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Phrynocephalus frontalis STRAUCH, 1876

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Higher TaxaAgamidae (Agaminae), Sauria, Iguania, Squamata (lizards)
Subspecies 
Common NamesE: Shansi Toadhead Agama
G: Shansi-Krötenkopfagame 
SynonymPhrynocephalus frontalis STRAUCH 1876: 15
Phrynocephalus frontalis — BOULENGER 1885: 375
Phrynocephalus frontalis — BEDRIAGA 1909: 404
Phrynocephalus frontalis — NIKOLSKY 1915: 217
Phrynocephalus frontalis — POPE 1935: 470
Phrynocephalus frontalis — WERMUTH 1967: 78
Phrynocephalus frontalis — WANG et al. 2003
Phrynocephalus frontalis — SINDACO & JEREMČENKO 2008
Phrynocephalus frontalis — LIAN et al. 2012
Phrynocephalus frontalis — MILTO & BARABANOV 2012
Phrynocephalus frontalis — ZENG et al. 2016
Phrynocephalus frontalis — DUNAYEV et al. 2021 
DistributionChina (Gansu, Ningxia, Shaanxi, Nei Mongol, Hebei)

Type locality: in provicia Chinensi, Ordos dicta (= Ordos desert, fide POPE 1935)  
ReproductionOviparous. 
TypesHolotype: ZISP 3920 
DiagnosisPholidosis. 24 – 29 scales across cap; 3 – 5 interna- sals; 13 – 15 (rarely 9) scales between parietal and supra- nasals; 11 – 17 supra- and infralabials on each side of head; 3 – 4 rows of infraorbitals; 24 – 30 scales on out- side of fourth finger of hindlimb; 1 – 2 (rarely 3) edges on subdigital plates, 4 – 5 in joint regions.

Color in life. Dorsum of body Isabel (isabellinus), ocher (pallido-ochraceus), yellowish-brown (flavido-fus- catus), smoky (fumosus), murine-gray (murinus), pale- blue-gray (cyaneo-griseus), brown (alutaceus) or dark- smoky (saturate fumosus). Bright spots in axillary area of red shades: pinkish (roseolo) with a violet (violaceus) hue on inner edge, dark-gray-violet (lividus), violet-car- mine (violaceo-carminatus), crimson, orange-red (auran- tius), or dark orange (atro-auranticus) (Fig. 6A). Speci- mens from East Alashan (38°40¢ N 105°37¢ E, No. R-13220 ZMMU) do not have an apparent axillary spot of red shades (Fig. 6C – E). Perhaps this is due to the hy- bridization of Phrynocephalus frontalis with Ph. prze- walskii, or the fact that these populations are close to the ancestral form of Ph. frontalis — przewalskii, which is clearly seen in Fig. 7. Top of head is covered in light speckles, which often merge into lines with a border of individual dark spots and stripes. Straight or arcuate dark
transverse bands or elongated spots occur in supraorbital region. Short dark longitudinal stripes located behind eyes, along sides of neck and in parietal region. Dark spots also present in area above supralabials. In young in- dividuals, top of head usually covered in scarce dark spots and has a transverse supraorbital line. Pattern on dorsal part of body consists of 8 – 10 pairs of irregular black, olive-gray (olivaceo-griseus), or olive (olivaceus) transverse bands or spots located on either side of chine (but not above vertebral column). In individuals living on more firm ground, number of pairs can be reduced to three major ones (suprascapular, in front of hips and be- tween them in the middle of back). They can be black or can have background color of back or brighter, with a thin black edging. However, many individuals may have traces of intermediate bands or spots between these well-marked bands, which markedly reduced in size and much less distinctly pronounced. Spots similar to afore- mentioned intermediate ones occur in cervical region and in proximal part of tail. In a number of cases (young indi- viduals and lizards that live on wind-blown barkhans), size of main pairs of spots (suprascapular, in front of hips and in the middle of back) may not differ from intermedi- ate ones. In this case, pattern of dorsal surface looks like a longitudinal series of small gray spots, in the area of which light tops of several of them interspersed. In many cases three main dark transverse bands extend to sides of back in form of similar, but independent spots. Dorso-lat- eral spots present and located opposite to the almost re- duced intermediate bands. As a result, on sides of back (above the line of white spots between hip and shoulder) occurs a series of four dark spots. Round or oval maculae often scattered randomly (small whitish spots, lined with a thin black or gray line) on upper side of body (on back, limbs, and base of tail), they often lined into short stripes. Transverse or oblique stripes visible on upper surface of limbs. Up to 2 – 3 dark transverse bands or pairs of spots on top of tail. White line with thin gray or black edging runs along vertebral column of tail. Between fore and hind limbs occurs an unclear light (whitish) band or a se- ries of spots with a thin dark edging. Contrasting border between lateral and abdominal surfaces wavy due to the white spots along it with a thin black border along upper edge.
Throat white, in mature males often with small rare black speckles, abdomen and lower limbs monochro- matic white. In mature males, lower part of thighs during the mating period yellow (flavus) or sulfuric yellow (sulphureus). Distal half of tail underside in young indi- viduals has 3 – 4 black transverse bands (widest bands are usually located distally, narrowest located proxi- mally, occasionally transforming into a spot). Contrast of dark transverse bands on underside of tail decreases with age. Cloacal region of immature individuals chromo-or- ange (aurantiacus), dark orange (atro-aurantiacus) or rusty (ferrugineus), this color also extends between prox- imal transverse black bands (Fig. 4C). Black distal end of tail. Females retain juvenile type of tail underside color- ing for longer. 
CommentSynonymy: BARABANOV & ANANJEVA 2007 list this species as synonym of Phrynocephalus przewalskii.

Distribution: Wang & Fu (2004) suggested to apply the name P. versicolor only to the populations in the northwestern part of its distribution area. Populations from Tengger Desert, Ordos highland, and further east include P. frontalis, P. przewalskii, and P. versicolor and should be called P. przewalskii. Some populations may contain a new species that remains to be described. Guo & Wang (2007) suggested that the Lanzhou population of P. frontalis is part of P. przewalskii.

Relationships: P. frontalis is closely related to P. przewalskii. 
References
  • 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
  • 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
  • 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
  • Dunayev, Evgeny A.; Evgeniya N. Solovyeva, Nikolay A. Poyarkov 2021. Systematics, Phylogeny, and Evolution of Phrynocephalus (Superspecies przewalskii) (Reptilia: Agamidae). Russian Journal of Herpetology 28 (1): 43-59 - get paper here
  • JI, Shengnan; Xue LIAN, Zhigang JIANG, Lili LI, Junhuai BI, and Chunwang LI 2017. Effects of Sand Grain Size on Habitat Selection in Steppe Toad-headed Lizard (Phrynocephalus frontalis). Asian Herpetological Research 8 (2): 123-130; DOI:10.16373/j.cnki.ahr.160014 - get paper here
  • JI, Shengnan; Xue LIAN; Zhigang JIANG; ; Lili LI; Junhuai BI; and Chunwang LI; ; 2019. Effects of Sand Grain Size on Habitat Selection in Steppe Toad-headed Lizard (Phrynocephalus frontalis). Asian Herpetological Research 10 (4): 123-130 - get paper here
  • Jin, Yuanting; Yubin Wo, Haojie Tong, Sen Song, Lixun Zhang and Richard P. Brown 2018. Evolutionary analysis of mitochondrially encoded proteins of toad-headed lizards, Phrynocephalus, along an altitudinal gradient. BMC Genomics (2018) 19:185 - get paper here
  • LIAN, Xue; Zhigang JIANG, Xiaoge PING, Songhua TANG, Junhuai BI and Chunwang LI 2012. Spatial Distribution Pattern of the Steppe Toad-headed Lizard (Phrynocephalus frontalis) and Its Influencing Factors. Asian Herpetological Research 3 (1): - 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
  • 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]
  • Pope, CLIFFORD H. 1935. The Reptiles of China. Turtes, Crocodilians, Snakes, Lizards. Amer. Mus. Nat. Hist., New York, Nat. Hist. Central Asia, 10: lii, 1-604 - get paper here
  • Sindaco, R. & Jeremcenko, V.K. 2008. The reptiles of the Western Palearctic. Edizioni Belvedere, Latina (Italy), 579 pp. - get paper here
  • Strauch in: PRZEWALSKI, N.M. 1876. Mongoliya i Strana Tangutov. Tryokhletneye puteshestviye v Vostochnoj Nagoruoj Asii (= Mongolia and the Tangut Country. A Three-Year Travel in Eastern High Asia). Volumes 1-2. Imper. Russ. Geogr. Soc., St. Petersburg (in Russian; English edition, London 1876, French edition, Paris 1880, German edition, Jena 1887; abridged Russian edition with added notes, Moscow 1946). - get paper here
  • Wermuth, Heinz 1967. Liste der rezenten Amphibien und Reptilien. Agamidae. Das Tierreich 86: 1-127
  • Zeng, Zhi-Gao; Jun-Huai Bi , Shu-Ran Li , Yang Wang , Travis R. Robbins , Shao-Yong Chen , and Wei-Guo Du 2016. Habitat Alteration Influences a Desert Steppe Lizard Community: Implications of Species-Specific Preferences and Performance. Herp. Monographs 30 (1): 34-48 - get paper here
 
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