Abstract
The scincid taxa connected to the genus Panaspis sensu lato have undergone various taxonomic changes based mainly on morphological studies in the last century, but their true relationships are still far from clear. In the present study we applied Bayesian, maximum parsimony, and maximum likelihood methods of phylogenetic inference to mitochondrial 12S rRNA and 16S rRNA gene fragments to examine the degree of genetic variation within the genus. We found considerable genetic differentiation between the different proposed subgenera. Similarly, we could show that two species considered to be part of the former subgenus Leptosiaphos are in reality genuine members of the genus Lacertaspis. Based on our results we propose the recognition of the four former subgenera as full genera, and we identify several additional, possibly subgeneric, lineages within the different newly recognized full genera.References
Angel, F. (1978) Zur Herpetofaunistik des Senegal. Bonner zoologische Beiträge, 29, 360–417.
Angel, F. (1940) Deux nouvelles espèces de Lygosoma du Caméroun [sic]. Matériaux de la mission P. Lepesme, R. Paulian et A. Villiers (1re note). Bulletin du Muséum d’Histoire naturelle, Paris, 12 (2ème sér.), 82–85.
Angel, F. (1944) Un lézard nouveau du Mont Nimba (Haute-Guinée Francaise) appartenant au genre Lygosoma (matériaux de la mission Lamotte au Mont Nimba). Bulletin du Muséum d’Histoire naturelle, Paris, 5 (2èmesér.), 293–294.
Angel, F., Guibé, J. & Lamotte, M. (1954) La Réserve Naturelle Intégrale du Mont Nimba. Fasciule II (Extrait): Lézards. Mémoire d’Institut francais Afrique Noire, 40, 371–379.
Barnett, L.K., Emms, C. & Santoni, C. (2001) The herpetofauna of Abuko Nature Reserve, The Gambia. Herpetological Bulletin, 77, 5–14.
Bauer, A.M., Shea, G. & Günther, R. (2003) An annoted catalogue of the types of scincid lizards (Reptilia, Squamata, Scincidae) in the collection of the Museum für Naturkunde der Humboldt-Universität zu Berlin (ZMB). Mitteilungen des Museums für Naturkunde, Berlin, Zoologische Reihe, 79 (2), 253–321.
Bocage, J.V.B. (1866) Reptiles nouveaux ou peu connus recueillis dans les possessions portugaises de l'Afrique occidentale, que se trouvent au Muséum de Lisbonne. Jornal de Sciencias Mathematicas, Physicas e Naturaes Lisboa, 1, 57–78.
Böhme, W. (1975) Zur Herpetofaunistik Kameruns, mit Beschreibung eines neuen Scinciden. Bonner zoologische Beiträge, 26, 2–48.
Böhme, W. & Nikolaus, G. (1989) Herpetological specimens from the Gotel Mountains and Mambilla Plateau, Nigeria. Tauraco Research Report, 1, 28–30.
Böhme, W. & Schmitz, A. (1996) A new lygosomine skink (Lacertilia: Scincidae: Panaspis) from Cameroon. Revue suisse de Zoologie, 103, 767–774.
Böhme, W., Schmitz, A. & Ziegler, T. (2000) A review of the West African skink genus Cophoscincopus Mertens (Reptilia: Scincidae: Lygosominae): Resurrection of C. simulans (Vaillant, 1884) and description of a new species. Revue suisse de Zoologie, 107, 777–791.
Boulenger, G.A. (1887) Catalogue of the lizards in the British Museum (Natural History). Second edition. Vol. III. Taylor and Francis, London, 575 pp.
Boulenger, G.A. (1900) A list of the Batrachians and Reptiles of the Gaboon (French Congo), with the description of new genera and species. Proceedings of the zoological Society, London, 1900, 433–456.
Boulenger, G.A. (1907) Descriptions of new lizards in the British Museum. Annals and Magazine of Natural History, 19, 486–489.
Boulenger, G.A. (1909) List of reptiles [and batrachians] collected by Capt. G. Ferrari at Jumbo, lower Juba [Bardera]. Annali del Museo civico di storia naturale Giacomo Doria, 44, 308–309 [310–311].
Boulenger, G.A. (1913) Descriptions of new lizards in the collection of the British Museum. Annals and Magazine of Natural History, 12, 563–566.
Brandley, M.C., Schmitz, A. & Reeder, T.W. (2005) Partitioned Bayesian Analyses, Partition Choice, and the Phylogenetic Relationships of Scincid lizards. Systematic Biology, in press.
Broadley, D.G. (1988) A checklist of the reptiles of Zimbabwe, with synoptic keys. Arnoldia Zimbabwe, 9, 369–430.
Broadley, D.G. (1989) A reappraisal of the genus Panaspis Cope, with the description of a new species of Leptosiaphos (Reptilia: Scincidae) from Tanzania. Arnoldia Zimbabwe, 9, 439–449.
Broadley, D.G. (1991) The herpetofauna of northern Mwinilunga District, Northwestern Zambia. Arnoldia Zimbabwe, 9, 519–538.
Broadley, D.G. & Blake, D.K. (1979) A check list of the reptiles of the national parks and other conservation areas of Zimbabwe Rhodesia. Arnoldia Rhodesia, 35, 1–15.
Broadley, D.G. & Howell, K.M. (1991) A check list of the reptiles of Tanzania, with synoptic keys. Syntarsus, 1, 1–70.
Chabanaud, P. (1917) Énumération des Reptiles non encore étudiés de l'Afrique occidentale, appartenant aux Collections du Muséum, avec la description des espèces nouvelles. Bulletin du Muséum d’Histoire naturelle, Paris, 23, 83–105.
Cope, E.D. (1862) On Lacerta echinata and Tiliqua dura. Proceedings of the Academy of Natural Sciences Philadelphia, 1862, 189–191.
Cope, E.D. (1868) Observations on reptiles of the Old World. Art. II. Proceedings of the Acadamy of Natural Sciences Philadelphia, 1868, 316–323.
Cope, E.D. (1970) Contribution à la systématique des lygosominés africains (Reptilia, Scincidae). I. Les espèces attibuées au genre Ablepharus. Revue Roumaine de Biologie, Zoologie, 15, 379–393.
Cope, E.D. (1972). Révision du phylum forestier du genre Panaspis Cope (Reptilia, Scincidae, Lygosominae). Revue Roumaine de Biologie, Zoologie, 17, 257–271.
Daniels, S.R., Heideman, N., Hendricks, M. & Willson, B. (2002) A molecular phylogeny for the South African limbless lizard taxa of the subfamily Acontinae (Sauria: Scincidae) with spezial emphasis on relationships within Acontias. Molecular Phylogenetics and Evolution, 24, 315–323.
De Witte, G.-F. (1933) Reptiles récoltés au Conge Belge par le Dr. H. Schouteden et par M. G.-F. Witte. Annales du Musée du Congo belge, Zoologie, 1, 53–100.
Duméril, A.M.C. & Bibron, G. (1839) Erpétologie générale ou Histoire naturelle complète des Reptiles. Lib. encycl. Roret, Paris.
Eniang, E.A. & Luiselli, L. (2002) Ikpan wetland rainforest: an area of high biodiversity importance in south-eastern Nigeria. Revue Écologie (Terre Vie), 57, 19–28.
Frank, N. & Ramus, E. (1996) A Complete Guide to the Scientific and Common Names of Reptiles and Amphibians of the World. N G Publishing Inc., Pottsville, 377 pp.
Fuhn, I.E. (1969) Revision and redefinition of the genus Ablepharus Lichtenstein, 1823 (Reptilia, Scincidae). Revue Roumaine de Biologie, Zoologie, 14, 23–41.
Grandison, A.G.C. (1956) On a collection of lizards from West Africa. Bulletin d’Institut francais Afrique Noire, Dakar, 18 (sér A), 224–245.
Gray, J.E. (1839) Catalogue of the slender-tongued saurians, with descriptions of many new genera and species. Annals and Magazine of Natural History, 1939, 287–337.
Gray, J.E. (1845) Catalogue of the specimens of lizards in the collection of the British Museum. Trustees of die British Museum/Edward Newman, London.
Greer, A.E. (1974) The generic relationships of the scincid lizard genus Leiolopisma and its relatives. Australian Journal of Zoology, Supplement Series, 31, 1–67.
Greer, J.E. (1977). The systematics and evolutionary relationships of the scincid lizard genus Lygosoma. Journal of natural History, 11, 515–540.
Haft, J. (1993) Ein Beitrag zur Biologie der Echsen der Insel São Tomé (Golf von Guinea), mit näherer Betrachtung zur Systematik von Leptosiaphos africana (GRAY). Faunistische Abhandlungen des Museums für Tierkunde Dresden, 19, 59–70.
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.
Hardwicke, T. & Gray, J.E. (1827) A synopsis of the species of saurian reptiles, collected in India by Major-General Hardwicke. Zoological Journal, London, 1827.
Huelsenbeck, J.P. & Crandall, K.A. (1997) Phylogeny estimation and hypothesis testing using maximum likelihood. Annual Review of Ecology and Systematics, 28, 437–466.
Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback. J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294, 2310–2314.
Illies, J. (1968) Die Gattung als ökologische Grundeinheit. Faunistisch-Ökologische Mitteilungen, 3, 369–372.
Ineich, I., Schmitz, A., Chirio, L. & LeBreton, M. (2004) New species of Leptosiaphos (Scincidae) from Adamaoua Massif, Central-Northern Province, Cameroon. Journal of Herpetology, 38 (4), 546–550.
Jacobsen, N.H.G. & Broadley, D.G. (2000) New species of Panaspis Cope (Reptilia: Scincidae) from southern Africa. African Journal of Herpetology, 49, 61–71.
Joger, U. (1990) The herpetofauna of the Central African Republic, with description of a new species of Rhinotyphlops (Serpentes: Typhlopidae). In: Peters, G. & Hutterer, R. (Eds.) Vertebrates in the tropics. Museum Alexander Koenig, Bonn, 1990, pp. 85–102.
Jukes, T.H. & Cantor, C.R. (1969) Evolution of protein molecules. In: Munro, H. (Ed) Mammalian Protein Metabolism. Academic Press, New York, U.S.A., pp. 21–123.
Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A. C. (1989) Dynamics of mitchondrial DNA evolution in animals: Amplification and sequencing with conserved primers. Proceedings of the National Acadamy of Sciences, USA, 86, 6196–6200.
Larget, B. & Simon, D.L. (1999) Markov chain Monte Carlo algorithms for the Bayesian analysis of phylogenetic trees. Molecular Biology and Evolution, 16, 750–759.
Laurent, R. F. (1982) Deux lézards intéressants de la cuvette centrale zairoise. Revue zoologique africain, 96, 439–444.
LeBreton, M. (1999) A working checklist of the herpetofauna of Cameroon. With localities for species occurring in southern Cameroon and a list of herpetofauna for the Dja Faunal Reserve. IUCN, Netherlands, 160 pp.
Leviton, A.E., Gibbs jr., R.H., Heal, E. & Dawson, C.E. (1985) Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia, 1985 (3), 802–832.
Lichtenstein, H. (1823) Verzeichniss der Doubletten des zoologischen Museums der Königl. Universität zu Berlin. Museum Berlin, Berlin.
Loveridge, A. (1952) Mission A. Villiers au Togo et au Dahomey (1950), XII. Tortoises and Lizards. Bulletin d’Institut fondamental d’Afrique noire, 15, 229–242.
Mau, B., Newton, M. & Larget, B. (1999) Bayesian phylogenetic inference via Markov chain Monte carlo methods. Biometrics, 55, 1–12.
Mausfeld, P. & Schmitz, A. (2003) Molecular phylogeography, intraspecific variation and speciation of the Asian scincid lizard genus Eutropis Fitzinger, 1843, (Squamata: Reptilia: Scincidae): taxonomic and biogeographic implications. Organisms, Diversity and Evolution, 3, 161–171.
Mausfeld, P., Schmitz, A., Böhme, W., Misof, B., Vrcibradic, D. & Rocha, C.F.D. (2002) Phylogenetic affinities of Mabuya atlantica Schmidt, 1945, endemic to the Atlantic Ocean archipelago of Fernando de Noronha (Brazil): Necessity of partitioning the genus Mabuya Fitzinger, 1826 (Scincidae: Lygosominae). Zoologischer Anzeiger, 241, 281–293.
Mausfeld, 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 (1/2), 159–177.
Mertens, R. (1964) Die Reptilien von Fernando Poo. Bonner zoologische Beiträge, 15, 211–238.
Mertens, R. (1968). Zur Kenntnis der Herpetofauna von Kamerun und Fernando Poo. Bonner zoologische Beiträge, 19, 69–84.
Misof, B., Rickert, A.M., Buckley, T.R., Fleck, G. & Sauer, K.P. (2001) Phylogenetic signal and its decay in mitochondrial SSU and LSU rRNA gene fragments of dragonflies. Molecular Biology and Evolution, 18 (1), 27–37.
Mittleman, M.B. (1952) A generic synopsis of the lizards of the subfamily Lygosominae. Smithsonian miscellaneous Collections, 117, 1–35.
Monard, A. (1949) Vertébrés nouveaux du Cameroun. Revue suisse de Zoologie, 56, 731–745.
Müller, L. (1910) Beiträge zur Herpetologie Kameruns. Abhandlungen der mathematischen-physikalischen Klasse der Königlichen-Bayrischen Akadademie der Wissenschaften, München, 24, 544–626.
Nieden, F. (1913) Neues Verzeichnis der Kriechtiere (ausser den Schlangen) von Deutsch-Ostafrika. I Teil, Reptilia. Mitteilungen des Zoologischen Museums Berlin, 7, 51–100.
Nylander, J.A.A. (2002) MrModeltest 1.1b, Uppsala, Sweden.
Palumbi, S.R., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool's guide to PCR. Department of Zoology and Kewalo Marine Laboratory, Hawai, 47 pp.
Perret, J.-L. (1973) Contribution à l'étude des Panaspis (Reptilia, Scincidae) d'Afrique occidentale avec la description de deux espèces nouvelles. Revue suisse de Zoologie, 80, 595–630.
Perret, J.-L. (1975a) La différenciation dans le genre Panaspis Cope (Reptilia, Scincidae). Bulletin de la Societé neuchâteloise Sciences Naturelles, 98, 5–16.
Perret, J.-L. (1975b) Revision critique de quelques types de reptiles et batraciens africains. Revue suisse de Zoology, 82, 185–192.
Perret, J.-L. (1982) Le sous-genre Leptosiaphos (Lacertilia, Scincidae) et ses implications. Bulletin de la Societé neuchâteloise Sciences Naturelles, 105, 107–121.
Perret, J.-L. & Wuest, J. (1982) La structure fine des écailles de quelques Lygosominae africains (Lacertilia, Scincidae), révélée par le microscope électronique à balayage. Revue suisse de Zoologie, 89, 269–280.
Perret, J.-L. & Wuest, J. (1983) La microstructure des écailles de quelques Scincidés africains et paléarctiques (Lacertilia), observée au microscope électronique à balayage, II. Revue suisse de Zoologie, 90, 913–928.
Peters, W. (1873) Über eine neue Schildkrötenart, Cinosternon effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der königlichen Akademie der Wissenschaften Berlin, 1873, 603–618.
Peters, W. (1874) Über neue Reptilien (Peropus, Agama, Euprepes, Lygosoma, Typhlops, Heterolepis) der herpetologischen Sammlung des Berliner zoologischen Museums. Monatsberichte der königlichen Akademie der Wissenschaften Berlin, 1874, 159–164.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Rannala, B. & Yang, Z.H. (1996) Probability distribution of molecular evolutionary trees: A new method of phylogenetic inference. Journal of Molecular Evolution, 43, 304–311.
Reeder, T.W. (2003) A phylogeny of the Australian Sphenomorphus group (Scincidae: Squamata) and the phylogenetic placement of the crocodile skinks (Tribolonotus): Bayesian approaches to assessing congruence and obtaining confidence in maximum likelihood inferred relationships. Molecular Phylogenetics and Evolution, 27, 384–397.
Rödel, M.-O., Grabow, K., Hallermann, J. & Böckheler, C. (1997) Die Echsen des Comoé-Nationalparks, Elfenbeinküste. Salamandra, 33, 225–240.
Schmidt, K.P. (1943) Amphibians and reptiles from the Sudan. Publications of the Field Museum of natural History, Zoological Series, 24, 331–338.
Schmitz, A. (2001) Der Tchabal Mbabo — unerforschtes Juwel an der nördlichen Grenze der Kamerunbergkette. Mitteilungsblatt AG Chamäleons, 23, 6–8.
Schmitz, A., Brandley, M.C., Mausfeld, P., Vences, M., Glaw, F., Nussbaum, R.A. & Reeder, T.W. (2005) [2004] Opening the black box: A molecular phylogeny of the Malagasy fossorial lizards of the subfamily Scincinae. Molecular Phylogenetics and Evolution, 34, 118–133.
Schmitz, A., Mausfeld, P. & Embert, D. (2004) Molecular studies on the genus Eumeces Wiegmann, 1834: phylogenetic relationships and taxonomic implications. Hamadryad, 28 (1/2), 73–89.
Sjöstedt, Y. (1897) Zwei neue Eidechsen aus West-Afrika. Zoologischer Anzeiger, 20, 56.
Smith, A. (1849) Illustrations of the zoology of South Africa, Reptilia. Smith, Elder & Co., London (1847–1849).
Smith, M.A. (1935) The Fauna of British India Including Ceylon and Burma. Reptilia and Amphibia Volume II.-Sauria. Taylor and Francis, London.
Smith, M.A. (1937) A review of the genus Lygosoma (Scincidae: Reptilia) and its allies. Records of the Indian Museum, 39, 213–234.
Spawls, S., Howell, K.M., Drewes, R.C. & Ashe, J. (2002) A field guide to the reptiles of East Africa. Academic Press, London, 543 pp.
Stejneger, L. (1891) Description of a new scincoid lizard from East Africa. Proceedings of the United States Natural Museum, XIV, 405–406.
Sternfeld, R. (1912) IV. Zoologie II. Lfg. Reptilia. In: Wissenschaftliche Ergebnisse der Deutschen Zentral-Afrika Expedition 1907–1908, unter Führung A. Friedrichs, Herzogs zu Mecklenburg. R. Schubotz. (ed.). Klinkhard und Biermann, Leipzig.
Sternfeld, R. (1918) Zur Tiergeographie Papuasiens und der pazifischen Inselwelt. Abhandlungen senckenbergianischen Naturforschenden Gesellschaft, 36, 375–436.
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts, USA.
Swofford, D.L., Olsen, G.J., Waddell, P.J. & Hillis, D.M. (1996) Phylogenetic inference. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.) Molecular Systematics. 2nd edition. Sinauer Associates, Sunderland, Massachusetts, U.S.A., pp. 407–543.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin F. & Higgins, D.G. (1997) The ClustalX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 24, 4876–4882.
Welch, K.G. (1982) Herpetology of Africa: a checklist and bibliography of the orders Amphisbaenia, Sauria and Serpentes. Robert E. Krieger Publishing Company Malaber, Florida.
Werner, F. (1902) Über westafrikanische Reptilien. Verhandlungen der Zoologischen Botanischen Gesellschaft Wien, 52, 332–348.
Werner, F. (1909) Beschreibung neuer Reptilien aus dem Kgl. Naturalienkabinett in Stuttgart. Jahreshefte des Vereins für vaterländische Naturkunde Württemberg, 65, 55–63.
Werner, F. (1919) Wissenschaftliche Ergebnisse der mit Unterstützung der Kaiserlichen Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommenen zoologischen Expedition nach dem Anglo-Aegyptischen Sudan (Kordofan) 1914. IV. Bearbeitung der Fische, Amphibien und Reptilien. Denkschriften der Akademie der Wissenschaften Wien, Mathematisch-Naturwissenschaftliche Klasse, 96, 1–73.
Wiegmann, A.F.A. (1834) Herpetologia Mexicana, seu Descriptio Amphibiorum Novae Hispaniae, quae Itineribus Comitis de Sack, Ferdinandi Deppe et Chr. Guil. Schiede in Museum Zoologicum Berolinense Pervenerunt. C. G. Lüderitz, Berlin.
Wilcox, T.P., Zwickl, D.J., Heath, T.A. & Hillis, D.M. (2002) Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25, 361–371.
Wilgenbusch, J. & de Queiroz, K. (2000) Phylogenetic relationships among the phrynosomatid sand lizards inferred from mitochondrial DNA sequences generated by heterogeneous evolutionary processes. Systematic Biology, 49, 592–612.
Yang, Z.H., Goldman, N. & Friday, A. (1994) Comparison of Models for Nucleotide Substitution Used in Maximum-Likelihood Phylogenetic Estimation. Molecular Biology and Evolution, 11, 316–324.