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Phrynocephalus putjatai BEDRIAGA, 1909

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
Common NamesChinese: 贵德沙蜥 
SynonymPhrynocephalus putjatai BEDRIAGA 1909: 302
Phrynocephalus putiatae — TZAREVSKY 1929: 419 (unjustified emendation).
Phrynocephalus putjatai — WERMUTH 1967: 86
Phrynocephalus putjaitai — ZHAO 1997: 31 (unjustified emendation)
Phrynocephalus putjatia — WANG et al., 2002: 372 (unjustified emendation).
Phrynocephalus putjatia — GUO & WANG 2007
Phrynocephalus (Oreosaura) putjatai — BARABANOV & ANANJEVA 2007
Phrynocephalus putjatae — JI et al. 2009
Phrynocephalus guinanensis JI, WANG & WANG 2009
Phrynocephalus putjatia — JIN et al. 2013
Phrynocephalus putjatia — WANG et al. 2017
Phrynocephalus putjatia — JIN et al. 2018
Phrynocephalus guinanensis — JIN et al. 2018
Phyrnocephalus guinanensis — JIN & BROWN 2019 (in error)
Phrynocephalus guinanensis — JIN & BROWN 2019
Phrynocephalus putjatia — JIN & BROWN 2019 
DistributionChina (Guide, Qinghai Province)

Type locality: “Gui-dui am Hoang-ho” [= Guide County (36º00' N, 101º 40' E), southeast of Qinghai (Kuku-Nor) Lake, Qinghai Province, China].

guinanensis: China (Guinan, Qinghai); Type locality: Senduo (35°34’N, 101°05’E; elevation 3350 m ASL), Guinan County, Qinghai Province, China.  
TypesLectotype: ZISP 7447.1 (designated by Peters, 1984: 44).
Holotype: NNU (Nanjing Normal University) P 2004.006, adult male (Fig. 1A, C); collected by Xiang Ji on 17 May 2004 [guinanensis] 
DiagnosisDiagnosis (guinanensis): A large viviparous species of Phrynocephalus, distinguishable from congeners using the following combination of characters: up to 84 mm SVL; dorsal ground color of head, neck, trunk, limbs and tail brown with weak light brown mottling; lateral ground color of head, neck, trunk and tail light black with weak white-gray mottling in adult males, and green with weak white-gray mottling in adult females; ventral ground color of tail white-gray to black in the distal part of the tail in adult males, and totally white-gray in adult females; ventral surfaces of hind-limbs white-gray; ventral surfaces of fore-limbs brick-red in adult males, and white-gray in adult females; ventral ground color of trunk and head black in the center but, in the periphery, brick-red in adult males and white-gray in adult females, and totally white-gray in adult females; 5− 8 nasal scales; 5−8 internasal scales; 8−12 scales around parietal eye; 13−18 supraocular scales; 126−164 dorsal scales; 101−136 ventral scales; 144−186 scales around mid-body; 21−31 gular folds; 15−19 subdigital lamellae of the 4th finger; 22−29 subdigital lamellae of the 4th toe. 
CommentSynonymy: Bayesian species delimitation analysis did not support division of P. putjatia and P. guinanensis into separate species (JIN et al. 2013). JIN et al. 2018 confirmed their close relationship. However, Jin & Brown 2019 recognize guinanensis as a separate species again. However, despite some genetic analysis their study does not attempt to show the phylogenetic relationship between different populations of these species. The original description of P. guinanensis also remains a bit dubious and seems to be primarily based on size and color variation (only guinanensis shown, no other species).

Phrynocephalus putjatae has been considered as a synonym of P. vlangalii. P. vlangalii and P. putjatae hybridize in the Qinghai region (JIN & LIU 2008). Of uncertain status fide BARABANOV & ANANJEVA 2007 but considered as a valid species by GUO et al. 2012.

Habitat (guinanensis): sand dunes with very sparse vegetation. 
EtymologyNamed after Mr. D.W. Putjata, “für seine amphibologischen Zuwendungen an das Museum”.
The specific epithet of P. guinanensis is a latinization of the name Guinan County, Qinghai Province, China. 
  • 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
  • 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
  • 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
  • GUO, Xianguang; Li LIU and Yuezhao WANG 2012. Phylogeography of the Phrynocephalus vlangalii Species Complex in the Upper Reaches of the Yellow River Inferred from mtDNA ND4-tRNALEU Segments. Asian Herpetological Research 3 (1): - get paper here
  • Ji, Xiang; Yue-Zhao Wang & Zheng Wang 2009. New species of Phrynocephalus (Squamata, Agamidae) from Qinghai, Northwest China. Zootaxa 1988: 61-68 - get paper here
  • Jin, Y-T., R.P. Brown 2013. Species history and divergence times of viviparous and oviparous Chinese toad-headed sand lizards (Phrynocephalus) on the Qinghai-Tibetan Plateau. Molecular Phylogenetics and Evolution 68 (2):259–268 - get paper here
  • Jin, Yuanting; Liu, Naifa 2008. Introgression of mt DNA between two Phrynocephalus lizards in Qinghai Plateau [in Chinese]. Acta zool. Sinica 54 (1): 111-121
  • Jin, Yuanting; Yubin Wo, Haojie Tong, Sen Song, Lixun Zhang and Richard P. Brown 2018. Evolutionary analysis of mitochondrially encoded proteins of toad-headed lizards, Phrynocephalus, along an altitudinal gradient. BMC Genomics (2018) 19:185 - get paper here
  • Jin, Yuanting; Zhisong Yang, Richard P. Brown, Pinghu Liao, Naifa Liu 2013. Intraspecific lineages of the lizard Phrynocephalus putjatia from the Qinghai-Tibetan Plateau: Impact of physical events on divergence and discordance between morphology and molecular markers. Molecular Phylogenetics and Evolution - 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
  • Noble, D.W.A.; Qi, Y; Fu, J. 2010. Species delineation using Bayesian model-based assignment tests: a case study using Chinese toad-headed agamas (genus Phrynocephalus). BMC Evolutionary Biology 10: 197 - 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, Yue Zhao; ZENG, Xiao Mao; FANG, Z| Li; WU Guan Fu; LIU, Zhi Jun; Theodore J.Papenfuss & J.Robert Macey 2002. A VALID SPECIES OF THE GENUS PHRYNOCEPHALUS: P. PUTJATIA AND A DISCUSSION ON TAXONOMY OF PHRYNOCEPHALUS HONGYUANENSIS (SAURIA:AGAMIDAE). Acta Zootaxonomica Sinica 27 (2): 372-383 - 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
  • Wermuth, Heinz 1967. Liste der rezenten Amphibien und Reptilien. Agamidae. Das Tierreich 86: 1-127
  • ZHANG, Kailong; Haojie TONG, Yubin WO, Naifa LIU and Yuanting JIN 2018. Sex Ratio and Sexual Size Dimorphism in a Toad-headed Lizard, Phrynocephalus guinanensis. Asian Herpetological Research 9 (1): 35-42 - get paper here
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