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Phrynocephalus guttatus (GMELIN, 1789)

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Higher TaxaAgamidae (Agaminae), Sauria (lizards) 
SubspeciesPhrynocephalus guttatus alpherakii BEDRIAGA 1906
Phrynocephalus guttatus guttatus (GMELIN 1789)
Phrynocephalus guttatus melanurus EICHWALD 1831
Phrynocephalus guttatus salsatus GOLUBEV, GORELOV, DUNAYEV & KOTENKO 1995 
Common NamesE: Spotted Toadhead Agama; melanurus: Saissan toad-headed agama; arcellazzii: Central Asian Toadhead Agama; salenskyi: Salensky's Toadhead Agama
G: Gefleckter Krötenkopf 
SynonymLacerta guttata GMELIN in LINNAEUS 1789: 1078
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 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 cauivolovulus var. incerta VON BEDRIAGA 1907: 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 caudivolvulus var. moltschanowi NIKOLSKY 1913: 34
Phrynocephalus guttatus moltschanowi — NIKOLSKY 1913
Phrynocephalus caudivolvulus moltschanowi — NIKOLSKY 1915: 201
Phrynocephalus bedriagai TSAREVSKY 1926 (fide KHALIKOV, pers. comm.)
Phrynocephalus frontalis STRAUCH 1876 (fide KHALIKOV, pers. comm.)
Phrynocephalus versicolor STRAUCH 1876 (fide KHALIKOV, pers. comm.)
Phrynocephalus guttatus guttatus — WERMUTH 1967: 80
Phrynocephalus moltschanovi — SEMENOV & SHENBROT 1982: 1194
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 guttatus guttatus — BARABANOV & ANANJEVA 2007
Phrynocephalus incertus — MILTO & BARABANOV 2012: 159
Phrynocephalus caudivolvulus moltschanowi — MILTO & BARABANOV 2012: 160

Phrynocephalus guttatus alpherakii (BEDRIAGA 1907)
Phrynocephalus alpherakii BEDRIAGA in NIKOLSKY 1905: 470
Phrynocephalus alpherakii BEDRIAGA 1907: 11
Phrynocephalus acutirostris BEDRIAGA 1907: 339
Phrynocephalus acutirostris — BEDRIAGA 1909
Phrynocephalus caudivolvulus alpherakii — TSAREVSKY 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 guttatus alpherakii — BARABANOV & ANANJEVA 2007
Phrynocephalus alpherakii — DUJSEBAYEVA 2010

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: 229
Phrynocephalus Haeckeli BEDRIAGA 1907: 237
Phrynocephalus zalenskyi — BEDRIAGA 1907: 141, 148 (unjustified emend.).
Phrynocephalus Isseli BEDRIAGA 1907: 229
Phrynocephalus Grum-Grzimailoi BEDRIAGA 1907
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 — TSAREVSKY 1926: 213.
Phrynocephalus bedrjagai Tsarevsky 1926: 214
Phrynocephalus bedriagai — TSAREVSKY 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 grumgrzimalailoi — Zhao, 1997: 28 (lapsus calami)
Phrynocephalus arcellazzii — BARTS & WILMS 2003
Phrynocephalus melanurus — BARTS & WILMS 2003
Phrynocephalus albolineatus — WANG et al. 2003
Phrynocephalus guttatus melanurus — BARABANOV & ANANJEVA 2007
Phrynocephalus grumgrzimailoi — SINDACO & JEREMČENKO 2008
Phrynocephalus melanurus — SINDACO & JEREMČENKO 2008
Phrynocephalus grumgrzimailoi — JI et al. 2009
Phrynocephalus kuschakewitschi — DUJSEBAYEVA 2010
Phrynocephalus melanurus — DUJSEBAYEVA 2010
Phrynocephalus grumgrzimailoi — LIU et a. 2012
Phrynocephalus kuschakewitschi — MILTO & BARABANOV 2012

Phrynocephalus guttatus salsatus GOLUBEV, GORELOV, DUNAYEV & KOTENKO 1995
Phrynocephalus guttatus salsatus — BARABANOV & ANANJEVA 2007 
DistributionSoviet Union, Russia (from NW coast of Caspian Sea eastward to E Kazakhstan, Dagestan, Astrakhan Oblast and Volgograd Oblast), Turkmenistan,
SE Europe, W Asia, Eritrea ? (P. isseli) , Ethiopia ? (P. isseli),
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].

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. Map legend:
Type locality - Type locality.
TDWG region - Region according to the TDWG standard, not a precise distribution map.

NOTE: TDWG regions are generated automatically from the text in the distribution field and not in every cases it works well. We are working on it.
 
TypesNeotype: 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 (designated by Peters, 1984). ZISP 10038.3, “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] 
CommentDistribution from previous entry including P. guttatus moltschanowi. Phrynocephalus guttatus moltschanowi NIKOLSKY 1913 and Phrynocephalus guttatus salenskyi are treated as a full species here.

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, Syntypes: ZMB: 787, 788 (designated as type species of the genus Phrynocephalus KAUP 1825, by Fitzinger, 1843: 18 = Lacerta guttata Gmelin IUCN 1964, art 69a iv).

Types: No types are known for salenskyi fide 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) 
EtymologyPhrynocephalus 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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