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Phrynocephalus mystaceus (PALLAS, 1776)

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards) 
SubspeciesPhrynocephalus mystaceus mystaceus (PALLAS 1776)
Phrynocephalus mystaceus khorasanus SOLOVYEVA, DUNAYEV, NAZAROV, RADJABIZADEH & POYARKOV 2018 
Common NamesE: Secret Toadhead Agama
G: Bärtiger Krötenkopf
E: Khorasan Secret Toad-headed Agama [khorasanus]
Russian: Ушастая круглоголовка 
SynonymLacerta mystacea PALLAS 1776: 702
Lacerta aurita — SHAW & NODDER 1803: plate 594
Phrynocephalus mystacia — KAUP 1827
Phrynocephalus mystaceus — BOULENGER 1885: 379
Phrynocephalus mystaceus mystaceus natio dagestanica KRASSOWSKY 1932
Phrynocephalus mystaceus galli KRASSOWSKY 1932
Megalochilus mystaceus — ANANJEVA 1986
Phrynocephalus mystaceus — WERMUTH 1967: 84
Phrynocephalus mystaceus — ENGELMANN et al 1993
Phrynocephalus mystaceus — ŠMÍD et al. 2014

Phrynocephalus mystaceus mystaceus (PALLAS 1776)
Lacerta aurita GMELIN 1789: 1073 (nom. subst. pro Lacerta mystacea)
Gekko auritus — LATREILLE 1801: 61.
Lacerta lobata SHAW 1802: 244 (nom. subst. pro Lacerta aurita)
Agama aurita — DAUDIN 1802: 429
Agama mystacea — MERREM 1820: 53
Agama aurita — LIECHTENSTEIN 1823: 142
Megalochilus auritus — EICHWALD 1831: 185
Phrynocephalus auritus — EVERSMANN 1834: 360
Phrynocephalus auritus — DUMÉRIL & BIBRON 1837: 524
Saccostoma auritum — FITZINGER 1843: 87
Phrynocephalus mystaceus — BOETTGER 1880: 260
Phrynocephalus mystaceus mystaceus — KRASSOWSKY 1932: 227
Phrynocephalus mystaceus Galli KRASSOWSKY 1932: 227
Phrynocephalus mystaceus mystaceus — WERMUTH 1967: 84
Phrynocephalus mystaceus aurantiacaudatus SEMENOV & SHENBROT 1990: 79
Phrynocephalus mystaceus mystaceus — MANTHEY & SCHUSTER 1999: 90
Phrynocephalus (Phrynocephalus) mystaceus mystaceus — BARABANOV & ANANJEVA 2007
Phrynocephalus (Phrynocephalus) mystaceus galli — BARABANOV & ANANJEVA 2007
Phrynocephalus mystaceus aurantiacaudatus — SINDACO & JEREMČENKO 2008
Phrynocephalus mystaceus galli — SINDACO & JEREMČENKO 2008
Phrynocephalus mystaceus galli — MILTO & BARABANOV 2012

Phrynocephalus mystaceus khorasanus SOLOVYEVA, DUNAYEV, NAZAROV, RADJABIZADEH & POYARKOV 2018 
DistributionSoviet Union, NW cost of Caspian Sea to E Kazakhstan, Russia (Dagestan, Astrakhan Oblast), Kazakhstan, Turkmenistan, Tajikistan, Uzbekistan, Kyrgyzstan, NE Iran, Afghanistan, NW China (W Xinjiang)

aurantiacaudatus: Kazakhstan
galli: Type locality: Repeteka, Turkestan.

galli: Turkmenistan, Uzbekistan, Kazakhstan, northeastern and eastern Iran and adjacent Afghanistan and Xinjiang, China; Terra typica: “Aus Repeteka andetrifft” [= vicinity of Repetek station, approximately 38º35' N, 63º11' E, Lebapsky Region, Turkmenistan].

khorasanus: Iran (Khorasan Razavi Province); Type locality: Iran, Khorasan historical area, Khorasan Razavi Province (ostan), environs of Gonabad, the right bank of the Kale-Shur River; sand dunes (see Fig. 9 in Solovyeva et al. 2018); N34°39', E58°43'; elevation 850 m.

Type locality: East Caucasus. Restricted to “Naryn-Steppe, an der Nordküste des kaspischen Maeeres” by MERTENS & MÜLLER 1928; Restricted by neotype designation to Ryn Peski (Ryn Sands), Uralskaya Region, north-western Kazakh-
stan, by BARABANOV & ANANJEVA 2007.  
ReproductionOviparous. 
TypesNeotype: ZISP 8735.1, adult male, designated by BARABANOV & ANAJEVA 2007
Holotype. ZMMU R-11913 (adult female); paratypes: ZMMU [khorasanus]
Lectotype: ZMMU Re-6413 (designated by Semenov & Shenbrot, 1990: 78) [galli] 
CommentSynonymy and subspecies mainly after KHALIKOV & ANANJEVA (pers. comm.), WERMUTH 1967, and BARABANOV & ANANJEVA 2007. SEMENOV & SHENBROT 1990 considered galli as a synonym of P. mystaceus. SOLOVYEVA et al. consider P. m. galli as a synonym of mystaceus.

Distribution: See map in SMID et al. 2014 for distribution in Iran.

NCBI taxon ID: 662673 [aurantiacaudatus]
NCBI taxon ID: 662675 [galli]
NCBI taxon ID: 2172839 [khorasanus]
NCBI taxon ID: 662674 [mystaceus]

Diagnosis (khorasanus). A member of Ph. mystaceus species complex based on the following combination of morphological attributes: (1) a large-sized Phrynocephalus species with SVL up to 97.5 mm, tail shorter than SVL; (2) pair of cutaneous flaps present at mouth corners with numerous spiny scales along flap edges; (3) distinctly flattened body and tail; (4) toes with fringes formed by triangular scales; subdigital lamellae on toes III and IV with ridges. Phrynocephalus mystaceus khorasanus ssp. n. can be distin- guished from the nominative subspecies of Ph. mystaceus by the following combination of two diagnostic morphological characteristics: (1) 24–27 lamellae on toe IV; (2) few supralabial scales (less than 14). In life, the new subspecies can be further distinguished from the nominative subspecies by the orange color of the lower surface of tail in young specimens (lemon to yellowish in Ph. m. mystaceus except the populations from Eastern Kazakhstan and western China, formerly described as Ph. m. aurantiacocau- datus). MtDNA sequences of Phrynocephalus mystaceus khorasanus ssp. n. are markedly distinct from those in all other populations of Ph. mystaceus with sequence divergence in the range of 6.84–7.28% between them. The new subspecies is notably smaller than the representatives of southern populations of Ph. m. mystaceus from Uzbekistan and Turkmenistan, formerly described as Ph. m. galli, which can reach SVL up to 122.7 mm (Anderson 1999), whereas for Iranian population Anderson (1999) reported the largest specimen of Ph. mystaceus to have SVL up to 77.7 mm. SVL in the largest specimen in our sampling reached 86.0 mm, while Molavi et al. (2014) recorded a specimen with SVL of 97.5 from Semnan Province.

Comparisons of khorasanus with other subspecies. Comparisons of the new subspecies from Khorasan Razavi and Semnan provinces of Iran with the nominative subspecies Ph. m. mystaceus sensu lato from Middle Asia, Caspian basin, and westernmost Xinjiang (China) are summarized below. In preservative, the new subspecies can be differentiated from specimens of Ph. m. mystaceus by the following combination of morphological attributes: lower number of subdigital lamellae on the IVth toe (SLIV 25.7 (24–27; N = 7) in vs. 30.2 (25–35; N = 70) in Ph. m. mystaceus sensu lato); comparatively lower number of supralabials (SL 12.1 (10–14; N = 7) vs. 14.9 (10-19; N = 70) in Ph. m. mys- taceus sensu lato) and by the comparatively shorter black distal part on the tail ventral surface (TL-black/TL 0.38 (0.36–0.40; N = 7) vs. 0.42 (0.32–0.48; N = 70) in Ph. m. mystaceus sensu lato). In life, juvenile and young specimens of the new subspecies can be further distinguished from Middle Asian / Caspian Basin populations of Ph. mystaceus by is rusty orange color of the proximal part of tail ventral surface (vs. lemon-yellow in Ph. m. mystaceus sensu stricto), but is similar to orange tail coloration in juveniles of East Kazakhstan – western China populations described as Ph. m. aurantiacocaudatus.
Solovyeva et al. 2018 do not recognize Ph. m. galli as a separate subspecies due to the absence of stable genetic and morphological differences of this subspecies from Ph. m. mystaceus (see above). The Phrynocephalus mystaceus dagestanica form from Daghestan (Ananjeva, “1986” 1987) is very close to the populations from the Volga River basin and was considered a synonym of Ph. m. mystaceus by several authors (Semenov and Shenbrot 1990; Barabanov and Ananjeva 2007). Our molecular and morphometric data do not support monophyly or significant differentiation of Ph. m. aurantiacocaudatus from Eastern Kazakhstan and western China. The only stable difference between this popu- lation and Ph. m. mystaceus sensu stricto is the tail coloration in juveniles. We consider that additional genetic and morphological data is needed to clarify taxonomic status of East Kazakhstan Ph. mystaceus populations.

Etymology (khorasanus). The name of the new subspecies khorasanus is a Latinized toponymic adjective, derived from Khorasan, the name of the historic area and a Khorasan Razavi Province in the northeast Iran, where the new subspecies was found. 
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