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
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).
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).
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External links	IUCN Red List National Center for Biotechnology Information http://dinets.travel.ru/deserts.htm http://reptile.fisek.com.tr/phrynocephalus.htm http://www.seh-herpetology.org/images/atlas/pdf/squamata/phrgut.pdf http://www.seh-herpetology.org/sites/seh-herpetology.org/files/uploads… https://www.facebook.com/groups/stshc/posts/358811189676819 https://www.facebook.com/groups/stshc/posts/567357975488805/ Google images

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