Phrynocephalus guttatus (GMELIN, 1789)

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Higher Taxa	Agamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
Subspecies	Phrynocephalus guttatus alpherakii BEDRIAGA 1906 Phrynocephalus guttatus guttatus (GMELIN 1789) Phrynocephalus guttatus melanurus EICHWALD 1831 Phrynocephalus guttatus moltschanowi NIKOLSKY 1913 Phrynocephalus guttatus salsatus GORELOV, DUNAYEV & KOTENKO in GOLUBEV et al. 1995
Common Names	E: Spotted Toadhead Agama; melanurus: Saissan toad-headed agama; arcellazzii: Central Asian Toadhead Agama; salenskyi: Salensky's Toadhead Agama G: Gefleckter Krötenkopf Russian: Круглоголовка-вертихвостка
Synonym	Lacerta guttata GMELIN in LINNAEUS 1789: 1078 Stincus guttatus — MEYER 1795: 30 (in error) Agama guttata — DAUDIN 1802: 426 Phrynocephalus guttatus KAUP 1827 Phrynocephalus Pallasii GRAY 1831: 59 Phrynocephalus guttatus — WANG et al. 2003 Phrynocephalus acutirostris — WANG et al. 2003 Phrynocephalus guttatus — BARTS & WILMS 2003 Phrynocephalus guttatus — SZCZERBAK 2003 Phrynocephalus guttatus — SOLOVYEVA et al. 2023 Phrynocephalus guttatus guttatus (GMELIN 1789) Lacerta caudivolvula PALLAS 1814 Phrynocephalus nigricans VON EICHWALD 1831 (partim) Phrynocephalus caudivolvula — EVERSMANN 1834: 362 ? Phrynocephalus nigricans — DUMÉRIL & BIBRON 1837: 516 Phrynocephalus caudivolvulus — DUMÉRIL & BIBRON 1837: 522 Phrynocephalus caudivolvulus — BOULENGER 1885: 375 Phrynocephalus suschkini BEDRIAGA in NIKOLSKY 1905: 472 Phrynocephalus cauivolovulus var. incerta BEDRIAGA in NIKOLSKY 1905: 475 Phrynocephalus isseli VON BEDRIAGA 1907 Phrynocephalus versicolor paraskiwi VON BEDRIAGA 1907 Phrynocephalus suschkini VON BEDRIAGA 1907: 14 (223) Phrynocephalus caudivolovulus var. incerta VON BEDRIAGA 1907: 18 (227) Phrynocephalus cauivolovulus var. ocellata VON BEDRIAGA 1909: 368 Phrynocephalus caudivolvulus (PALLAS 1814) (fide KHALIKOV, pers. comm.) Phrynocephalus bergi ELPATJEVSKY 1907 (fide TERENTJEW & TSCHERNOW 1949) Phrynocephalus bedriagai TZAREVSKY 1926 (fide KHALIKOV, pers. comm.) Phrynocephalus versicolor STRAUCH 1876 (fide KHALIKOV, pers. comm.) Phrynocephalus guttatus guttatus — WERMUTH 1967: 80 Phrynocephalus guttatus kalmykus BADMAYEVA & SZCZERBAK 1983 Phrynocephalus guttatus — ENGELMANN et al 1993 Phrynocephalus guttatus kalmykus— MANTHEY & SCHUSTER 1999 Phrynocephalus guttatus — MANTHEY & SCHUSTER 1999: 89 Phrynocephalus versicolor paraskiwi — SEMENOV et al. 1987 (fide LAMBERT 2002) Phrynocephalus (Phrynocephalus) guttatus guttatus — BARABANOV & ANANJEVA 2007: 40 Phrynocephalus guttatus incerta — DUNAYEV 2009: 284 Phrynocephalus incertus — MILTO & BARABANOV 2012: 159 Phrynocephalus incertus — SOLOVYEVA et al. 2023 Phrynocephalus guttatus moltschanowi NIKOLSKY 1913 Phrynocephalus caudivolvulus var. moltschanowi NIKOLSKY 1913: 34 Phrynocephalus guttatus moltschanowi — NIKOLSKY 1913 Phrynocephalus caudivolvulus moltschanowi — NIKOLSKY 1915: 201 Phrynocephalus moltschanovi — SEMENOV & SHENBROT 1982: 1194 Phrynocephalus guttatus moltschanowi — DUNAYEV 2009 Phrynocephalus guttatus moltschanowi — MILTO & BARABANOV 2012: 160 Phrynocephalus moltschanowi — SOLOVYEVA et al. 2023 Phrynocephalus guttatus alpherakii (BEDRIAGA 1907) Phrynocephalus alpherakii BEDRIAGA in NIKOLSKY 1905: 470 Phrynocephalus alpherakii BEDRIAGA 1907: 11 (220) Phrynocephalus acutirostris BEDRIAGA 1907: 339 Phrynocephalus acutirostris — BEDRIAGA 1909 Phrynocephalus caudivolvulus alpherakii — TZAREVSKY 1926: 212. Phrynocephalus acutirostris — WERMUTH 1967: 76 Phrynocephalus versicolor paraskiwi SEMONOV et al. 1987 Phrynocephalus versicolor paraskivi — SHENBROT & SEMONOV, 1987: 269 (unjustified emendation) Phrynocephalus guttatus alpherakii — GOLUBEV 1992: 37. Phrynocephalus (Phrynocephalus) guttatus alpherakii — BARABANOV & ANANJEVA 2007: 41 Phrynocephalus alpherakii — DUNAYEV 2009 Phrynocephalus alpherakii — DUJSEBAYEVA 2010 Phrynocephalus alpherakii — SOLOVYEVA et al. 2023 Phrynocephalus guttatus melanurus EICHWALD 1831 Phrynocephalus melanurus EICHWALD 1831: 186 Phrynocephalus melanurus — DUMÉRIL & BIBRON 1837: 516 Phrynocephalus caudivolvulus var. incerta BEDRIAGA In NIKOLSKY 1905: 475 Phrynocephalus kuschakewitschi BEDRIAGA In NIKOLSKY 1905: 476 Phrynocephalus salenskii BEDRIAGA “1905” 1906: 167 (nomen nudum) Phrynocephalus haeckeli BEDRIAGA “1905” 1906: 167 (nomen nudum) Phrynocephalus isseli BEDRIAGA “1905” 1906: 167 (nomen nudum) Phrynocephalus bergi ELPATJEVSKY 1906 "1907": 51 Phrynocephalus Salenskyi BEDRIAGA 1907: 141, 213 Phrynocephalus kuschakewitschi BEDRIAGA 1907: 20 (229) Phrynocephalus Haeckeli BEDRIAGA 1907: 237 Phrynocephalus zalenskyi — BEDRIAGA 1907: 141, 148 (unjustified emend.). Phrynocephalus Isseli BEDRIAGA 1907: 229 Phrynocephalus grum-grzimailoi BEDRIAGA 1907: 240 Phrynocephalus arcellazzii BEDRIAGA 1909 Phrynocephalus caudivolvulus var ocellata — BEDRIAGA, 1909: 367. Phrynocephalus Grum-Grzimailoi BEDRIAGA, 1909: 420 Phrynocephalus helioscopus levis KASCENKO 1909: 122 Phrynocephalus arcellazzii — NIKOLSKY 1915: 224 Phrynocephalus helioscopus laevis — NIKOLSKY, 1915: 185 (unjustified emendation). Phrynocephalus caudivolvulus ocellatus — NIKOLSKY, 1915: 200 Phrynocephalus caudivolvulus incertus — NIKOLSKY, 1915: 200 Phrynocephalus salenskii — NIKOLSKY, 1915: 232 (unjustified emendation). Phrynocephalus caudivolvulus kuschakewitschi — TZAREVSKY 1926: 213. Phrynocephalus bedrjagai TZAREVSKY 1926: 214 Phrynocephalus bedriagai — TZAREVSKY 1929: 416 (unjustified emendation) Phrynocephalus guttatus kuschakewitschi — TERENTJEV & CHERNOV 1949: 157 Phrynocephalus guttatus kuschakewitschi — WERMUTH 1967: 80 Phrynocephalus arcellazzii — WERMUTH 1967: 76 Phrynocephalus haeckeli — WERMUTH 1967 Phrynocephalus salenskyi — WERMUTH 1967: 88 Phrynocephalus albolineatus ZHAO 1979: 113 Phrynocephalus guttatus salenski — Mezhzherin & GOLUBEV 1993: 60 (unjustified emendation) Phrynocephalus guttatus kuschakewitschi — BRUSHKO 1995: 16 Phrynocephalus melanurus — BRUSHKO 1995: 44 Phrynocephalus melanurus — DUNAYEV 1996 Phrynocephalus kuschakewitschi — DUNAYEV 1996 Phrynocephalus grumgrzimalailoi — ZHAO 1997: 28 (lapsus calami) Phrynocephalus arcellazzii — BARTS & WILMS 2003 Phrynocephalus melanurus — BARTS & WILMS 2003 Phrynocephalus albolineatus — WANG et al. 2003 Phrynocephalus melanurus — ANANJEVA et L. 2006: 57 Phrynocephalus (Phrynocephalus) guttatus melanurus — BARABANOV & ANANJEVA 2007: 41 Phrynocephalus grumgrzimailoi — SINDACO & JEREMČENKO 2008: 151 Phrynocephalus melanurus — SINDACO & JEREMČENKO 2008: 156 Phrynocephalus grumgrzimailoi — JI et al. 2009 Phrynocephalus melanurus — DUNAYEV 2009 Phrynocephalus kuschakewitschi — DUNAYEV 2009 Phrynocephalus kuschakewitschi — DUJSEBAYEVA 2010 Phrynocephalus melanurus — DUJSEBAYEVA 2010 Phrynocephalus grumgrzimailoi — LIU et a. 2012 Phrynocephalus kuschakewitschi — MILTO & BARABANOV 2012 Phrynocephalus grumgrizimailoi — SHUANG et al. 2014 (in error) Phrynocephalus kuschakewitschi — SOLOVYEVA et al. 2023 Phrynocephalus guttatus salsatus GORELOV, DUNAYEV & KOTENKO in GOLUBEV et al. 1995 Phrynocephalus (Phrynocephalus) guttatus salsatus — BARABANOV & ANANJEVA 2007: 42 Phrynocephalus guttatus salsatus — DUNAYEV 2009 Phrynocephalus guttatus salsatusTHEY & SCHUSTER 1999: 89 Phrynocephalus versicolor paraskiwi — SEMENOV et al. 1987 (fide LAMBERT 2002) Phrynocephalus (Phrynocephalus) guttatus guttatus — BARABANOV & ANANJEVA 2007: 40 Phrynocephalus guttatus incerta — DUNAYEV 2009: 284 Phrynocephalus incertus — MILTO & BARABANOV 2012: 159 Phrynocephalus incertus — SOLOVYEVA et al. 2023 Phrynocephalus guttatus moltschanowi NIKOLSKY 1913 Phrynocephalus caudivolvulus var. moltschanowi NIKOLSKY 1913: 34 Phrynocephalus guttatus moltschanowi — NIKOLSKY 1913 Phrynocephalus caudivolvulus moltschanowi — NIKOLSKY 1915: 201 Phrynocephalus moltschanovi — SEMENOV & SHENBROT 1982: 1194 Phrynocephalus guttatus moltschanowi — DUNAYEV 2009 Phrynocephalus guttatus moltschanowi — MILTO & BARABANOV 2012: 160 Phrynocephalus moltschanowi — SOLOVYEVA et al. 2023 Phrynocephalus guttatus alpherakii (BEDRIAGA 1907) Phrynocephalus alpherakii BEDRIAGA in NIKOLSKY 1905: 470 Phrynocephalus alpherakii BEDRIAGA 1907: 11 (220) Phrynocephalus acutirostris BEDRIAGA 1907: 339 Phrynocephalus acutirostris — BEDRIAGA 1909 Phrynocephalus caudivolvulus alpherakii — TZAREVSKY 1926: 212. Phrynocephalus acutirostris — WERMUTH 1967: 76 Phrynocephalus versicolor paraskiwi SEMONOV et al. 1987 Phrynocephalus versicolor paraskivi — SHENBROT & SEMONOV, 1987: 269 (unjustified emendation) Phrynocephalus guttatus alpherakii — GOLUBEV 1992: 37. Phrynocephalus (Phrynocephalus) guttatus alpherakii — BARABANOV & ANANJEVA 2007: 41 Phrynocephalus alpherakii — DUNAYEV 2009 Phrynocephalus alpherakii — DUJSEBAYEVA 2010 Phrynocephalus alpherakii — SOLOVYEVA et al. 2023 Phrynocephalus guttatus melanurus EICHWALD 1831 Phrynocephalus melanurus EICHWALD 1831: 186 Phrynocephalus melanurus — DUMÉRIL & BIBRON 1837: 516 Phrynocephalus caudivolvulus var. incerta BEDRIAGA In NIKOLSKY 1905: 475 Phrynocephalus kuschakewitschi BEDRIAGA In NIKOLSKY 1905: 476 Phrynocephalus salenskii BEDRIAGA “1905” 1906: 167 (nomen nudum) Phrynocephalus haeckeli BEDRIAGA “1905” 1906: 167 (nomen nudum) Phrynocephalus isseli BEDRIAGA “1905” 1906: 167 (nomen nudum) Phrynocephalus bergi ELPATJEVSKY 1906 "1907": 51 Phrynocephalus Salenskyi BEDRIAGA 1907: 141, 213 Phrynocephalus kuschakewitschi BEDRIAGA 1907: 20 (229) Phrynocephalus Haeckeli BEDRIAGA 1907: 237 Phrynocephalus zalenskyi — BEDRIAGA 1907: 141, 148 (unjustified emend.). Phrynocephalus Isseli BEDRIAGA 1907: 229 Phrynocephalus grum-grzimailoi BEDRIAGA 1907: 240 Phrynocephalus arcellazzii BEDRIAGA 1909 Phrynocephalus caudivolvulus var ocellata — BEDRIAGA, 1909: 367. Phrynocephalus Grum-Grzimailoi BEDRIAGA, 1909: 420 Phrynocephalus helioscopus levis KASCENKO 1909: 122 Phrynocephalus arcellazzii — NIKOLSKY 1915: 224 Phrynocephalus helioscopus laevis — NIKOLSKY, 1915: 185 (unjustified emendation). Phrynocephalus caudivolvulus ocellatus — NIKOLSKY, 1915: 200 Phrynocephalus caudivolvulus incertus — NIKOLSKY, 1915: 200 Phrynocephalus salenskii — NIKOLSKY, 1915: 232 (unjustified emendation). Phrynocephalus caudivolvulus kuschakewitschi — TZAREVSKY 1926: 213. Phrynocephalus bedrjagai TZAREVSKY 1926: 214 Phrynocephalus bedriagai — TZAREVSKY 1929: 416 (unjustified emendation) Phrynocephalus guttatus kuschakewitschi — TERENTJEV & CHERNOV 1949: 157 Phrynocephalus guttatus kuschakewitschi — WERMUTH 1967: 80 Phrynocephalus arcellazzii — WERMUTH 1967: 76 Phrynocephalus haeckeli — WERMUTH 1967 Phrynocephalus salenskyi — WERMUTH 1967: 88 Phrynocephalus albolineatus ZHAO 1979: 113 Phrynocephalus guttatus salenski — Mezhzherin & GOLUBEV 1993: 60 (unjustified emendation) Phrynocephalus guttatus kuschakewitschi — BRUSHKO 1995: 16 Phrynocephalus melanurus — BRUSHKO 1995: 44 Phrynocephalus melanurus — DUNAYEV 1996 Phrynocephalus kuschakewitschi — DUNAYEV 1996 Phrynocephalus grumgrzimalailoi — ZHAO 1997: 28 (lapsus calami) Phrynocephalus arcellazzii — BARTS & WILMS 2003 Phrynocephalus melanurus — BARTS & WILMS 2003 Phrynocephalus albolineatus — WANG et al. 2003 Phrynocephalus melanurus — ANANJEVA et L. 2006: 57 Phrynocephalus (Phrynocephalus) guttatus melanurus — BARABANOV & ANANJEVA 2007: 41 Phrynocephalus grumgrzimailoi — SINDACO & JEREMČENKO 2008: 151 Phrynocephalus melanurus — SINDACO & JEREMČENKO 2008: 156 Phrynocephalus grumgrzimailoi — JI et al. 2009 Phrynocephalus melanurus — DUNAYEV 2009 Phrynocephalus kuschakewitschi — DUNAYEV 2009 Phrynocephalus kuschakewitschi — DUJSEBAYEVA 2010 Phrynocephalus melanurus — DUJSEBAYEVA 2010 Phrynocephalus grumgrzimailoi — LIU et a. 2012 Phrynocephalus kuschakewitschi — MILTO & BARABANOV 2012 Phrynocephalus grumgrizimailoi — SHUANG et al. 2014 (in error) Phrynocephalus kuschakewitschi — SOLOVYEVA et al. 2023 Phrynocephalus guttatus salsatus GORELOV, DUNAYEV & KOTENKO in GOLUBEV et al. 1995 Phrynocephalus (Phrynocephalus) guttatus salsatus — BARABANOV & ANANJEVA 2007: 42 Phrynocephalus guttatus salsatus — DUNAYEV 2009 Phrynocephalus guttatus salsatus — SHESTOPAL & SCHCHERBINA 2024
Distribution	Soviet Union, Russia (from NW coast of Caspian Sea eastward to E Kazakhstan, Dagestan, Astrakhan Oblast and Volgograd Oblast), Turkmenistan, SE Europe, W Asia, W China (N Xinjiang), Mongolia Subspecies including synonyms: albolineatus: China (N Xinjiang); Type locality: 45 km south of Tacheng Co., Xinjiang Uygur Autonomous Region, China. acutirostris: only known from its type locality (fide WERMUTH 1967); Type locality: Dschyn-Cho south of Ebi-Nor and in Chuldscha”. alpheraki: Kazakhstan (Ili River Hollow), China arcellazzii: C Asia (Dsungarei); Type locality: “Gaschun, in der Nähe von Gutschen in der Dschungarei”. grumgrzimailoi: W China (Xinjiang); Type locality (restricted by lectotype designation). “Gutschen” [= Qitai County, Xinjiang Uygur Autonomous Region, China]. isseli: China; Type locality: “in der Wüste in der Nähe der Berge Ssalburty” [in Saur mountains of the Tarbagatay range, Tacheng district, northern Xinjiang Uygur Auto- nomous Region, China] (Peters 1984: 31) kushakewitschi: valley of lake Alu-kulj north of the Lepsa river, adjacent Dsungarei; Kazakhstan; Type locality: “Kreis von Sergiopo” (fide WERMUTH 1967). melanurus: Sibiria, Russia, E Kazakhstan; Type locality: in Sibiria australi, versus Altaicos montes. moltschanowi: Kazakhstan, Uzbekistan; Type locality: mouth of the Amudarya River. Gebirge Bel Tau, Wüste Amu-Daria (fide WERMUTH 1967). salenskyi: C Asia (south of the Altai Mountains, north of Tien-schan), Kazakhstan; Type locality: “am Fluss Urungu, am Uljungur und im Bulun-Tschi am Uljungur” (fide WERMUTH 1967). salsatus: Turkmenistan; Type locality: North Turkmeniya, salt hollow Kazakhlyshor between sands Kumsebshen and the chink of Ustjurt (40º17' N, 55º46' E) Type locality: Ural River Valley, North Kazakhstan (“in deserto Uralensi” = area of the lower Ural River fide WERMUTH 1967). Restricted by neotype designation to vicinity of Atyrau (Gurjev), approximately 47o 06' N, 51o 54' E, Atyrau Province, north-western Kazakhstan by BARABANOV & ANANJEVA 2007.
Reproduction	Oviparous. Copulations of a P. salenskyi male with a P. versicolor female has been observed in the Kiev Zoo (Golubev 1993)
Types	Neotype: ZISP 4910.1, adult male, designated by BARABANOV & ANANJEVA 2007. Holotype, ZISP 8166 [arcellazzii] Lectotype by present designation, ZISP 8178.1, adult male, SVL 56.1 mm, TL 70.5 mm [grumgrzimailoi] Lectotype: ZISP 10038.3, (designated by Peters, 1984), “Chorgos” [Korgas river, tributary of the upper Ily river, western Xinjiang Uygur Autonomous Region, China]. Leg: S. N. Alpheraky, 1881; Paralectotypes. ZISP 5788.1–5788.5, 10038.1– 10038.18, 23 specimens, “Chorgos.” Leg: S. N. Alphera- ky, 1881. — ZISP 5795, 1 specimen, “Tardshi.” Leg: S. N. Alpheraky, 1881 [alpherakii] Syntypes: ZMB 787, "Kleine Tartarei", ZMB 788 "Bucharei;" leg. Eversmann; Terra typica restricta, fide MERTENS & WERMUTH (1960): Jaikische Steppen, Gebiet des unteren Uralflusses (no types (restriction invalid, fide TILLACK, pers. comm. 22 Jan 2014) [caudivolvula] Types: unknown fide BARABANOV & ANANJEVA 2007 [salenskyi] Holotype: ZISP 5164 [isseli] Neotype: ZMMU Re-5611 (designated by Semenov et al., 1987: 105) [melanurus] Lectotype: ZISP 19585 (designated by Semenov & Shenbrot, 1982) [moltschanowi] Holotype: NMNHU Re-21/1 (given as IZANU by Barabanov & Ananjeva 2007) [salsatus]
Diagnosis	Diagnosis (genus). “Kopf rundlich, breit. Körper dorsoventral stark abgeplattet. Präanal- oder Femoralporen fehlen. Längs der Rücken- und Schwanzmitte kein Kamm oder angedeuteter Schuppenfirst ausgebildet. Kehlsack fehlt. Trommelfell unter der Haut verborgen, unsichtbar. Zehen seitlich mit Fransenschuppen ausgestattet, die an der dritten und vierten Hinterzehe besonders deutlich differenziert sind; oft sind sie nur an einer Zehenseite erkennbar, mitunter von dreieckiger Form. Schädel stark verknöchert. Das Articulare ist - im Gegensatz zu Agama - caudad nicht verlängert. Bezahnung heterodont, der Grad der Heterodontie reflektiert die verschiedene Ernährungsweise der jeweiligen Arten. Für alle Phrynocephalus-Arten charakteristisch ist eine deutliche altersbedingte Variation in der Bezahnung (Ananjeva 1977). Schwanz nicht autotomiefähig, was offenbar mit seiner großen Bedeutung als Kommunikationsorgan korreliert ist. Dabei ist die Fähigkeit, mit dem Schwanz ein- und ausrollende Spiralbewegungen durchzuführen, ebenfalls gattungstypisch.” (Ananjeva 1981: 178). Additional details (968 characters) are available for collaborators and contributors. Please contact us for details.
Comment	Synonymy after WERMUTH 1967 and BARABANOV & ANANJEVA 2007. WERMUTH cited PALLAS 1814 as opposed to PALLAS 1827 and who recognized only 2 subspecies, guttatus and kushakewitschi; WERMUTH also synonymized alpherakii with guttatus guttatus. Note that Phrynocephalus acutirostris is listed as valid species by WANG et al. 2003. P. melanurus is listed as synonym of Phrynocephalus versicolor, with a question mark, by WERMUTH 1967: 89. Type species: Lacerta caudivolvula PALLAS 1814 is the type species of the genus Phrynocephalus KAUP 1825 ((designated by Fitzinger, 1843: 18, see also IUCN 1964, art 69a iv). NCBI taxon ID: 662669 [salenskyi] Group: the guttatus group includes P. alpheraki; P. guttatus; P. incertus; P. kuschakewitschii; P. melanurus; P. moltschanovi, fide SOLOVYEVA et al. 2023.
Etymology	The name guttata is Latin for "spotted" and refers to the numerous light spots and blotches on the back of the species. The genus has been after the Greek phrynos = toad and “kephale” = head. Phrynocephalus arcellazzii BEDRIAGA 1909 was named after T. Arcellazzi, who may have been an Italian artist who made drawings for Bedriaga (at least Michael Watkins, pers. comm., thinks that).
References	Ananjeva, N. & Orlov, N. 2005. Lizards of North Eurasia. Reptilia (GB) (38): 54-63 - get paper here 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 Ananjeva, Natalija B. 1981. Phrynocephalus guttatus - Gefleckter Krötenkopf. In: Böhme,W. (ed.), Handbuch der Reptilien und Amphibien Europas, Band I., Echsen I. Akademische Verlagsgesellschaft, Wiesbaden, pp. 180-190 Ananjeva, NB, Orlov NL; Khalikov RG; Darevsky IS; Ryabov IS & Barabanov AV 2006. The Reptiles of North Eurasia. Taxonomic Diversity, Distribution, Conservation Status [this comprises the territory of the former Soviet Union and Mongolia]. Pensoft Series Faunistica 47, 250 pp. Bakiev, A., Kirillov, A., Kirillova, N., Ruchin, A., Klenina, A., Gorelov, R., & Kostina, N. 2020. Reptile occurrences data in the Volga River basin (Russia). Biodiversity Data Journal, 8: e58033 - 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 Bedriaga, JACQUES VON 1907. Verzeichnis der von der Central-Asiatischen Expedition unter Stabs-Kapitän W. Roborowski in den Jahren 1893-1895 gesammelten Reptilien. Annu. Mus. Zool. Acad. Impér. Sci. St.-Petersbourg, 10 (3-4): 159-200 [1905] - get paper here Bedriaga, JACQUES VON 1907. Neue Saurier aus Russisch-Asien. Annu. Mus. Zool. Acad. Impér. Sci. St.-Petersbourg, 10 [1905]: 210-243 - get paper here Boistel, Renaud;, Anthony Herrel; Renaud Lebrun, Gheylen Daghfous, Paul Tafforeau, Jonathan B. Lososô and Bieke Vanhooydonck\|\| 2011. Shake Rattle and Roll: The Bony Labyrinth and Aerial Descent in Squamates. Integrative and Comparative Biology, doi:10.1093/icb/icr034 - get paper here Borkin, L.J. 2022. EARLY STUDIES ON THE HERPETOFAUNA OF KAZAKHSTAN (XVIII CENTURY) [in russian] Almaty: Institute of Zoology of the Republic of Kazakhstan, 2022. - 156 pp. 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 Brushko, Z.K. 1995. Lizards of desert regions of Kazakhstan. [in Russian] Kazakhstan National Academy of Sciences, Institute of Zoology, Almaty, 232 pp. Chetanov N. A., Litvinov N. A., Ganshchuk S. V., Yugov M. V. 2023. Some features of thermobiology of steppe-runner Eremias arguta (Gmelin, 1789) and spotted toadhead agama Phrynocephalus guttatus (Gmelin, 1789) (Squamata, Reptilia) in cohabitation in the Northern Pre-Caspian. Current Studies in Herpetology 23 (3): 164 - get paper here Dujsebayeva, T. N. (ed.) 2010. Short review of last changes in the checklist of amphibians and reptiles of Kazakhstan. [in Russian]. In: Dujsebayeva, T. N. (ed.) Herpetological Researches in Kazakhstan and adjacent countries, p. 37-52 Duméril, A. M. C. and G. Bibron. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Encyclopédique Roret, Paris, 570 pp. - get paper here Dunaev E.A., Orlova V.F. 2017. Amphibians and reptiles of Russia. Atlas and determination. 2nd. ed. Moscow: Phyton XXI, 328 p Dunayev, E. A. 1996. On the possible use of the Ethological features in the taxonomy and phylogeny of toad agamas, Phrynocephalus (Reptilia, Agamidae). Russ. J. Herpetol. 3 (1): 32-38 - get paper here Dunayev, E. A. 2009. SYSTEMATICS AND PALEOGEOGRAPHY: CONCEPTUAL SYNTHESIS BY THE EXAMLE OF PHRYNOCEPHALUS (SUPERSPECIES GUTTATUS) (REPTILIA: AGAMIDAE) [in Russian] Arkh. Zool. Muz. MGU, 50: 275 – 298 Eichwald, E. 1831. Zoologia specialis, quam expositis animalibus tum vivis, tum fossilibus potissimuni rossiae in universum, et poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi. Zawadski, Vilnae. - get paper here Elpatjevsky, VLADIMIR S. 1907. Izvest. Taschkent Otd. russ. geogr. Obschtsch. 4 (7): 51 Engelmann, W.E. et al. 1993. Lurche und Kriechtiere Europas. Neumann Verlag (Radebeul, Germany), 440 pp. Eversmann,E. 1834. Lacertae Imperii Rossici variis in itineribus meis observatae. [Lacerta sylvicola]. Nouv. Mém. Soc. imp. Natur. (Moscow) 3: 339-358 - get paper here Fitzinger, L. 1823. in Lichtenstein: in Evermann's Reise [von Orenburg] nach Buchara. E. H. G. Christiani, Berlin. - get paper here Gmelin, J.F. 1789. Caroli a Linné Systema naturae. 13. ed., Tom 1 Pars 3. G. E. Beer, Lipsiae. 1033-1516 pp. - get paper here Golubev, M.L. 1989. Vest. Zool., 5: 38 Golubev, M.L. 1991. About the name Agama ocellata Lichtenstein in Eversmann, 1823 (Reptilia, Agamidae) with redescription of the types. Herpetological Researches (1):12-17. 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 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 with English summary] Bulletin of the Moscow Society Ispytatelej Prirody, Otd. Biologichesky, 100(3): 31–39 Gravenhorst, J. L. C. 1838. Beiträge zur genaueren Kenntniss einiger Eidechsengattungen. Nova Acta Acad. Caes. Leop.-Carol. 18: 712-784 [1837] 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 Kaup, J. 1827. Zoologische Monographien. Isis von Oken 20: 610—625 [columns] - get paper here Kidov, A. A., Khayrutdinov, I. Z., Ivanov, A. A., & Kidova, E. A. 2020. AGE STRUCTURE AND GROWTH OF THE SPOTTED TOADHEAD AGAMA, PHRYNOCEPHALUS GUTTATUS (REPTILIA, SAURIA, AGAMIDAE) IN TEREK SAND MASSIF (EASTERN CISCAUCASIA). [in russian] ИЗВЕСТИЯ ВЫСШИХ УЧЕБНЫХ ЗАВЕДЕНИЙ ПОВОЛЖСКИЙ РЕГИОН, 57 - get paper here Lambert, M.R.K. 2002. Preliminary observations on herpetofaunal diversity in the Almaty region, Southern Kazakhstan (September 1998). Herpetological Bulletin (79): 7-13 - get paper here LIU, Yang; Yucheng SONG, Wenrong LI, and Lei SHI 2012. Reproductive Strategy and Cycle of the Toad-headed Agama Phrynocephalus grumgrzimailoi (Agamidae) in Xinjiang, China. Asian Herpetological Research 3 (3): 198–204 - get paper here 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 Manthey,U. & SCHUSTER,N. 1999. Agamen, 2. Aufl. Natur und Tier Verlag (Münster), 120 pp. - get paper here Mertens, R., and L. Müller. 1940. Liste der Amphibien und Reptilien Europas. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft. Frankfurt am Main. 451: 1-62 - get paper here Meyer, F.A.A. 1795. Synopsis Reptilium, novam ipsorum sistens generum methodum, nec non Gottingensium huius ordinis animalium enumerationem. Vandenhoeck und Ruprecht, Göttingen, 32 pp [for current names see Rösler & Böhme 2021] - get paper here Mezhzherin, Sergei;Golubev, Michael 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 - 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 Neymark, L. A. and B. D. Vasiliev 2016. ECOLOGY OF SEMIDESERT LIZARDS (REPTILIA, SQUMATA) UNDER PLANT COLONIZATION OF SANDS IN THE LOWER REACHES OF THE VOLGA RIVER (in Russian). ЗООЛОГИЧЕСКИЙ ЖУРНАЛ 95 (6): 720–727 - get paper here Nikolsky, A. M. 1905. Herpetologia rossica. Mem. Acad. Imp. Sci., Phys.-Math. Classe, St. Pétersbourg, XVII (1): 1-518 Nikolsky, A. M. 1913. Herpetologia caucasica [in Russian]. Tiflis (Museum caucasicum), 272 pp. - 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] Okshtein I. L. 2016. The thermal biology of the Phrynocephalus guttatus (Gmelin, 1789) in the Astrakhan region. PROCEEDINGS OF THE 6th CONGRESS OF THE A. M. NIKOLSKY HERPETOLOGICAL SOCIETY pp. 107-112 - get paper here Okshteyn, I. L.; A. A. Korneeva, <br>A. V. Mendelevich,L. N. Vasil'eva, <br>A. V. Sadovskaya, P. A. Galkina 2019. Growth rate of Phrynocephalus guttatus guttatus in the Astrakhan region. PROCEEDINGS OF THE VII CONGRESS OF THE NIKOLSKY HERPETOLOGICAL SOCIETY, in "University proceedings. Volga region. Natural sciences” 1: 126-137 - get paper here Pallas, P. S. 1814. Zoographia Rosso-Asiatica, sistens omnium in extenso Imperio Rossico , et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum. Vol. 3. Caesarea Academie Scientiarum; Petropoli. vii + 428 + cxxv pages. [1811-1831] - get paper here 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 Roitberg, E. S., Mazanaeva, L.F., Ilyina, E. & Orlova, V.F. 2000. Die Echsen D<br>agestans (Nordkaukasus, Russland): Artenliste und aktuelle Verbreitungsdaten (Reptilia: Sauria: Gekkonidae, Agamidae, Anguidae, Scincidae et Lacertidae). Faunist. Abh. Staatl. Mus. Tierk. Dresden 22 (8) - get paper here Rösler, Herbert & Wolfgang Böhme 2021. Das Leben von Friedrich Albrecht Anton Meyer (1768 – 1795) und die Herpetologie in seinen Schriften, unter besonderer Berücksichtigung der „Synopsis Reptilium“. Sekretär 21: 83-129 Shenbrot, G. I.; Semenov, D. V. 1987. Present distribution and taxonomy of Phrynocephalus guttatus (Reptilia, Agamidae) [in Russian]. Zool. J. (Moscow) 66:259-272 Shestopal A. A., Shcherbina A. A. 2024. Distribution of reptiles in Northwestern Turkmenistan. [in Russian] Current Studies in Herpetology 24 (1-2): 102 - get paper here Sindaco, R. & Jeremcenko, V.K. 2008. The reptiles of the Western Palearctic. Edizioni Belvedere, Latina (Italy), 579 pp. - 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 Szczerbak, N.N. 2003. Guide to the Reptiles of the Eastern Palearctic. Krieger, Malabar, FL, 260 pp. - get paper here Tabachishin, Vasily G. and Evgeni V. Zavialov 2005. Present distribution and taxonomic status of the Spotted Toad-headed Agama Phrynocephalus guttatus (Gmelin, 1789) in the Volga region, Russia. Herpetozoa 18 (3/4): 141-146. - get paper here Terentjev, P.V. & Chernov, S.A. 1949. Opredelitel Presmykatushchikhsya i Zemnovodnykh. [Encyclopedia of reptiles and amphibians, 2nd ed., in Russian]. Moscow Tzarevsky, S. (as Carevskij), S. 1929. Contribution to the classification and distribution of the lizards of the genus Phrynocephalus. Comptes Rendus de l’Académie des Sciences de l’URSS 1929: 415-419 Vedmederya, Valeriy; Oleksandr Zinenko, and Andrei Barabanov 2009. An Annotated Type Catalogue of Amphibians and Reptiles in the Museum of Nature at V. N. Karazin Kharkiv National University (Kharkiv, Ukraine). Russ. J. Herpetol. 16 (3): 203-212 - get paper here Wittig, W. 2002. Haltung und Nachzucht von Phrynocephalus guttatus. Elaphe 10 (3): 29-31 Zeng, Xiaomao; Wang, Yaozhao; Liu, Zhijun;Yu, Pin 1994. A COMPARATIVE STUDY ON THE KARYOTYPE OF THREE SPECIES IN Phrynocephalus (Agamidae). Acta Zootaxonomica Sinica 1994 (2) - 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 Zhao,Ken-Tang 1979. Studies on the classification and distibution of sand lizards in China (in Chinese). Acta Univ. Inner Mongolia, Hohhot 1979 (2): 111-121
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