Abstract
A new species of lacertid lizard of the genus Psammodromus is described from the Iberian Peninsula. Genetic and recentlypublished phenotypic data support the differentiation of Psammodromus hispanicus into three, and not as previously sug-gested two, distinct lineages. Age estimates, lineage allopatry, the lack of mitochondrial and nuclear haplotype sharingbetween lineages, ecological niche divergence, and the current biogeographic distribution, indicated that the three lineagescorrespond to three independent species. Here, we describe a new species, Psammodromus occidentalis sp. n., which isgenetically different from the other sand racers and differentiated by the number of femoral pores, number of throat scales,snout shape, head ratio, green nuptial coloration, and number of supralabial scales below the subocular scale. We also pro-pose to upgrade the two previously recognized subspecies, Psammodromus hispanicus hispanicus Fitzinger, 1826 fromcentral Spain and Psammodromus hispanicus edwardsianus (Dugès, 1829) from eastern Spain, to the species level: Psam-modromus hispanicus stat. nov. and Psammodromus edwardsianus stat. nov. Given that the holotype of Psammodromushispanicus was lost, we designate a neotype. We also analysed museum specimens of P. blanci, P. microdactylus and P. algirus to describe differentiation of the Psammodromus hispanicus lineages/species from their closest relatives.References
Akaike, H. (1973) Information theory as an extension of the maximum likelihood principle. In: Petrov B. N. & Csaki, F. (Eds.). Second international symposium of information theory. Akademiai Kiado, Budapest, Hungary.
Arevalo, E., Davis, S. K. & Sites, J. W. (1994) Mitochondrial-DNA Sequence Divergence and Phylogenetic-Relationships among 8 Chromosome Races of the Sceloporus-Grammicus Complex (Phrynosomatidae) in Central Mexico. Systematic Biology, 43, 387–418.
Arnold, E. N. (1989) Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. Bulletin of the British Museum of Natural History, 55, 209–257.
Böhme, W. (1981) Psammodromus algirus (Linnaeus 1766) - Algerischer Sandläufer, p. 479-491. In: Böhme W. (Ed.). Handbuch der Reptilien und Amphibien Europas. Akademische Verlagsgesellschaft, Wiesbaden. Vol. 1.
Bons, J. (1989) Psammodromus hispanicus. In: Bons, J., Castanet, J. & Guyetant, R. (Eds.). Atlas de repartition des amphibiens et reptiles de France. Société Herpétologique de France, Paris, pp. 144–145.
Boulenger, G. A. (1921) Psammodromus In: Boulenger, G. A. (Ed.) Monograph of the Lacertidae. Johnson Reprint Corporation, London, Vol. 2, pp. 163–179.
Carranza, S., Harris, D. J., Arnold, E. N., Batista, V. & De la Vega, J. P. G. (2006) Phylogeography of the lacertid lizard, Psammodromus algirus, in Iberia and across the Strait of Gibraltar. Journal of Biogeography, 33, 1279–1288.
Dugès, A. (1829) Mémoire sur les espèces indigènes du genre Lacerta. Annales des Sciences naturelles, 16, 337–339.
Duméril, A. M. C. & Bibron, G. (1839) Erpétologie Générale ou Histoire Naturelle Complète des Reptiles. Librairie Encyclopédique de Roret, Paris, 253.
Felsenstein, J. (1981) Evolutionary trees from DNA sequences: a maximum likelihood approach. Journal of Molecular Evolution, 17, 368–376.
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791.
Fitze, P. S. & Richner, H. (2002) Differential effects of a parasite on ornamental structures based on melanins and carotenoids. Behavioral Ecology 13, 401–407.
Fitze, P. S., Gonzalez-Jimena, V., San-Jose, L. M., San Mauro, D., Aragón, P., Suarez, T. & Zardoya, R. Drivers of Diversification in Psammodromus hispanicus (Squamata: Lacertidae). BMC Evolutionary Biology, 11, 347.
Fitzinger, L. I. (1826) Neue Classification der Reptilien nach ihren natürlichen Verwandtschaften. Heubner, J. G., Wien, 66 pp.
Huelsenbeck, J. P. & Ronquist, F. R. (2000) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
ICZN (2000) International code of Zoological Nomenclature, 4th edition, International Trust for Zoological Nomenclature.
Lessells, C. M. A. M. (1987) Unrepeatable repeatabilites: a common mistake. The Auk, 104, 116–121.
Mertens, R. (1925) Amphibien und Reptilien aus dem nördlichen und östlichen Spanien, Abhandlungen der Senckenbergishen Naturforschenden Gesellschaft, 39, 27–129.
Mertens, R. & Müller, L. (1928) Liste der Amphibien und Reptilien Europas. Abhandlungen der Senckenbergischen Naturforschenden Gelleschaft, 41, 1–62.
Pérez-Mellado, V. (1998) Psammodromus hispanicus Fitzinger, 1826. In: Reptiles. Salvador, A. (Coord.) Fauna Ibérica, Vol. 10. Ramos, M. A. et al., (Eds). Museo Nacional de Ciencias Naturales-CSIC, Madrid, pp. 318–326.
Pérez-Mellado, V. & Gosá, A. (1988) Biometría y folidosis en Lacertidae (Sauria, Reptilia), algunos aspectos metodológicos. Revista Española de Herpetología, 3, 103–119.
Posada, D. & Crandall, K. A. (1998): MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Ronquist, F. & Huelsenbeck, J. P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19,1572–1574.
Schleich, H. H., Kästle W. & Kabisch K. (1996). Amphibians and reptiles of North Africa. Koeltz, Königstein. 630 pp.
Stamatakis, A. (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
Stamatakis, A., Blagojevic, F., Nikolopoulos, D. & Antonopoulos, C. (2007) Exploring new search algorithms and hardware for phylogenetics: RAxML meets the IBM Cell. The Journal of VLSI Signal Processing, 48, 271–286.
Thompson, J. D., Gibson, T. J., Plewniak, F., Jeanmougin, J. & Higgins, D. G. (1997) The CLUSTAL X windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
Zulueta, A. (1909) Nota sobre reptiles de Melilla (Marruecos). Boletin de la Real Sociedad Española de Historia Natural. 351–354.