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Taxonomy of the genus Phrynocephalus Biodiversity of the family Agamidae

Bibliography of the genus
Phrynocephalus (Toad-headed Agamas)

(Reptilia: Sauria: Agamidae)

Note: In order to limit redundancy, relevant literature indexed in the related bibliographies in the left column may not have been included in this page. For a comprehensive search of literature, these bibliographies should therefore also be consulted.


Phrynocephalus in general

Ananjeva, N.B. 1986. [On the validity of Megalochilus mystaceus (Pallas, 1776)]. (In Russian, English summary). Trudy Zoologicheskogo Instituta 157: 4-13.

Ananjeva, N.B. 1998. The skull structure of some arid Asian agamids of Phrynocephalus genus (Agamidae, Sauria). Russian Journal of Herpetology 5(1): 29-35.

Ananjeva, N.B. 2012. [To viviparity in Agama lizards (Sauria, Acrodonta, Agamidae)]. (In Russian). Zoologicheskii Zhurnal 91(11): 1358-1365.

Ananjeva, N.B.; Sokolova, T.M. 1990. [The position of the genus Phrynocephalus Kaup, 1825 in agamids system]. (In Russian, English summary). Trudy Zoologicheskogo Instituta 207: 12-21, 233.

Ananjeva, N.B.; Tuniev, B.S. 1992. Historical biogeography of the Phrynocephalus species of the USSR. Asiatic Herpetological Research 4: 76-98.

Arnold, E.N. 1999. Modes of ear reduction in iguanian lizards (Reptilia, Iguania); different paths to similar ends. Bulletin of the Natural History Museum Zoology Series 65(2): 165-171.

Arnold, E.N. 1999. Phylogenetic relationships of toad-headed lizards (Phrynocephalus, Agamidae) based on morphology. Bulletin of the Natural History Museum Zoology Series 65(1): 1-13.

Badmaeva, V.I.; Onaeva, N.S. 1977. [The circadian activity of Phrynocephalus in eastern Manych]. pp. 19-20. In: Darevsky, I.S. [Fourth All-Union Herpetological Conference. Questions in Herpetology]. Akademiya Nauk SSSR, Zoologicheskij Institut. Izdatel'stvo 'Nauka', Leningrad. 243 pp. (In Russian).

Barabanov, A.V.; Ananjeva, N.B. 2007. Catalogue of the available scientific species-group names for lizards of the genus Phrynocephalus Kaup, 1825 (Reptilia, Sauria, Agamidae). Zootaxa 1399: 1-56.

Böhme, W. (ed.) 1981. Handbuch der Reptilien und Amphibien Europas. Band 1. Echsen (Sauria) 1. (Gekkonidae, Agamidae, Chamaeleonidae, Anguidae, Amphisbaenidae, Scincidae, Lacertidae 1). Akademische Verlagsgesellschaft, Wiesbaden. 520 pp.

Bondarenko, D.A. 1992. [Influence of the toad agamas activities (Sauria, Phrynocephalus) upon evaluation of their density]. (In Russian). Ekologiya (Moscow) 6: 79-82.

Darevsky, I.S. 1957. Hautdrüsen auf dem Rücken einiger agamider Eidechsen der Gattung Phrynocephalus. Zoologische Jahrbücher (Abteilung für Systematik, Geographie und Biologie der Tiere) (Jena) 85: 301-304.

Dunayev, E.A. 1996. [Nomenclature and distribution of toad-agams, Phrynocephalus (Reptilia, Agamidae) in Iliyskaya Hollow]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 101(3): 36-41.

Dunayev, E.A. 1996. On the possible use of the ethological features in the taxonomy and phylogeny of toad agamas, Phrynocephalus (Reptilia, Agamidae). Russian Journal of Herpetology 3(1): 32-38.

Dunayev, E.A. 2008. Phylogeny of lizards of Phrynocephalus genus (Reptilia: Agamidae): history of study and methodic approaches. pp. 117-126. In: Ananjeva, N.B.; Danilov, I.G.; Dunayev, E.A.; Ischenko, V.G.; Lada, G.A.; Litvinchuk, S.N.; Orlova, V.F.; Smirina, E.M.; Tuniyev, B.S. & Khalikov, R.G. (eds.). The problems of herpetology. Proceedings of the 3rd meeting of the Nikolsky Herpetological Society, 9-13 October 2006, Putschino. Russian Academy of Sciences, St. Petersburg. 467 pp.

Golubev, M.L. 1989. [Phrynocephalus guttatus (Gmel.) or Ph. versicolor Str. (Reptilia, Agamidae): which Phrynocephalus species occurs in Kazakhstan? (In Russian, English summary). Vestnik Zoologii 1989(5): 38-46.

Golubev, M.L. 1998. A new subspecies of Phrynocephalus ornatus Boulenger (Reptilia: Agamidae) from eastern Iran, with a key to south-western and middle Asian microphrynocephalids. Hamadryad 23(2): 162-168.

Guo, X.G.; Liu, L.; Wang, Y. 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): 52-68.

Guo, X.G.; 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 45(2): 643-662.

Hoser, R.T. 2012. A five-way division of the agamid genus Laudakia Gray, 1845 (Squamata: Sauria: Agamidae). Australasian Journal of Herpetology 14: 17-23.

Jin, Y.T.; Brown, R.P. 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.

Jin, Y.T.; Li, J.Q.; Liu, N.F. 2013. Elevation-related variation in life history traits among Phrynocephalus lineages on the Tibetan Plateau: do they follow typical squamate ecogeographic patterns. Journal of Zoology (London) 290(4): 293-301.

Jin, Y.T.; Liu, N.F.; Luo, S.; Ma, X.N. 2007. [A study of taxonomy and phylogeography of Phrynocephalus vlangalii complex in Qinghai plateau with the evidence of mtDNA. pp. 9-14]. (In Chinese). In: Ji, X. Herpetologica Sinica 11. China Scientific Book Services, Beijing. 283 pp.

Kamalova, Z.Y. 1985. [Species composition of insects in the diet of lizard entomophages]. (In Russian). Trudy Tashkentskogo Sel'skokhozyaistvennogo Instituta 113: 92-96.

Khan, M.S. 1999. A checklist and key to the phrynocephalid lizards of Pakistan, with ethnological notes (Squamata: Agamidae). Pakistan Journal of Zoology 31(1): 17-24.

Leroy, P. 1939. Les Phrynocephales de Mongolie et du NW. Chinois. Peking Nat. Hist. Bull. 14(2): 139-146.

Li, J.; Guo, X.G.; Wang, Y. 2010. [Progress in the biogeography of Phrynocephalus (Agamidae)]. (In Chinese). Sichuan Journal of Zoology 29(1): 144-150.

Licent, E. 1933. Note sur le genre Phrynocephalus. Pub. Mus. Hoang Pai Ho TienTsin 23: 10-12.

Macey, J.R.; Schulte, J.A.; Ananjeva, N.B.; Dyke, E.T. van; Wang, Y.; Orlov, N.; Shafiei, S.; Robinson, M.D.; Dujsebayeva, T.; Freund, G.S.; Fischer, C.M.; Liu, D.; Papenfuss, T. 2018. A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. Zootaxa 4467(1): 1–81.

Manilo, V.V. 2000. [Description of karyotypes of some species and subspecies of the genus Phrynocephalus (Sauria, Agamidae) from Central Asia]. (In Russian, English summary). Vestnik Zoologii 34(6): 113-118, 123.

Manilo, V.V. 2001. Cytogenetic review and evolution of karyotypes in the species of the genus Phrynocephalus Kaup, 1825 (Sauria, Agamidae) from the eastern Palaearctic. Hamadryad 26(2): 227-234.

Manilo, V.V.; Golubev, M.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.

Manilo, V.V.; Golubyev, M.L. 1994. [Peculiarities of karyotypes of Phrynocephalus species from the group "guttatus" sensu lato (Sauria, Agamidae)]. (In Russian, English summary). Tsitologiya i Genetika 28(2): 66-70.

Mezhzherin, S.; Golubev, M.L. 1993. Allozyme variation and genetic relationships within the Phrynocephalus guttatus species group (Sauria: Agamidae) in the former USSR. Asiatic Herpetological Research 5: 59-64.

Mezhzherin, S.V.; Golubev, M.L. 1989. [Genetic divergence of gen. Phrynocephalus Kaup (Reptilia: Agamidae) of the USSR fauna]. (In Ukrainian, English summary). Dopovidi Akademiyi Nauk Ukrayins'koyi RSR Seriya B Heolohichni Khimichni Ta Biolohichni Nauky 1989(11): 68-70.

Mishagina, J.V. 2008. Ecological groups of preys in Eastern Kara Kum psammobiont lizards, Eremias (Lacertidae) and Phrynocephalus (Agamidae). pp. 298-307. In: Ananjeva, N.B.; Danilov, I.G.; Dunayev, E.A.; Ischenko, V.G.; Lada, G.A.; Litvinchuk, S.N.; Orlova, V.F.; Smirina, E.M.; Tuniyev, B.S. & Khalikov, R.G. (eds.). The problems of herpetology. Proceedings of the 3rd meeting of the Nikolsky Herpetological Society, 9-13 October 2006, Putschino. Russian Academy of Sciences, St. Petersburg. 467 pp.

Pang, J.; Wang, Y.; Yang, Z.; Hoelzel, A.R.; Papenfuss, T.J.; Zeng, Z.; Ananjeva, N.B.; Zhang, Y.P. 2003. A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 27(3): 398-409.

Peters, G. 1984. Die Krotenkopfagamen Zentralasiens (Agamidae: Phrynocephalus). Mitteilungen aus dem Zoologischen Museum in Berlin 60(1): 23-67.

Rogovin, K.A.; Semenov, D.V. 2003. [On methods of age determination in desert lizards: Natural longevity in toad-headed lizards (Phrynocephalus) (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 82(7): 862-864.

Semenov, D.V. 1983. [On homing in the desert lizards]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 88(3): 50-52.

Semenov, D.V.; Brushko, Z.K.; Kubykin, R.A.; Shenbrot, G.I. 1987. [Taxonomic position and protective status of the round-headed lizard (Reptilia, Agamidae) in the territory of the USSR]. (In Russian, English summary). Zoologicheskii Zhurnal 66(1): 98-109.

Semenov, D.V.; Shenbrot, G.I. 1982. [On species independence of Phrynocephalus moltschanovi (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 61(8): 1194-1204.

Semenov, D.V.; Shenbrot, G.I. 1985. [An estimate of absolute density of lizard populations taking into account the marginal effect]. (In Russian, English summary). Zoologicheskii Zhurnal 64(8): 1246-1253.

Semenov, D.V.; Shenbrot, G.I. 1986. [Data to the shedding in lizards of genus Phrynocephalus]. (In Russian). Ekologiya (Moscow) 1986(2): 80-82.

Semyonov, D.V.; Dunayev, Y.A. 1989. [Morphology of the hemipenis and classification of lizards of the genus Phrynocephalus (Reptilia: Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 68(10): 56-64.

Shenbrot, G.I.; Semenov, D.V. 1986. Some biological peculiarities of Phrynocephalus (Agamidae). pp. 579-582. In: Rocek, Z. (ed.). Studies in herpetology. Proceedings of the European Herpetological Meeting (3rd Ordinary General Meeting of the Societas Europaea Herpetologica) Prague 1985. Charles University, Prague. 754 pp.

Sokolovsky, V.V. 1974. A comparative karyological study of lizards of the family Agamidae. 1. Chromosome complements of 8 species of the genus Phrynocephalus (Reptilia, Agamidae). Tsitologiya 16(7): 920-925.

Solovyeva, E.N.; Poyarkov, N.A.; Dunaev, E.A.; Dujsebayeva, T.N.; Bannikova, A.A. 2011. Molecular differentiation and taxonomy of the sunwatcher toad-headed agama species complex Phrynocephalus superspecies helioscopus (Pallas 1771) (Reptilia: Agamidae). Russian Journal of Genetics 47(7): 842-856.

Tertyshnikov, M.F.; Gorovaya, V.I. 1984. [On the study of western populations of Phrynocephalus]. pp. 43-44. (In Russian). In: Ishchenko, V.G. (ed.). [The species and its productivity within its distribution range. Part 5. Questions of herpetology. Material from the 4th All-Union Symposium 3-7 April 1984]. Urals Scientific Centre, Academy of Sciences USSR, Sverdlovsk. 54 pp.

Tzarewsky, S. 1927. Les lezards du genre Phrynocephalus, collectionnes par les expeditions de P. Koslov au Tibet en 1899-1901 et en Mongolie et Sze-Tschwan 1907-1909. C. R. Acad. Sci. Russ. Leningrad 1927(A. 19): 303-306.

Tzarewsky, S. 1927. Taxonomy and distribution of lizards of the genus Phrynocephalus. C. R. Acad. Sci. U.R.S.S. 2: 23-20.

Tzarewsky, S. 1929. Contribution to the classification and distribution of the lizards of the genus Phrynocephalus. C. R. Acad. Russ. Leningrad 1929(17): 415.

Tzarewsky, S.T.; Tzarewsky, S.F. 1928. Contributions a la systematique et la distribution des lezards du genre Phrynocephalus. C. R. Acad. Sci. Russ. Leningrad 1926: 119-122, 211-214.

Ushakov, B.P. 1962. [Cytophysiological analysis of intraspecific differentiation of takyr toad lizards]. (In Russian). Doklady Akademii Nauk SSSR (Biological Sciences) 144: 645-647.

Wang, H.; Tang, X.; Wang, Y.; Feng, Y.; Pu, P.; Men, S.; Zhao, Y.; Peng, Z.; Chen, Q. 2018. Function of lactate dehydrogenase in cardiac and skeletal muscle of Phrynocephalus lizard in relation to high-altitude adaptation. Asian Herpetological Research 9(4): 258-273.

Wang, Y. 1987. [The skeletal anatomy of four species of Phrynocephalus with a discussion on their interspecific relationships]. (In Chinese, English summary). Acta Herpetologica Sinica 6(4): 27-34.

Wang, Y.; Fu, J. 2004. Cladogenesis and vicariance patterns in the toad-headed lizard Phrynocephalus versicolor species complex. Copeia 2004(2): 199-206.

Wang, Y.; Zeng, X.M.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, J.R. 1999. [Study on the relationships of classification, phylogenetics and distribution of the genus Phrynocephalus spp. (Sauria: Agamidae) with the paleogeographical changes during Cenozoic era in Tibet Plateau]. (In Chinese, English summary). Zoological Research 20(3): 178-185.

Yadamsuren, O.; Murdoch, J.D.; Chuluunbat, S.; Purevee, E.; Munkhbayar, M.; Jargalsaikhan, A.; Purevjargal, Z.; Khorloo, M.; Khayankhyarvaa, T. 2018. Estimating occupancy and detectability of Toad Headed Agamas at the periphery of their range in Mongolia. Journal of Herpetology 52(3): 361-368.

Yadamsuren, O.; Murdoch, J.D.; Chuluunbat, S.; Purevee, E.; Munkhbayar, M.; Jargalsaikhan, A.; Purevjargal, Z.; Khorloo, M.; Khayankhyarvaa, T. 2018. Estimating occupancy and detectability of Toad Headed Agamas at the periphery of their range in Mongolia. Journal of Herpetology 52(3): 361-368.

Yadgarov, T.; Makhmudov, B. 1991. [Changes in the distributional range and abundance of several species of toad agamas from Uzbekistan]. (In Russian). Uzbekskii Biologicheskii Zhurnal 1: 51-57, 84.

Ye, X.; Zhang, J.D.; Dai, J.H.; Wang, Y. 2005. [A summarisation of systematic research of Phrynocephalus (Agamidae)]. (In Chinese, English summary). Sichuan Journal of Zoology 24(3): 421-427.

Zamolodchikov, D.G. 1993. [Reptile population dynamics: a trial of systems analysis]. (In Russian, English summary). Zhurnal Obshchei Biologii 54(5): 592-608.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

Zhao, K. 1997. [Toad-headed agamids in China]. (In Chinese, English summary). Chinese Journal of Zoology 32(1): 15-19.

Phrynocephalus ahvazicus

Melnikov, D.A.; Melnikova, E.; Nazarov, R.; Rajabizadeh, M.; Al-Johany, A.; Amr, Z.S.; Ananjeva, N.B. 2014. Taxonomic revision of Phrynocephalus arabicus Anderson, 1984 complex with description of a new species from Ahvaz, south-western Iran. Russian Journal of Herpetology 21(2): 149-159.

Phrynocephalus arabicus

Al-Sadoon, M.K.; Abdo, N.M. 1994. Comparative metabolic rate-temperature curves of Phrynocephalus arabicus Anderson, 1894 and Agama (Stellio) stellio brachydactyla Hass, 1951 (Agamidae, Sauria, Reptilia). Journal of Arid Environments 28(3): 249-256.

Al-Shammari, A.M. 2012. Additional records of lizards in Ha'il Province, Saudi Arabia. Russian Journal of Herpetology 19(4): 287-291.

Al-Sirhan, A.R.; Brown, G. 2010. The status of the two toad-headed agamas, Phrynocephalus arabicus (Anderson, 1894) and P. maculatus (Anderson, 1872), in Kuwait (Sauria: Agamidae). Zoology in the Middle East 51: 23-30.

Amr, Z.S.; Al-Oran, R.; Disi, A.M. 1994. Reptiles of southern Jordan. The Snake 26(1): 41-49.

Anderson, J. 1894. On two new species of agamoid lizards from the Hadramut, southeastern Arabia. Annals and Magazine of Natural History (Ser. 6) 14: 376-378.

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 2: 273-332.

Haas, G. 1957. Some amphibians and reptiles from Arabia. Proceedings of the California Academy of Sciences (Ser. 4) 29(4): 47-86.

Haas, G.; Battersby, J.C. 1959. Amphibians and reptiles from Arabia. Copeia 1959: 196-202.

Londei, T. 2015. Arabian sand boa Eryx jayakari (Squamata: Boidae) preying on Arabian toad-headed agama Phrynocephalus arabicus (Squamata: Agamidae): a nocturnal-to-diurnal species interaction. Herpetology Notes 8: 155-156.

Melnikov, D.A.; Melnikova, E.; Nazarov, R.; Rajabizadeh, M.; Al-Johany, A.; Amr, Z.S.; Ananjeva, N.B. 2014. Taxonomic revision of Phrynocephalus arabicus Anderson, 1984 complex with description of a new species from Ahvaz, south-western Iran. Russian Journal of Herpetology 21(2): 149-159.

Ross, W. 1995. Body language in agamids of eastern Saudi Arabia. Tropical Fish Hobbyist 43(10): 124, 127-129, 131-133.

Ross, W. 1995. Tail signalling in populations of Phrynocephalus arabicus, Anderson, 1894 (Reptilia: Agamidae). Zoology in the Middle East 11: 63-71.

Wittenberg, J. 1992. First records of Arabian toad-head agamid, (Phrynocephalus arabicus) (Sauria: Agamidae) in Jordan. Zoology in the Middle East 7: 59-63.

Phrynocephalus axillaris

Autumn, K.; Wang, Y. 1988. Preliminary observations on the ecology of Phrynocephalus axillaris and Eremias velox in the Turpan Depression, Xinjiang Uygur Autonomous Region, China. Chinese Herpetological Research 2(1): 6-13.

Golubev, M.L.; Dunayev, E.A. 1995. Phrynocephalus nasatus (Reptilia, Agamidae), a new species of toad agama from western China. Russian Journal of Herpetology 2(1): 5-9.

Li, J.; Quo, X.G.; Wang, Y. 2010. [Population genetic differentiation of Phrynocephalus axillaris in East of Xinjiang Uygur Autonomous Region based on sequence variation of mitochondrial ND4-tRNALeu gene]. (In Chinese). Zoological Research 31(4): 361-369.

Sun, Y.Y.; Du, Y.; Yang, J.; Fu, T.B.; Lin, C.X.; Ji, X. 2012. Is the evolution of viviparity accompanied by a relative increase in maternal abdomen size in lizards. Evolutionary Biology (New York) 39(3): 388-399.

Wang, Z.; Li, M.; Shao, M.; Ji, X. 2013. Differences in incubation length and hatchling morphology among five species of oviparous Phrynocephalus lizards (Agamidae) from China. Asian Herpetological Research 4(4): 225-232.

Wu, M.; Li, P.; Zhao, Y.; Ji, X. 1989. [A study of the karyotype of three species of Phrynocephalus from Xinjiang]. (In Chinese). Zoological Research 10(2): 114,153.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

Zhang, Q.; Xia, L.; He, J.B.; Wu, Y.; Fu, J.; Yang, Q. 2010. Comparison of phylogeographic structure and population history of two Phrynocephalus species in the Tarim Basin and adjacent areas. Molecular Phylogenetics and Evolution 57(3): 1091-1104.

Zhao, K. 1995. [Description of a new species of the genus Phrynocephalus from Inner Mongolia]. (In Chinese, English summary). Sichuan Journal of Zoology 14 (Supplement): 47-50.

Phrynocephalus bannikovi

Macey, J.R.; Schulte, J.A.; Ananjeva, N.B.; Dyke, E.T. van; Wang, Y.; Orlov, N.; Shafiei, S.; Robinson, M.D.; Dujsebayeva, T.; Freund, G.S.; Fischer, C.M.; Liu, D.; Papenfuss, T. 2018. A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. Zootaxa 4467(1): 1–81.

Shammakov, S. 1977. [On the ecology of Phrynocephalus reticulatus bannicovi]. p. 230. In: Darevsky, I.S. [Fourth All-Union Herpetological Conference. Questions in Herpetology]. Akademiya Nauk SSSR, Zoologicheskij Institut. Izdatel'stvo 'Nauka', Leningrad. 243 pp. (In Russian).

Phrynocephalus clarkorum

Anderson, S.C.; Leviton, A.E. 1967. A new species of Phrynocephalus (Sauria: Agamidae) from Afghanistan, with remarks on Phrynocephalus ornatus Boulenger. Proceedings of the California Academy of Sciences (Ser. 4) 35: 227-234.

Clark, R.J. 1992. Notes on the distribution and ecology of Phrynocephalus clarkorum Anderson & Leviton 1967 and Phrynocephalus ornatus Boulenger 1887 in Afghanistan. Herpetological Journal 2(4): 140-142.

Phrynocephalus erythrurus

Jin, Y.T.; Liu, N.F. 2010. Phylogeography of Phrynocephalus erythrurus from the Qiangtang Plateau of the Tibetan Plateau. Molecular Phylogenetics and Evolution 54(3): 933-940.

Tang, X.; Ying, X.; Wang, H.; Li, W.; Zhang, Y.; Liang, S.; He, J.; Wang, N.B.; Ma, M.; Chen, Q. 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.

Yang, Y.; Wang, L.; Han, J.; Tang, X.; Ma, M.; Wang, K.; Zhang, X.; Ren, Q.; Chen, Q.; Qiu, Q. 2015. Comparative transcriptomic analysis revealed adaptation mechanism of Phrynocephalus erythrurus, the highest altitude lizard living in the Qinghai-Tibet Plateau. BMC Evolutionary Biology 15: 101.

Yuan, H. 1988. [A survey on the herpetofauna of northern Xizang]. (In Chinese). Acta Herpetologica Sinica 1988(2): 148.

Phrynocephalus forsythii

Sun, Y.Y.; Du, Y.; Yang, J.; Fu, T.B.; Lin, C.X.; Ji, X. 2012. Is the evolution of viviparity accompanied by a relative increase in maternal abdomen size in lizards. Evolutionary Biology (New York) 39(3): 388-399.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

Zhang, Q.; Xia, L.; He, J.B.; Wu, Y.; Fu, J.; Yang, Q. 2010. Comparison of phylogeographic structure and population history of two Phrynocephalus species in the Tarim Basin and adjacent areas. Molecular Phylogenetics and Evolution 57(3): 1091-1104.

Phrynocephalus golubewii

Shenbrot, G.; Semyonov, D.V. 1990. [A new species of the genus Phrynocephalus (Reptilia, Agamidae) from Turkmenia]. (In Russian, English summary). Zoologicheskii Zhurnal 69(9): 154-156.

Phrynocephalus guinanensis

Ji, X.; Wang, Y.; Wang, Z. 2009. New species of Phrynocephalus (Squamata, Agamidae) from Qinghai, Northwest China. Zootaxa 1988: 61-68.

Jin, Y.; Yang, Z.; Brown, R.P.; Liao, P.; Liu, N.F. 2014. 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 71: 288-297.

Wang, Z.; Lu, H.; Li, M.; Ji, X. 2013. Differences in thermal preference and tolerance among three Phrynocephalus lizards (Agamidae) with different body sizes and habitat use. Asian Herpetological Research 4(3): 214-220.

Zhang, K.; Liu, Y.; Tong, H.; Yu, X.; Jin, Y. 2017. [Correlation between the spatial distribution of active cave and insect resources in Phrynocephalus guinanensis (Lacertilia Agamidae)]. (In Chinese). Acta Ecologica Sinica 37(16): 5550-5555

Zhang, K.; Tong, H.; Wo, Y.; Liu, N.; Jin, Y. 2018. Sex ratio and sexual size dimorphism in a Toad-headed Lizard, Phrynocephalus guinanensis. Asian Herpetological Research 9(1): 35-42.

Phrynocephalus guttatus

Ananjeva, N.B. 1981. Phrynocephalus guttatus (Gmelin 1789) - Gefleckter Krötenkopf. pp. 180-190. In: Böhme, W. (ed.). Handbuch der Reptilien und Amphibien Europas. Band 1. Echsen (Sauria) 1. (Gekkonidae, Agamidae, Chamaeleonidae, Anguidae, Amphisbaenidae, Scincidae, Lacertidae 1). Akademische Verlagsgesellschaft, Wiesbaden. 520 pp.

Ananjeva, N.B.; Smirina, E.M.; Nikitina, N.G. 2003. Dentition of Phrynocephalus melanurus: Does tooth number depend on body size and/or age? Russian Journal of Herpetology 10(1): 1-6.

Badmaeva, V.I.; Szczerbak, N.N. 1983. [Phrynocephalus guttatus kalmykus ssp. n. (Sauria, Agamidae) from Kalmykia]. (In Russian, English summary). Vestnik Zoologii 1983(6): 34-37.

Dunaev, E.A.; Belov, V.V. 1989. [Study of feed spectrum of the Zaisan lizard Phrynocephalus melanurus Eichwald, 1831 (Reptilia: Agamidae) based on excrements]. (In Russian). Ekologiya (Moscow) 1989(5): 81-83.

Dunaev, E.A.; Semenov, D.V. 1988. [The lizard species Phrynocephalus melanurus]. (In Russian). Priroda (Moscow) 1988(4): 54-55.

Dunayev, E.A. 1989. [Taxonomic status and peculiarities of ecology and behaviour of Phrynocephalus melanurus Eichwald, 1831 (Reptilia, Agamidae)]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 94(4): 41-53.

Golubev, M.L. 1989. [Phrynocephalus guttatus (Gmel.) or Ph. versicolor Str. (Reptilia, Agamidae): which Phrynocephalus species occurs in Kazakhstan? (In Russian, English summary). Vestnik Zoologii 1989(5): 38-46.

Golubev, M.L.; Gorelov, Y.K.; Dunayev, E.A.; Kotenko, T.I. 1995. [On the finding of Phrynocephalus guttatus (Gmel.) (Sauria, Agamidae) in Turkmeniya and its taxonomic status]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 100(3): 31-39.

Kubykin, R.A. 1977. [Behaviour of the lizard Phrynocephalus guttatus in the lower reaches of the River Ili (southern Pribalkhash'e).] pp. 204-206. (In Russian). In: Manteifel', B.P. (ed.). [Animal group behaviour]. Izdatel'stvo Nauk, Moscow. 452 pp.

Kubykin, R.A. 1977. [Ecological observations on tagged Phrynocephalus guttatus in the lower reaches of the River Il'ya, southern Pribalkhash.] pp. 122-123. In: Darevsky, I.S. [Fourth All-Union Herpetological Conference. Questions in Herpetology]. Akademiya Nauk SSSR, Zoologicheskij Institut. Izdatel'stvo 'Nauka', Leningrad. 243 pp. (In Russian).

Kulikova, G.S.; Semenov, D.V.; Shenbrot, G.I. 1986. [Ecological and behavioural patterns of Phrynocephalus moltschanovi Nik. (Reptilia, Agamidae)]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 91(3): 36-40.

Lantz, L.A. 1922. Revision des reptiles decrits dans le "Journal de Voyage" d'Iwan Lepechin. Bulletin de la Société Zoologique de France 47: 191-194.

Liu, Y.; Shi, L. 2009. [Ontogenetic shifts of sexual dimorphism in Phrynocephalus grumgrzimailoi (Agamidae)]. (In Chinese). Sichuan Journal of Zoology 28(5): 710-713.

Liu, Y.; Song, Y.; Li, W.; Shi, L. 2012. Reproductive strategy and cycle of the Toad-headed Agama Phrynocephalus grumgrzimailoi (Agamidae) in Xinjiang, China. Asian Herpetological Research 3(3): 198-204.

Liu, Y.; Wang, S.; Shi, L. 2011. [Effects of age and sex on diet composition of Phrynocephalus grumgrzimailoi]. (In Chinese). Chinese Journal of Zoology 46(6): 111-117.

Manilo, V.V.; Golubev, M.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.

Manilo, V.V.; Golubyev, M.L. 1994. [Peculiarities of karyotypes of Phrynocephalus species from the group "guttatus" sensu lato (Sauria, Agamidae)]. (In Russian, English summary). Tsitologiya i Genetika 28(2): 66-70.

Mezhzherin, S.; Golubev, M.L. 1993. Allozyme variation and genetic relationships within the Phrynocephalus guttatus species group (Sauria: Agamidae) in the former USSR. Asiatic Herpetological Research 5: 59-64.

Nakaryonok, E.G. 2003. Habitats and ecological peculiarities of Spotted Toad-headed Agama (Phrynocephalus guttatus) of the sandy deserts of the northern Caspian area. Russian Journal of Ecology 2003(1): 77-81.

Polynova, G.V.; Bazshynova, A.V. 2012. [Assessment of the Phrynocephalus guttatus guttatus population number with regard to daily activity of animals]. (In Russian). Zoologicheskii Zhurnal 91(11): 1411-1414.

Rogovin, K.A.; Semenov, D.V. 2002. Demography of translocated population of toad-headed agama Phrynocephalus guttatus in Southern Kalmykya. Russian Journal of Herpetology 9(2): 169-176.

Rogovin, K.A.; Semenov, D.V. 2004. [The toad-headed agama Phrynocephalus guttatus (Reptilia: Agamidae): the correlates of survival in a translocated population in the southwestern Kalmyk Republic]. (In Russian, English summary). Ekologiya (Moscow) 6: 436-441. (Serial normally translated cover to cover in: Russian Journal of Ecology).

Semenov, D.V. 1986. [Individual plots in the lizard Phrynocephalus guttatus (Reptilia, Agamidae)]. (In Russian). Ekologiya (Moscow) 1986(1): 82-85.

Semenov, D.V.; Brushko, Z.K.; Kubykin, R.A.; Shenbrot, G.I. 1987. [Taxonomic position and protective status of the round-headed lizard (Reptilia, Agamidae) in the territory of the USSR]. (In Russian, English summary). Zoologicheskii Zhurnal 66(1): 98-109.

Semenov, D.V.; Rogovin, K.A. 2009. [Application of aposematic elements of body pattern for individual identification of Squamata populations: a case of the toad-headed agama (Phrynocephalus guttatus, Reptilia, Agamidae)]. (In Russian). Zoologicheskii Zhurnal 88(2): 228-233.

Semenov, D.V.; Shenbrot, G.I. 1982. [On species independence of Phrynocephalus moltschanovi (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 61(8): 1194-1204.

Shenbrot, G.I.; Semenov, D.V. 1987. [Present distribution and taxonomy of Phrynocephalus guttatus (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 66(2): 259-272.

Smirina, E.M.; Ananjeva, N.B. 2001. On short-term life span of small desert lizards (by the example of agamids). Zoologicheskii Zhurnal 80(1): 115-117.

Tabachishin, V.G.; Kaybeleva, E.I.; Zavialov, E.V. 2008. Karyotype of spotted toad-headed agama (Phrynocephalus guttatus) from Volgograd region. Selevinia 2007: 155-156.

Tabachishin, V.G.; Kaybeleva, E.I.; Zavialov, E.V.; Shlyakhtin, G.V. 2005. [Ecologo-caryological peculiarities of spotted toad-headed Agama (Phrynocephalus guttatus) in north of the Lower Volga region]. (In Russian, English summary). Povolzhskii Ekologicheskii Zhurnal 2: 180-184.

Tabachishin, V.G.; Zavialov, E.V. 2005. Present distribution and taxonomic status of the spotted toad-headed agama Phrynocephalus guttatus (Gmelin, 1789) in the Volga region, Russia (Squamata: Sauna: Agamidae). Herpetozoa 18(3-4): 141-146.

Veselovsky, M.V.; Vasilevskaya, H.I. 2001. [Investigation of the peripheral nervous system in representatives of genus Phrynocephalus (Reptilia, Agamidae)]. (In Russian, English summary). Vestnik Zoologii 35(1): 63-74, 102.

Wang, Y.; Zeng, X.; Wu, G.F.; Liu, Z.j.; Fang, Z.l.; Papenfuss, T.J.; Macey, R.J. 1996. [A newly recorded species of Phrynocephalus from China]. (In Chinese). Acta Zootaxonomica Sinica 21(1): 9.

Wang, Z.; Li, M.; Shao, M.; Ji, X. 2013. Differences in incubation length and hatchling morphology among five species of oviparous Phrynocephalus lizards (Agamidae) from China. Asian Herpetological Research 4(4): 225-232.

Wittig, W. 2002. Haltung und Nachzucht von Phrynocephalus guttatus. Elaphe 10(3): 29-31.

Wu, M.; Li, P.; Zhao, Y.; Ji, X. 1989. [A study of the karyotype of three species of Phrynocephalus from Xinjiang]. (In Chinese). Zoological Research 10(2): 114,153.

Zeng, X.; Wang, Y.; Liu, Z.; Yu, P. 1994. [A comparative study on the karyotype of three species in Phrynocephalus (Agamidae)]. (In Chinese, English summary). Zoological Research 15(2): 80-84.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

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(2): 535-538.

Phrynocephalus helioscopus

Anonymous. 2015. [Sexual dimorphism and female reproduction of Phrynocephalus helioscopus (Agamidae)]. (In Chinese). Shengtaixue Zazhi 34(6): 1602-1606.

Aliev, T.R.; Akhmedov, S.B.; Gasanov, N.B. 1988. [New data on the herpetofauna of the south eastern part of Azerbaidjan]. (In Russian). Izvestiya Akademii Nauk Azerbaidzhanskoi SSR Seriya Biologicheskikh Nauk 1988(5): 70-73.

Ananjeva, N.B. 1981. Phrynocephalus helioscopus (Pallas 1771) - Sonnengucker. pp. 191-202. In: Böhme, W. (ed.). Handbuch der Reptilien und Amphibien Europas. Band 1. Echsen (Sauria) 1. (Gekkonidae, Agamidae, Chamaeleonidae, Anguidae, Amphisbaenidae, Scincidae, Lacertidae 1). Akademische Verlagsgesellschaft, Wiesbaden. 520 pp.

Ananjeva, N.B.; Tuniyev, B.; Aghasyan, A.; Orlov, N.L.; Tuniyev, S.; Anderson, S.C. 2010. Phrynocephalus helioscopus. The IUCN Red List of Threatened Species 2010: e.T157259A5063556. http://dx.doi.org/10.2305/IUCN.UK.2010-4.RLTS.T157259A5063556.en.

Baran, I.; Kasparek, M.; Oz, M. 1989. On the distribution of four species of agama (Agamidae) in Turkey. Zoology in the Middle East 3: 37-46.

Berezovikov, N.N. 2004. [About record of Trapelus sanguinolentus and Phrynocephalus helioscopus in Malaysary mountains (Dzhungar Alatau)]. (In Russian). Selevinia 2003: 218.

Berezovikov, N.N. 2007. [New findings of Alsophylax pipiens and Phrynocephalus helioscopus in the south-western foothills of Southern Altay]. (In Russian). Selevinia 2006: 213.

Bondarenko, B.M.; Zamolodchikov, D.G. 1991. [Efficiency of different population estimation methods as applied to a model population of Phrynocephalus helioscopus]. (In Russian). Vestnik Zoologii 1991(5): 78-81.

Clemann, N.; Melville, J.; Ananjeva, N.B.; Scroggie, M.P.; Milto, K.; Kreuzberg, E. 2008. Microhabitat occupation and functional morphology of four species of sympatric agamid lizards in the Kyzylkum Desert, central Uzbekistan. Animal Biodiversity and Conservation 31(2): 51-62.

Ivanova, S.A. 1993. [Reproductive cycles of Phrynocephalus reticulatus and Phrynocephalus helioscopus (Sauria, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 72(7): 116-130.

Kühnemann, K. 1985. In meinem Terrarium: Der Sonnengucker - Phrynocephalus helioscopus (Pallas 1771). Elaphe 7(4): 73-75.

Laister, A.F. 1912. [Zur Frage über die gegenseitigen Beziehungen zwischen Phrynocephalus helioscopus Pall. und Phrynocephalus persicus De Fil., sowie deren Verbreitung im Kaukasus]. (In Russian). Charikov Trav. Soc. Nat. 45 [1911-1912]: 73-93.

Manylo, V.V.; Golubev, M.L.; Sattorov, T. 1991. [The karyotype of Phrynocephalus helioscopus saidalievi (Sauria, Agamidae) of the Ferghana Valley]. (In Russian, English summary). Vestnik Zoologii 1991(2): 79-81.

Melkumyan, L.S. 1972. Reproduction period and seasonal changes of lizard body weight in the Aracs river basin. Soviet Journal Ecology 3(3): 279-280.

Meyer, F.; Zinke, O. 1992. Zur Ökologie von Phrynocephalus helioscopus (Pallas, 1771) in der Dschungarischen Gobi (Reptilia; Agamidae). Bonner Zoologische Beiträge 43(1): 131-143.

Obst, F.J. 1959. Krötenköpfe. DATZ (Die Aquarien- und Terrarien-Zeitschrift) 12: 187-188.

Ostroumoff, A. 1889. Zur Entwickelungsgeschichte der Eidechsen (Phrynocephalus helioscopus, Pall.). Trudui Kazan Univ. 19(3): 122 pp.

Peters, G.; Semenov, D.V.; Borkin, L.J. 1990. [On the sunwatcher, Phrynocephalus helioscopus (Pallas, 1771) in Mongolia]. (In Russian, English summary). Trudy Zoologicheskogo Instituta 207: 224-229, 235.

Petzold, H.G. 1960. Interessante mittelasiatische Eidechsenarten im Tierpark Berlin. Aquarien Terrarien 6: 373-377.

Rogovin, K.A. 1991. [Social behavior of Phrynocephalus helioscopus and Ph. reticulatus (Reptilia, Agamidae) and their relationships in joint settlements]. (In Russian, English summary). Zoologicheskii Zhurnal 70(3): 61-72.

Rustamov, A.K.; Shammakov, S. 1967. Ecology of Phrynocephalus helioscopus helioscopus Pallas in Turkmenia. Zoologicheskii Zhurnal 46: 741-748.

Saparova, A. 1989. [On behaviour of Phrynocephalus h. helioscopus in clay desert of south-western Turkmenistan]. (In Russian, English summary). Izvestiya Akademii Nauk Turkmenskoi SSR Seriya Biologicheskikh Nauk 1989(5): 33-41.

Sattarov, T.S. 1981. [Phrynocephalus helioscopus saidalievi ssp. n. (Sauria, Reptilia) - a new subspecies of Ph. helioscopus from the Ferghana Valley]. (In Russian, English summary). Vestnik Zoologii 1981(1): 73-75.

Sattarov, T.S. 1984. [On the ecology and distribution of Phrynocephalus helioscopus saidaliewi Sattorow, 1981 in northern Tadzhikistan]. (In Russian). Izvestiya Akademii Nauk Tadzhikskoi SSR Otdelenie Biologicheskikh Nauk 1984(2): 71-76.

Schleich, H.H. 1976. Über Phrynocephalus helioscopus aus Persien (Reptilia, Sauria, Agamidae). Salamandra 12(4): 189-193.

Schwenk, K.; Bell, D.A. 1988. A cryptic intermediate in the evolution of chameleon tongue projection. Experientia (Basel) 44(8): 697-700.

Schwenk, K.; Greene, H.W. 1987. Water collection and drinking in Phrynocephalus helioscopus: a possible condensation mechanism. Journal of Herpetology 21(2): 134-139.

Semenov, D.V. 2007. [The comparative aspects of spatial ecology of lizards exemplified by the toad-headed lizards (Reptilia, Agamidae, Phrynocephalus)]. (In Russian, English summary). Zhurnal Obshchei Biologii 68(5): 379-393.

Semenov, D.V.; Brushko, Z.K.; Kubykin, R.A.; Shenbrot, G.I. 1987. [Taxonomic position and protective status of the round-headed lizard (Reptilia, Agamidae) in the territory of the USSR]. (In Russian, English summary). Zoologicheskii Zhurnal 66(1): 98-109.

Shenbrot, G.I. 1987. [Dynamics of the Phrynocephalus helioscopus and Ph. reticulatus populations (Reptilia, Agamidae) in the south of Bukhara district]. (In Russian, English summary). Zoologicheskii Zhurnal 66(5): 787-789.

Shenbrot, G.I.; Antonova, G.S. 1981. [Comparative ecology of Phrynocephalus helioscopus and P. reticulatus in the conditions of their common habitat, south of Bukhara region (Uzbek SSR)]. pp. 156-157. In: Darevsky, I.S.; Ananjeva, N.B.; Barkagan, Z.S.; Borkin, L.Y.; Sokolova, T.M. & Szczerbak, N.N. (eds.). The problems of herpetology: abstracts. Nauka, Leningrad. 163 pp. (abstract, in Russian).

Sokolov, V.E.; Darevsky, I.S.; Kotova, E.L.; Chernova, O.F. 1997. [Specialized skin organs of Phrynocephalus helioscopus (Reptilia, Squamata, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 76(4): 466-472.

Solovyeva, E.N.; Dunayev, E.A.; Poyarkov, N.A. 2012. [Interspecific taxonomy of sunwatcher toadhead agama species complex (Phrynocephalus helioscopus, Squamata)]. (In Russian). Zoologicheskii Zhurnal 91(11): 1377-1396.

Wang, Z.; Li, M.; Shao, M.; Ji, X. 2013. Differences in incubation length and hatchling morphology among five species of oviparous Phrynocephalus lizards (Agamidae) from China. Asian Herpetological Research 4(4): 225-232.

Zander, A. 1895. Einige transkaspische Reptilien. Zoologische Garten 36: 297-305.

Phrynocephalus hongyuanensis

Jiang, Y.; Huang, Q.; Zhao, E. 1980. [A new subspecies of Phrynocephalus vlangalii (Strauch) and preliminary observations on its ecology]. (In Chinese, English summary). Acta Zoologica Sinica 26(2): 178-183.

Liu, L.; Guo, X.G.; Wang, Y. 2008. [Genetic variation and diversity of Phrynocephalus vlangalii hongyuanensis in Zoige Wetland inferred from ND4-tRNAleu gene]. (In Chinese). Zoological Research 29(2): 121-126.

Macey, J.R.; Schulte, J.A.; Ananjeva, N.B.; Dyke, E.T. van; Wang, Y.; Orlov, N.; Shafiei, S.; Robinson, M.D.; Dujsebayeva, T.; Freund, G.S.; Fischer, C.M.; Liu, D.; Papenfuss, T. 2018. A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. Zootaxa 4467(1): 1–81.

Wan, H.; Qi, Y.; Guo, X.G.; Wang, Y. 2011. [Preliminary study on sexual dimorphism and seasonal variation of the belly patch and tail-tip badge of the adult toad-headed lizard Phrynocephalus vlangalii hongyuanensis]. (In Chinese). Sichuan Journal of Zoology 30(1): 54-58.

Wang, Y.; Jiang, Y. 1992. [Study on the taxonomy of Phrynocephalus vlangalii hongyuanensis - a new species of the genus of Phrynocephalus (Reptilia: Agamidae). pp. 111-115]. (In Chinese, English summary). In: Jiang, Y. (ed.). Collected papers on herpetology. Sichuan Publishing House of Science and Technology, Chengdu. 157 pp.

Wang, Y.; Zeng, X.M.; Fang, Z.L.; Wu, G.; Liu, Z.J.; Papenfuss, T.J.; Macey, J.R. 2002. [A valid species of the genus Phrynocephalus: P. putjatia and a discussion on taxonomy of Phrynocephalus hongyuanensis (Sauria: Agamidae)]. (In Chinese, English summary). Acta Zootaxonomica Sinica 27(2): 372-383.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

Phrynocephalus horvathi

Cicek, K.; Dincer, A.; Varol, T.C.; Tayhan, Y. 2011. Data on food composition of Phrynocephalus horvathi Mehely, 1894 (Reptilia: Agamidae) in Mount Ararat (Northeastern Anatolia, Turkey). Ecologia Balkanica 3(1): 69-73.

Cicek, K.; Kumas, M.; Ayaz, D.; Tok, C.V. 2012. Preliminary data on the age structure of Phrynocephalus horvathi in Mount Ararat (Northeastern Anatolia, Turkey). Biharean Biologist 6(2): 112-115.

Melnikov, D.A.; Ananjeva, N.B.; Agasyan, A.L.; Rajabizadeh, Z. 2008. Historical background and taxonomic status of the Persian Toadhead agama, Phrynocephalus persicus de Filippi, 1863 and Horvath’s Sun Watcher Toadhead Agama Phrynocephalus helioscopus horvathi Mehely, 1894. pp. 286-297. In: Ananjeva, N.B.; Danilov, I.G.; Dunayev, E.A.; Ischenko, V.G.; Lada, G.A.; Litvinchuk, S.N.; Orlova, V.F.; Smirina, E.M.; Tuniyev, B.S. & Khalikov, R.G. (eds.). The problems of herpetology. Proceedings of the 3rd meeting of the Nikolsky Herpetological Society, 9-13 October 2006, Putschino. Russian Academy of Sciences, St. Petersburg. 467 pp.

Tosunoglu, M.; Gul, C.; Topyildiz, H.; Uysal, I. 2011. Notes on distribution, ecology, and morphological characters of Phrynocephalus helioscopus horvathi Mehely, 1894 from Northeast Anatolia. Russian Journal of Herpetology 18(4): 247-252.

Yenmis, M.; Dinçer, A.; Sherbrooke, W.C.; Vesely, M. 2016. A comparative behavioural and structural study of rain-harvesting and non-rain-harvesting agamid lizards of Anatolia (Turkey). Zoomorphology (Berlin) 135(1): 137-148.

Phrynocephalus interscapularis

Cherlin, V.A. 1989. [Age differences in the thermobiological characteristics of some lizards]. (In Russian). Izvestiya Akademii Nauk Turkmenskoi SSR Seriya Biologicheskikh Nauk 1989(1): 35-38.

Cherlin, V.A.; Muzychenko, I.V. 1983. [Thermobiology of Eremius grammica, Phrynocephalus mystaceus and Ph. interscapularis in east Karakumy in summer]. (In Russian, English summary). Zoologicheskii Zhurnal 62(6): 897-908.

Cherlin, V.A.; Muzychenko, I.V. 1984. [A case study of determination methods of body temperature critical maximum in some Middle Asia lizards]. (In Russian, English summary). Izvestiya Akademii Nauk Turkmenskoi SSR Seriya Biologicheskikh Nauk 1984(5): 73-76.

Cherlin, V.A.; Muzychenko, I.V. 1988. [Seasonal variations in thermobiological indices of some arid lizards]. (In Russian, English summary). Zoologicheskii Zhurnal 67(3): 406-416.

Clemann, N.; Melville, J.; Ananjeva, N.B.; Scroggie, M.P.; Milto, K.; Kreuzberg, E. 2008. Microhabitat occupation and functional morphology of four species of sympatric agamid lizards in the Kyzylkum Desert, central Uzbekistan. Animal Biodiversity and Conservation 31(2): 51-62.

Dujsebayeva, T.N. 1994. [The topography and numerical distribution of skin sense organs in the skin of Phrynocephalus interscapularis (Lacertilia: Agamidae)]. (In Russian, English summary). Selevinia 2(4): 10-14.

Kamalova, Z.Y. 1978. [On the ecology of Phrynocephalus interscapularis L. in south west Kyzyl-Kum]. (In Russian). Uzbekskii Biologicheskii Zhurnal 1978(2): 52-54.

Kukikova, G.S.; Semenov, D.V. 1984. [On the life-span and growth of Phrynocephalus interscapularis]. (In Russian, English summary). Zoologicheskii Zhurnal 63(5): 786-787.

Kulik, I.L. 1971. A study of home ranges of desert lizards. Zoologicheskii Zhurnal 50: 300-302.

Mishagina, Z.V. 1988. [Trophic connections of Phrynocephalus interscapularis, Agamidae in the east Karakum]. (In Russian, English summary). Problemy Osvoeniya Pustyn' 1988(2): 48-55.

Orlov, V.N.; Milishnikov, A.N.; Pobedonostseva, E.Y.; Likhnova, O.P. 1988. [Data obtained in the study of the population structure of Phrynocephalus interscapularis (Agamidae) by morphological and electrophoretic methods]. (In Russian, English summary). Zoologicheskii Zhurnal 67(11): 1689-1698.

Pawley, R. 1994. Herping in the former USSR. Vivarium (Lakeside) 6(2): 30-37.

Rogovin, K.A. 1989. [Communicative behaviour of Phrynocephalus interscapularis]. (In Russian, English summary). Zoologicheskii Zhurnal 68(7): 97-109.

Rogovin, K.A.; Semenov, D.V.; Gallina, S.; Mauri, M.E. 1982. [On the niche overlap in time and space in lizard communities of the Repetek Biosphere Reserve]. (In Russian). Doklady Akademii Nauk SSSR 264(4): 1016-1018.

Semenov, D.V.; Kulikova, G.S. 1983. [Home ranges and migrations of Phrynocephalus interscapularis (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 62(8): 1209-1220.

Serebrennikov, M.K. 1933. Biologische Skizzen über die Wirbeltiere des Südwestlichen Kizilkum. Aquarium (Berlin) 1933: 8-13, 50-54.

Smirina, E.; Ananjeva, N.B. 2017. On the longevity, growth and reproductive characteristics of Lichtenstein’s Toadhead Agama, Phrynocephalus interscapularis Lichtenstein, 1856 (Agamidae, Sauria). Amphibia-Reptilia 38(1): 31–39.

Ulmasov, K.; Zatsepina, O.; Molodtsov, V.; Evgenjev, M. 1999. Natural body temperature and kinetics of heat-shock protein synthesis in the toad-headed agamid lizard Phrynocephalus interscapularis. Amphibia-Reptilia 20(1): 1-9.

Ulmasov, K.; Zatsepina, O.G.; Rybtsov, S.A.; Dzhumageldyev, B.T.; Evgenev, M.B. 1997. [Some aspects of the state of heat shock system in lizards of different ecological niches]. (In Russian, English summary). Izvestiya Rossiiskoi Akademii Nauk Seriya Biologicheskaya 2: 133-141.

Zander, A. 1895. Einige transkaspische Reptilien. Zoologische Garten 36: 297-305.

Zatsepina, O.G.; Ulmasov, K.A.; Beresten, S.F.; Molodtsov, V.B.; Rybtsov, S.A.; Evgenjev, M.B. 2000. Thermotolerant desert lizards characteristically differ in terms of heat-shock system regulation. Journal of Experimental Biology 203(6): 1017-1025.

Phrynocephalus longicaudatus

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Phrynocephalus lutensis

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Phrynocephalus luteoguttatus

Chandra, H.; Charan, S. 1983. Some lizard predators of acridids of southwest Afghanistan. Plant Protection Bulletin (Faridabad) 33(3-4): 153-155.

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Phrynocephalus maculatus

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Phrynocephalus mystaceus

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Phrynocephalus nejdensis

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Phrynocephalus ornatus

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Phrynocephalus persicus

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Xue, L.; Jiang, Z.; Ping, X.; Tang, S.; Bi, J.; Li, C. 2012. Spatial distribution pattern of the Steppe Toad-headed Lizard (Phrynocephalus frontalis) and its influencing factors. Asian Herpetological Research 3(1): 46-51.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

Zeng, Z.G.; Zhao, J.M.; Sun, B. 2013. Life history variation among geographically close populations of the toad-headed lizard (Phrynocephalus przewalskii): exploring environmental and physiological associations. Acta Oecologica 51: 28-33.

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(2): 535-538.

Zhao, W.; Liu, N.F. 2013. Sexual dimorphism of head size in Phrynocephalus przewalskii: testing the food niche divergence hypothesis. Asian Herpetological Research 4(4): 242-247.

Zhao, W.; Liu, N.F. 2014. The proximate causes of sexual size dimorphism in Phrynocephalus przewalskii. PLoS ONE 9(1): e85963.

Zhao, W.; Wang, Y.J.; Nan-Nan, Y.; Liu, N.F. 2010. [Reproduction of two Phrynocephalus przewalskii populations. pp. 327-331]. (In Chinese). In: Ji, X. Herpetologica Sinica 12. Chinese Scientific Books, Beijing. 409 pp.

Zhao, W.; Yu, N.; Wang, Y.; Ji, W.; Liu, N.F. 2011. Female reproductive cycles of Phrynocephalus przewalskii (Lacertilia: Agamidae) in the Tengger Desert, China. Asian Herpetological Research 2(1): 30-35.

Zhao, X.; Bi, J.H. 2014. [Sexual dimorphism in juvenile toad-headed lizard (Phrynocephalus frontalis)]. (In Chinese). Chinese Journal of Zoology 49(2): 274-279.

Phrynocephalus putjatai

Jin, Y.; Yang, Z.; Brown, R.P.; Liao, P.; Liu, N.F. 2014. 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 71: 288-297.

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): 1-15.

Wang, Y.; Zeng, X.M.; Fang, Z.L.; Wu, G.; Liu, Z.J.; Papenfuss, T.J.; Macey, J.R. 2002. [A valid species of the genus Phrynocephalus: P. putjatia and a discussion on taxonomy of Phrynocephalus hongyuanensis (Sauria: Agamidae)]. (In Chinese, English summary). Acta Zootaxonomica Sinica 27(2): 372-383.

Yang, Y.; Wang, L.; Han, J.; Tang, X.; Ma, M.; Wang, K.; Zhang, X.; Ren, Q.; Chen, Q.; Qiu, Q. 2015. Comparative transcriptomic analysis revealed adaptation mechanism of Phrynocephalus erythrurus, the highest altitude lizard living in the Qinghai-Tibet Plateau. BMC Evolutionary Biology 15: 101.

Phrynocephalus raddei

Semenov, D.V. 2007. [The comparative aspects of spatial ecology of lizards exemplified by the toad-headed lizards (Reptilia, Agamidae, Phrynocephalus)]. (In Russian, English summary). Zhurnal Obshchei Biologii 68(5): 379-393.

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Shammakov, S.; Saparova, A. 1986. [Individual plots and homing of Phrynocephalus raddei and Phrynocephalus maculatus]. (In Russian, English summary). Izvestiya Akademii Nauk Turkmenskoi SSR Seriya Biologicheskikh Nauk 1986(1): 74-77.

Zander, A. 1895. Einige transkaspische Reptilien. Zoologische Garten 36: 297-305.

Phrynocephalus reticulatus

Baytin, A.D. 1990. [Influence of an anthropogenic factor on reptiles from south-west Tadzhikistan]. (In Russian). Izvestiya Akademii Nauk Tadzhikskoi SSR Otdelenie Biologicheskikh Nauk 1990(2): 48-49.

Golubev, M.L. 1991. [About the name Agama ocellata Lichtenstein in Eversmann, 1823 (Reptilia, Agamidae) with redescription of the types]. (In Russian, English summary). Gerpetologicheskie Issledovaniya 1: 12-17, 171.

Ivanova, S.A. 1993. [Reproductive cycles of Phrynocephalus reticulatus and Phrynocephalus helioscopus (Sauria, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 72(7): 116-130.

Kamalova, Z.Y. 1971. Feeding habits of Phrynocephalus reticulatus Strauchi Nik. in the sands of the Fergana Valley during the summer season. Ecology 1970(5): 457-458.

Mezhzherin, S.; Golubev, M.L. 1998. [Allozyme variation and genetic differentiation of toad agama species group Phynocephalus [Phrynocephalus] ocellatus (=reticulatus) (Reptilia: Agamidae)]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 103(6): 9-16.

Nuridjanov, D.; Nazarov, R.; Shestapol, A.; Ananjeva, N.B. 2017. Phrynocephalus reticulatus (errata version published in 2018). The IUCN Red List of Threatened Species 2017: e.T47755693A128947879. http://dx.doi.org/10.2305/IUCN.UK.2017-2.RLTS.T47755693A47755699.en.

Rogovin, K.A. 1991. [Social behavior of Phrynocephalus helioscopus and Ph. reticulatus (Reptilia, Agamidae) and their relationships in joint settlements]. (In Russian, English summary). Zoologicheskii Zhurnal 70(3): 61-72.

Sattarov, T.S. 1981. [On species independence of Phrynocephalus strauchi Nik. (Reptilia, Sauria) from the Ferghana Valley]. (In Russian, English summary). Vestnik Zoologii 1981(6): 82-85.

Sattarov, T.S.; Said-Aliev, S.A. 1980. [New data on distribution & ecology of lizards in northern Tadzhikistan]. (In Russian). Izvestiya Akademii Nauk Tadzhikskoi SSR Otdelenie Fiziko-tekhnicheskikh I Khimicheskikh Nauk 1980(1): 106-109.

Sattarov, T.S.; Said-Aliev, S.A. 1980. [New data on distribution and ecology of Phrynocephalus reticulatus in north Tadzhikistan]. (In Russian). Izvestiya Akademii Nauk Tadzhikskoi SSR Otdelenie Biologicheskikh Nauk 1980(1): 106-109.

Semenov, D.V. 2007. [The comparative aspects of spatial ecology of lizards exemplified by the toad-headed lizards (Reptilia, Agamidae, Phrynocephalus)]. (In Russian, English summary). Zhurnal Obshchei Biologii 68(5): 379-393.

Semenov, D.V.; Shenbrot, G.I. 1982. [On species independence of Phrynocephalus moltschanovi (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 61(8): 1194-1204.

Shenbrot, G.I. 1987. [Dynamics of the Phrynocephalus helioscopus and Ph. reticulatus populations (Reptilia, Agamidae) in the south of Bukhara district]. (In Russian, English summary). Zoologicheskii Zhurnal 66(5): 787-789.

Shenbrot, G.I.; Antonova, G.S. 1981. [Comparative ecology of Phrynocephalus helioscopus and P. reticulatus in the conditions of their common habitat, south of Bukhara region (Uzbek SSR)]. pp. 156-157. In: Darevsky, I.S.; Ananjeva, N.B.; Barkagan, Z.S.; Borkin, L.Y.; Sokolova, T.M. & Szczerbak, N.N. (eds.). The problems of herpetology: abstracts. Nauka, Leningrad. 163 pp. (abstract, in Russian).

Vashetko, E.V.; Kamalova, Z.Y.; Yadgarov, T. 1974. [Systematic review of vertebrates. Reptilia.] pp. 60-84. (In Russian). In: Sultanov, G.S. (Ed.) (Vertebrates of the Fergana Valley). Fan, Tashkent. 217 pp.

Phrynocephalus rossikowi

Ataev, C.A.; Shammakov, S.M. 1990. [New data on the distribution and abundance of some reptiles at the limit of their range in Turkmenistan]. (In Russian). Izvestiya Akademii Nauk Turkmenskoi SSR Seriya Biologicheskikh Nauk 1990(3): 69-71.

Bondarenko, D.A. 1982. [Summer-autumn ecology of Phrynocephalus rossikowi]. (In Russian). Vestnik Zoologii 1982(5): 55-59.

Bondarenko, D.A. 1984. [Phrynocephalus rossikowi]. (In Russian). Priroda (Moscow) 1984(4): 48-49.

Golubev, M.L.; Manilo, V.V.; Tokar, A. 1994. Geographic variability of Phrynocephalus rossikowi Nik. (Reptilia: Agamidae) in Turkmenistan and adjacent regions. Monographiae Biologicae 72: 351-364.

Nikolsky, A.M. 1898. On two new lizards from Russia. Annuaire Mus. St. Petersb. 1898: 284-288.

Szczerbak, N.N.; Golubev, M.L. 1979. [A new subspecies of Phrynocephalus rossikowi shammakowi ssp. n. (Reptilia, Sauria, Agamidae) from the central Kara-Kum Desert]. (In Russian, English summary). Vestnik Zoologii 1979(6): 81-83.

Phrynocephalus sakoi

Melnikov, D.; Melnikova, E.; Nazarov, R.; Al-Johany, A.; Ananjeva, N.B. 2015. A new species of Phrynocephalus (Agamidae, Sauria) from Al Sharqiyah Sands, northeastern Oman, dedicated to the memory of Sako Tuniyev (1983-2015). Russian Journal of Herpetology 22(4): 301-309.

Phrynocephalus scutellatus

Kafash, A.; Malakoutikhah, S.; Yousefi, M.; Ataei, F.; Heidari, H.; Rastegar-Pouyani, E. 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): 47-54.

Mocquard, F. 1910. Rectification du nom specifique de Phrynocephalus olivieri Dum. et Bibr. Bulletin du Muséum National d’Histoire Naturelle (Paris) 1910: 13-15.

Rahimian, H.; Shafiei, S.; Rastegar-Pouyani, N.; Rastegar-Pouyani, E. 2015. Phylogenetic relationships of the gray-toad agama, Phrynocephalus scutellatus (Olivier, 1807), species complex from Iran. Zootaxa 3990(3): 369–380.

Sharma, R.C.; Singh, C. 1976. Records of some little known reptiles from Afghanistan. Newsletter of the Zoological Survey of India 2(2): 70-74.

Phrynocephalus sogdianus

Macey, J.R.; Schulte, J.A.; Ananjeva, N.B.; Dyke, E.T. van; Wang, Y.; Orlov, N.; Shafiei, S.; Robinson, M.D.; Dujsebayeva, T.; Freund, G.S.; Fischer, C.M.; Liu, D.; Papenfuss, T. 2018. A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. Zootaxa 4467(1): 1–81.

Rzepakovsky, V.T. 1981. [Material on the study of ecology of Phrynocephalus sogdianus in southwest Tadzhikistan]. pp. 114-115. In: Darevsky, I.S.; Ananjeva, N.B.; Barkagan, Z.S.; Borkin, L.Y.; Sokolova, T.M. & Szczerbak, N.N. (eds.). The problems of herpetology: abstracts. Nauka, Leningrad. 163 pp. (abstract, in Russian).

Phrynocephalus strauchi

Bondarenko, D.A.; Peregontsev, E.A.; Abdunazarov, B.B.; Sudarev, V.O. 2010. [On the recent state of herpetofauna of sand massif in the Ferghana Valley (Uzbekistan)]. (In Russian). pp. 246-248. In: Duisebayeva, T.N.; Zima, I.A. & Chirikova, M.A. (eds.). Gerpetologicheskie Issledovaniya v Kazakhstane i sopredelnykh stranakh [Herpetological researches in Kazakhstan and adjacent countries]. ASBK & SOPKI, Almaty. 255 pp. (In Russian).

Dunayev, E.A. 1995. Reviewed description of the types of Phrynocephalus strauchi Nikolsky, 1899 (Squamata, Agamidae) and materials on the history of its study, distribution, and variability. Russian Journal of Herpetology 2(2): 87-94.

Sattarov, T.S. 1981. [On species independence of Phrynocephalus strauchi Nik. (Reptilia, Sauria) from the Ferghana Valley]. (In Russian, English summary). Vestnik Zoologii 1981(6): 82-85.

Phrynocephalus theobaldi

Barabanov, A.V.; Ananjeva, N.B.; Papenfuss, T.J.; Wang, Y. 2002. A new name for Phrynocephalus theobaldi orientalis Wang, Papenfuss et Zeng, 1999. Russian Journal of Herpetology 9(1): 80.

Huang, Y. 1979. [Herpetofauna of Ali, Tibet]. pp. 73-79, illustr. (In Chinese). In: Qinghai Sheng Shengwu Yanjiusuo (ed.). Xizang ali diqu dongzhiwu Kaocha. [Plants and animals of Ali region, Xizang.] Science Press, Peking. 227 pp.

Jin, Y.; Liao, P. 2015. An elevational trend of body size variation in a cold-climate agamid lizard, Phrynocephalus theobaldi. Current Zoology 61(3): 444-453.

Jin, Y.; Liu, N.; Brown, R.P. 2017. The geography and timing of genetic divergence in the lizard Phrynocephalus theobaldi on the Qinghai-Tibetan plateau. Scientific Reports 7: 2281

Wang, Y.; Zeng, X.; Fang, Z.; Liu, Z.; Wu, G.; Papenfuss, T.J.; Macey, R.J. 1996. [A new species of the genus Phrynocephalus - Phrynocephalus zetanensis sp. nov]. (In Chinese, English summary). Zoological Research 17(1): 27-29.

Zeng, X.; Wang, Y.; Liu, Z.; Yu, P. 1994. [A comparative study on the karyotype of three species in Phrynocephalus (Agamidae)]. (In Chinese, English summary). Zoological Research 15(2): 80-84.

Zeng, X.; Wang, Y.; Liu, Z.J.; Fang, Z.L.; Wu, G.F.; Papenfuss, T.J.; Macey, R.J. 1997. [Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus]. (In Chinese, English summary). Acta Zoologica Sinica 43(4): 399-410.

Phrynocephalus turcomanus

Macey, J.R.; Schulte, J.A.; Ananjeva, N.B.; Dyke, E.T. van; Wang, Y.; Orlov, N.; Shafiei, S.; Robinson, M.D.; Dujsebayeva, T.; Freund, G.S.; Fischer, C.M.; Liu, D.; Papenfuss, T. 2018. A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. Zootaxa 4467(1): 1–81.

Solovyeva, E.N.; Dunayev, E.A.; Poyarkov, N.A. 2012. [Interspecific taxonomy of sunwatcher toadhead agama species complex (Phrynocephalus helioscopus, Squamata)]. (In Russian). Zoologicheskii Zhurnal 91(11): 1377-1396.

Phrynocephalus versicolor

Ananjeva, N.B. 1986. [Bau und Morphometrie des Schädels der Buntkrötenkopfagame (Phrynocephalus versicolor Str.) aus Echijn-Gol der Mongolei]. pp. 179-196, 236. (In Russian, German summary). In: Vorobeva, E.I. (ed.). Gerpetologicheskie issledovaniya v mongolskoi narodnoi respublike. [Herpetological studies in Mongolia.] Acad. Sci. USSR, Moscow. 247 pp.

Berdibaeva, Z.S. 1986. [Short notes on Phrynocephalus versicolor]. (In Russian). In: Gvozdev, E.V. (ed.). Rare species of animals of Kazakhstan. Nauka, Alma Ata, Kazakhstan. 246 pp. Pp. 178 pp.

Borkin, L.Y.; Semenov, D.V. 1986. [The temperature and the daily activity in the motley round-headed lizard, Phrynocephalus versicolor. (Reptilia, Agamidae) in the Transaltai Gobi Desert]. (In Russian, English summary). Zoologicheskii Zhurnal 65(11): 1655-1663.

Brushko, Z.K.; Kubykin, R.A. 1983. [Reproduction of a lizard Phrynocephalus versicolor at the plain of the upper reaches of the Ili River]. (In Russian). Ekologiya (Moscow) 14(4): 70-72.

Chang, C.; Chen, Q.; Liu, N. 1993. [The coexist mechanisms of two species of lizards in the same community]. (In Russian, English summary). Journal of Lanzhou University Natural Sciences 29(3): 180-183.

Golubev, M.L. 1989. [Phrynocephalus guttatus (Gmel.) or Ph. versicolor Str. (Reptilia, Agamidae): which Phrynocephalus species occurs in Kazakhstan? (In Russian, English summary). Vestnik Zoologii 1989(5): 38-46.

Golubev, M.L. 1992. [Variegated toad agama Phrynocephalus versicolor (Reptilia, Agamidae) of the Djungar Gate (East Kazakhstan), with notes on systematics of the species]. (In Russian, English summary). Vestnik Zoologii 1992(2): 31-38.

Golubev, M.L. 1993. The variegated toad agama in Djungar Gate (eastern Kazakstan [sic]) with notes on certain systematic problems of Phrynocephalus versicolor str. (Reptilia: Agamidae). Asiatic Herpetological Research 5: 51-58.

Guo, L.; Zhao, H.Y. 2008. [Comparison of the morpha and structure of two populations of Phrynocephalus immaculatus in Inner Mongolia]. (In Chinese). NeiMongol Daxue Xuebao Ziran Kexue Ban 39(5) (Sum 161): 552-554.

Guo, L.; Zhao, H.Y. 2009. [A study on the interspecific relationship of lizards in Urad Houqi county desert in Inner Mongolia]. (In Chinese). NeiMongol Daxue Xuebao Ziran Kexue Ban 40(6) (Sum 168): 694-698.

Kubykin, R.A.; Brushko, Z.K. 1990. [Important representatives of rare species. Phrynocephalus versicolor. pp. 217-228]. (In Russian). In: Kovshar, A.F. (ed.). Redkie zhivotnye pustyn': problemy sokhraneniya genofonda pozvonochnykh Kazakhstana. Rare animals of desert regions: problems of protecting Kazakhstan vertebrates genofond [sic]. Nauka Kazakhstan, Alma Ata. 254 pp.

Milishnikov, A.N.; Lichnova, O.A. 1986. [Einige Besonderheiten der genetischen Variabilität in der Populationen von Buntkrötenkopfagame (Phrynocephalus versicolor Str.) der Mongolei und Tuwa. pp. 143-151, 239-240]. (In Russian, German summary). In: Vorobeva, E.I. (ed.). Gerpetologicheskie issledovaniya v mongolskoi narodnoi respublike. [Herpetological studies in Mongolia.] Acad. Sci. USSR, Moscow. 247 pp.

Murdoch, J.D.; Davie, H.; Galbadrah, M.; Donovan, T.; Reading, R.P. 2013. Do Siberian marmots influence toad-headed agama occupancy? Examining the influence of marmot colonies and three steppe habitats in Mongolia. Journal of Arid Environments 92: 76-80.

Murdoch, J.D.; Suuri, B.; Reading, R.P. 2010. Estimates of toad headed agama density in three steppe habitats of Mongolia. Erforschung Biologischer Ressourcen der Mongolei 11: 383-389.

Obst, F.J. 1958. Hinweise zur Pflege einiger mittelasiatischer Echsen. DATZ (Die Aquarien- und Terrarien-Zeitschrift) 11: 149-151.

Oddie, K.R.; Ariunaa, T.; Enkhnasan, D. 2003. Ecological sampling design and observer bias: an example from toad-headed agama (Phrynocephalus versicolor) in the Southern Gobi, Mongolia. Mongolian Journal of Biological Sciences 1(1): 49-54.

Qu, Y.F.; Gao, J.F.; Mao, L.X.; Ji, X. 2011. Sexual dimorphism and female reproduction in two sympatric toad-headed lizards, Phrynocephalus frontalis and P. versicolor (Agamidae). Animal Biology (Leiden) 61(2): 139-151.

Qu, Y.F.; Li, H.; Gao, J.F.; Xu, F.; Ji, X. 2011. Thermal preference, thermal tolerance and the thermal dependence of digestive performance in two Phrynocephalus lizards (Agamidae), with a review of species studied. Current Zoology 57(6): 684-700.

Qu, Y.F.; Li, H.; Zhou, T.; Gao, J.F.; Ji, X. 2005. [Inter- and intra-specific differences in morphological traits in three species of toad-headed lizards from Inner Mongolia]. (In Chinese). In: Zhou, K. & Ji, X. (eds.). Herpetologica Sinica 10. Quinghai People's Press. 398 pp.

Rogovin, K.A.; Semenov, D.V.; Shenbrot, G.I. 2001. Lizards of the northern Mongolian deserts: densities and community structure. Asiatic Herpetological Research 9 [2000]: 113-121.

Semenov, D.V. 1985. [The behaviour of Phrynocephalus versicolor (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 64(10): 1545-1555.

Semenov, D.V. 1986. [Feeding of the motley round-headed lizard Phrynocephalus versicolor (Reptilia, Agamidae) in south Mongolia]. (In Russian, English summary). Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii 91(6): 16-28.

Semenov, D.V. 1986. [Materialien zur Variabilität und innerartlicher Systematik der Buntkrötenkopfagame (Phrynocephalus versicolor Str.). pp. 157-173, 241]. (In Russian, German summary). In: Vorobeva, E.I. (ed.). Gerpetologicheskie issledovaniya v mongolskoi narodnoi respublike. [Herpetological studies in Mongolia.] Acad. Sci. USSR, Moscow. 247 pp.

Semenov, D.V. 1991. [Variability of spatial distribution parameters of individuals in population of the lizard Phrynocephalus versicolor Str. (Reptilia, Agamidae)]. (In Russian). Ekologiya (Moscow) 1991(3): 77-81.

Semenov, D.V.; Borkin, L.Y. 1985. [Migrations and individual ranges in Phrynocephalus versicolor (Reptilia, Agamidae) in Transaltaian Gobi, Mongolia]. (In Russian, English summary). Zoologicheskii Zhurnal 64(2): 252-263.

Semenov, D.V.; Brushko, Z.K.; Kubykin, R.A.; Shenbrot, G.I. 1987. [Taxonomic position and protective status of the round-headed lizard (Reptilia, Agamidae) in the territory of the USSR]. (In Russian, English summary). Zoologicheskii Zhurnal 66(1): 98-109.

Semenov, D.V.; Shenbrot, G.I. 1982. [On species independence of Phrynocephalus moltschanovi (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 61(8): 1194-1204.

Semenov, D.V.; Smirina, E.M. 1988. [Population dynamics, demographic structure and reproductive parameters of the lizard species Phrynocephalus versicolor in Mongolia]. pp. 110-112. (In Russian). In: Anonymous. Ekologiya populyatsii. Chast 1. Tezisy dokladov vsesoyuznogo soveshchaniya (4-6 Oktyabrya 1988, Novosibirsk). [Population ecology. Part 1. Abstracts and short papers from an All-Union Conference (4-6th Oct. 1988, Novosibirsk)]. Acad. Sci. USSR, Moscow. 304 pp.

Semyonov, D.V.; Shenbrot, G.I. 1989. [Taxonomic position of Phrynocephalus versicolor (Reptilia, Agamidae) of Mongolia]. (In Russian, English summary). Zoologicheskii Zhurnal 68(12): 79-87.

Smirina, E.M.; Semenov, D.V. 1985. [New data on movements of Phrynocephalus versicolor (Reptilia, Agamidae)]. (In Russian, English summary). Zoologicheskii Zhurnal 64(8): 1272-1274.

Tang, X.; Yue, F.; Ma, M.; Wang, N.B.; He, J.; Chen, Q. 2012. Effects of thermal and hydric conditions on egg incubation and hatchling phenotypes in two Phrynocephalus lizards. Asian Herpetological Research 3(3): 184-191.

Terbisch, H. 1986. [Ernährungsbiologie der Buntkrötenkopfagame (Phrynocephalus versicolor Str.) in der Westmongolei. pp. 77-84, 242]. (In Russian, Germany summary). In: Vorobeva, E.I. (ed.). Gerpetologicheskie issledovaniya v mongolskoi narodnoi respublike. [Herpetological studies in Mongolia.] Acad. Sci. USSR, Moscow. 247 pp.

Veselovsky, M.V.; Vasilevskaya, H.I. 2001. [Investigation of the peripheral nervous system in representatives of genus Phrynocephalus (Reptilia, Agamidae)]. (In Russian, English summary). Vestnik Zoologii 35(1): 63-74, 102.

Wang, Y.; Wang, H. 1993. Geographical variation and diversity in three species of Phrynocephalus in the Tengger Desert, western China. Asiatic Herpetological Research 5: 65-73.

Wang, Y.; Zeng, X.M.; Fang, Z.L.; Wu, G.; Liu, Z.J.; Papenfuss, T.J.; Macey, J.R. 2002. [A valid species of the genus Phrynocephalus: P. putjatia and a discussion on taxonomy of Phrynocephalus hongyuanensis (Sauria: Agamidae)]. (In Chinese, English summary). Acta Zootaxonomica Sinica 27(2): 372-383.

Wu, M.; Li, P.; Zhao, Y.; Ji, X. 1989. [A study of the karyotype of three species of Phrynocephalus from Xinjiang]. (In Chinese). Zoological Research 10(2): 114,153.

Zamolodchikov, D.G.; Gilmanov, T.G. 1990. [Ecology of Phrynocephalus versicolor (Reptilia, Agamidae) in the Ubsunur Valley]. (In Russian, English summary). Zoologicheskii Zhurnal 69(4): 148-151.

Zhao, K. 1995. [Description of a new species of the genus Phrynocephalus from Inner Mongolia]. (In Chinese, English summary). Sichuan Journal of Zoology 14 (Supplement): 47-50.

Phrynocephalus vindumi

Golubev, M.L. 1998. A new subspecies of Phrynocephalus ornatus Boulenger (Reptilia: Agamidae) from eastern Iran, with a key to south-western and middle Asian microphrynocephalids. Hamadryad 23(2): 162-168.

Macey, J.R.; Schulte, J.A.; Ananjeva, N.B.; Dyke, E.T. van; Wang, Y.; Orlov, N.; Shafiei, S.; Robinson, M.D.; Dujsebayeva, T.; Freund, G.S.; Fischer, C.M.; Liu, D.; Papenfuss, T. 2018. A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. Zootaxa 4467(1): 1–81.

Phrynocephalus vlangalii

Ding, G.H.; Li, Y.D.; Shen, Y.M.; Ji, X. 2007. [Selected body temperature shifts upward after feeding in the lizard, Phrynocephalus vlangalii (Agamidae) - physiological status affects the set-point of thermoregulation. pp. 146-152]. (In Chinese). In: Ji, X. Herpetologica Sinica 11. China Scientific Book Services, Beijing. 283 pp.

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