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Phrynocephalus scutellatus (OLIVIER, 1807)

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
Subspecies 
Common NamesE: Gray Toadhead Agama 
SynonymAgama scutellata OLIVIER 1807 (III): 110
Agama scutellata — OLIVIER 1807 (V): 196
Phrynocephalus olivieri DUMÉRIL & BIBRON 1837: 517 (fide SMITH 1935)
Phrynocephalus tickelii GRAY 1845: 260 (fide BOULENGER 1885)
Phrynocephalus olivieri — ANDERSON 1872: 386
Phrynocephalus olivieri — BOULENGER 1885: 370
Phrynocephalus olivieri — ALCOCK & FINN 1897
Phrynocephalus olivieri var. brevipes NIKOLSKY 1906
Phrynocephalus olivieri var. carinipes NIKOLSKY 1906
Phrynocephalus scutellatus — SMITH 1935: 229
Phrynocephalus scutellatus — LEVITON 1959: 450
Phrynocephalus scutellatus — WERMUTH 1967: 88
Phrynocephalus scutellata — DAS 1996: 45
Phrynocephalus scutellatus — KHAN 1999
Phrynocephalus (Phrynocephalus) scutellatus — BARABANOV & ANANJEVA 2007
Phrynocephalus scutellatus — SINDACO & JEREMČENKO 2008
Phrynocephalus scutellatus — NASRABADI et al. 2017
Phrynocephalus (Phrynosaurus) scutellatus — SOLOVYEVA et al. 2023
Phrynocephalus (Phrynosaurus) brevipes — SOLOVYEVA et al. 2023
Phrynocephalus (Phrynosaurus) carinipes — SOLOVYEVA et al. 2023
Phrynocephalus (Phrynosaurus) tickelii — SOLOVYEVA et al. 2023 
DistributionIran (incl. Kavir desert), Afghanistan, SW Pakistan (Baluchistan)

Type locality: “... au pied de la montagne (nommée Sophia ou Sophissar, en vue d’Ispahan)” [= Mt. Sophia, near Esfahan, Esfahan Province, Iran].

brevipes: Type locality: “Naim-abad (Damysan) in Chorosano occident” [Naim-Abad, Razavi Khorasan Province, Iran, 36.14 N 58.92 E].

carinipes: Type locality: “Pudeschk-Kupa” [Tudeshg, Isfahan Province, Iran, 32.72 N 52.67 E].  
Reproductionoviparous 
TypesHolotype: MNHN-RA 6947
Lectotype: ZISP 10235, adult female, (designated by Barabanov & Ananjeva 2007) [brevipes]
Lectotype: ZISP 10236 (designated by Barabanov & Ananjeva, 2007) [carinipes] 
DiagnosisDiagnosis: Dorsal scales heterogeneous; enlarged scales nail-like, with free posterior margin often tubercular; more than 16 scales across head between eyes; width of space between nostrils equal to or less than half distance between nostril and preocular ridge, side of back of head and neck without long, flat, upturned fringe-like scales (but sometimes with short, spiny scales); nasals large, in contact, or rarely separated by single series of scales; crossbars on tail most intense (black) and always present ventrally, though usually quite dark dorsally as well. Tail 118--157 percent of snout-vent length (Anderson 1999: 97). 
CommentSynonymy mostly after WERMUTH 1967. We list the species suggested by COI barcoding in Solovyeva et al. 2023 as synonyms, given that they are based on a single 654 bp fragment. The localities of the sampled specimens is not clear from their map even though their range maybe defined in the text (if anywhere).

Distribution: See map in SMID et al. 2014 for distribution in Iran. 
EtymologyThe epithets scutellatus (masculine) and scutellata (feminine) refer to the scut- dorsal shield-like form of the trunk of the species. Hence scutellatus literally means ‛having (-at) a small (-ell-) shield (scut-)’ (Mikaili & Shayegh 2011). 
References
  • ANANJEVA, NATALIA B.; KONSTANTIN D. MILTO, ANDREI V. BARABANOV, EVGENY A. GOLYNSKY 2020. An annotated type catalogue of amphibians and reptiles collected by Nikolay A. Zarudny in Iran and Middle Asia. Zootaxa 4722 (2): 101–128 - get paper here
  • Anderson, J. 1872. On some Persian, Himalayan, and other Reptiles. Proc. Zool. Soc. London 1872: 371-404 - get paper here
  • Anderson, Steven C 1999. The lizards of Iran. Contributions to Herpetology Volume 15, Society for the Study of Amphibians and Reptiles, Saint Louis, Missouri: i-vii, 1-442 [review in Copeia 2000 (4): 1144] - get paper here
  • BARABANOV, A.V. & N.B. ANANJEVA 2007. Catalogue of the available scientific species-group names for lizards of the genus Phrynocephalus Kaup, 1825 (Reptilia, Sauria, Agamidae). Zootaxa 1399: 1-56 - get paper here
  • Boulenger, G.A. 1885. Catalogue of the lizards in the British Museum (Nat. Hist.) I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. London: 450 pp. - get paper here
  • Boulenger, George A. 1890. The Fauna of British India, Including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis, London, xviii, 541 pp. - get paper here
  • Clark, R. 1990. A report on herpetological observations in Afghanistan. British Herpetological Society Bulletin (33): 20-42
  • Damadi, Ehsan; Rasoul Karamiani; Nasrullah Rastegar-Pouyani; Ali Gholamifard 2017. A checklist of lizards from southeastern part of the Sistan and Baluchestan Province in southeastern Iran. Iranian Journal of Animal Biosystematics, DOI: 10.22067/ijab.v13i1.50398 - get paper here
  • Duméril, A. M. C. and G. Bibron. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Encyclopédique Roret, Paris, 570 pp. - get paper here
  • Ebrahimipour, Farnaz; Eskandar Rastegar-Pouyani, Batoul Ghorbani 2016. A Preliminary Study of the Reptile’s Fauna in Northwestern Yazd Province, Iran. Russ. J. Herpetol. 23 (4): 243-248 - get paper here
  • Forcart, Lothar 1950. Amphibien und Reptilien von Iran. Verh. naturf. Ges. Basel 61: 141-156
  • Frynta,D. et al. 1997. Results of the Czech biological expedition to Iran. Part 1. Notes on the distribution of amphibians and reptiles. Acta Soc. Zool. Bohem. 61: 3-17
  • Ghaffari, S. M., Mahmoudi, M., Salehi, H., & Sari, A. 2022. Karyological stucies of four agamid lizards from Semnane province of Iran. Iranian Journal of Animal Biosystematics - get paper here
  • Haas G. and Werner, Y. L. 1969. Lizards and snakes from Southwestern Asia, collected by Henry Field. Bull. Mus. Comp. Zool. Harvard 138: 327-406. - get paper here
  • Kafash, Anooshe; Shima Malakoutikhah, Masoud Yousefi, Farhad Ataei, Hamid Heidari, and Eskandar Rastegar-Pouyani 2018. The Gray Toad-headed Agama, Phrynocephalus scutellatus, on the Iranian Plateau: The degree of niche overlap depends on the phylogenetic distance. Zoology in the Middle East 64 (1) - get paper here
  • Kamali, Kamran 2020. A Field Guide to the Reptiles and Amphibians of Iran. Edition Chimaira, Frankfurt am Main, Germany (www.chimaira.de). 574 pp.
  • Khan, M. S. 1999. A checklist and key to the phrynocephalid lizards of Pakistan, with ethnological notes (Squamata: Agamidae). Pakistan J. Zool. 31 (1): 17-24 - get paper here
  • Khan, M.S. 2003. CHECKLIST AND KEY TO THE LIZARDS OF PAKISTAN. Pakistan J. Zool. Suppl. Ser. (1): 1-25 - get paper here
  • Leviton, A.E. 1959. Systematics and Zoogeography of Philippine Snakes. Unpublished PhD Thesis.
  • Leviton, A.E.; Anderson, S.C. 1961. Further Remarks on the Amphibians and Reptiles of Afghanistan. Wasmann Journal of Biology 19 (2): 269-276 - get paper here
  • Leviton, Alan E. 1959. Report on a collection of reptiles from Afghanistan. Proc. Cal. Acad. Sci. 29 (12): 445-463 - get paper here
  • NASRABADI, REZA; NASRULLAH RASTEGAR-POUYANI, ESKANDAR RASTEGAR-POUYANI, AHMAD GHARZI 2017. A revised key to the lizards of Iran (Reptilia: Squamata: Lacertilia). Zootaxa 4227 (3): 431–443 - get paper here
  • Nikolsky, A. M. 1897. Les reptiles, amphibiens et poissons recueillis (part.) Mr. N. Zaroudny dans la Perse orientale [in Russian]. Annuaire Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg 2: 306-348 - get paper here
  • Nikolsky, A. M. 1915. Faune de la Russie, Reptiles Vol. 1 Chelonia et Sauria(Reptiles of Russia and adjacent countries. Reptiles (Reptilia) Vol. 1) [in Russian]. Imper. Acad. Sci., Petrograd, 532 pp. [Translated from Russian by the Israel Program for Scientific Translations 1963]
  • Nilson,G. & Andren,C. 1981. Die Herpetofauna des Kavir-Schutzgebietes, Kavir-Wüste, Iran. Salamandra 17 (3-4): 130-146 - get paper here
  • Olivier, G.A. 1807. Voyage dans l’Empire Othoman, l’Egypte et la Perse, Tome V. Agasse, Paris, 485 pp. (8th ed.) - get paper here
  • Olivier, G.A. 1807. Voyage dans l’Empire Othoman, l’Egypte et la Perse, Tome III. Agasse, Paris, 566 pp. (4th ed.) - get paper here
  • RAHIMIAN, HASSAN; SOHEILA SHAFIEI, NASRULLA RASTEGAR POUYANI & ESKANDAR RASTEGAR POUYANI 2015. Phylogenetic relationships of the gray-toad agama, Phrynocephalus scutellatus (Olivier, 1807), species complex from Iran. Zootaxa 3990 (3): 369–380 - get paper here
  • RASTEGAR-POUYANI, NASRULLAH; HAJI GHOLI KAMI, MEHDI RAJABZADEH, SOHEILA SHAFIEI AND STEVEN CLEMENT ANDERSON 2008. Annotated Checklist of Amphibians and Reptiles of Iran. Iranian Journal of Animal Biosystematics 4 (1): 7-30
  • Schmidt,K.P. 1939. Reptiles and amphibians from Southwestern Asia. Publ. Field Mus. nat. Hist., zool. Ser., 24: 49-92 - get paper here
  • Sindaco, R. & Jeremcenko, V.K. 2008. The reptiles of the Western Palearctic. Edizioni Belvedere, Latina (Italy), 579 pp. - get paper here
  • ŠMÍD, JIŘÍ; JIŘÍ MORAVEC, PETR KODYM, LUKÁŠ KRATOCHVÍL, SEYYED SAEED HOSSEINIAN YOUSEFKHANI, ESKANDAR RASTEGAR-POUYANI & DANIEL FRYNTA 2014. Annotated checklist and distribution of the lizards of Iran. Zootaxa 3855 (1): 001–097 - get paper here
  • 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.
  • Solovyeva, E. N., Dunayev, E. A., Nazarov, R. A., Bondarenko, D. A., & Poyarkov, N. A. 2023. COI-Barcoding and Species Delimitation Assessment of Toad-Headed Agamas of the Genus Phrynocephalus (Agamidae, Squamata) Reveal Unrecognized Diversity in Central Eurasia. Diversity 2023, 15, x - get paper here
  • Wettstein, O. von & Löffler, H. 1951. Ergebnisse der österreichischen Iran-Expedition 1949l50. Sitzungsber. Akad. Wiss. Wien, math.-nat. Kl., Abt. 1, 160: 427-448 - get paper here
  • Zadhoush, B., Mousavi, S., Mousavi, S. J., & Rajabizadeh, M. 2022. A preliminary study of the lizard fauna (Reptilia: Squamata: Lacertilia) and their habitats in Bafq Protected Area and Darreh-Anjir Wildlife Refuge, Yazd Province, Central Iran. Journal of Wildlife and Biodiversity, 6(X) - get paper here
 
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