Abstract
The genus Hemidactylus is one of the most species-rich and widely distributed of all reptile genera, being found in thetropical and subtropical regions of the world and hundreds of continental and oceanic islands. Despite having already 111species, the number of species described in recent years is very high. This has been facilitated, in part, by the use ofmolecular techniques, which in most cases have been employed to confirm the differentiation at the DNA level of somemorphologically variable forms and to discover some cryptic lineages.Preliminary analyses indicate that some Hemidactylus species from Oman are quite variable in their morphologyand may include more than one species. In order to test this hypothesis we inferred a molecular phylogeny including 131Hemidactylus (20 species) using 1385 base pairs of mitochondrial DNA (353 bp 12S; 302 bp cytb; 588 bp nd4 and 142bp tRNAs) and 1481 bp of nuclear DNA (403 bp c-mos; 668 bp mc1r and 410 bp rag2) and analyzed 226 specimens (15species) for several meristic and pholidotic characters of which we took 3103 photographs that have been deposited inMorphoBank (project 483). Our results indicate the presence of eight new species of Hemidactylus geckos in Arabia: H.luqueorum sp. nov. and H. hajarensis sp. nov. from North Oman; H. masirahensis sp. nov. from Masirah Island; H.inexpectatus sp. nov. from one locality on coastal Central Oman; H. alkiyumii sp. nov., H. festivus sp. nov. and H.paucituberculatus sp. nov. from Dhofar, Southern Oman; and finally H. endophis sp. nov. probably from North Omanand described on the basis of morphology alone. An identification key to the genus Hemidactylus from Oman is alsopresented. With these descriptions, the number of Hemidactylus species found in Oman increases from 7 to 13 and thenumber of endemic Hemidactylus from 0 to 6. The description of three new species endemic to the Hajar Mountains inNorth Oman highlights the importance of this mountain range as a biodiversity hotspot that, up to now, includes 12reptile species that are found nowhere else in the World. Another hotspot of Hemidactylus biodiversity is the DhofarMountain range, in the extreme Southwestern corner of Oman and East Yemen. As a result of its particular geographicsituation, orography and the effect of the Southwest Monsoons, this mountain range presents a diverse variety of habitatswith different species of Hemidactylus adapted to them.With the exception of H. flaviviridis and H. leschenaultii, which belong to the Tropical Asian clade of Hemidactylus,all Arabian Hemidactylus for which DNA sequence is available are members of the Arid clade of Hemidactylus.Relatively recent dispersal appears to have taken place within Arabia in the H. turcicus group, with the South Arabian H.lemurinus occurring far from other confirmed members of this assemblage. Hemidactylus flaviviridis and a clade of H. robustus are genetically uniform, widespread in Arabia and beyond and occur around human habitations, suggesting thatmuch of their large distributions are anthropogenic, as appears to be so in several other Hemidactylus species outsideArabia.The way in which species of Arabian Hemidactylus separate ecologically is surprisingly varied. They may occur atsimilar altitudes but replace each other geographically, or if they are sympatric there may be altitudinal separation.Humidity may also be an important factor, and when animals exist within a few meters of each other, structural nichemay be significant. While four native species occur close together in Dhofar, most Hemidactylus communities in Arabiaconsist of only one or two species, although climbing geckos belonging to other genera, such as Asaccus and Ptyodactylus, may also be present.غReferences
Akaike, H. (1973) Information theory and an extension of the maximum likelihood principle. In: Petrov, B.N. & Csaki, F. (Eds.), Information theory and an extension of the maximum likelihood principle. Akademiai Kiado, Budapest, pp. 267–281.
Amr, Z.S., Modrý, D., Abu Baker, M., Qarqas, M., Al Zaidanyen, J. & Moravec, J. (2007) First record of Hemidactylus mindiae Baha El Din, 2005 from Jordan. Herpetozoa, 20, 73–75.
Ancochea, E., Fuster, J.M., Ibarrola, E., Cendrero, A., Coello, J., Hernán, F., Cantagrel, J.M. & Jamond, C. (1990) Volcanic evolution of the island of Tenerife (Canary Islands) in the light of new K–Ar data. Journal of Volcanology and Geothermal Research, 44, 231–249.
Ancochea, E., Hernán, F., Huertas, M.J., Brändle, J.L. & Herrera, R. (2006) A new chronostratigraphical and evolutionary model for La Gomera: implications for the overall evolution of the Canarian Archipelago. Journal of Volcanology and Geothermal Research, 157, 271–293.
Anderson, J. (1872) On some Persian, Himalayan, and other Reptiles. Proceedings of the Zoological Society of London, 1872, 371–404.
Anderson, J. (1895). On a collection of reptiles and batrachians made by Colonel Yerbury at Aden and its neighbourhood. Proceeding of the Zoological Society of London, 1895, 635–663.
Anderson, S.C. (1999) The Lizards of Iran. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 442 pp.
Arévalo, E., Davis, S.K. & Sites, J.W.Jr. (1994) Mitochondrial DNA sequence divergence and phylogenetic relationships among eight chromosome races of the Sceloporus grammicus complex (Phrynosomatidae) in central Mexico. Systematic Biology, 43, 387–418.
Arnold, E.N. (1972). Lizards with northern affinities from the mountains of Oman. Zoologische Mededelingen, 47, 111–128
Arnold, E.N. (1977) The scientific results of the Oman flora and fauna survey 1975. Little-known geckoes (Reptilia: Gekkonidae) from Arabia with descriptions of two new species from the Sultanate of Oman. Journal of Oman studies special report, 1, 81–110.
Arnold, E.N. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The reptiles and amphibians of Dhofar, southern Arabia. Journal of Oman studies special report, 2, 273–332.
Arnold, E.N. (1986) A key and annotated checklist to the lizards and amphisbaenians of Arabia. Fauna of Saudi Arabia, 8, 385–435.
Arnold, E.N. (2009) Relationships, evolution and biogeography of Semaphore geckos, Pristurus (Squamata, Sphaerodactylidae) based on morphology. Zootaxa, 2060, 1–21.
Arnold, E.N. & Gallagher, M.D. (1977) The scientific results of the Oman flora and fauna survey 1975. Reptiles and amphibians from the mountains of northern Oman with special reference to the Jebel Akhdar region. Journal of Oman studies special report, 1, 59–80.
Arnold, E.N & Gardner, A.S. (1994) A review of the Middle Eastern leaf-toed geckoes (Gekkonidae: Asaccus) with descriptions of two new species from Oman. Fauna of Saudi Arabia 14, 424–441.
Arnold, E.N., Robinson, M.D. & Carranza, S. (2009) A preliminary analysis of phylogenetic relationships and biogeography of the dangerously venomous carpet vipers, Echis (Squamata, Serpentes, Viperidae) based on mitochondrial DNA sequences. Amphibia-Reptilia, 30, 273–282.
Arnold, E.N., Vasconcelos, R., Harris, D.J., Mateo, J.A. & Carranza, S. (2008) Systematics, biogeography and evolution of the endemic Hemidactylus geckos (Reptilia, Squamata, Gekkonidae) of the Cape Verde Islands: based on morphology and mitochondrial and nuclear DNA sequences. Zoologica Scripta, 37, 619–636.
Babocsay, G. (2004) A new species of saw-scaled viper of the Echis coloratus complex (Ophidia: Viperidae) from Oman, Eastern Arabia. Systematics and Biodiversity, 1, 503–514.
Baha El Din, S.M. (2005) A new species of Hemidactylus (Squamata: Gekkonidae) from Egypt. African Journal of Herpetology 52, 39–47.
Baha El Din, S.M. (2005) An overview of Egyptian species of Hemidactylus (Gekkonidae), with the description of a new species from the high mountains of South Sinai. Zoology in the Middle East, 34, 27–34.
Bauer, A.M., Vyas, R., Jackman, T.R., Lajmi, A. & Giri, V. Hemidactylus porbanadarensis Sharma, 1981 is a synonym of Hemidactylus robustus Heyden, 1827. Hamadryad, in press.
Blanford, W.T. (1874) Descriptions of new lizards from Persia and Baluchistàn. Annals and Magazine of Natural History, 13, 453–455.
Blanford, W.T. (1881) Notes on the lizards collected in Socotra by Prof. I. Bayley Balfour. Proceedings of the Zoological Society of London, 1881, 464–469.
Blondel, J. & Aronson, J. (1999) Biology and wildlife of the Mediterranean region. Oxford University Press, New York, 327 pp.
Bosworth, W., Huchon, P. & McClay, K. (2005) The Red Sea and Gulf of Aden Basins. Journal of African Earth Sciences, 43, 334–378.
Boulenger, G.A. (1885) Catalogue of the lizards in the British Museum (Natural History). Vol. I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Trustees of the British Museum, London, XII + 436 pp. + pls I–XXXII. Boulenger, G.A. (1887) Catalogue of the Lizards of the Collection of the British Museum. Vol. III. Lacertidæ, Gerrhosauridæ, Scincidæ, Anelytropidæ, Dibamidæ, Chameleontidæ Trustees of the British Museum (Natural History), London, xii + 575 pp + 40 pl.
Boulenger, G.A. (1887) A list of the reptiles and batrachians obtained near Muscat, Arabia, and presented to the British Museum by Surgeon-Major A.S.G. Jayakar. Annals and Magazine of Natural History, 20, 407–408.
Boulenger, G.A. (1888) Description of a new snake from Muscat, Arabia. Annals and Magazine of Natural History, 2, 508–509.
Boulenger, G.A. (1889) Second account of the fishes obtained by Surgeon-Major A. S. G. Jayakar at Muscat, east coast of Arabia. Proceedings of the Zoological Society of London, 1889 (pt 2), 236–246.
Boulenger, G.A. (1896) Report on Capt. Bottego's second collection of reptiles and batrachians from Somaliland. Annali del Museo Civico di Storia Naturale di Genova, 17, 15–23.
Boulenger, G.A. (1900) Description of a new sea-horse (Hippocampus) from Muscat. Annals and Magazine of Natural History, 6, 51–52.
Boulenger, G.A. (1912) Missione per la frontiera Italo-Etiopica sotto il comando del Capitano Carlo Citerni. Risultati zoologici. List of the reptiles and batrachians. Annali del Museo Civico di Storia Naturale di Genova, 5, 329–332
Brown, R.P. & Pestano, J. (1998) Phylogeography of skinks (Chalcides) in the Canary Islands inferred from mitochondrial DNA sequences. Molecular Ecology, 7, 1183–1191.
Brown, R.P., Terrasa, B., Pérez-Mellado, V., Castro, J.A., Hoskisson, P.A., Picornell, A. & Ramon, M.M. (2008) Bayesian estimation of post-Messinian divergence times in Balearic Island lizards. Molecular Phylogenetics and Evolution, 48, 350–358.
Brown, R.P. & Yang, Z. (2010) Bayesian Dating of Shallow Phylogenies with a Relaxed Clock. Systematic Biology, 59, 119–131.
Busais, S.M. & Joger, U. (2011a) Three new species and one new subspecies of Hemidactylus Oken, 1817 from Yemen (Squamata, Gekkonidae). Vertebrate Zoology, 61, 267–280.
Busais, S.M. & Joger, U. (2011b) Molecular phylogeny of the gecko genus Hemidactylus Oken, 1817 on the mainland of Yemen. Zoology in the Middle East, 53, 25–34.
Buttiker, W. & Gallagher, M.D. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). First records of ophthalmotropic behaviour of Lepidoptera in Oman. Journal of Oman studies special report, 2, 217–221.
Carracedo, J.C., Day, S., Guillou, H., Rodríguez-Badiola, E., Canas, J.A. & Pérez Torrado, F.J. (1998) Hotspot volcanism close to a passive continental margin: the Canary Islands. Geological Magazine, 135, 591–604.
Carranza, S. & Arnold, E.N. (2006) Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 38, 531–545.
Carranza, S., Arnold, E.N., Geniez, P., Roca, J. & Mateo, J.A. (2008a) Radiation, multiple dispersal and parallelism in the skinks, Chalcides and Sphenops (Squamata: Scincidae), with comments on Scincus and Scincopus and the age of the Sahara Desert. Molecular Phylogenetics and Evolution, 46, 1071–1094.
Carranza, S., Arnold, E.N., Mateo, J. & Lopez-Jurado, L. (2000) Long-distance colonization and radiation in gekkonid lizards, Tarentola (Reptilia: Gekkonidae), revealed by mitochondrial DNA sequences. Proceedings of the Royal Society B: Biological Sciences, 267, 637–649.
Carranza, S., Arnold, E.N., Mateo, J.A. & Geniez, P. (2002) Relationships and evolution of the North African geckos, Geckonia and Tarentola (Reptilia: Gekkonidae), based on mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution, 23, 244–256.
Carranza, S., Romano, A., Arnold, E.N. & Sotgiu, G. (2008b) Biogeography and evolution of European cave salamanders, Hydromantes (Urodela: Plethodontidae), inferred from mtDNA sequences. Journal of Biogeography, 35, 724–738.
Carranza, S. & Wade, E. (2004) Taxonomic revision of Algero-Tunisian Pleurodeles (Caudata : Salamandridae) using molecular and morphological data. Revalidation of the taxon Pleurodeles nebulosus (Guichenot, 1850). Zootaxa, 488, 1–24.
Castresana, J. (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution, 17, 540–552.
Clement, M., Posada, D. & Crandall, K.A. (2000) TCS: a computer program to estimate gene genealogies. Molecular Ecology, 9, 1657.
Coello, J., Cantagrel, J.M., Hernán, F., Fuster, J.M., Ibarrola, E., Ancochea, E., Casquet, C., Jamond, C., Deteran, J.R.D. & Cendrero, A. (1992) Evolution of the Eastern volcanic ridge of the Canary-Islands based on new K-Ar Data. Journal of Volcanology and Geothermal Research, 53, 251–274.
Cox, S.C., Carranza, S. & Brown, R.P. (2010) Divergence times and colonization of the Canary Islands by Gallotia lizards. Molecular Phylogenetics and Evolution, 56, 747–757.
Doondorf, J.A. (1798) Amphibien und Fische, vol. Dritter Band. Zoologisches Beyträge zur XIII. Ausgabe des Linneischen Natursystems. Weidmannschen Buchhandlung, Leipzig, Germany, vi + 980 + 1 pp.
Drummond, A. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214.
Duggen, S., Hoernle, K., van den Bogaard, P., Rupke, L. & Morgan, J.P. (2003) Deep roots of the Messinian salinity crisis. Nature, 422, 602–606.
Duméril, A.M.C. & Bibron, G. (1836) Erpetologie Générale ou Histoire Naturelle Complete des Reptiles. Vol.3. Libr. Encyclopédique Roret, Paris, 528 pp.
Escoriza, D., Comas, M.M., Donaire, D. & Carranza, S. (2006) Rediscovery of Salamandra algira Bedriaga, 1833 from the Beni Snassen Massif (Morocco) and phylogenetic relationships of North African Salamandra. Amphibia-Reptilia, 27, 448–455.
Felsenstein, J. (1985) Confidence-limits on phylogenies – an approach using the bootstrap. Evolution, 39, 783.
Flot, J.F. (2010) seqphase: a web tool for interconverting phase input/output files and fasta sequence alignments. Molecular Ecology Resources, 10, 162–166.
Gallagher, M.D. & Rogers, T.D. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). On some birds of Dhofar and other parts of Oman. Journal of Oman studies special report, 2, 347–385.
Gamble, T., Bauer, A.M., Greenbaum, E. & Jackman, T. R. (2008). Evidence of Gondwanan vicariance in an ancient clade of geckos. Journal of Biogeography, 35, 88–104.
Gardner, A.S. (1992) Hemidactylus leschenaultii (Bark Gecko) Herpetological Review, 23, 123.
Gardner, A.S. (1994). A new species of Asaccus (Gekkonidae) from the mountains of northern Oman. Journal of Herpetology, 28, 141–145.
Giri, V.B. (2008) A new rock-dwelling Hemidactylus (Squamata: Gekkonidae) from Maharashtra, India. Hamadryad, 32, 25–33.
Giri, V.B. & Bauer, A.M. (2008) A new ground-dwelling Hemidactylus (Squamata: Gekkonidae) from Maharashtra, with a key to the Hemidactylus of India. Zootaxa, 1700, 21–34.
Giri, V.B., Bauer, A.M., Vyas, R. & Patil, S. (2009) New species of rock-dwelling Hemidactylus (Squamata: Gekkonidae) from Gujarat, India. Journal of Herpetology, 43, 385–393.
Glennie, K. (2006) Oman's Geological Heritage. Stacey International Publishers, London, 247 pp.
Gómez-Díaz, E., Sindaco, R., Pupin, F., Fasola, M. & Carranza, S. Origin and in situ diversification in Hemidactylus geckos of the Socotra Archipelago. Molecular Ecology, in press,
Greathead, D.J. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). Beeflies (Bombyliidae, Diptera) from Oman. Journal of Oman studies special report, 2, 233–250.
Guillou, H., Carracedo, J.C. & Duncan, R.A. (2001) K–Ar, 40Ar–39Ar ages and magnetostratigraphy of Brunhes and Matuyama lava sequences from La Palma Island. Journal of Volcanology and Geothermal Research, 106, 175–194.
Guillou, H., Carracedo, J.C., Paris, R. & Torrado, F.J. (2004) Implications for the early shield-stage evolution of Tenerife from K/Ar ages and magnetic stratigraphy. Earth and Planetary Science Letters, 222, 599–614.
Guillou, H., Carracedo, J.C., Pérez Torrado, F.J. & Rodriguez Badiola, E. (1996) K–Ar ages and magnetic stratigraphy of a hotspot-induced, fast grown oceanic island: El Hierro, Canary Islands. Journal of Volcanology and Geothermal Research, 73, 141–155.
Günther, A. (1894) Report on the collection of reptiles and fishes made by Dr. J. W. Gregory during his expedition to Mount Kenia. Proceedings of the Zoological Society of London, 1894, 84-–91.
Harrigan, R.J., Mazza, M.E. & Sorenson, M.D. (2008) Computation vs. cloning: evaluation of two methods for haplotype determination. Molecular Ecology Resources, 8, 1239–1248.
Harrison, D.L. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The mammals obtained in Dhofar by the 1977 Oman flora and fauna survey. Journal of Oman studies special report, 2, 387–397.
Heyden, C.H.G. von (1827) Atlas zu der Reise im nördlichen Afrika von Eduard Rüppel. Reptilien. Brönner, Frankfurt am Main, 24 pp.
Ho, S.Y.W., Phillips, M.J., Cooper, A. & Drummond, A.J. (2005) Time dependency of molecular rate estimates and systematic overestimation of recent divergence times. Molecular Biology and Evolution, 22, 1561–1568.
Hoogstraal, H. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). Ticks (Ixodoidea) from Oman. Journal of Oman studies special report, 2, 265–272.
Hsü, K., Montadert, J., Beernouilli, L.D., Cita, M.B., Erickson, A., Garrison, R.E., Kidd, R.B., Melieres, F., Muller, C. & Wright, R. (1977) History of the Mediterranean Salinity Crisis. Nature, 267, 399–403.
Hsü, K.J., Ryan, W.B.F. & Cita, M.B. (1973) Late Miocene desiccation of the Mediterranean. Nature, 242, 240–244.
Huelsenbeck, J.P. & Rannala, B. (2004) Frequentist properties of Bayesian posterior probabilities of phylogenetic trees under simple and complex substitution models. Systematic Biology, 53, 904–913.
Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Katoh, K. & Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics, 9, 286–298.
Kocher, T.D., Thomas, W.K., Meyer, A., Edwards, S.V., Paabo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences USA, 86, 6196–6200.
Kraus, F. (2009) Alien Reptiles and Amphibians, a Scientific Compendium and Analysis. Springer Verlag, Dordrecht, xii + 567 pp., CD ROM.
Krijgsman, W., Hilgen, F.J., Raffi, I., Sierro, F.J. & Wilson, D.S. (1999) Chronology, causes and progression of the Messinian Salinity Crisis. Nature, 400, 652–655.
Largen, M. & Spawls, S. (2010) The Amphibians and Reptiles of Ethiopia and Eritrea. Chimaira, Frankfurt, 693 pp.
Larsen, T.B. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The butterflies of Dhofar and their zoogeographic composition. Journal of Oman studies special report, 2, 153–186.
Laughton, A.S. (1966) The Gulf of Aden. Philosophical Transactions of the Royal Society of London Series A, Mathematical and Physical Sciences, 259, 150–171.
Leviton, A.E., Anderson, S.C., Adler, K. & Minton, S.A. (1992) Handbook to Middle East amphibians and reptiles. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 252 pp.
Linnaeus, C. (1758). Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae. 10th Edition, 824 pp.
Ljubisavljevic, K., Arribas, O., Dzukic, G. & Carranza, S. (2007) Genetic and morphological differentiation of Mosor rock lizards, Dinarolacerta mosorensis (Kolombatovic, 1886), with the description of a new species from the Prokletije Mountain Massif (Montenegro) (Squamata: Lacertidae). Zootaxa, 1613, 1–22.
Mahony, S. (2009) A new species of gecko of the genus Hemidactylus (Reptilia: Gekkonidae) from Andhra Pradesh, India. Russian Journal of Herpetology, 16, 27–34.
Minton, S.A. (1966) A contribution to the herpetology of West Pakistan. Bulletin of the American Museum of Natural History, 134, 27–184.
Moravec, J., Kratochvíl, L., Amr, Z.S., Jandzik, D., Smíd, J. & Gvozdík, V. (2011) High genetic differentiation within the Hemidactylus turcicus complex (Reptilia: Gekkonidae) in the Levant, with comments on the phylogeny and systematics of the genus. Zootaxa, 2894, 21–38.
Moravec, J. & Böhme, W. (1997). A new subspecies of the Mediterranean gecko, Hemidactylus turcicus from the Syrian lava desert. (Squamata: Sauria: Gekkonidae). Herpetozoa 10, 121–128.
Pestano, J., Brown, R.P., Suarez, N.M. & Fajardo, S. (2003) A mtDNA study of phylogeography and systematics of the Canary Island bats Pipistrellus and Hypsugo. Molecular Phylogenetics and Evolution, 26, 56–63.
Pinho, C., Rocha, S., Carvalho, B.M., Lopes, S., Mourão, S., Vallinoto, M., Brunes, T.O., Haddad, C.F.B., Gonçalves, H., Sequeira, F. & Ferrand, N. (2010) New primers for the amplification and sequencing of nuclear loci in a taxonomically wide set of reptiles and amphibians. Conservation Genetic Resources, 2, 181–185.
Pleguezuelos, J.M., Fahd, S. & Carranza, S. (2008) El papel del Estrecho de Gibraltar en la conformación de la actual fauna de anfibios y reptiles en el Mediterráneo Occidental. Boletín de la Asociación Hereptológica Española, 19, 2–17.
Posada, D. (2008) jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution, 25, 1253–1256.
Rambaut, A. & Drummond, A. (2007) Tracer v1.4. available from: http://beast.bio.ed.ac.uk/Tracer.
Rato, C., Carranza, S. & Harris, D.J. (2011) When selection deceives phylogeographic interpretation: the case of the Mediterranean house gecko, Hemidactylus turcicus (Linnaeus, 1758). Molecular Phylogenetics and Evolution, 58, 365–373.
Rato, C., Carranza, S., Perera, A., Carretero, M.A. & Harris, D.J. (2010) Conflicting patterns of nucleotide diversity between mtDNA and nDNA in the Moorish gecko, Tarentola mauritanica. Molecular Phylogenetics and Evolution, 56, 962–971.
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Rüppell, E. (1835) Neue Wirbelthiere zu der Fauna von Abyssinien gehörig. [vol. 3] Amphibien. S. Schmerber, Frankfurt am Main, 18 pp.
Sale, J.B. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The environment of the mountain region of Dhofar. Journal of Oman studies special report, 2, 17–23.
Samuel, M.A., Harbury, N., Bott, R. & Manan, A. (1997) Field observations from the Socotran platform: Their interpretation and correlation to Southern Oman. Marine and Petroleum Geology, 14, 661–673.
Schatti, B. & Desvoignes, A. (1999) The herpetofauna of southern Yemen and the Sokotra Archipelago. Gilbert-E. Huguet, Genève, 1–179 pp.
Sindaco, R. & Jeremcenko, V.K. (2008) The reptiles of the Western Palearctic. Annotated checklist and distributional atlas of the turtles, crocodiles, amphisbaenians and lizards of Europe, North Africa, Middle East and Central Asia. Monografie della Societas Herpetologica Italica – I. 579 pp.
Sindaco, R., Metallinou, M., Pupin, F., Fasola, M. & Carranza, S. Forgotten in the ocean: systematics, biogeography and evolution of the Trachylepis skinks of the Socotra Archipelago. Zoologica Scripta, in press,
Sindaco, R., Razzetti, E., Ziliani, U., Wasonga, V., Carugati, C. & Fasola, M. (2007) A New Species of Hemidactylus from Lake Turkana, Northern Kenya (Squamata: Gekkonidae). Acta Herpetologica 2, 37–48.
Sindaco, R., Ziliani, U., Razzetti, E., Carugati, C., Grieco, C., Pupin, F., Al-Aseily, B.A., Pella, F. & Fasola, M. (2009) A misunderstood new gecko of the genus Hemidactylus from Socotra Island, Yemen (Reptilia: Squamata: Gekkonidae). Acta Herpetologica, 4, 83–98.
Smith, M.A. (1935) The fauna of British India, including Ceylon and Burma. Reptiles and Amphibia, Vol. II. Sauria. Taylor and Francis, London, 440 pp.
Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
Stephens, M., Smith, N.J. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. The American Journal of Human Genetics, 68, 978–989.
Talavera, G. & Castresana, J. (2007) Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Systematic Biology, 56, 564–577.
Tamura, K., Peterson, D., Peterson, N., Stecher, G., Nei, M. & Kumar, S. (2011) MEGA5: Molecular evolutionary genetics analysis using Maximum Likelihood, evolutionary distance, and Maximum Parsimony methods. Molecular Biology and Evolution, 28, 2731–2739.
Thomas, O. (1894) Preliminary description of a new goat of the Genus Hemitragus, from South-eastern Arabia. Annals and Magazine of Natural History, 13, 365-–366.
Thorpe, R.S., McGregor, D.P., Cumming, A.M. & Jordan, W.C. (1994) DNA evolution and colonization sequence of island lizards in relation to geological history: mtDNA RFLP, cytochrome b, cytochrome oxidase, 12SrRNA sequence, and nuclear RAPD anaylsis. Evolution, 48, 230–240.
Torki, F., Manthey, U. & Barts, M. (2011) A new Hemidactylus Gray, 1825 from Lorestan Province, western Iran, with notes on Hemidactylus robustus Heyden, 1827 (Reptilia: Squamata: Gekkonidae). Sauria, 33, 47–56.
Uetz, P. (2012) The Reptile Database. Available at http://wwwreptile-databaseorg (Accessed on 10 February 2012),
Ullenbruch, K., Grell, O. & Boehme, W. (2010) Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus (Gekkonidae), and a country-wide checklist. Bonn Zoological Bulletin, 57, 31–54.
van der Kooij, J. (2000) The herpetofauna of the Sultanate of Oman. Part 2: the geckos. Podarcis, 1, 105–120.
Vyas, R., Giri, V. & Bauer, A. (2006) First records of Hemidactylus persicus Anderson, 1872 (Squamata: Sauria: Gekkonidae) from the republic of India, with notes on its distribution. Hamadryad, 30, 211–213.
Waterston, A.R. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The dragonflies (Odonata) of Dhofar. Journal of Oman studies special report, 2, 149–151.
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.
Wilms, T.M. & Schmitz, A. (2007) A new polytypic species of the genus Uromastyx Merrem, 1820 (Reptilia: Squamata: Agamidae: Leiolepidinae) from southwestern Arabia. Zootaxa, 1394, 1–23.
Wiltshire, E.P. (1980) The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The larger moths of Dhofar and their zoogeographic composition. Journal of Oman studies special report, 2, 187–216.