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Phrynocephalus przewalskii STRAUCH, 1876

IUCN Red List - Phrynocephalus przewalskii - Least Concern, LC
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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
Subspecies 
Common NamesE: Przewalski's Toadhead Agama; birulai: Tsarewsky's Toadhead Agama; steindachneri: Steindachner's Toadhead Agama; parvulus: Slight Toadhead Agama
Chinese: 荒漠沙蜥 
SynonymPhrynocephalus Przewalskii STRAUCH 1876: 10
Phrynocephalus affinis STRAUCH 1876: 10
Phrynocephalus przewalskii — BOULENGER 1885: 377
Phrynocephalus affinis — BOULENGER 1885: 377
Phrynocephalus affinis — BOULENGER 1885: 377
Phrynocephalus steindachneri BEDRIAGA 1907: 207
Phrynocephalus carinilabris BEDRIAGA 1909: 412
Phrynocephalus elegans TZAREVSKY 1927
Phrynocephalus birulai TZAREVSKY 1927: 304
Phrynocephalus carinatus ZAREVSKY 1927
Phrynocephalus parvulus TZAREVSKY 1927
Phrynocephalus przewalskii — POPE 1935: 470
Phrynocephalus przewalskii — WERMUTH 1967: 85
Phrynocephalus przewalskii — WERMUTH 1967: 86
Phrynocephalus vlangalii — WERMUTH 1967: 91
Phrynocephalus steindachneri — WERMUTH 1967: 89
Phrynocephalus przewalskii — WANG et al. 2003
Phrynocephalus affinis — BARTS & WILMS 2003
Phrynocephalus elegans — BARTS & WILMS 2003
Phrynocephalus birulai — BARTS & WILMS 2003
Phrynocephalus steindachneri — BARTS & WILMS 2003
Phrynocephalus parvulus — BARTS & WILMS 2003
Phrynocephalus (Phrynocephalus) przewalskii — BARABANOV & ANANJEVA 2007
Phrynocephalus suschkinianus ZAREVSKY 1927 (? syn. fide POPE 1935)
Phrynocephalus bogdanowi — MANILO et al. 1993
Phrynocephalus przewalskii — SINDACO & JEREMČENKO 2008
Phrynocephalus birulai — MILTO & BARABANOV 2012
Phrynocephalus parvulus — MILTO & BARABANOV 2012: 162
Phrynocephalus przewalskii — MILTO & BARABANOV 2012: 162
Phrynocephalus steindachneri — MILTO & BARABANOV 2012: 163
Phrynocephalus przewalskii — DUNAYEV et al. 2021
Phrynocephalus przewalskii — SOLOVYEVA et al. 2023 
DistributionChina (Nei Mongol = Inner Mongolia), Ningxia, Gansu, Qinghai), elevation 1000 to 1500m

affinis: China (Inner Mongolia: Ala-Shan, Ordos deserts); Type locality: “in provincia Chinensi, Ordos dicta, nec non in deserto Alaschanico”.

birulai: China (Inner Mongolia: southern Ala-Shan desert); Type locality: “südliches Alashan an der Grenze von Kansu und längs des Flusses Tangar, östliches Nan-shan” (fide WERMUTH 1967).

elegans: China (Inner Mongolia, NE Ala-Shan desert); Type locality: “östlicher Ala-shan, zwischen Tingyuan-ying und Scharuzan-sumé, ning-sia” (fide WERMUTH 1967).

parvulus: China (Inner Mongolia: Eastern Ala-Shan desert);Type locality: östliche Alashan, zwischen Tingyuangying and Scharuzan-sume, Ningsía (fide WERMUTH 1967: 90)

steindachneri: C Asia (northern Nan-schan); Type locality: “Gaotai, am Nordrande des Nan-schan” (only known from the type locality fide WERMUTH 1967). No type known fide BARABANOV & ANANJEVA 2007

Type locality: in deserto Alaschanico (= Alashan desert).  
ReproductionOviparous 
TypesHolotype: ZISP 3928
Holotype: ZISP 8157, “in GaoTaï, also am Nordrande des Nan-schan” [Ruo river, northern limit of Nan-shan mountains near Gaotai, approx. 39°22’ N 99°48’ E, Gansu province, China]. Leg: G. E. Grum-Grzimailo, 1891 [steindachneri] 
Diagnosis 
CommentSynonymy after BARABANOV & ANANJEVA 2007 and WERMUTH 1967 who lists Phrynocephalus carinatus and P. elegans as synonyms with question marks. WERMUTH 1967 gives POPE 1935 as source for this synonymization. P. parvulus was listed as synonym of Phrynocephalus versicolor by WERMUTH 1967 (with a question mark). POPE 1935 suggested to synonymize Phrynocephalus affinis with Phrynocephalus przewalskii. Phrynocephalus frontalis STRAUCH 1876 has been considered as a synonym of Phrynocephalus przewalskii by BARABANOV & ANANJEVA 2007 but treated as valid by more recent authors.

Distribution: 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) suggested that the Lanzhou population of P. frontalis is part of P. przewalskii.

Group: The przewalskii group includes P. frontalis; P. hispidus; P. kulagini; P. przewalskii (including P. versicolor), fide SOLOVYEVA et al. 2023. 
EtymologyNamed after General Nikolai Mikhailovitch Przewalski (1839-1888), a Russian Cossack naturalist who explored Central Asia (Beolens et al. 2009).

Etymology [parvulus]: Named after Latin “parvulus, -a, -um” = very small. 
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
  • 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
  • Barts, M. & Wilms, T. 2003. Die Agamen der Welt. Draco 4 (14): 4-23 - 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
  • Beolens, Bo; Michael Watkins, and Michael Grayson 2009. The Eponym Dictionary of Mammals. Johns Hopkins University Press, 592 pp. - get paper here
  • Beolens, Bo; Michael Watkins, and Michael Grayson 2011. The Eponym Dictionary of Reptiles. Johns Hopkins University Press, Baltimore, USA - 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
  • 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
  • Gao, Wei; Yan-Bo Sun, Wei-Wei Zhou, Zi-Jun Xiong, Luonan Chen, Hong Li, Ting-Ting Fu, Kai Xu, Wei Xu, Li Ma, Yi-Jing Chen, Xue-Yan Xiang, Long Zhou, Tao Zeng, Si Zhang, Jie-Qiong Jin, Hong-Man Chen, Guojie Zhang, David M. Hillis, Xiang Ji, Ya-Ping Zh 2019. Genomic and transcriptomic investigations of the evolutionary transition from oviparity to viviparity. PNAS, February 11, 2019 - get paper here
  • Gozdzik, Agnes 2005. Species, Speciation, and Hybridization in Toad-Headed Lizards of the Phrynocephalus przewalskii Complex. Unpublished MSc thesis, University of Guelph, Canada.
  • Gozdzik, Agnes and Jinzhong Fu 2009. Are Toad-Headed Lizards Phrynocephalus przewalskii and P. frontalis (Family Agamidae) the Same Species? Defining Species Boundaries with Morphological and Molecular Data. Russ. J. Herpetol. 16 (2): 107-118 - 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
  • LIAN, Xue; Zhigang JIANG, Xiaoge PING, Songhua TANG, Junhuai BI and Chunwang LI 2012. Spatial Distribution Pattern of the Steppe Toad-headed Lizard (Phrynocephalus frontalis) and Its Influencing Factors. Asian Herpetological Research 3 (1): - get paper here
  • Macey, J. Robert;Papenfuss, Theodore J.;Zhao, Ermi 1988. The snakes of Ningxia Hui autonomous region as an indication of a herpetofaunal corridor. Chinese Herpetological Research 2 (1): 4-5
  • 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
  • Milto, Konstantin D.; Andrei V. Barabanov 2012. A Catalogue of the Agamid and Chamaeleonid Types in the Collection of the Zoological Institute, Russian Academy of Sciences, St. Petersburg. Russ. J. Herpetol. 19 (2): 155-170 - 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]
  • Pang, Junfeng; Yuezhao Wang, Yang Zhong, A. Rus Hoelzel, Theodore J. Papenfuss, Xiaomao Zeng, Natalia B. Ananjeva and Ya-ping Zhang 2003. A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 27: 398-409 - get paper here
  • Peters G. 1984. Die Krötenkopfagamen Zentralasiens (Agamidae: Phrynocephalus). Mitt. Zool. Mus. Berlin 60(1): 23-67
  • Pietschmann, V. 1919. Franz Steindachner. Annalen des Naturhistorischen Museums in Wien 33: 47-48 - get paper here
  • Pope, CLIFFORD H. 1935. The Reptiles of China. Turtes, Crocodilians, Snakes, Lizards. Amer. Mus. Nat. Hist., New York, Nat. Hist. Central Asia, 10: lii, 1-604 - get paper here
  • 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
  • Strauch in: PRZEWALSKI, N.M. 1876. Mongoliya i Strana Tangutov. Tryokhletneye puteshestviye v Vostochnoj Nagoruoj Asii (= Mongolia and the Tangut Country. A Three-Year Travel in Eastern High Asia). Volumes 1-2. Imper. Russ. Geogr. Soc., St. Petersburg (in Russian; English edition, London 1876, French edition, Paris 1880, German edition, Jena 1887; abridged Russian edition with added notes, Moscow 1946). - get paper here
  • Tang X, Xin Y, Wang H, Li W, Zhang Y, et al. 2013. Metabolic Characteristics and Response to High Altitude in Phrynocephalus erythrurus (Lacertilia: Agamidae), a Lizard Dwell at Altitudes Higher Than Any Other Living Lizards in the World. PLoS One 8 (8): e71976. doi:10.1371/journal.pone.0071976 - get paper here
  • TANG, Xiaolong; Feng YUE, Ming MA, Ningbo WANG, Jianzheng HE and Qiang CHEN 2012. Effects of Thermal and Hydric Conditions on Egg Incubation and Hatchling Phenotypes in Two Phrynocephalus Lizards. Asian Herpetological Research 3 (3): 184–191 - get paper here
  • Tzarevsky [= Carevskij], S. 1927. Les lézards du genre Phrynocephalus, collectionnés par les expéditions de P. Koslov au Tibet en 1899-1901 et en Mongolie et Sze-Tschwan 1907-1909 [in Russian with French title]. Comptes Rendus de l’Académie des Sci- <br />ences de l’URSS, 1927: 303–306
  • Urquhart, J.; Bi, K.; Gozdzik, A. & Fu, J. 2005. Isolation and characterization of microsatellite DNA loci in the toad-headed lizards, Phrynocephalus przewalskii complex. Molecular Ecology Notes 5: 928 –930 - get paper here
  • Wang, Kai; Jinlong Ren, Hongman Chen, Zhitong Lyu, Xianguang Guo Ke Jiang, Jinmin Chen, Jiatang Li, Peng Guo, Yingyong Wang, Jing Che 2020. The updated checklists of amphibians and reptiles of China. Biodiversity Science 28 (2): 189-218 - get paper here
  • Wang, Y. & Fu, J. 2004. Cladogenesis and vicariance patterns in the toad-headed lizard Phrynocephalus versicolor species complex. Copeia 2004 (2): 199-206 - get paper here
  • WANG, Yajuan; Weihong JI, Wei ZHAO, Nannan YU and Naifa LIU 2011. Geographic Variation in Clutch and Egg Size for the Lizard Phrynocephalus przewalskii (Squamata: Agamidae). Asian Herpetological Research 2 (2): 97-102 - get paper here
  • Wang, Yuezhao; Wang, Huizhao 1993. Geographic variation and diversity in three species of Phrynocephalus in the Tengger Desert, Western China. Asiatic Herpetological Research 5: 65-73 - get paper here
  • Wang, Zheng; Li Ma, Min Shao, Xiang Ji 2017. Are viviparous lizards more vulnerable to climate warming because they have evolved reduced body temperature and heat tolerance? Oecologia - get paper here
  • Zhan, A. & Fu, J. 2009. Microsatellite DNA markers for three toad-headed lizard species (Phrynocephalus vlangalii, P. przewalskii and P. guttatus). Molecular Ecology Resources 9: 535–538 - get paper here
 
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