

    Malnate, E.V. (1971) A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proceedings of the Academy of Natural Sciences of Philadelphia, 123, 345–375.

    Malnate, E.V. (1972) Observations on the vertebral hypapophyses and associated musculature in some snakes, with special reference to the Colubridae. Zoologische Mededelingen, 47 (18), 225–239.

    Marques, M.P., Ceríaco, L.M.P., Blackburn, D.C. & Bauer, A.M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola: Atlas of historical and bibliographic records (1840–2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement II), 1–501.

    Meier, J. & White, J. (Eds.) (1995) Handbook of Clinical Toxicology of Animal Venoms and Poisons. CRC Press, Boca Raton, New York, London and Tokyo, 752 pp.

    Meirte, D. (1992) Clés de détermination des serpents d’Afrique. Muséum Royal d’Afrique Centrale, Tervuren Belgique Annual Series Octavo Science Zoologique, 267, 1–152.

    Menegon, M., Loader, S.P., Marsden, S.J., Branch, W.R., Davenport, T.R.B. & Ursenbacher, S. (2014) The genus Atheris (Serpentes: Viperidae) in East Africa: Phylogeny and the role of rifting and climate in shaping the current pattern of species diversity. Molecular Phylogenetics and Evolution, 79, 12–22.

    https://doi.org/10.1016/j.ympev.2014.06.007

    Menzies, J.I. (1961) Correspondence: bat-eating snakes. The Nigerian Field, 26, 44.

    Menzies, J.I. (1966) The snakes of Sierra Leone. Copeia, 1966 (2), 169–179.

    https://doi.org/10.2307/1441123

    Mertens, R. (1938) Herpetologische Ergebnisse einer Reise nach Kamerun. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 442, 1–52.

    Mertens, R. (1941) Zur Kenntnis der Herpetofauna von Fernando-Poo. 3. Beitrag zu den wissenschaftlichen Ergebnissen der Forschungreise H. Eidmann nach Spanisch-Guinea 1939/40. Zoologischer Anzeiger, 135, 275–281.

    Mertens, R. (1964) Die Reptilien von Fernando Poo. Bonner zoologische Beiträge, 15 (3/4), 211–238.

    Mocquard, F. (1887) Sur les ophidiens repportés du Congo par la Mission de Brazza. Bulletin de la Société Philomathique de Paris, Série 7, 11, 62–92.

    Mocquard, F. (1896) Notes sur quelques ophidiens de Conakry. Bulletin du Muséum Paris, 20, 59–60.

    Monard, A. (1951) Résultats de la mission zoologique suisse au Cameroun. Reptiles. Batraciens. Mémoires de l’Institut Français d’Afrique Noire, Sciences Naturelles, (1), 123–170.

    Morley, R.J. & Kingdon, J. (2013) Africa’s environmental and climatic past. In: Kingdon, J., Happold, D., Hoffmann, M., Butynski, T., Happold, M. & Kalina, J. (Eds.), Mammals of Africa. Volume I. Introductory Chapters and Afrotheria. Bloomsbury, London, New Delhi, New York and Sydney, pp. 43–56.

    Müller, F. (1885) Vierter nachtrag zum Katalog der herpetologischen Sammlung der Basler Museums. Verhandlungen der Naturforschenden Gesellschaft in Basel (1887–1890), 7, 668–717.

    https://doi.org/10.5962/bhl.part.2388

    Nagy, Z.T., Chifundera Kusamba, Z., Gembu Tungaluna, G.-C., Lotana Lokasola, A., Kolby, J. & Kielgast, J. (2011) Foraging acrobatics of Toxicodryas blandingii in the Democratic Republic of the Congo. Herpetology Notes, 4, 91–92.

    Onadeko, A.B. & Rödel, M.-O. (2009) Anuran surveys in south-western Nigeria. Salamandra, 45 (1), 1–14.

    Parker, H.W. (1936) Dr. Karl Jordan’s expedition to South-West Africa and Angola: Herpetological collection. Novitates Zoologicae, 40, 115–146.

    Parker, H.W., Moreau, R.E. & Pakenham, R.H.W. (1940) XXV.—Notes on the herpetology of the East African Islands. Journal of Natural History, 5 (27), 309–314.

    https://doi.org/10.1080/00222934008527048

    Pasqual, J.R.H. (1962) Correspondence: West African snakes. The Nigerian Field, 27, 189–190.

    Pauwels, O.S.G., Albert, J.-L., Arrowood, H., Mvele, C., Casanova, M., Dodane, J.-B., Morgan, J., Primault, L., Thepenier, L. & Fenner, J.N. (2017a) Miscellanea Herpetologica Gabonica X. Bulletin of the Chicago Herpetological Society, 52 (8), 133–138.

    Pauwels, O.S.G., Biyogho Bi Essono, T, Carlino, P., Chirio, L., Huijbregts, B., Leuteritz, T.E.J., Rousseaux, D., Tobi, E., Vigna, C. & Van Neer, W. (2017b) Miscellanea Herpetologica Gabonica VII. Bulletin of the Chicago Herpetological Society, 52 (1), 1–7.

    Pauwels, O.S.G., Carlino, P., Chirio, L., Daversa, D.R., Lips, J., Oslisly, R. & Testa, O. (2019a) Amphibians and reptiles found in caves in Gabon, western Equatorial Africa. Cave and Karst Science, 46 (1), 3–12.

    Pauwels, O.S.G., Carlino, P., Chirio, L., Meunier, Q., Vivien Okouyi Okouyi, J., Orbell, C., Rousseaux, D. & Testa, O. (2017c) Miscellanea Herpetologica Gabonica IX. Bulletin of the Chicago Herpetological Society, 52 (6), 97–102.

    Pauwels, O.S.G., Kamdem Toham, A. & Chimsunchart, C. (2002a) Recherches sur l’herpétofaune du Massif du Chaillu, Gabon. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Biologie, 72, 47–57.

    Pauwels, O.S.G., Kamdem Toham, A. & Chimsunchart, C. (2002b) Recherches sur l’herpétofaune des Monts de Cristal, Gabon. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Biologie, 72, 59–66.

    Pauwels, O.S.G., Morelle, S., Albert, J.-L., Carlino, P., Rahola, N. & Trape, J.-F. (2019b) New reptile records from Lékédi Park and Haut-Ogooué Province, southeastern Gabon. Amphibian & Reptile Conservation, 13 (1), 143–161.

    Pauwels, O.S.G., Pauly, A., Araldi, A., Ndonda Makemba, R., Fonteyn, D., Oslisly, R. & Whittaker, A.M. (2020) Miscellanea Herpetologica Gabonica XVI. Bulletin of the Chicago Herpetological Society, 55 (5), 93–99.

    Pauwels, O.S.G. & Vogel, G. (2011). Ein Fall von Thanatose (Totstellreflex) bei Boiga blandingii (Squamata, Serpentes, Colubridae) in Gabun. Elaphe, 4, 24–25.

    Pel, H.S. (1852) Over de jagt aan de Goudkust, volgens eene tienjarige eigene ondervinding. Nederlandsch Tijdschrift voor Liefhebbers der Jagt und Visscherei, 1, 149–173.

    Penner, J., Adum, G.B., McElroy, M.T., Doherty-Bone, T., Hirschfeld, M., Sandberger, L., Weldon, C., Cunningham, A.A., Ohst, T., Wombwell, E., Portik, D.M., Reid, D., Hillers, A., Ofori-Boateng, C., Oduro, W., Plötner, J., Ohler, A., Leaché, A. & Rödel, M.-O. (2013) West Africa—a safe haven for frogs? A sub-continental assessment of the chytrid fungus (Batrachochytrium dendrobatidis). PLoS ONE, 8 (2), e56236.

    https://doi.org/10.1371/journal.pone.0056236

    Penner, J., Wegmann, M., Hillers, A., Schmidt, M. & Rödel, M.-O. (2011) A hotspot revisited–a biogeographical analysis of West African amphibians. Diversity and Distributions, 17, 1077–1088.

    https://doi.org/10.1111/j.1472-4642.2011.00801.x

    Perret, J.-L. (1961) études herpétologiques africaines III. Bulletin de la Société de Sciences Naturelles de Neuchâtel, 84, 133–138.

    Peters, W.C.H. (1877) übersicht der Amphibien aus Chinchoxo (Westafrika), welche von der Afrikanischen Gesellschaft dem Berliner zoologischen Museum übergeben sind. Monatsberichte der Königlichen Preussischen Akademie der Wissenschaften zu Berlin, 1877, 611–621.

    Pitman, C.R.S. (1938) A guide to the snakes of Uganda. Part XI. (Conclusion). The Uganda Journal, 5, 160–244, pls. XV–XVIII.

    Pitman, C.R.S. (1958) Snake and lizard predators of birds. Bulletin of the British Ornithologists’ Club, 78, 82–86.

    Pitman, C.R.S. (1962) More snake and lizard predators of birds. Bulletin of the British Ornithologists’ Club, 82, 45–55.

    Pitman, C.R.S. (1974) A Guide to the Snakes of Uganda, Revised Edition. Wheldon & Wesley, Ltd., Codicote, xxii + 290 pp.

    Portik, D.M., Leaché, A.D., Rivera, D., Barej, M.F., Burger, M., Hirschfeld, M., Rödel, M.-O., Blackburn, D.C. & Fujita, M.K. (2017) Evaluating mechanisms of diversification in a Guineo-Congolian tropical forest frog using demographic model selection. Molecular Ecology, 26 (19), 5245–5263.

    https://doi.org/10.1111/mec.14266

    Portillo, F., Branch, W.R., Conradie, W., Rödel, M.-O., Penner, J., Barej, M.F., Kusamba, C., Muninga, W.M., Aristote, M.M., Bauer, A.M., Trape, J.-F., Nagy, Z.T., Carlino, P., Pauwels, O.S.G., Menegon, M., Burger, M., Mazuch, T., Jackson, K., Hughes, D.F., Behangana, M., Zassi-Boulou, A.-G. & Greenbaum, E. (2018) Phylogeny and biogeography of the African burrowing snake subfamily Aparallactinae (Squamata: Lamprophiidae). Molecular Phylogenetics and Evolution, 127, 288–303.

    https://doi.org/10.1016/j.ympev.2018.03.019

    Portillo, F., Branch, W.R., Tilbury, C.R., Nagy, Z.T., Hughes, D.F., Kusamba, C., Muninga, W.M., Aristote, M.M., Behangana, M. & Greenbaum, E. (2019a) A cryptic new species of Polemon (Squamata: Lamprophiidae, Aparallactinae) from the miombo woodlands of Central and East Africa. Copeia, 107 (1), 22–35.

    https://doi.org/10.1643/CH-18-098

    Portillo, F., Stanley, E.L., Branch, W.R., Conradie, W., Rödel, M-O., Penner, J., Barej, M.F., Kusamba, C., Muninga, W.M., Aristote, M.M., Bauer, A.M., Trape, J-F., Nagy, Z.T., Carlino, P., Pauwels, O.S.G., Menegon, M., Ineich, I., Burger, M., Zassi-Boulou, A-G., Mazuch, T., Jackson, K., Hughes, D.F., Behanga, M. & Greenbaum, E. (2019b) Evolutionary history of burrowing asps (Squamata: Lamprophiidae: Atractaspidinae) with emphasis on fang evolution and prey selection. PLoS ONE, 14 (5), e0214889.

    https://doi.org/10.1371/journal.pone.0214889

    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

    Rasmussen, J.B. (1979) An intergeneric analysis of some boigine snakes—Bogert’s (1940) Group XIII and XIV (Boiginae, Serpentes). Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjøbenhavn, 141, 97–155.

    Rasmussen, J.B. (1997a) Afrikanske slanger (10). Boiga blandingii. Nordisk Herpetologisk Forening, 40 (3), 97–103.

    Rasmussen, J.B. (1997b) Afrikanske slanger (11). Boiga pulverulenta. Nordisk Herpetologisk Forening, 40 (4), 106–110.

    Rödel, M.-O., Emmrich, M., Penner, J., Schmitz, A. & Barej, M.F. (2014) The taxonomic status of two West African Leptopelis species: L. macrotis Schiøtz, 1967 and L. spiritusnoctis Rödel, 2007 (Amphibia: Anura: Arthroleptidae). Zoosystematics and Evolution, 90 (1), 21–32.

    https://doi.org/10.3897/zse.90.7120

    Rödel, M.-O. & Glos, J. (2019) Herpetological surveys in two proposed protected areas in Liberia, West Africa. Zoosystematics and Evolution, 95(1), 15–35.

    https://doi.org/10.3897/zse.95.31726

    Rödel, M.-O., Kouadio, K. & Mahsberg, D. (1999) Die Schlangenfauna des Comoé-Nationalparks, Elfenbeinküste: Ergänzungen und Ausblick. Salamandra, 35 (3), 165–180.

    Rödel, M.-O. & Mahsberg, D. (2000). Vorläufige Liste der Schlangen des Tai-Nationalparks/Elfenbeinküste und angrenzender Gebiete. Salamandra, 36 (1), 25–38.

    Roux-Estève, R. (1963) Les serpents de la région de la Maboké-Boukoko. Cahiers de la Maboke, 3 (1), 51–92.

    Roux-Estève, R. (1969) Les serpents de la région de Lamto (Côte-d’Ivoire). Annales de l’Université d’Abidjan, écologie, 2 (1), 81–140.

    Sabaj, M.H. (2016) Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 6.5. 16 August 2016. Electronically accessible. American Society of Ichthyologists and Herpetologists, Washington, D.C. Available from: http://www.asih.org/ (accessed 7 April 2021)

    Savage, J.M. & McDiarmid, R.W. (2017) The Herpetological Contributions of Giorgio Jan (1791–1866): With an Introduction, Annotated Bibliography, Synopsis of Herpetological Taxa, and a Comprehensive Guide to the Iconographie générale des Ophidiens. Society for the Study of Amphibians and Reptiles, Villanova, Pennsylvania, viii + 926 pp.

    Schmidt, K.P. (1923) Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909–1915. Part II.—Snakes. Bulletin of the American Museum of Natural History, 49, 1–146, pls. I–XXII.

    Segniagbeto, G.H., Trape, J.F., David, P., Ohler, A., Dubois, A. & Glitho, I.A. (2011) The snake fauna of Togo: Systematics, distribution and biogeography, with remarks on selected taxonomic problems. Zoosystema, 33 (3), 325–360.

    https://doi.org/10.5252/z2011n3a4

    Shirley, M.H., Vliet, K.A., Carr, A.N. & Austin, J.D. (2014) Rigorous approaches to species delimitation have significant implications for African crocodilian systematics and conservation. Proceedings of the Royal Society B, 281, 20132483.

    https://doi.org/10.1098/rspb.2013.2483

    Skinner, H.A. (1973) Snakes and Us: An Introduction to East African Herpetology. East African Literature Bureau, Nairobi, Kampala and Dar es Salaam, xvi + 146 pp.

    Smith, H.M., Smith, R.B. & Sawin, H.L. (1977) A summary of snake classification (Reptilia, Serpentes). Journal of Herpetology, 11 (2), 115–121.

    https://doi.org/10.2307/1563131

    Spawls, S. (1979) Sun, Sand & Snakes. William Morrow and Company, Inc., New York, New York, 254 pp.

    Spawls, S. & Branch, B. (1995) The Dangerous Snakes of Africa: Natural History, Species Directory, Venoms and Snakebite. Ralph Curtis—Books, Sanibel Island, Florida, 192 pp.

    Spawls, S. & Branch, B. (2020) The Dangerous Snakes of Africa. Bloomsbury, London, Oxford, New York, New Delhi and Sydney, 336 pp.

    Spawls, S., Howell, K., Hinkel, H. & Menegon, M. (2018) Field Guide to East African Reptiles, 2nd Edition. Bloomsbury, London, New Delhi, New York and Sydney, 624 pp.

    Sternfeld, R. (1909) Die Fauna der deutschen Kolonien, Herausgegeben mit Unterstützung des Reichs-Kolonialamtes vom Zoologischen Museum in Berlin. Reihe I. Kamerun. Heft I. Die Schlangen Kameruns. R. Friedländer & Sohn, Berlin, iv + 28 pp.

    Stucki-Stirn, M.C. (1979) A Comparative Study of the Herpetological Fauna of the former West Comeroon [sic]/Africa: With a Classification and Synopsis of 95 Different Snakes and Description of Some New Sub-species. Snake Report 721. Herpeto-Verlag, Teuffenthal, Switzerland, 650 pp.

    Swiecicki, A.W. (1965) Snakes and snake bite in the western region, Ghana. Journal of Tropical Medicine and Hygiene, 68 (12), 300–304.

    Takemoto, H., Kawamoto, Y. & Furuichi, T. (2015) How did bonobos come to range south of the Congo River? Reconsideration of the divergence of Pan paniscus from other Pan populations. Evolutionary Anthropology, 24 (5), 170–184.

    https://doi.org/10.1002/evan.21456

    Taub, A.M. (1967) Comparative histological studies on Duvernoy’s gland of colubrid snakes. Bulletin of the American Museum of Natural History, 138 (1), 1–50.

    Taylor, E.H. & Weyer, D. (1958) Report on a collection of amphibians and reptiles from Harbel, Republic of Liberia. The University of Kansas Science Bulletin, 38 (14), 1191–1229.

    https://doi.org/10.5962/bhl.part.10973

    Thys van den Audenaerde, D.F.E. (1965) Les serpents des environs de Léopoldville. Revue de zoologie et de botanique africaines, 72 (3–4), 366–388.

    Trape, J.-F. (1985) Les serpents de la région de Dimonika (Mayombe, République Populaire du Congo). Revue de Zoologie Africaine, 99, 135–140.

    Trape, J.-F. & Mané, Y. (2006) Guide des Serpents d’Afrique Occidentale: Savane et Désert. IRD éditions, Paris, 226 pp.

    https://doi.org/10.4000/books.irdeditions.37382

    Uetz, P., Cherikh, S., Shea, G., Ineich, I., Campbell, P.D., Doronin, I.V., Rosado, J., Wynn, A., Tighe, K.A., McDiarmid, R., Lee, J.L., Köhler, G., Ellis, R., Doughty, P., Raxworthy, C.J., Scheinberg, L., Resetar, A., Sabaj, M., Schneider, G., Franzen, M., Glaw, F., Böhme, W., Schweiger, S., Gemel, R., Couper, P., Amey, A., Dondorp, E., Ofer, G., Meiri, S. & Wallach, V. (2019) A global catalog of primary reptile type specimens. Zootaxa, 4695 (5), 438–450.

    https://doi.org/10.11646/zootaxa.4695.5.2

    Uetz, P., Freed, P. & Hoŝek, J. (Eds.) (2020) The Reptile Database. Available from: http://www.reptile-database.org (accessed 12 December 2020)

    Ullenbruch, K. & Böhme, W. (2017) Six new records of Afrotropical lizard and snake species (Reptilia: Squamata) from the Republic of South Sudan. Bonn zoological Bulletin, 66 (2), 139–144.

    Underwood, G. (1967) A Contribution to the Classification of Snakes. British Museum (Natural History), London, x + 179 pp.

    Villiers, A. (1950a) Initiations Africaines. II. Les Serpents de l’Ouest Africain. Institut Français d’Afrique Noire, Dakar, Senegal, 148 pp.

    Villiers, A. (1950b) Catalogues. IV. La Collection de Serpents de l’I.F.A.N. Institut Français d’Afrique Noire, Dakar, Senegal, 155 pp.

    Villiers, A. (1951) Mission A. Villiers au Togo et au Dahomey (1950). II.—Ophidiens. études Dahoméennes, 5, 17–43.

    Villiers, A. (1955) Note sur quelques Ixodidae et Gamasidae parasites des Vertébrés rencontrés en Afrique Occidentale française. Bulletin de l’Institut fondamental d’Afrique noire, Série A, 17 (2), 444–454.

    Villiers, A. (1956) La collection de serpents de l’I.F.A.N. (Acquisitions 1954–1955). Bulletin de l’Institut fondamental d’Afrique noire, Série A, 18 (3), 878–883.

    Voelker, G., Marks, B.D., Kahindo, C., A’genonga, U., Bapeamoni, F., Duffie, L.E., Huntley, J.W., Mulotwa, E., Rosenbaum, S.A. & Light, J.E. (2013) River barriers and cryptic biodiversity in an evolutionary museum. Ecology and Evolution, 3 (3), 536–545.

    https://doi.org/10.1002/ece3.482

    Wakeman, B.N. (1966) Uganda’s poisonous snakes: Further observations on feeding habits. The Uganda Journal, 30, 101–103

    Wallach, V. (1998) The lungs of snakes. In: Gans, C. & Gaunt, A.S. (Eds.), Biology of the Reptilia. Vol. 19. Morphology G (Visceral Organs). Society for the Study of Amphibians and Reptiles, Ithaca, New York, pp. 93–295.

    Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World: A Catalogue of Living and Extinct Species. CRC Press, Boca Raton, Florida, xxvii + 1209 pp.

    Weinell, J.L., Barley, A.J., Siler, C.D., Orlov, N.L., Ananjeva, N.B., Oaks, J.R., Burbrink, F.T. & Brown, R.M. (2021) Phylogenetic relationships and biogeographic range evolution in cat-eyed snakes, Boiga (Serpentes: Colubridae). Zoological Journal of the Linnean Society. [in press]

    https://doi.org/10.1093/zoolinnean/zlaa090

    Weinstein, S.A. & Kardong, K.V. (1994) Properties of Duvernoy’s secretions from opisthoglyphous and aglyphous colubrid snakes. Toxicon, 32 (10), 1161–1185.

    https://doi.org/10.1016/0041-0101(94)90347-6

    Weinstein, S.A. & Smith, L.A. (1993) Chromatographic profiles and properties of Duvernoy’s secretions from some boigine and dispholidine colubrids. Herpetologica, 49 (1), 78–94.

    Weinstein, S.A., Warrell, D.A., White, J. & Keyler, D.E. (2011) “Venomous” Bites from Non-Venomous Snakes: A Critical Analysis of Risk and Management of “Colubrid” Snake Bites. Elsevier, Amsterdam, Boston, Heidelburg, London, New York, Oxford, Paris, San Diego, San Francisco, Singapore, Sydney and Tokyo, xxvii + 336 pp.

    Welch, K.R.G. (1982) Herpetology of Africa: A Checklist and Bibliography of the Orders Amphisbaenia, Sauria and Serpentes. Robert E. Krieger Publishing Company, Malabar, Florida, 293 pp.

    Werner, F. (1897) Ueber Reptilien und Batrachier aus Togoland, Kamerun und Tunis aus dem kgl. Museum für Naturkunde in Berlin. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 47, 395–408.

    https://doi.org/10.5962/bhl.part.11464

    Werner, F. (1899) Ueber Reptilien und Batrachier aus Togoland, Kamerun und Deutsch-Neu-Guinea, grösstentheils aus dem kgl. Museum für Naturkunde in Berlin. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 49, 132–157.

    https://doi.org/10.5962/bhl.part.24106

    Wickler, V.W. & Uhrig, D. (1969) Verhalten und ökologische Nische der Gelbflügelfledermaus, Lavia frons (Geoffroy) (Chiroptera, Megadermatidae). Zeitschrift für Tierpsychologie, 26 (6), 726–736.

    https://doi.org/10.1111/j.1439-0310.1969.tb01972.x

    Wiens, J.J. & Penkrot, T.A. (2002) Delimiting species using DNA and morphological variation and discordant species limits in spiny lizards (Sceloporus). Systematic Biology, 51 (1), 69–91.

    https://doi.org/10.1080/106351502753475880

    Woodward, S.F. (1960) A bat-eating snake. The Nigerian Field, 25, 172–174.

    Wüster, W., Chirio, L., Trape, J-F., Ineich, I., Jackson, K., Greenbaum, E., Kusamba, C., Barron, C., Nagy, Z., Storey, R., Hall, C., Wüster, C., Barlow, A. & Broadley, D.G. (2018) Integration of nuclear and mitochondrial gene sequences and morphology reveal unexpected diversity in the forest cobra (Naja melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae). Zootaxa, 4455 (1), 68–98.

    https://doi.org/10.11646/zootaxa.4455.1.3

    Young, R.A. (1983) Feeding behavior of captive Boiga. Bulletin of the Chicago Herpetological Society, 18 (3–4), 89–91.

    Zaher, H., Murphy, R.W., Arredondo, J.C., Graboski, R., Machado-Filho, P.R., Mahlow, K., Montingelli, G.G., Quadros, A.B., Orlov, N.L., Wilkinson, M., Zhang, Y.-P. & Grazziotin, F.G. (2019) Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes). PLoS ONE, 14 (5), e0216148.

    https://doi.org/10.1371/journal.pone.0216148

    Zimkus, B.M., Bala, V., Belasen, A.M., Bell, R.C., Channing, A., Doumbia, J., Fokam, E.B., Gonwouo, L.N., Greenbaum, E., Gvoždík, V., Hirschfeld, M., Jackson, K., James, T.Y., Kusamba, C., Larson, J.G., Mavoungou, L.-B., Rödel, M.-O., Zassi-Boulou, A.-G. & Penner, J. (2020) Chytrid pathogen (Batrachochytrium dendrobatidis) in African amphibians: A continental analysis of occurrences and modeling of its potential distribution. Herpetologica, 76 (2), 201–215.

    https://doi.org/10.1655/0018-0831-76.2.201