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Phrynocephalus versicolor STRAUCH, 1876

IUCN Red List - Phrynocephalus versicolor - Least Concern, LC
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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
SubspeciesPhrynocephalus versicolor doriai BEDRIAGA 1909
Phrynocephalus versicolor siebenrocki BEDRIAGA 1909
Phrynocephalus versicolor versicolor STRAUCH 1876 
Common NamesE: Tuvan Toad-headed Agama
Russian: Пестрая круглоголовка
Chinese: 变色沙蜥 
SynonymPhrynocephalus versicolor STRAUCH 1876: 18
Phrynocephalus versicolor — BOULENGER 1885: 374
Phrynocephalus guentheri BEDRIAGA 1906: 168
Phrynocephalus blanfordi BEDRIAGA 1907
Phrynocephalus ciliaris BEDRIAGA 1907
Phrynocephalus versicolor var. Doriai BEDRIAGA 1909: 332
Phrynocephalus versicolor var. Bogdanowi BEDRIAGA 1909: 334
Phrynocephalus helioscopus var. levis KASHTCHENKO 1909: 122
Phrynocephalus Potanini BEDRIAGA 1909: 389
Phrynocephalus pewzowi
Phrynocephalus versicolor versicolor — NIKOLSKY 1915: 188
Phrynocephalus versicolor doriai — NIKOLSKY 1915: 190
Phrynocephalus versicolor bogdanowi — NIKOLSKY 1915: 191
Phrynocephalus rostralis ZAREVSKY 1930
Phrynocephalus parvulus ZAREVSKY 1927 (fide POPE 1935)
Phrynocephalus stejnegeri BANNIKOV 1958: 78 (nomen nudum)
Phrynocephalus versicolor — WERMUTH 1967: 89
Phrynocephalus immaculatus ZHAO 1995: 47
Phrynocephalus versicolor — WANG et al. 2003
Phrynocephalus (Phrynocephalus) versicolor versicolor — BARABANOV & ANANJEVA 2007
Phrynocephalus immaculatus — ANANJEVA 2011
Phrynocephalus versicolor — DUNAYEV et al. 2021
Phrynocephalus przewalskii — SOLOVYEVA et al. 2023: 26

Phrynocephalus versicolor siebenrocki BEDRIAGA 1909
Phrynocephalus versicolor var. siebenrocki BEDRIAGA 1909: 232
Phrynocephalus versicolor var. Siebenrocki — BEDRIAGA 1909: 333
Phrynocephalus versicolor siebenrocki — NIKOLSKY 1915: 191
Phrynocephalus versicolor siebenrocki — MILTO & BARABANOV 2012: 165 
DistributionW China (Xinjiang, Gansu, Ningxia, Nei Mongol = Inner Mongolia), (Siberia), SE Russia (Tuva), Mongolia

doriai: Chian (Zinjiang); Terra typica restricta: "See Ebi-nor" [= Ebinur Lake, Xinjiang Uygur Autonomous Region, China].

Type locality: Alashan Desert, N China  
Reproductionoviparous 
TypesLectotype: ZISP 3929.1 (designated by Peters, 1984: 36)
Lectotype: ZISP 5117.1 (designated by Peters, 1984). “See Ebinor” [= Ebinur lake, Xinjiang Uygur Autonomous Region, China]. Leg: N. M. Przewalsky, 1878; however, Barabanov & Ananjeva 2007: 20 state that there is no type [doriai]
Holotype: ZISP 5156, “zwischen Metschin-Ola und der Stadt Barkul” [between Mieh-Ch’eng-O-La Shan (about 30 km northeast of Barkol) and Barkol, approx. 43°55’ N 93°00 E, Barkol county, Xinjiang Uygur Autonomous Region, China]. Leg: G. N. Potanin, 1878 [siebenrocki] 
Diagnosis 
CommentSubspecies: Wang & Fu (2004) suggested to apply the name P. versicolor only to the populations in the northwestern part of its distribution area. Populations from Tengger Desert, Ordos highland, and further east include P. frontalis, P. przewalskii, and P. versicolor and should be called P. przewalskii. Some populations may contain a new species that remains to be described. Guo & Wang (2007) recognized populations in Kuytun, Xinjiang Uygur Autonomous Region as P. guttatus instead of P. versicolor. Dunayev et al. 2021 elevated hispidus and kulagini to full species status and thus removed them from versicolor, but they did not comment on either doriai or siebenrocki. Ganbold et al. 2022 considered kulagini and hispidus as subspecies of versicolor, but showed in their phylogenetic analysis that they are paraphyletic with versicolor, hence we keep them as valid species (note, however, that their phylogeny is only based on a 522 bp fragment of ND2).

Synonymy partly after BARABANOV & ANANJEVA (2007) who consider doriai and hispidus as synonyms of versicolor (and thus recognize only 2 subspecies: versicolor and kulagini), but see subspecies above. 
References
  • ANANJEVA, Natalia B.; Xianguang GUO and Yuezhao WANG 2011. Taxonomic Diversity of Agamid Lizards (Reptilia, Sauria, Acrodonta, Agamidae) from China: A Comparative Analysis. Asian Herpetological Research 2 (3): 117-128 - 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
  • 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
  • Bedriaga, J. von 1907. Wissenschaftliche Resultate der von N. M. Przewalski nach Central-Asien unternommenen Reisen.Zoologischer Theil, Amphibien und Reptilien, Band 3, Abt. 1, Lieferung 2: Lacertilia (Gekkonidae, Agamidae). Kaiserl. Akad. Wissensch., St. Petersburg, pages 71-278
  • Bedriaga, J.V. 1909. Amphibien und Reptilien. In: Wissenschaftliche Resultate der Reisen N.M. Przewalskijs durch Zentralasien. Zoologische Teil. Band 3. Part 1. Lacertilia. (in Russian). Sankt-Petersburg, Vol 3 (1) 3: 73-102, 367 - 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
  • Buehler, Matthew D.; Purevdorj Zoljargal, Erdenetushig Purvee, Khorloo Munkhbayar, Munkhbayar Munkhbaatar, Nyamsuren Batsaikhan, Natalia B. Ananjeva, Nikolai L. Orlov, Theordore J. Papenfuss, Diego Roldán-Pińa, - Douchindorj, Larry Lee Grismer, Jamie 2021. The Results of Four Recent Joint Expeditions to the Gobi Desert: Lacertids and Agamids. Russian Journal of Herpetology 28 (1): 15-32 - get paper here
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  • Dunayev, Evgeny A.; Evgeniya N. Solovyeva, Nikolay A. Poyarkov 2021. Systematics, Phylogeny, and Evolution of Phrynocephalus (Superspecies przewalskii) (Reptilia: Agamidae). Russian Journal of Herpetology 28 (1): 43-59 - get paper here
  • Eifler, Douglas A; Maria A Eifler 2019. Movement and habitat use by adult and juvenile Toad-headed Agama lizards (Phrynocephalus versicolor Strauch, 1876) in the eastern Gobi Desert, Mongolia. Herpetology Notes 12: 717-719 - get paper here
  • Ganbold, Onolragchaa; Erdenetushig Purevee, Munkhbayar Khorloo, Ariunbold Jargalsaikhan, Terbish Khayankhyarvaa, Altangoo Ochirbat, Zoljargal Purevdorj & Munkhbaatar Munkhbayar 2022. Strong Population Genetic Structure of Phrynocephalus versicolor in Mongolia Herpetologica 78(1): 21–29. - get paper here
  • Golubev, M.L. 1993. The variegated toad agama in Djungar Gate (Eastern Kazakstan) with notes on certain systematic problems of Phrynocephalus versicolor Str. (Reptilia, Agamidae). Asiatic Herpetological Research 5: 51-58 - get paper here
  • Guo, X. & Wang, Y. 2007. Partitioned Bayesian analyses, dispersal–vicariance analysis, and the biogeography of Chinese toad-headed lizards (Agamidae: Phrynocephalus): A re-evaluation. Molecular Phylogenetics and Evolution <br />45 (2): 643-662 - get paper here
  • Kashtchenko, Nikolai F. [as KASCENKO] 1909. The reptiles and amphibians collected by the expeditions of 1902-6 and 1908 by Prof. V. V. Saposnikov in Central Asia. [in Russian]. Annu. Mus. Zool. Acad. Imper. Sci.,<br />St. Petersburg, 14(1-2): 119-130 - get paper here
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  • Manilo, Valantina V.;Golubev, Michael L. 1993. Karyotype information on some toad agamas of the Phrynocephalus guttatus species group (Sauria, Agamidae) of the former USSR. Asiatic Herpetological Research 5: 105-108 - get paper here
  • MCGRATH, CASEY. 2022. Highlight: Yellow is the New Black for Gobi Desert Lizards. Genome Biology and Evolution 14(7): evac095. - get paper here
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  • Munkhbayar, Munkhbaatar; Terbish Khayankhyarvaa, Onolragchaa Ganbold, Zoljargal Purevdorj, Burnee Mundur, Gurragchaa Jargalsaikhan, Munkhbayar Khorloo 2020. A Herpetofaunal Survey of Northwestern Mongolia with the First Country Record of the Moorfrog, Rana arvalis Nilsson 1842. IRCF Reptiles & Amphibians 27 (2): 154-160 - get paper here
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  • 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
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  • Tong H, Li J, Wo Y;Gang Shao; Wei Zhao; Diana Aguilar‐Gómez & Yuanting Jin 2019. Effects of substrate color on intraspecific body color variation in the toad‐headed lizard, Phrynocephalus versicolor. Ecol Evol. 2019: 00:1–10 - get paper here
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