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Published: 2017-06-13

Systematic revision of Acanthodactylus busacki (Squamata: Lacertidae) with a description of a new species from Morocco

The Steinhardt Museum of Natural History, Israel National Center for Biodiversity Studies, Tel Aviv University, 6997801 Tel-Aviv, Israel Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, 6997801 Tel Aviv, Israel
EPHE, PSL Research University, CNRS, UM, SupAgro, IRD, INRA, UMR 5175 Centre d’Écologie Fonctionnelle et Évolutive (CEFE), F-34293 Montpellier cedex 5, France
CIBIO/InBIO, Centro de Investigacão em Biodiversidade e Recursos Genéticos da Universidade do Porto, R. Padre Armando Quintas, 4485-661 Vairão, Portugal
CEFE UMR 5175, CNRS - Université de Montpellier - Université Paul-Valéry Montpellier - EPHE, 1919 Route de Mende, 34293 Montpellier cedex 5, France
Reptilia Atlas Mountains evolution lizards pardalis species-group phylogeny taxonomy


Recent molecular phylogenies of the Acanthodactylus pardalis species-group have revealed a deep genetic divergence within the nominal species A. busacki from north-west Africa. The species is phylogenetically separated into northern and southern lineages, which correspond to a previously observed morphological differentiation between the northern and southern populations of this species. Based on morphological comparisons of the type material and location of the type locality, the nomen Acanthodactylus busacki Salvador, 1982 is assigned here to the southern lineage, known from the northern Saharan Atlantic coastal desert. The northern lineage, described here as Acanthodactylus margaritae sp. nov., is prominently characterized by weakly keeled dorsal scales and a characteristic colour pattern. The new species is endemic to Morocco and confined to arid and semi-arid bioclimatic areas between the High Atlas and Anti-Atlas Mountains, from around Tamri in the north to Tiznit in the south and the Souss valley in the east.



  1. Arnold, E.N. (1983) Osteology, genitalia and the relationships of Acanthodactylus (Reptilia: Lacertidae). Bulletin of the British Museum (Natural History), Zoology, 44, 291–339.

    Bandelt, H.J., Forster, P. & Röhl, A. (1999) Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16, 37–48.

    Barata, M., Perera, A., Harris, D.J., Van der Meijden, A., Carranza, S., Ceacero, F., García-Muñoz, E., Gonçalves, D., Henriques, S., Jorge, F. & Marshall, J.C. (2011) New observations of amphibians and reptiles in Morocco, with a special emphasis on the eastern region. British Herpetological Society Herpetological Bulletin, 116, 4–14.

    Bhiry, N. & Occhietti, S. (2004) Fluvial sedimentation in a semi-arid region: the fan and interfan system of the Middle Souss Valley, Morocco. Proceedings of the Geologists' Association, 115, 313–324.

    Bons, J. & Geniez, P. (1996) Amphibiens et reptiles du Maroc. Asociación Herpetológica Española, Barcelona, 319 pp.

    Brito, J.C., Rebelo, H., Crochet, P.-A. & Geniez, P. (2008) Data on the distribution of amphibians and reptiles from North and West Africa, with emphasis on Acanthodactylus lizards and the Sahara Desert. Herpetological Bulletin, 105, 19–27.

    Carretero, M.A., Fonseca, M.M., Garcia-Munoz, E., Brito, J.C. & Harris, D.J. (2011) Adding Acanthodactylus beershebensis to the mtDNA phylogeny of the Acanthodactylus pardalis group. North-Western Journal of Zoology, 7, 138–142.

    Castresana, J. (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Phylogenetics and Evolution, 17, 540–552.

    Crochet, P.-A., Geniez, P. & Ineich, I. (2003) A multivariate analysis of the fringe-toed lizards of the Acanthodactylus scutellatus group (Squamata: Lacertidae): systematic and biogeographical implications. Zoological Journal of the Linnean Society, 137, 117–155.

    Crochet, P.-A., Leblois, R. & Renoult, J.P. (2015) New reptile records from Morocco and Western Sahara. Herpetology Notes, 8, 583–588.

    Dewey, J.F., Helman, M.L., Turco, E., Knott, S.D. & Hutton, D.H.W. (1989) Kinematics of the western Mediterranean. Geological Society, London, Special Publications, 45, 265–283.

    Donaire, D., Mateo, J.A., Hasi, M. & Geniez, P. (2000) Nuevos datos sobre la fauna reptiliana de la Hamada de Tinduf (Argelia). Boletín de la Asociación Herpetológica Española, 11, 8–12.

    Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29, 1969–1973.

    Fitzinger L.J.F.J. (1834). Acanthodactylus. In: Wiegman A.F.A. (Ed.), Herpetologia mexicana, seu Descriptio amphibiorum Novae Hispaniae quae itineribus comitis De Sack, Ferdinandi Deppe et Chr. Guil. Schiede in Museum zoologicum Berolinense pervenerunt. Pars prima saurorum species amplectens, adjecto systematis saurorum prodromo, additisque multis in hunc amphibiorum ordinem observationibus. C. G. Luderitz, Berolini, pp. 10.

    Flot, J.F. (2010) SeqPHASE: a web tool for interconverting PHASE input/output files and FASTA sequence alignments. Molecular Ecology Resources, 10.1, 162–166.

    Fonseca, M.M., Brito, J.C., Paulo, O.S., Carretero, M.A. & Harris, D.J. (2009) Systematic and phylogeographical assessment of the Acanthodactylus erythrurus group (Reptilia: Lacertidae) based on phylogenetic analyses of mitochondrial and nuclear DNA. Molecular Phylogenetics and Evolution, 51, 131–142.

    Fonseca, M.M., Brito, J.C., Rebelo, H., Kalboussi, M., Larbes, S., Carretero, M.A. & Harris, D.J. (2008) Genetic variation among spiny-footed lizards in the Acanthodactylus pardalis group from North Africa. African Zoology, 43, 8–15.

    Geniez, P., Mateo, J.A., Geniez, M. & Pether, J. (2004) The amphibians and reptiles of the Western Sahara (former Spanish Sahara) and adjacent regions. Edition Chimaira, Frankfurt, 228 pp.

    Geniez, P., Mateo Miras, J.A., Joger, U., Pleguezuelos, J., Slimani, T. & El Mouden, E.H. (2010) Acanthodactylus busacki. The IUCN Red List of Threatened Species 2010: e.T178569A7572735. Available from: (accessed 24 April 2017)

    Gomez, F., Beauchamp, W. & Barazangi, M. (2000) Role of the Atlas Mountains (northwest Africa) within the African-Eurasian plate-boundary zone. Geology, 28, 775–778.<775:ROTAMN>2.0.CO;2

    Harris, D.J. & Arnold, E.N. (2000) Elucidation of the relationships of spiny footed lizards, Acanthodactylus spp. (Reptilia: Lacertidae) using mitochondrial DNA sequence, with comments on their biogeography and evolution. Journal of Zoology, 252, 351–362.

    Harris, D.J., Batista, V. & Carretero, M. (2004) Assessment of genetic diversity within Acanthodactylus erythrurus (Reptilia: Lacertidae) in Morocco and the Iberian Peninsula using mitochondrial DNA sequence data. Amphibia-Reptilia, 25, 227–232.

    Harris, D.J., Perera, A., Barata, M., Tarroso, P. & Salvi, D. (2010) New distribution notes for terrestrial herpetofauna from Morocco. North-Western Journal of Zoology, 6, 309–315.

    Huelsenbeck, J.P. & Rannala, B. (2004) Frequentist properties of Bayesian posterior probabilities of phylogenetic trees under simple and complex substitution models. Systematic Biology, 53.6, 904–913.

    Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.

    Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33, 1870–1874.

    Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2016) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34, 772–773.

    Librado, P. & Rozas, J. (2009) DnaSP v5: A software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452.

    Mellado, J. & Dakki, M. (1988) Inventaire commenté des amphibiens et reptiles du Maroc. Bulletin de l’Institut Scientifique, Rabat, 12, 171–181.

    Mellado, J. & Olmedo, G. (1990) El género Acanthodactylus en Marruecos: problemas de identificación en los grupos de especies A. pardalis y A. scutellatus. Amphibia-Reptilia, 11, 131–146.

    Pyron, R.A., Burbrink, F.T. & Wiens, J.J. (2013) A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology, 13, 93.

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v1.6. Available from: (accessed 24 April 2017)

    Salvador, A. (1982) A revision of the lizards of the genus Acanthodactylus (Sauria: Lacertidae). Bonner Zoologisches Monographien, Nr. 16, Zoologisches Forschungsinstitut und Museum Alexander Koenig, 1–167.

    Schleich, H.H., Kästle, W. & Kabisch, K. (1996) Amphibians and reptiles of North Africa. Koeltz, Koenigstein, 627 pp.

    Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution, 12.4, 335–337.

    Sindaco, R. & Jeremčenko, 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. Edizioni Belvedere, Latina, 579 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., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57, 758–771.

    Stephens, M. & Scheet, P. (2005) Accounting for decay of linkage disequilibrium in haplotype inference and missing-data imputation. The American Journal of Human Genetics, 76.3, 449–462.

    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.

    Tamar, K., Carranza, S., Sindaco, R., Moravec, J. & Meiri, S. (2014) Systematics and phylogeography of Acanthodactylus schreiberi and its relationships with Acanthodactylus boskianus (Reptilia: Squamata: Lacertidae). Zoological Journal of the Linnean Society, 172, 720–739.

    Tamar, K., Carranza, S., Sindaco, R., Moravec, J., Trape, J.-F. & Meiri, S. (2016) Out of Africa: Phylogeny and biogeography of the widespread genus Acanthodactylus (Reptilia: Lacertidae). Molecular Phylogenetics and Evolution, 103, 6–18.

    Trape, J.-F., Trape, S. & Chirio, L. (2012) Lézards, crocodiles et tortues d'Afrique occidentale et du Sahara. IRD Orstom, Marseille, 503 pp.

    Uetz, P. & Hošek, J. (2017) The Reptile Database. Available from: (accessed 25 March 2017)

    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 Biology and Evolution, 25, 361–371.