Open Access Open Access  Restricted Access Subscription or Fee Access

The “Cobra-preta” of São Tomé Island, Gulf of Guinea, is a new species of
Naja Laurenti, 1768 (Squamata: Elapidae)



The Cobra-Preta (black snake in Portuguese) of Sao Tomé Island in the Gulf of Guinea has historically been referred to as Naja (Boulengerina) melanoleuca (Squamata: Elapidae). Its presence on the island has been traditionally explained as an introduction from the mainland by Portuguese settlers, supposedly to control the rat population. This explanation has been widely accepted by local authorities and even international conservation agencies. The taxonomic identity of this snake has remained undisputed by all taxonomists who have published about it, with the exception of L. Capocaccia in 1961. Arguments supporting the human introduction hypothesis are weak and are contradicted by historical, morphological and molecular data. Further, the biogeographic history of the Gulf of Guinea oceanic islands and recent insights on the taxonomic identity and evolutionary history of other taxonomic groups occurring there suggest that the Cobra-Preta, in fact, represents a distinct lineage of the melanoleuca group, endemic to São Tomé. We here describe the Cobra Preta as a new species. The new species differs from N. (B.) melanoleuca, its sister species, by a distinct coloration ventral pattern and the type of contact of the sublingual scales. Data on the toxicology, distribution, ecology, folklore and conservation status of the new species are presented.



Reptilia, Naja, Boulengerina, taxonomy, São Tomé & Príncipe, conservation, Naja (Boulengerina) peroescobari sp. nov.

Full Text:



Albuquerque, L. (1989) Introdução à História dos Descobrimentos Portugueses. Europa-América, Lisboa, 294 pp.

Anonymous (2008) Bulletin ECOFAC/RAPAC São Tomé et Príncipe. Compte rendu du mois de Août 2008. Available from: (accessed 4 July 2016)

Bedriaga, J. (1893) Note sur les Amphibiens et Reptiles recueillis par M. Adolphe Moller aux îles de la Guinée. O Instituto, 40, 299–303, 432–440.

Bell, R.C., Drewes, R.C., Channing, A., Gvozdik, V., Kielgast, J., Lotters, S., Stuart, B.L. & Zamudio, K.R. (2015) Overseas dispersal of Hyperolius reed frogs from Central Africa to the oceanic islands of São Tomé and Príncipe. Journal of Biogeography, 42 (1), 65–75.

Bell, R.C. (2016) A new species of Hyperolius (Amphibia: Hyperoliidae) from Príncipe Island, Democratic Republic of São Tomé and Príncipe. Herpetologica, 72 (4), 343–351.

Bersacola, E., Svensson, M.S., Bearder, S.K. & Mills, M. (2014) Hunted in Angola. Surveying the Bushmeat Trade. Swara, 38, 31–36.

Bocage, J.V.B. (1866) Lista dos reptis das possessões portuguesas d’Africa ocidental que existem no Museu de Lisboa. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 1 (1), 37–56.

Bocage, J.V.B. (1879) Subsidios para a Fauna das possessões portuguezas d’Africa occidental. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 7 (26), 85–96.

Bocage, J.V.B. (1886) Reptis e Amphibios de S. Thomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 11 (42), 65–70.

Bocage, J.V.B. (1889) Breves considerações sobre a fauna de S. Thomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 1 (1), 33–36.

Bocage, J.V.B. (1895) A doninha de São Tomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 4 (13), 24–27.

Bocage, J.V.B. (1905) Contribution à la Faune des quatre îles du golfe de Guinée (suite). Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 7, 65–96.

Boulenger, G.A. (1906) Report on the reptiles collected by the late L. Fea in West Africa. Annali del Museo Civico di Storia Naturale di Genova, 3 (2), 196–216.

Broadley, D.G. (1968) A review of the African cobras of the genus Naja (Serpentes: Elapinae). Arnoldia (Rhodesia), 3, 1–14.

Broadley, D.G. (1983) FitzSimons’ Snakes of Southern Africa (Revised Edition). Delta Books, Johannesburg, 376 pp.

Broadley, D.G. (1990) FitzSimons’ Snakes of Southern Africa (Revised Edition). Jonathan Ball & Ad. Donker Publishers, Parklands, 387 pp., 84 color pls., Addendum.

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

Broadley, D.G. & Blaylock, R. (2013) The Snakes of Zimbabwe and Botswana. Edition Chimaira, Frankfurt am Main, 387 pp.

Broadley, D.G. & Wüster, W. (2004) A review of the Southern African ‘non-spitting’ cobras (Serpentes: Elapidae: Naja). African Journal of Herpetology, 53 (2), 101–122.

Capocaccia, L. (1961) Contributo allo studio dei serpenti delle isole del Golfo di Guinea. Annali del Museo Civico di Storia Naturale “Giacomo Doria, 72, 285–309.

Carleton, M.D. & Olson, S.L. (1999) Amerigo Vespucci and the rat of Fernando de Noronha: a new genus and species of Rodentia (Muridae, Sigmodontinae) from a volcanic Island off Brazil’s continental shelf. American Museum Novitates, 3256, 1–58.

Ceríaco, L.M.P. (2012) Human attitudes towards herpetofauna: the influence of folklore and negative values on the conservation of amphibians and reptiles in Portugal. Journal of Ethnobiology and Ethnomedicine, 8, 8.

Ceríaco, L.M.P. (2015) Lost in the middle of the sea, found in the back of the shelf: A new giant species of Trachylepis (Squamata: Scincidae) from Tinhosa Grande islet, Gulf of Guinea. Zootaxa, 3973 (3), 511–527.

Ceríaco, L.M.P., Marques, M., Jacquet, F., Nicolas, V., Colyn, M., Denys, C., Sardinha, P. & Bastos-Silveira, C. (2015) Descritption of Crocidura fingui, a new endemic species of shrew (Mammalia, Soricomorpha) from Príncipe Island (Gulf of Guinea). Mammalia, 79 (3), 325–341.

Ceríaco, L.M.P., Marques, M.P. & Bauer, A.M. (2016) A review of the genus Trachylepis (Sauria: Scincidae) from the Gulf of Guinea, with the descriptions of two new species in the Trachylepis maculilabris (Gray, 1845) species complex. Zootaxa, 4109 (3), 284–314.

Chippaux, J.-P., Williamns, V. & White, J. (1991) Snake venom variability: Methods of study, results and interpretation. Toxicon, 29 (11), 1279–1303.

Chippaux, J.-P. (2006) Les serpents d’Afrique occidentale et centrale. Institut de Recherche pour le Développement (IRD), Paris, 311 pp., 16 pls.

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

Chirio, L. & Lebreton, M. (2007) Atlas des reptiles du Cameroun. Muséum national d’Histoire naturelle/ Institut de Recherche pour le Développement (IRD), Paris, 688 pp.

Correia, J.G. (1928) Unpublished field notebook, housed in the Ornithology. Department of the American Museum of Natural History, New York.

Dallimer, M., King, T. & Atkinson, R.J. (2009) Pervasive threats within a protected area: conserving the endemic birds of São Tomé, West Africa. Animal Conservation, 12, 209–219.

Felsenstein, J. (1985) Phylogenies and the Comparative Method. The American Naturalist, 125 (1), 1–15.

Ferreira, J.B. (1902) Lista dos Reptis e Batrachios da colecção do Sr. Newton. Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 6, 281–284.

FitzSimons, V. (1962) Snakes of Southern Africa. Purnell & Sons, Cape Town, 423 pp.

Fritz, U., Branch, W.R., Hofmeyr, M.D., Maran, J., Prokop, H., Schleicher, A., Siroky, P., Stuckas, H., Vargas-Ramírez, M., Vences, M. & Hundsdorfer, A. (2011) Molecular phylogeny of African hinged and helmeted terrapins (Testudines: Pelomedusidae: Pelusios and Pelomedusa). Zoologia Scripta, 40, 115–125.

Greef, R. (1884) Ueber die Fauna der Guinea-Inseln S. Thomé und Rolas. Sitzungsberichte der Gesellschaft zur Beforderung der gesammten Naturwissenschaften zu Marburg, 1884 (2), 41–80.

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

Henriques, J. (1917) A ilha de S. Tomé sob o ponto de vista histórico-natural e agrícola. Boletim da Sociedade Broteriana, 27, 1–197.

Hofer, D. (2002) The São Tomé and Principe Handbook. D. Hofer Verlag, Bern, 152 pp.

Huelsenback, J.P. & Hillis, D.M. (1993) Success of phylogenetic methods in the four-taxon case. Systematic Biology, 42, 247–264.

Jesus, J., Nagy, Z., Branch, W.B., Wink, M., Brehm, A. & Harris, D. J. (2009) Phylogenetic relationships of African green snakes (genera Philothamnus and Hapsidophrys) from São Tomé, Príncipe and Annobon islands based on mtDNA sequences and comments on their colonization and taxonomy. The Herpetological Journal, 19, 41–48.

Kaiser, H. (2014) Best practices in herpetological taxonomy: errata and addenda. Herpetological Review, 45, 257–268.

Kaiser, H., Crother, B.I., Kelly, C.M.R., Luiselli, L., O’Shea, M., Ota, H., Passos, P., Schleip, W.D. & Wüster, W. (2013) Best practices: In the 21st Century, taxonomic decisions in herpetology are acceptable only when supported by a body of evidence and published via peer-review. Herpetological Review, 44, 8–23.

Kelly, C.M.R., Barker, N.P., Villet, M.H. & Broadley, D.G. (2009) Phylogeny, biogeography and classification of the snake superfamily Elapoidea: a rapid radiation in the late Eocene. Cladistics, 25, 38–63.

Laurent, R. (1955) Diagnoses préliminaires de quelques Serpents venimeux. Revue de Zoologie et Botanique Africaine, 51, 127–139.

Laurent, R. (1956) Contribution à l’Herpétologie de la Région des Grands Lacs de l’Afrique centrale. Annales du Musée Royal du Congo Belge, Série in 8°, Sciences Zoologiques, 48, 1–390, 31 pls.

Lee, D.C., Halliday, A.N., Fitton, J.G. & Poli, G. (1994) Isotopic variations with distance and time in the volcanic islands of the Cameroon line: evidence for a mantle plume origin. Earth and Planetary Science Letters, 123, 119–138.

Leventis, A.P. & Olmos, F. (2009) As aves de São Tomé e Príncipe. Um Guia Fotográfico / The Birds of São Tomé and Príncipe. A Photoguide. Editora Aves e Fotos, São Paulo, 142 pp.

Machado, A.M., Sousa, A.F., Marques, R.M. & Pereira, S.M. (2013) Guia Turístico de São Tomé e Príncipe. Pocket Tropics, Évora, 288 pp.

Manaças, S. (1958) Anfíbios e Répteis das ilhas de São Tomé e do Príncipe e do Ilhéo das Rolas. In: Anonymous. Conferência Internacional dos Africanistas Ocidentais, Vol IV, Junta Investigação do Ultramar, Lisboa, pp. 179–192.

Manaças, S. (1973) Alguns dos anfíbios e répteis da província de S. Tomé e Príncipe. In: Anonymous. Livro de Homenagem ao Prof. Fernando Frade. Junta Investigação do Ultramar, Lisboa, pp. 219–230.

Matos, R.J.C. (1916) Corografia história das Ilhas de São Tomé, Príncipe, Ano Bom e Fernando Pó. Imprensa Nacional, Lisboa, 120 pp., 20 pls.

Measey, J. (2013) Taxonomic publishing, vandalism and best practice: African Journal of Herpetology makes changes that will safeguard authors. African Herp News, 60, 2–4.

Measey, G.J., Vences, M., Drewes, R.C., Chiari, Y., Melo, M. & Bourles, B. (2007) Freshwater paths across the ocean: molecular phylogeny of the frog Ptychadena newtoni gives insights into amphibian colonization of oceanic islands. Journal of Biogeography, 34, 7–20.

Melo, M., Warren, B.H. & Jones, P.J. (2011) Rapid parallel evolution of aberrant traits in the diversification of the Gulf of Guinea white-eyes (Aves, Zosteropidae). Molecular Ecology, 20, 4953–4967.

Meirte, D. (1992) Clés de determination des serpents d’Afrique. Annales Sciences Zoologiques, 267, 1–152.

Meshaka W.E., Loftus, W.F. & Steiner, T. (2000) The herpetofauna of Everglades National Park. Florida Scientist, 63, 84–103.

Miller, E.C., Sellas, A.B. & Drewes, R.C. (2012) A new species of Hemidactylus (Squamata: Gekkonidae) from Príncipe Island, Gulf of Guinea, West Africa with comments on the African–Atlantic clade of Hemidactylus geckos. African Journal of Herpetology, 61 (1), 40–57.

Munson, P. & Ronghui, Z. (2012) Feeding the Dragon: Managing Chinese resources acquisition in Africa. Seattle Journal of Environmetal Law, 2 (1), 343–382.

Nagy, Z.T., Vidal, N., Vences, M., Branch, W.R., Pauwels, O.S.G., Wink, M. & Joger, U. (2005) Molecular systematics of African Colubroidea (Squamata: Serpentes). In: Huber, B.A., Sinclair, B.J. & Lampe, K.H. (Eds.), African biodiversity: molecules, organisms, ecosystems. Springer Verlag, Bonn, pp. 221–228.

Nill, T. (1993) Die Schlangen der Insel São Tomé (Golf von Guinea) (Reptilia: Serpentes). Faunistische Abhandlungen Staatliches Museum für Tierkunde Dresden, 19 (7), 71–73.

Padial, J., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16.

Pauwels, O.S.G. & Vande Weghe, J.P. (2008) Reptiles du Gabon. Smithsonian Institution, Washington D.C., 272 pp.

Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14 (9), 817–818.

Rodda, G.H. & Savidge, J.A. (2007) Biology and Impacts of Pacific Island Invasive Species. 2. Boiga irregularis, the Brown Tree Snake (Reptilia: Colubridae). Pacific Science, 61 (3), 307–324.[307:BAIOPI]2.0.CO;2

Rodrigues, M., Bos, A.R., Schembri, P.J., Lima, R.F., Lymberakis, P., Parpal, L., Cento, M., Ruette, S., Ozkurt, S.O., Santos-Reis, M., Merila, J. & Fernandes, C. (2017) Origin and introduction history of the least weasel (Mustela nivalis) on Mediterranean and Atlantic islands inferred from genetic data. Biological invasions, 19 (1), 399–421.

Schätti, B. & Loumont, C. (1992) Ein Beitrag zur Herpetofauna von São Tomé (Golf von Guinea) (Amphibia et Reptilia). Zoologische Abhandlungen des Staatliches Museum für Tierkunde Dresden, 47 (1), 23–36.

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.

Silva-Rocha, I., Salvi, D., Sillero, N., Mateo, J.A. & Carretero, M.A. (2015) Snakes on the Balearic Islands: An Invasion Tale with Implications for Native Biodiversity Conservation. PLoS ONE, 10 (4), e0121026.

Silvestro, D. & Michalak, I. (2012) RaxmlGUL: a graphical front-end for RAxML. Organismal Diversity and Evolution, 12, 335–337.

Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H.-M. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701.

Slowinski, J.B. & Keogh, S. (2000) Phylogenetic relationships of elapid snakes based on cytochrome b mtDNA sequences. Molecular Phylogenetics and Evolution, 15, 157–164.

Stamatakis, A. (2006) RaxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.

Stucki-Stirn, M.C. (1979) Snake Report 721: A Comparative Study of the Herpelogical Fauna of the Former West Cameroon/Africa, with a Classification and Synopsis of 95 Different Snakes and Descriptions of Some New Sub-species. Herpeto-Verlag, Teuffenthal, 650 pp.

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.

Themido, A.A. (1941) Répteis e batráquios das colónias portuguesas. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 119, 1–31.

Trape, J.-F. & Mané, Y. (2006) Guide des Serpentes d’Afrique Occidentale. IRD Éditions, Paris, 226 pp.

Trape, J.-F., Chirio, L., Broadley, D.G. & Wüster, W. (2009) Phylogeography and systematic revision of the Egyptian cobra (Serpentes: Elapidae Naja haje) species complex, with the description of a new species from West Africa. Zootaxa, 2236, 1–25.

Underwood, G. (1979) Classification and distribution of venomous snakes in the world. In: Lee, C.Y. (Ed.), Handbook of Experimental Pharmacology. Springer-Verlag, Berlin, pp. 41–60.

Uyeda, J.C., Drewes, R.C. & Zimkus, B.M. (2007) The California Academy of Sciences Gulf of Guinea Expeditions (2001, 2006) VI. A new species of Phrynobatrachus from the Gulf of Guinea Islands and reanalysis of Phrynobatrachus dispar and P. feae (Anura: Phrynobatrachidae). Proceedings of the California Academy of Sciences, 58, 367–385.

Vieira, L. (1886) Noticia ácerca dos productos zoologicos trazidos da ilha de S. Thomé para o Muzeu Zoologico da Universidade de Coimbra pelo Sr. Adolpho F. Moller, em 1885. O Instituto, 34, 235–241.

Villiers, A. (1950a) La collection de serpents de L’I.F.A.N. Institut Français d’Afrique Noir, Dakar, 155 pp.

Villiers, A. (1950b) Les serpents de l’Ouest Africain. Institut Français d’Afrique Noir, Dakar, 148 pp.

Villiers, A. (1965) Les serpents de l’Ouest Africain. 2nd Edition. Institut Français d’Afrique Noir, Dakar, 190 pp.

Wallach, V., Wüster, W. & Broadley, D.G. (2009) In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae). Zootaxa, 2236, 26–36.

Wüster, W. & Broadley, D.G. (2003) A new species of spitting cobra (Naja) from north-eastern Africa (Serpentes: Elapidae). Journal of Zoology, 259, 345–359.

Wüster, W. & Broadley, D.G. (2007) Get an eyeful of this: a new species of giant spitting cobra from eastern and north-eastern Africa (Squamata: Serpentes: Elapidae: Naja). Zootaxa, 1532, 51–68.

Wüster, W., Crookes, S., Ineich, I., Mané, Y., Pook, C.E., Trape, J.-F. & Broadley, D.G. (2007) The phylogeny of cobras inferred from mitochondrial DNA sequences: Evolution of venom spitting and the phylogeoggraphy of the African spitting cobras (Serpentes: Elapidae: Naja nigricollis complex). Molecular Phylogenetics and Evolution, 45, 437–453.

Zheng, H. & Zhang, Q. (2000) Medicinal using status and conservation strategy of snakes in China. In: Zhang, E. & Zheng, H. (Eds.), Conservation of endangered wild medicinal fauna and Xora resources in China. Second Military Medical University Press, Shanghai, pp. 96–101.

Zhou, Z. & Jiang, Z. (2005) Identifying snake species threatened by economic exploitation and international trade in China. Biodiversity and Conservation, 14, 3525–3536.



  • There are currently no refbacks.

ISSN 1175-5326 (Print Edition) & ISSN 1175-5334 (Online Edition)
Published by Magnolia Press, Auckland, New Zealand