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Phrynocephalus guttatus (GMELIN, 1789)

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
SubspeciesPhrynocephalus 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 NamesE: Spotted Toadhead Agama; melanurus: Saissan toad-headed agama; arcellazzii: Central Asian Toadhead Agama; salenskyi: Salensky's Toadhead Agama
G: Gefleckter Krötenkopf
Russian: Круглоголовка-вертихвостка 
SynonymLacerta 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 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 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 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 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
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 guttatus salsatus GORELOV, DUNAYEV & KOTENKO in GOLUBEV et al. 1995
Phrynocephalus (Phrynocephalus) guttatus salsatus — BARABANOV & ANANJEVA 2007: 42
Phrynocephalus guttatus salsatus — DUNAYEV 2009 
DistributionSoviet 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.  
ReproductionOviparous. Copulations of a P. salenskyi male with a P. versicolor female has been observed in the Kiev Zoo (Golubev 1993) 
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: 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] 
DiagnosisDiagnosis (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).

Diagnosis (species). “Unterscheidet sich von P. helioscopus und P. mystaceus durch das Kopfprofil, wobei der Schnauzenschädel relativ flach nach vorn abfällt, so daß die Nasenlöcher von oben sichtbar bleiben. Von P. mystaceus auch durch die geringere Körpergröße und das Fehlen von Hautlappen in den Mundwinkeln verschieden. Von P. helioscopus durch das Fehlen von Halsfalten und gefärbten Nackenflecken sowie durch die homogene Rückenbeschuppung und die wirtlig angeordneten Schwanz­ schuppen unterschieden.
Schädel zarter gebaut als bei P. helioscopus und besonders bei P. mystaceus. Gegenüber P. helioscopus sind außerdem noch der geringere Ossifikations­grad und ein deutlich größeres Foramen parietale charakteristisch” (Ananjeva 1981: 180).
CommentSynonymy 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] 
EtymologyThe 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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