Abstract
The scincid genus Trachylepis is represented in the oceanic islands of the Gulf of Guinea by four species, Trachylepis maculilabris, T. affinis, T. adamastor and T. ozorii. Here we describe two new species, Trachylepis thomensis sp. nov., endemic to São Tomé Island and Rolas Islet, and Trachylepis principensis sp. nov., endemic to Príncipe Island. Phylogenetic analysis using the mitochondrial gene 16S shows that both new species are genetically divergent and reciprocally monophyletic, and confirms evidence for the uniqueness of these lineages presented in previous studies. Morphological data (scalation and morphometry) identify consistent phenotypic differences between these two island species. We were also able to confirm that the T. affinis population of Príncipe Island is conspecific with the African mainland population and most probably the result of recent introductions. These findings raise the number of known Trachylepis species in the Gulf of Guinea islands group to five, four of which are endemic, although the phylogenetic and phylogeographic relationships of T. adamastor and T. ozorii remain unknown and require further investigation.
References
Albuquerque, L. (1989) Introdução à História dos Descobrimentos Portugueses. Europa–América, Lisboa, 294 pp.
Angel, M.F. (1920) Sur une collection de reptiles et de batraciens de l'Ile de San Thomé et de l'Ile du Prince et description d'une espèce nouvelle du genre Typhlops. Bulletin du Muséum national d'Histoire naturelle, 26, 197–199.
Bauer, A.M. (2003) On the identity of Lacerta punctata Linnaeus, 1758, the type species of the genus Euprepis Wagler, 1830, and the generic assignment of Afro–Malagasy skinks. African Journal of Herpetology, 52, 1–7.
http://dx.doi.org/10.1080/21564574.2003.9635472Bauer, A.M., Shea, G. & Gunther, R. (2003) An annotated catalogue of the type of scincid lizards (Reptilia, Squamata, Scincidae) in the collection of the Museum fur Naturkunde der Humboldt-Universitat zu Berlin (ZMB). Mitteilungen aus dem Museum fur Naturkunde in Berlin, 79, 253–321.
http://dx.doi.org/10.1002/mmnz.20030790204Bedriaga, J. (1892) Note sur les Amphibiens et Reptiles recueillis par M. Adolphe Moller aux îles de la Guinée. O Instituto, 39, 498–507; 642–648; 736–742; 814–820; 901–907.
Bedriaga, J. (1893a) Note sur les Amphibiens et Reptiles recueillis par M. Adolphe Moller aux îles de la Guinée. O Instituto, 40, 299–303; 432–440.
Bedriaga, J. (1893b) Remarques supplémentaires sur les Amphibiens et Reptiles du Portugal et de l'île de St. Thomé. O Instituto, 40, 766–773; 848–856; 927–936.
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. (1872) Diagnoses de quelques espèces nouvelles de Reptiles d’Afrique ocidental. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 4 (13), 72–82.
Bocage, J.V.B. (1886a) Reptis e Amphibios de S. Thomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 11 (42), 65–70.
Bocage, J.V.B. (1886b) Reptiles et Batraciens nouveaux de l’île de S. Thomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 11 (42), 71–75.
Bocage, J.V.B. (1886c) Note additionelle sur les reptiles de S. Thomé. Jornal das Sciencias Mathematicas, Physicas e Naturaes, 11 (42), 103–104.
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. (1893) Diagnoses de deux reptiles nouveaux de l'ile de Anno-bom. Jornal de Sciencias Mathematicas, Physicas e Naturaes. Segunda Série, 3 (9), 47–48.
Bocage, J.V.B. (1895) Herpétologie d’Angola et du Congo. Imprimerie Nationale, Lisbonne, 203 pp., 20 pls.
Bocage, J.V.B. (1897) Mammiferos, reptis e batrachios d’Africa de que existem exemplares typicos no Museu de Lisboa. Jornal de Sciencias Mathematicas, Physicas e Naturaes. Segunda Série, 4 (16), 187–206.
Bocage, J.V.B. (1903) Contribution à la Faune des quatre îles du golfe de Guinée. Jornal das Sciencias Mathematicas, Physicas e Naturaes, Segunda Série, 7, 25–59.
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. (1903) Descriptions of new lizards in the collection of the British Museum. Annals and Magazine of Natural History, series 7 12, 429–435.
http://dx.doi.org/10.1080/00222930308678877Boulenger, 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. (1974) A review of the Mabuya maculilabris group in South–Eastern Africa (Sauria: Scincidae). Arnoldia (Rhodesia), 6 (23), 1–10.
Broadley, D.G. (2000) A review of the genus Mabuya in southeastern Africa (Sauria: Scincidae). African Journal of Herpetology, 49 (2), 87–110.
http://dx.doi.org/10.1080/21564574.2000.9635437Brygoo, E.R. (1981) Systematique des lézards scincides de la region Malgache VIII. Les Mabuya des iles de l’Océan Indien ocidental: Comores, Europa, Séchelles. Bulletin du Museum National d'Histoire Naturelle, 3, 911–930.
Capocaccia, L. (1961) Catalogo dei tipi di rettili del Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale “Giacomo Doria”, 72, 86–111.
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.
http://dx.doi.org/10.11646/zootaxa.3973.3.6Ceríaco, L.M.P., Bauer, A.M., Blackburn, D.C. & Lavres, A.C.F.C. (2014) The Herpetofauna of the Capanda Dam Region, Malanje, Angola. Herpetological Review, 45(4), 667–674.
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.
http://dx.doi.org/10.1515/mammalia-2014-0056Chirio, L., Ineich, I., Schmitz, A. & LeBreton, M. (2008) A new species of Trachylepis Fitzinger, 1843 (Squamata: Scincidae) from Central African forests. African Journal of Herpetology, 57 (1), 13–28.
http://dx.doi.org/10.1080/21564574.2008.9635565de Queiroz, K. (1999) 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, USA, pp. 57–75.
Felsenstein, J. (1985) Phylogenies and the Comparative Method. The American Naturalist, 125 (1), 1–15.
http://dx.doi.org/10.1086/284325Fischer, J.G. (1886) Über zwei neue Eidechsen des Naturistorischen Museums zu Hamburg. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 3 ,1–15 + plate 1.
Gray, J.E. (1839 [1838]) Catalogue of the slender-tongued saurians, with descriptions of many new genera and species. Annals and Magazine of Natural History, 2 (1), 287–293.
Gray, J.E. (1845) Catalogue of the specimens of lizards in the collection of the British Museum. Trustees of the British Museum, London, 289 pp.
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.
Hallermann, J. (1998) Annotated catalogue 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.
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.
Hoogmoed, M.S. (1974) Ghanese lizards of the genus Mabuya (Scincidae, Sauria, Reptilia). Zoologische Verhandelingen, 138, 1–62.
Huelsenback, J.P. & Hillis, D.M. (1993) Success of phylogenetic methods in the four-taxon case. Systematic Biology, 42, 247–264.
http://dx.doi.org/10.2307/2992463Jesus, J., Brehm, A. & Harris, J. (2003) The herpetofauna of Annobon island, Gulf of Guinea, West Africa. Herpetological Bulletin, 86, 20–22.
Jesus, J., Harris, D.J. & Brehm, A. (2005a) Relationships of scincid lizards (Mabuya spp.) from the islands of the Gulf of Guinea based on mtDNA sequence data. Amphibia–Reptilia, 26, 467–473.
http://dx.doi.org/10.1163/156853805774806331Jesus, J., Harris, D.J. & Brehm, A. (2005b) Phylogeography of Mabuya maculilabris (Reptilia) from São Tomé Island (Gulf of Guinea) inferred from mtDNA sequences. Molecular Phylogenetics and Evolution, 37, 503–510.
http://dx.doi.org/10.1016/j.ympev.2005.04.032Kingdon, R. & Spawls, S. (2010) A new gregarious Trachylepis (Reptilia: Sauria: Scincidae) from Lolui Island, Lake Victoria, Uganda, with a key to Ugandan Trachylepis. African Journal of Herpetology, 59 (2), 123–132.
http://dx.doi.org/10.1080/21564574.2010.508220Lee, 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.
http://dx.doi.org/10.1016/0012-821X(94)90262-3Lima, A., Harris, D.J., Rocha, S., Miralles, A., Glaw, F. & Vences, M. (2013) Phylogenetic relationship of Trachylepis skink species from Madagascar and the Seychelles (Squamata: Scincidae). Molecular Phylogenetics and Evolution, 67 (3), 615–620.
http://dx.doi.org/10.1016/j.ympev.2013.02.001MacArthur, R.H. & Wilson, E.O. (1967) The Theory of Island Biogeography. Princeton University Press, Princeton, 203 pp.
Manaças, S. (1958) Anfíbios e Répteis das ilhas de São Tomé e do Príncipe e do Ilhéo das Rolas. In: 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: Livro de Homenagem ao Prof. Fernando Frade. Junta Investigação do Ultramar, Lisboa, pp. 219–230.
Marzoli, A., Picirillo, E.M., Renne, P.R., Bellieni, G., Iacumin, M., Nyobe, J.B. & Tongwa, A.T. (2000) The Cameroon Volcanic Line revisited: petrogenesis of continental basaltic magmas from lithosperic and asthenospheric mantle sources. Journal of Petrology, 41, 87–109.
http://dx.doi.org/10.1093/petrology/41.1.87Mausfeld, P., Vences, M., Schmitz, A. & Veith, M. (2000) First data on the molecular phylogeography of scincid lizards of the genus Mabuya. Molecular Phylogenetics and Evolution, 17 (1), 11–14.
http://dx.doi.org/10.1006/mpev.2000.0809Mausfeld–Lafdhiya, P., Schmitz, A., Ineich, I. & Chirio, L. (2004) Genetic variation in two African Euprepis Species (Reptilia, Scincidae), based on Maximum–Likelihood and Bayesian Analyses: Taxonomic and Biogeographic conclusions. Bonner zoologische Beiträge, 52, 159–177.
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.
http://dx.doi.org/10.1111/j.1365-2699.2006.01589.xMiller, 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.
http://dx.doi.org/10.1080/21564574.2012.666680Peters, W.C.H. (1854) Diagnosen neuer Batrachier, welche zusammen mit der friiher (24. Juli und 17. August) gegebenen Übersicht der Schlangen und Eidechsen mitgetheilt werden. Bericht uber die zur Bekanntmachung geeigneten Verhandlungen der koniglich-preussischen Akademie der Wissen- schaften zu Berlin, 1854, 614–628.
Peters, W.C.H. (1879) Neue oder weniger bekannte Eidechsenarten aus der Familie der Scinci (Eumeces Guntheri, Euprepes notabilis, Ablepharus rutilus). Sitzungs-Berichte der Gesellschaft naturforschender Freunde zu Berlin, 1879, 35–37.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14 (9), 817–818.
http://dx.doi.org/10.1093/bioinformatics/14.9.817Rocha, S., Carretero, M.A. & Harris, D.J. (2010) Genetic diversity and phylogenetic relationship of Mabuya spp. (Squamata: Scincidae) from western Indian Ocean islands. Amphibia–Reptilia, 31, 375–385.
http://dx.doi.org/10.1163/156853810791769473Schätti, B. & Loumont, C. (1992) Ein Beitrag zur Herpetofauna von São Tomé (Golf von Guinea) (Amphibia et Reptilia). Zoologische Abhandlungen Staatliches Museum für Tierkunde Dresden, 47 (1), 23–36.
Sindaco, R., Metallinou, M., Pupin, F., Fasola, M. & Carranza, S. (2012) Forgotten in the ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago. Zoologica Scripta, 41, 346–362.
http://dx.doi.org/10.1111/j.1463-6409.2012.00540.xSternfeld, R. (1913) Reptilia. In: Schubotz, H. (Ed.), Wissenschaftliche Ergebnisse der deutschen Zentral-Afrika-Expedition 1907–1908 unter Führung Adolf Friedrichs, Herzogs zu Mecklenberg. Band IV. Zoologie II. (1912) Klinkhardt & Biermann, Leipzig, pp. 197–279.
http://dx.doi.org/10.5962/bhl.title.7048
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.
http://dx.doi.org/10.1093/molbev/mst197Themido, 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., Chirio, L. & Trape, S. (2012) Lézards, crocodiles et tortues d'Afrique occidentale et du Sahara. IRD, Marseille, 503 pp.
Uetz, P. & Hošek, J. (2015) The Reptile Database. Available from: http://www.reptile–database.org (accessed 14 February 2015)
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.