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Acanthodactylus margaritae TAMAR, GENIEZ, BRITO & CROCHET, 2017

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Higher TaxaLacertidae, Eremiadinae, Sauria, Lacertoidea, Squamata (lizards)
Subspecies 
Common Names 
SynonymAcanthodactylus margaritae TAMAR, GENIEZ, BRITO & CROCHET 2017
Acanthodactylus pardalis bedriagai — ARNOLD 1983: 319 (part.)
Acanthodactylus bedriagai — HARRIS & ARNOLD 2000: 352 (part.)
Acanthodactylus busacki — MELLADO & DAKKI 1988: 175 (part.)
Acanthodactylus busacki — MELLADO & OLMEDO 1990: 133 (part.)
Acanthodactylus busacki — BONS & GENIEZ 1996: 162 (part.)
Acanthodactylus busacki — SCHLEICH et al. 1996: 391 (part.)
Acanthodactylus busacki — DONAIRE et al. 2000: 10
Acanthodactylus busacki — GENIEZ et al. 2004: 102 (part.)
Acanthodactylus busacki — BRITO et al. 2008: 21 (part.)
Acanthodactylus busacki — SINDACO & JEREMČENKO 2008: 218 (part.)
Acanthodactylus busacki — FONSECA et al. 2008: 9 (part.)
Acanthodactylus busacki — GENIEZ et al. 2010: IUCN red list (part.)
Acanthodactylus busacki — HARRIS el al. 2010: 22 (part.)
Acanthodactylus busacki — CARRETERO et al. 2011: 139 (part.)
Acanthodactylus busacki — TRAPE et al. 2012: 302 (part.)
Acanthodactylus busacki — PYRON et al. 2013: 17 (part.)
Acanthodactylus busacki — TAMAR et al. 2016: 8 (part.) 
DistributionS Morocco

Type locality: 500 m north-east of Sidi Binzarne, approximately 3 km south-east of Sidi R’bat (Massa), Morocco, 30.0580°N, 9.6456°W (WGS84).  
Reproductionoviparous. 
TypesHolotype: MNHN RA-2016.0105 (formerly BEV.9439, tissue sample in the BEV tissue collection, code T1176), adult male, collected on the 11th of February 2008 by P.-A. Crochet (Fig. 6). Paratypes. BEV.9440 (tissue sample code T1177, adult female, collected in the Ademine forest 1.6 km past the Agadir-Taroudant road towards Biougra, Morocco, 30.31579°N 9.36004°W, on the 11th of February 2008 by P.- A. Crochet); MCCI-R.560 (adult male, collected from between Inchaden and Tifnite, Morocco, 30.15°N, 9.59°W, on the 9th of May 1993 by R. Sindaco and N. Fedrighini); MCCI-R.561 (adult male, collected from Souss valley, outskirts of Biougra, Morocco, 30.21°N, 9.38°W, on the 9th of May 1993 by R. Sindaco and N. Fedrighini); MCCI- R.562 (sub-adult female, collected from Souss valley, outskirts of Biougra, Morocco, 30.21°N 9.38°W, on the 9th of May 1993 by R. Sindaco and N. Fedrighini) (Fig. 7). 
DiagnosisDiagnosis: A new species of the pardalis species-group (i.e., small flat or carinated dorsal scales; three series of scales on the fingers; three supraoculars; 12 and sometimes 14 straight longitudinal row of ventrals; slightly pectinate toes; body pattern combining longitudinal rows of light ocelli and black reticulation) from between the High Atlas and Anti-Atlas Mountains characterized by the combination of the following characters: (1) maximum recorded snout-vent length 71 mm (54–71 mm in adult males and 54–64.7 mm in adult females); (2) three supraoculars, the first supraocular is either entire or fragmented with usually one row of granules between the supraoculars and the superciliaries; (3) 16–48 granules around the supraoculars (mean 29.7); (4) subocular with a distinct keel is situated between the fourth and fifth upper labials and does not contact the lip; (5) upper temporals small and pointed whereas the lower temporals are large and smooth; (6) 10–14 collar scales; (7) usually 12 longitudinal rows of ventral scales; (8) 27–34 (mean 31.2) ventral transverse rows; (9) 14–24 (mean 21.1) femoral pores on each side (higher number in males; 14–23 in females vs. 18–24 in males; mostly in contact in males, separated in females); (10) three rows of scales on fingers with slight lateral pectination, with 16–23 (mean 19.5) lamellae underneath the fourth toe; (11) dorsal scales weakly carinated, flat and imbricate; (12) dorsal colour pattern consists of three longitudinal pale lines or rows of whitish, elongated ocelli on each side (six longitudinal dorsal lines in total), with obvious black reticulation between the two outer ones and no or little reticulation between the inner ones on the middle of the dorsum; (13) the dorsal and ventral scales of the nape and body show yellowish colouration in both males and females during the spring, which fades later during the summer; (14) juveniles have blue colouration of the tail, and sub-adult females sometimes have a reddish ventral tail colour.


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CommentHabitat: ground-dwelling, inhabiting stony plains with fine grained soils, stable sands and fixed dunes, or hard clay grounds with scarce low vegetation but also, especially in the Souss valley, open argan tree forests. 
EtymologyThe specific epithet “margaritae”, a noun in the genitive case, honours Dr Margarita Metallinou who tragically lost her life during field-work in Africa in July 2015. The new species is dedicated to Margarita Metallinou from all the authors in recognition of her passion, interest and strong contribution to the study of reptile systematics (especially of geckoes of the genera Stenodactylus and Ptyodactylus) and to her friendship over the years. 
References
  • Arnold E.N. 1983. Osteology, genitalia and the relationships of Acanthodactylus (Reptilia: Lacertidae). Bulletin of the British Museum (Natural History) Zoology 44: 291–339 - get paper here
  • Bons, J. & Geniez.,P 1996. Amphibiens et Reptiles du Maroc (Sahara Occidental compris), Atlas biogéographique [trilingual in French, Spanish and English]. Asoc. Herpetol. Espanola, Barcelona [ISBN 84-921999-0-3], 320 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
  • Donaire D, Mateo JA, Hasi M, Geniez Ph. 2000. Nuevos datos sobre la fauna reptiliana de la Hamada de Tinduf (Algeria). Boletín de la Asociation Herpetológica Española<br>11: 8–12. - get paper here
  • Elbahi A, Lawton C, Oubrou W, El Bekkay M, Hermas J, Dugon M 2022. Reptile biodiversity in Souss-Massa National Park: an internationally important hotspot in the Mediterranean region. Biodiversity Data Journal 10: e79088 - get paper here
  • Fonseca, Miguel M.; José C. Brito, Hugo Rebelo, Mohsen Kalboussi, Saïd Larbes, Miguel A. Carretero, and D. James Harris 2008. Genetic variation among spiny-footed lizards in the Acanthodactylus pardalis group from North Africa. African Zoology 43 (1): 8-15 - get paper here
  • 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. [reviewed in Reptilia GB 44: 81]
  • 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
  • Harris,D.J. & Arnold,E.N. 2000. Elucidation of the relationships of spiny-footed lizards, Acanthodactylus ssp. (Reptilia: Lacertidae) using mitochondrial DNA sequence, with comments of their biogeography and evolution. Journal of Zoology 252: 351-362 - get paper here
  • Martínez del Mármol, Gabriel; D. James Harris, Philippe Geniez, Philip de Pous, and Daniele Salvi 2019. Amphibians and Reptiles of Morocco. Edition Chimaira, Frankfurt am Main, Germany, 478 pp - get paper here
  • Mellado, J. & Dakki,L.M. 1988. Inventaire commenté des amphibiens et reptiles du Maroc. Bull. Inst. Sci. 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 - get paper here
  • Pyron, R.A.; Frank T Burbrink, John J Wiens 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evol Biol 13: 93 - get paper here
  • Schleich, H.H., Kästle,W., Kabisch, K. 1996. Amphibians and Reptiles of North Africa. Koeltz, Koenigstein, 627 pp.
  • Sindaco, R. & Jeremcenko, V.K. 2008. The reptiles of the Western Palearctic. Edizioni Belvedere, Latina (Italy), 579 pp. - get paper here
  • Tamar, K; S Carranza, R Sindaco, J Moravec, JF Trape, S Meiri 2016. Out of Africa: Phylogeny and biogeography of the widespread genus Acanthodactylus (Reptilia: Lacertidae). Molecular Phylogenetics and Evolution, doi:10.1016/j.ympev.2016.07.003 - get paper here
  • TAMAR, KARIN; PHILIPPE GENIEZ, JOSÉ C. BRITO, PIERRE-ANDRÉ CROCHET 2017. Systematic revision of Acanthodactylus busacki (Squamata: Lacertidae) with a description of a new species from Morocco. Zootaxa 4276 (3): 357–386 - get paper here
  • Trape, J.F.; Trape, S. & Chirio, L. 2012. Lézards, crocodiles et tortues d'Afrique occidentale et du Sahara. IRD Orstom, 503 pp. - get paper here
 
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