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Phrynocephalus maculatus ANDERSON, 1872

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
SubspeciesPhrynocephalus maculatus longicaudatus HAAS 1957
Phrynocephalus maculatus maculatus ANDERSON 1872 
Common NamesE: Blacktail Toadhead Agama, Spotted Toad-headed Agama
G: Gefleckte Krötenkopfagame 
SynonymPhrynocephalus maculatus ANDERSON 1872: 389
Phrynocephalus maculatus — BOULENGER 1885: 377
Phrynocephalus maculatus — BOULENGER 1920
Phrynocephalus maculatus — SMITH 1935: 233
Phrynocephalus maculatus — WETTSTEIN & LÖFFLER 1951
Phrynocephalus maculatus — WERMUTH 1967: 83
Phrynocephalus maculatus — KHAN 1999
Phrynocephalus maculatus — ŠMÍD et al. 2014
Phrynocephalus maculatus — SOLOVYEVA et al. 2023

Phrynocephalus maculatus maculatus ANDERSON 1872
Phrynocephalus spiniventris NIKOLSKY 1896: 370
Phrynocephalus maculatus spiniventris — NIKOLSKY 1915: 221
Phrynocephalus maculatus maculatus — WERMUTH 1967: 83
Phrynocephalus maculatus maculatus — NILSON & ANDREN 1981: 136
Phrynocephalus (Phrynocephalus) maculatus maculatus — BARABAOV & ANANJEVA 2007
Phrynocephalus maculatus maculatus — SINDACO & JEREMČENKO 2008

Phrynocephalus maculatus longicaudatus HAAS 1957: 67
Phrynocephalus (Phrynocephalus) maculatus longicaudatus — BARABANOV & ANANJEVA 2007
Phrynocephalus maculatus longicaudatus — SINDACO & JEREMČENKO 2008
Phrynocephalus longicaudatus — SOLOVYEVA et al. 2023
Phrynocephalus longicaudatus — ALOUFI et al. 2023 
DistributionIran (incl. Kavir desert), Turkmenistan, Syria, Jordan, Iraq, Pakistan,
Afghanistan, Pakistan, N Saudi Arabia, Oman, United Arab Emirates (UAE), Kuwait

longicaudatus: N Arabia (along the coast). Type locality: Doha Dhalum, Saudi Arabia.

maculatus: Iran etc.; Type locality: Awada, Shiraz, Persia, corrected to Abadeh [31º10' N, 52º37' E; Fars Province, Iran]  
Reproductionoviparous 
TypesHolotype: ZSI 4825
Holotype: CAS 84449 [longicaudatus] 
DiagnosisDiagnosis: No cutaneous fold at angle of mouth; no fringe of scales on posterior border of thigh and sides of base of tail; sides of head and neck without projecting fringe-like scales (Fig. 47b); dorsal scales homogeneous; no enlarged scales along flanks; scales on region considerably larger than those on flanks; nasals separated by one to three scales; tall 140-158 percent of snout-vent length (Anderson 1999: 90). 
CommentSynonymy partly after WERMUTH 1967.

Distribution: See map in SMID et al. 2014 for distribution in Iran and map in Burriel-Carranza et al. 2019 for map in UAE.

Phylogenetics: Ebrahimipour et al. 2021 found populations of P. maculatus to be paraphyletic, especially including P. lutensis. Some populations of maculatus may need to be assigned to P. lutensis.

Group: Solovyeva et al. 2023 included the following 7 species in the P. maculatus group: P. ahvazicus; P. arabicus; P. longicaudatus; P. maculatus; P. nejdensis; P. sakoi. 
EtymologyNamed after the spots (”macula”) this species displays (like many others). 
References
  • Alcock, A. W. & FINN,F. 1897. An account of the Reptilia collected by Dr. F.P. Maynard, Captain A.H. McMahon, C.I.E., and the members of the Afghan-Baluch Boundary Comission of 1896. J. Asiat. Soc. Bengal 65: 550-566 [1896] - get paper here
  • Aloufi, A., Al Zayer, M., & Amr, Z. S. 2023. Reptiles and Amphibians along the Coastal Area of the Eastern Province, Saudi Arabia. Ecologica Montenegrina, 68, 66-87 - get paper here
  • Ananjeva, Natalia B.;Tuniyev, Boris S. 1992. Historical biogeography of the Phrynocephalus species of the USSR. Asiatic Herpetological Research 4: 76-98 - 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
  • 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 (No. 2): 273-332 - get paper here
  • Baker, Mohammad Abu, Pavel Siroky, Zuhair Amr and David Modry 2005. Discovery of a population of Phrynocephalus maculatus Anderson, 1872 in the Hashemite Kingdom of Jordan. Herpetozoa 18 (3/4): 107-113 - 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
  • Burriel-Carranza B, Pedro Tarroso, Johannes Els, Andrew Gardner, Pritpal Soorae, Ahmed Ali Mohammed, Sai Ravi Krishna Tubati, Mohamed Mustafa Eltayeb, Junid Nazeer Shah, Héctor Tejero-Cicuéndez, Marc Simó-Riudalbas, Juan Manuel Pleguezuelos, Daniel F 2019. An integrative assessment of the diversity, phylogeny, distribution, and conservation of the terrestrial reptiles (Sauropsida, Squamata) of the United Arab Emirates. PLoS ONE 14(5): e0216273 - get paper here
  • Carranza S, Xipell M, Tarroso P, Gardner A, Arnold EN, Robinson MD, et al. 2018. Diversity, distribution and conservation of the terrestrial reptiles of Oman (Sauropsida, Squamata). PLoS One 13 (2): e0190389 - get paper here
  • Dunayev, E. A. 1996. On the possible use of the Ethological features in the taxonomy and phylogeny of toad agamas, Phrynocephalus (Reptilia, Agamidae). Russ. J. Herpetol. 3 (1): 32-38 - get paper here
  • Ebrahimipour, F., Rastegar-Pouyani, N., Rastegar-Pouyani, E., Hosseinian Yousefkhani, S. S., & Kamali, K. 2021. Molecular phylogenetic relationships within the Phrynocephalus maculatus-arabicus species complex (Sauria: Agamidae) in Iran. Journal of Zoological Systematics and Evolutionary Research, 59, 1583–1588
  • 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
  • Eissa, S.M. & EL ASSY, Y.S. 1975. Record of certain reptilian species found in Kuwait. Jour. Univ. Kuwait (Sci.), 2: 123-146 - get paper here
  • Forcart, Lothar 1950. Amphibien und Reptilien von Iran. Verh. naturf. Ges. Basel 61: 141-156
  • Gardner, A.S. 2013. The amphibians and reptiles of Oman and the UAE. Edition Chimaira, 480 pp.
  • Gebhart, J. 2013. Oman, quer durch`s Weihrauchland - ein Reisetagebuch. L@CERTIDAE (Eidechsen Online) 2013 [8]: 142-159 - get paper here
  • Haas, Georg 1957. Some amphibians and reptiles from Arabia. Proc. Cal. Acad. Sci. 29 (3): 47-86 - get paper here
  • Jongbloed, M. 2000. Field Guide to the reptiles and amphibians of the UAE - Wild about reptiles. Barkers Trident Communications, 116 pp.
  • 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, Alan E. 1959. Report on a collection of reptiles from Afghanistan. Proc. Cal. Acad. Sci. 29 (12): 445-463 - get paper here
  • Leviton,A.E.; Anderson,S.C.; Adler, K.; Minton,S.A. 1992. Handbook to Middle East Amphibians and Reptiles. SSAR, Oxford, Ohio (Contr. to Herpetol. No. 8), 1-252
  • Nikolsky, A. M. 1896. Diagnoses reptilium et amphibiorum novorum in Persia orientali a N. Zarudny collectorum. Annuaire Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg, 1:369-372. - 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
  • 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
  • SCHOLZ, S. & SIEGENTHALER, F. & RADSPIELER, C. & WILMS, TH. M. 2012. A new locality record of Phrynocephalus maculatus ANDERSON, 1872, from Jordan [Short Note]. Herpetozoa 25 (3/4): 174-179 - 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
  • Szczerbak, N.N. 2003. Guide to the Reptiles of the Eastern Palearctic. Krieger, Malabar, FL, 260 pp. - get paper here
  • van der Kooij, Jeroen 2001. The herpetofauna of the Sultanate of Oman: Part 5: Checklist of the reptiles and amphibians of Oman, and bibliography. Podarcis 2 (3): 94-103 - 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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