Phrynocephalus KAUP, 1825

Krötenkopfagamen / Toad-headed Agamids

ANANJEVA, N.B. (1998): The skull structure of some arid Asian Agamids of Phrynocephalus genus (Agamidae, Sauria). – Russ. J. Herpet. 5 (1): 29-35.

The skull structure of some species of arid Asian agamids genus Phrynocephalus from the Middle Asia was studied. It differs from other lizards of this family by some specific characters. The construction of the short and wide skull forms its “bulldog”-like shape created by the strong protruding of the lower jaw together with quadratum bone. Of special comparative interest are the following cranial characters: the angles between maxillae and its appendices, the shift of the occipital junction, the dental formula and the size of the parietal foramen. In general the juvenile specimens have larger parietal foramen that decreases during their ontogeny. The differences in the size of the parietal foramen in several species of Phrynocephalus are noted and compared. The paedomorphic characters in the skull structure and dentition are discussed.

ANANJEVA, N.B., GUO, X. & Y. WANG (2011): Taxonomic Diversity of Agamid Lizards (Reptilia, Sauria, Acrodonta, Agamidae) from China: A Comparative Analysis. - Asian Herpetological Research 2 (3): 117-128.

ANANJEVA, N.B. & T.M. SOKOLOVA (1990): The position of the genus Phrynocephalus Kaup, 1825 in agamid systems. -. Trudy, Zoological Institute, Akademii Nauk USSR, Leningrad, 207: 12-21. (in Russisch)

ANANJEVA, N.B. & B.S. TUNIYEV (1992): Historical biogeography of the Phrynocephalus species of the USSR. – Asiatic Herpetological Research, 4: 76-98.

ARNOLD, E.N. Phylogenetic relationships of toad-headed lizards (Phrynocephalus, Agamidae) based on morphology. – Bull. nat. Hist. Mus. London (Zool.), 65: 1-13.

BADMAEVA, V.I. & N.S. ONAEVA (1977): The circadian activity of Phrynocephalus in eastern Manych. – In: Darevskij, I.S. (ed.): Fourth all-Union Herpetological Conference. Questions of Herpetology. – Akademiya Nauk SSSR, Zoologicheskij Institut. Izdatel´stvo ´Nauka´, Leningrad. 19-20. (in Russisch)

BARABANOV, A.V. & N.B. ANANJEVA (2007): Catalogue of available scientific species-group names for lizards of the genus Phrynocephalus KAUP, 1825 (Reptilia, Sauria, Agamidae). – Zootaxa, Auckland, 1399: 1-56.

CAREVSKIJ, S.P. (1929): Contribution to the classification and distribution of the lizards of the genus Phrynocephalus. – Reports of Academy of Sciences of USSR, 1929: 415-419. (in russisch)

DAREVSKIJ, I.S. (1957): Hautdrüsen auf dem Rücken einiger agamider Eidechsen der Gattung Phrynocephalus. – Zool. Jb. (Syst.), 85: 301-304.

DUNAYEV, E.A. (1996): Nomenclature and distribution of toad-agams, Phrynocephalus (Reptilia, Agamidae) in Iliyskaya Hollow. – Byulleten Moskovskogo Obshchestva Ispytalelei Prirody Otdel Biologicheskii, 101(3): 36-41. (in Russisch)

DUNAYEV, E.A. (1996): On the possible use of the ethological features in the taxonomy and phylogeny of toad agamas, Phrynocephalus (Reptilia, Agamidae). – Russian Journal of Herpetology, 3 (1): 32-38.

DUNAYEV, E.A. (2008): Phylogeny of lizards of Phrynocephalus genus (Reptilia: Agamidae): history of study and methodic approaches. – In: Voprosy gerpetologii / The Problems of Herpetology, Proceedings of the 3rd Meeting of the Nikolsky Herpetological Society 9-13 October 2006, Pushchino; St. Petersburg”: 117-126. (in Russian; English abstract).

The overview of literature allowed to conclude that Phrynocephalus species with the most archaic characteristics (Ph. arabicus, Ph. maculatus, Ph. mystaceus, Ph. scutellatus, Ph. ornatus, Ph. luteoguttatus, Ph. euptilopus and some others) are inhabited the territory of the Middle East, Fore and South Asia. They all have T-shaped frontale, short apical lobes of hemi-penises, plain repertory of tail demonstrations and single premaxillary tooth in most species. These taxons are also located at the base position at phylogenetic trees built on the results of electrophoresis and genetic data analysis and have 48 chromosomes diploid set. Mentioned facts can be considered as the proof of phylogenetic isolation of this group of species from other ones and allow to assume their present territory of living as the area of genus possible origin. This conclusion contradicts the opinion of R. S. Whiteman (1978) about Central Asian origin of Phrynocephalus. Taking into account above mentioned information it looks necessary to run more detailed investigation of phylogenetic picture of Phrynocephalus genus and its different levels clusterization. Premature separation of one or more locally grouped taxons into independent subgenus groups (Megalochilus, Oreosaurus) excessively complicates taxonomic structure of genus and hides understanding of phylogenetic relations within it.

GUO, X.-G. & Y.-Z. WANG (2007): Partitioned Bayesian analyses, dispersal-vicariance analysis, and the biogeography of Chinese toad-headed lizards (Agamidae: Phrynocephalus): a re-evaluation. – Molecular Phylogenetics and Evolution, 45: 643-662.

JIN, Y.T. & R.P. BROWN (2013): Species history and divergence times of viviparous and oviparous Chinese toad-headed sand lizards (Phrynocephalus) on the Qinghai-Tibetan Plateau. – Molec. Phylogen. Evol., 68 (2): 259-268.

The Qinghai-Tibetan Plateau (QTP) is an important biogeographical area and has recently become a focus for biodiversity studies. Phyrnocephalus lizards form a widespread Eurasian group with oviparous and viviparous reproductive modes, but two previous mtDNA studies of species from around the QTP have provided different phylogenetic hypotheses. We analysed three loci (mtDNA, RAG-1, AME) from all recognised Chinese Phrynocephalus species to reconstruct the speciation history of the group and to estimate species divergence times. The effects of mtDNA partitioning strategy on phylogenetic inference were examined. Bayes factor comparisons of marginal likelihoods (mLs) estimated using stepping-stone sampling revealed that partitioning strategy had a major impact on mL. Nevertheless, it had a negligible effect on the inferred tree topology. The impact of hard-bound uniform or equivalent soft-bound gamma speciation time calibration priors as well as the use of a fixed topology (as opposed to integration over all possible species histories) on divergence time estimation were also assessed, and found to have little impact on posterior estimates. All three gene trees and the species tree supported the hypothesis that the Chinese species form oviparous and viviparous sister clades. This was in agreement with an early mtDNA study but differed from a subsequent reanalysis of the mtDNA data. Inclusion of mtDNA from more widely distributed Phrynocephalus (from previous studies) indicates that the oviparous P. interscapularis from Central Asia lies outside the clade of Chinese viviparous and oviparous species, but that other Asian oviparous species lie within the Chinese oviparous clade. The median of the posterior on the divergence time of Chinese oviparous and viviparous species was 9.7 Ma ago (95% interval: 7.2–13.0 Ma ago), which coincides with major uplifting of the QTP and indicates that viviparity evolved when this clade became restricted to regions of high elevation. We also found that cladogenesis within the viviparous clade began around 5 Ma ago whereas those in the oviparous clade began around 8.6 Ma ago. We establish more robust estimates of divergence times and relationships within this important group and so provide improved insights into the origins of Phrynocephalus diversity across the QTP.

JIN, Y.T., LI, J.Q. & N.F. LIU (2013): Elevation-related variation in life history traits among Phrynocephalus lineages on the Tibetan Plateau: do they follow typical squamate ecogeographic patterns. – J. Zool. (London), 290 (4): 293-301.

We examined body size, litter size and evidence for Bergmann’s and Rensch’s rules among eight closely related viviparous Phrynocephalus (Agamidae) lineages from the Qinghai-Xizang (Tibetan) Plateau. Mean snout-vent length and (relative) abdomen length was greater in females than males, whereas absolute and relative mean head size, fore – and hindlimb length was larger in males. These patterns suggest that body size may reflect sexual evolutionary conservatism. The lizards are smaller at higher elevations or in colder climates, representing the converse of Bergmann’s rule. Absolute differences in female body size among taxa are not equal to the male differences among taxa. This results in a slope less than one when male size is regressed on female size, which allows rejection of Rensch’s rule for this group. The female body size elevation-related cline was steeper than the corresponding male cline. Litter sizes were both smaller and less variable at higher elevations. These elevational clines remained after application of phylogenetic comparative methods, indicating that ecological processes play a more important role than phylogeny in shaping patterns of size and reproductive variation in these lizards. It is suggested that seasonal activity and temperature are important environmental factors that contribute to the converse Bergmann’s cline, while fecundity selection in females and sex-specific differential-plasticity likely explain why patterns do not conform to Rensch’s rule.

JIN, Y., WO, Y., TONG, H., SONG, S., ZHANG, L. & R.P. BROWN (2018): Evolutionary analysis of mitochondrially encoded proteins of toad-headed lizards, Phrynocephalus, along an altitudinal gradient. – BMC Genomics, 19: 185.Background: Animals living at high altitude must adapt to environments with hypoxia and low temperatures, but relatively little is known about underlying genetic changes. Toad-headed lizards of the genus Phrynocephalus cover a broad altitudinal gradient of over 4000 m and are useful models for studies of such adaptive responses. In one of the first studies to have considered selection on mitochondrial protein-coding regions in an ectothermic group distributed over such a wide range of environments, we analysed nineteen complete mitochondrial genomes from all Chinese Phrynocephalus (including eight genomes sequenced for the first time). Initial analyses used site and branch-site model (program: PAML) approaches to examine nonsynonymous: synonymous substitution rates across the mtDNA tree.
Results: Ten positively selected sites were discovered, nine of which corresponded to subunits ND2, ND3, ND4, ND5, and ND6 within the respiratory chain enzyme mitochondrial Complex I (NADH Coenzyme Q oxidoreductase). Four of these sites showed evidence of general long-term selection across the group while the remainder showed evidence of episodic selection across different branches of the tree. Some of these branches corresponded to increases in altitude and/or latitude. Analyses of physicochemical changes in protein structures revealed that residue changes at sites that were under selection corresponded to major functional differences. Analyses of coevolution point to coevolution of selected sites within the ND4 subunit, with key sites associated with proton translocation across the mitochondrial membrane.
Conclusions: Our results identify mitochondrial Complex I as a target for environment-mediated selection in this group of lizards, a complex that frequently appears to be under selection in other organisms. This makes these lizards good candidates for more detailed future studies of molecular evolution.

KHAN, M.S. (1999): A checklist and key to the phrynocephalid lizards of Pakistan, with ethnological notes (Squamata: Agamidae). Pakistan J. Zool. 31 (1): 17-24.

LICHANOVA, O.P. (1992): Biochemical polymorphism, systematics, and phylogeny of lizards of genus Phrynocephalus (Agamidae, Reptilia). – Doctoral dissertation, Moscow State University, Moscow. 25 S. (in Russisch)

MACEY, J.R., ANANJEVA, N.B., ZHAO, E.M., WANG, Y.Z. & T.J. PAPENFUSS (1993): An allozyme-based phylogenetic hypothesis for Phrynocephalus (Agamidae) and its implication for the historical biogeography of arid Asia. – In: Zhao, E.M., Chen, P.H. & T.J. Papenfuss (eds.): Proceedings of the First Asian Herpetological Meeting. China Forestry Press, Beijing. 349-350.

MACEY, J.R., SCHULTE, J.A. II, ANAAJEVA, N.B., DYKE, E.T. van, WANG, Y., ORLOV, N., SHAFIEI, S., ROBINSON, M.D., DUJSEBAYEVA, T., FREUND, G.S., FISCHER, C.M., LIU, D. & T.J. PAPENFUSS (2018): A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics. – Zootaxa, 4467 (1): 1-81.

Phylogenetic relationships of the agamid lizard genus Phrynocephalus are described in the context of plate tectonics. A near comprehensive taxon sampling reports three data sets: (1) mitochondrial DNA from ND1 to COI (3’ end of ND1, tRNA^Gln, tRNA^Ile, tRNA^Met, ND2, tRNA^Trp, tRNA^Ala, tRNA^Asn, tRNA^Cys, tRNA^Tyr, and the 5’ end of COI) with 1761 aligned positional sites (1595 included, 839 informative), (2) nuclear RAG-1 DNA with 2760 aligned positional sites (342 informative), and (3) 25 informative allozyme loci with 213 alleles (107 informative when coded as presence/absence). It is hypothesized that Phrynocephalus phyletic patterns and speciation reflect fault lines of ancient plates now in Asia rejuvenated by the more recent Indian and Arabian plate collisions. Molecular estimates of lineage splits are highly congruent with geologic dates from the literature.  A southern origin for the genus in Southwest Asia is resolved in phylogenetic estimates and a northern origin is statistically rejected. On the basis of monophyly and molecular evidence several taxa previously recognized as subspecies are recognized as species: P. hongyuanensis, P. sogdianus, and P. strauchi as “Current Status”; Phrynocephalus bannikovi, Phrynocephalus longicaudatus, Phrynocephalus turcomanus, and Phrynocephalus vindumi are formally “New Status”. Phylogenetic evaluation indicates a soft substrate habitat of sand for the shared ancestor of modern Phrynocephalus. Size diversity maximally overlaps in the Caspian Basin and northwestern Iranian Plateau. The greatest species numbers of six in sympatry and regional allopatry are found in the southern Caspian Basin and southern Helmand Basin, both from numerous phylogenetic lineages in close proximity attributed to tectonic induced events.

MANILO, V.V. (2000): Description of karyotypes of some species and subspecies of the genus Phrynocephalus (Sauria, Agamidae) from Central Asia. – Vestn. Zool., 34 (6): 113-118. (In Russian).

MELNIKOV, D. & N. ANANJEVA (2010): Molecular studies of Phrynocephalus. Review. - Abstracts of the Second International Symposium on Agamid Lizards «DeAgamis2». - Current Studies in Herpetology, 10 (3/4): 149.

MEZHZHERIN, S.V. & M.L. GOLUBEV (1989): The genetic divergence of Phrynocephalus Kaup (Reptilia, Agamidae) of the USSR fauna. – Reports of Ukraine SSR Academy of Science. Series B Geology, Chemistry and Biological Sciences, 12: 72-74. (in Russisch)

MISHAGINA, J.V. (2008): Ecological groups of preys in Eastern Kara Kum Psammobiont lizards, Eremias (Lacertidae) and Phrynocephalus (Agamidae). – In: Voprosy gerpetologii / The Problems of Herpetology, Proceedings of the 3rd Meeting of the Nikolsky Herpetological Society 9-13 October 2006, Pushchino; St. Petersburg”: 298-307. (in Russian; English abstract).

Food habits of Eremias grammica, E. scripta (Lacertidae), Phrynocephalus mystaceus and Ph. interscapularis (Agamidae) from Eastern Kara Kum barchan - and - hillock and barchan sands (Repetek biosphere reserve, Turkmenistan, 1984— 1992) were examined by droppings analysis mainly. Preys (more than 15 thousand prey items from 685 lizards) were divided into numerous ecological groups of different rank. Variability of ecological structure of preys among 26 samples was illustrated by principal component analysis. Such the variability can be considered as a mirror of similarity and ecological isolation of sympatric species.

MOODY, M.S. (1982): Phylogenetic origins and relationships of terrestrial genera Agama, Phrynocephalus, and Uromastyx within the family Agamidae (Reptilia: Sauria). – Vertebrata Hungarica, Musei historico-naturalis hungarici, Budapest. 252.

NEYMARK, L.A. (2013): Prospects of faecal analysis as non-invasive method of trophic research: Evidence from Toad-headed Agamas (Phrynocephalus, Agamidae).- In: Ananjeva, N.B., Syromyatnikova, E.V. & I.V. Doronin (eds.): Modern herpetology: Problems and ways of their solutions. The first International Conference of the Young Herpetologists of Russia and Neighboring Countries 25–27 November 2013, Saint-Petersburg. Russian Academy of Sciences Zoological Institute of RAS A.M. Nikolsky Herpetological Society. pp. 118-120. (in Russisch)

NOBLE, D.W.A., QI, Y. & J. FU (2010): Species delineation using Bayesian model-based assignment tests: a case study using Chinese toad-headed agamas (genus Phrynocephalus). - BMC Evolutionary Biology 2010, 10:197.

Background: Species are fundamental units in biology, yet much debate exists surrounding how we should delineate species in nature. Species discovery now requires the use of separate, corroborating datasets to quantify independently evolving lineages and test species criteria. However, the complexity of the speciation process has ushered in a need to infuse studies with new tools capable of aiding in species delineation. We suggest that modelbased assignment tests are one such tool. This method circumvents constraints with traditional population genetic analyses and provides a novel means of describing cryptic and complex diversity in natural systems. Using toadheaded agamas of the Phrynocephalus vlangalii complex as a case study, we apply model-based assignment tests to microsatellite DNA data to test whether P. putjatia, a controversial species that closely resembles P. vlangalii morphologically, represents a valid species. Mitochondrial DNA and geographic data are also included to corroborate the assignment test results. Results: Assignment tests revealed two distinct nuclear DNA clusters with 95% (230/243) of the individuals being assigned to one of the clusters with > 90% probability. The nuclear genomes of the two clusters remained distinct in sympatry, particularly at three syntopic sites, suggesting the existence of reproductive isolation between the identified clusters. In addition, a mitochondrial ND2 gene tree revealed two deeply diverged clades, which were largely congruent with the two nuclear DNA clusters, with a few exceptions. Historical mitochondrial introgression events between the two groups might explain the disagreement between the mitochondrial and nuclear DNA data. The nuclear DNA clusters and mitochondrial clades corresponded nicely to the hypothesized distributions of P. vlangalii and P. putjatia. Conclusions: These results demonstrate that assignment tests based on microsatellite DNA data can be powerful tools for distinguishing closely related species and support the validity of P. putjatia. Assignment tests have the potential to play a significant role in elucidating biodiversity in the era of DNA data. Nonetheless, important limitations do exist and multiple independent datasets should be used to corroborate results from assignment programs.

PANG, J., WANG, Y., ZHONG, Y., HOELZEL, A.R., PAPENFUSS, T.J., ZENG, X., ANAJEVA, N.B. & Y. ZHANG (2003): A phylogeny of Chinese species in the genus Phrynocephalus (Agamidae) inferred from mitochondrial DNA sequences. – Molecular Phylogenetics and Evolution, 27: 398-409.

In this study, we report the complete mitochondrial genome of Phrynocephalus forsythii (Reptilia, Squamata, Agamidae), which is a circular molecule of 16,143 bp in size and consists of 13 protein-coding genes, 22 transfer RNAs, 2 ribosomal RNAs and 2 non-coding sequence (D-loop). The mitogenome of P. forsythii was similar to the typical mtDNA of vertebrates in gene arrangement and composition. The control region composed of two parts: one (348 bp) between tRNAPhe and the other (636 bp) between tRNAPro and 12S rRNA. The A + T content of overall base of the composition of H-strand is 62.0% (T: 25.6%, C: 25.7%, A: 36.3% and G: 12.3%). The whole mitogenomic sequence of P. forsythii provides powerful data to study of ist phylogenetic position within toad-headed lizards.

SHARMA, R.C. (1978): A new species of Phrynocephalus Kaup (Reptilia: Agamidae) from the Rajastan Desert, India with notes on its ecology. – Bulletin of the Zoological Survey of India, 1 (3): 291-294.

SOKOLOVSKY, V.V., (1975): Comparative and karyologic study of the lizards in the family Agamidae. 1. Chromosome complements of 8 species of the genus Phrynocephalus (Reptilia, Agamidae). – Tsitologiya, Leningrad, 17 (1): 91-93. (in Russisch)

SOLOVYEVA, E.N. (2013): Structure of genetic variation and phylogeny of the genus Phrynocephalus (Reptilia: Agamidae). Dissertation. Moscow. 23 pp (in Russisch)

SOLOVYEVA, E.N., DUNAYEV, E.A., NAZAROV, R.A., BONDARENKO, D.A. & N.A. POYARKOV (2023): COI-barcoding and species delimitation assessment of toad-headed agamas of the genus Phrynocephalus (Agamidae, Squamata) reveal unrecognized diversity in Central Eurasia. – Diversity 2023, 15, 149.

We provide a diversity assessment of the agamid genus Phrynocephalus Kaup, 1825. We analyze COI mtDNA barcodes from 385 individuals sampled all over Phrynocephalus range. We apply the ABGD, ASAP, bGMYC, mlPTP and hsPTP species delimitation algorithms to analyze the COI gene fragment variation and assess the species diversity in Phrynocephalus. Nine species groups are revealed in Phrynocephalus in agreement with earlier studies on the phylogenetic relationships of the genus. We demonstrate that the present taxonomy likely underestimates the actual diversity of the genus. Alternative species delimitation algorithms provide a confusingly wide range of possible number of Phrynocephalus species—from 54 to 103 MOTUs (molecular operational taxonomic units). The ASAP species delimitation scheme recognizing 63 MOTUs likely most closely fits the currently recognized taxonomic framework of Phrynocephalus. We also report on 13 previously unknown Phrynocephalus lineages as unverified candidate species. We demonstrate that the ASAP and the ABGD algorithms likely most closely reflect the actual diversity of Phrynocephalus, while the mlPTP and hsPTP largely overestimate it. We argue that species delimitation in these lizards based exclusively on mtDNA markers is insufficient, and call for further integrative taxonomic studies joining the data from morphology, mtDNA and nuDNA markers to fully stabilize the taxonomy of Phrynocephalus lizards.

SOLOVYEVA, E.N., POYARKOV, N.A., DUNAYEV, E.A., NAZAROV, R.A., LEBEDEV, V.S. & A.A. BANNIKOVA (2014): Phylogenetic Relationships and Subgeneric Taxonomy of Toad-headed Agamas Phrynocephalus (Reptilia, Squamata, Agamidae) as Determined by Mitochondrial DNA Sequencing. - Doklady Biological Sciences 455: 119–124.

SOLOVYEVA, E.N., SIMONOV, E.P., ORESHKOVA, N.V. & N.A. POYARKOV (2021): What the full mitochondrial genomes of the toad-headed agamas can tell us (Phrynocephalus, Agamidae). - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p. 240-242. (in Russisch)

WANG, Z., LU, H.-L., MA, L. & X. JI (2014): Viviparity in high-altitude Phrynocephalus lizards is adaptive because embryos cannot fully develop without maternal thermoregulation. – Oecologia, 174: 639-649.

Viviparous Phrynocephalus lizards (Agamidae) are mainly restricted to the Qinghai-Tibet Plateau of China. In this study, we used Phrynocephalus vlangalii females kept under seven thermal regimes for the whole gestation period to test the hypothesis that viviparity in high-altitude Phrynocephalus lizards is adaptive because embryos cannot fully develop without maternal thermoregulation. All females at 24 °C and 93 % of the females at 28 °C failed to give birth or produced stillborns, and proportionally fewer females gave birth at 29 or 35 °C than at 32 °C. Though the daily temperatures encountered were unsuitable for embryonic development, 95 % of the females in nature and 89 % of the females thermoregulating in the laboratory gave birth. There was no shift in the thermal preferences of females when they were pregnant. Although thermal conditions inside natural burrows were unsuitable for embryonic development, mass and sprint speed were both greater in neonates produced in nature. Our data show that (1) longterm exposure of P. vlangalii embryos to temperatures outside the range of 29–35 °C may result in the failure of development, but daily or short-term exposure may not necessarily increase embryonic mortality; (2) low gestation temperatures slow but do not arrest embryonic development, and females produce high-quality offspring in the shortest possible time by maintaining gestation temperatures close to the upper thermal limit for embryonic development; and (3) viviparity is currently adaptive at high elevations because embryos in nature cannot fully develop without relying on maternal thermoregulation. Our data validate the hypothesis tested.

WANG, Z., MA, L., SHAO, M. & X. JI (2013): Differences in Incubation Length and Hatchling Morphology among Five Species of Oviparous Phrynocephalus Lizards (Agamidae) from China. – Asian Herpetological Research 4 (4): 225-232.

We incubated eggs of five Phrynocephalus species (P. albolineatus, P. axillaries, P. grumgrzimailoi, P. helioscopus and P. przewalskii) at three constant temperatures (24 °C, 28 °C and 32 °C) to examine differences in incubation length and hatchling morphology among species and among temperature treatments. We combined data from this study with those reported previously for P. frontalis and P. versicolor to examine whether embryonic stage at laying is a causal factor for interspecific variation in incubation length, and whether the phylogenetic relationship inferred from hatchling morphology is consistent with the relationship based on mitochondrial DNA data. Mean values for incubation length differed among the five species studied herein and, in all these five species, incubation length decreased at a decreasing rate as temperature increased. In none of the five species did hatchling size (snout-vent length and body mass) and other morphological variables differ among the three temperature treatments. The seven oviparous Phrynocephalus lizards found in China differ from each other in hatchling morphology, and embryonic stage at laying is a causal factor of inter- and intra-specific variation in incubation length. The phylogenetic relationship inferred from hatchling morphology is not always consistent with the currently known relationship based on mitochondrial DNA data. Data from this study and those reported previously allow the conclusion that any Phrynocephalus species may have its unique position along the axis defined by hatchling morphology.

WANG, Y.Z. & J.R. MACEY (1993): On the ecologico-geographic differentiation of Chinese species of the genus Phrynocephalus. – In: Proceedings of the First Asian Herpetological Meeting. Forestry Press, China, Beijing. 147-153. (in chinesisch)

WANG, Y.Z., ZENG, X.M., FANG, Z.L., WU, G.F., PAPENFUSS, T.J. & J.R. MACEY (1999): Study on the relationships of classification, phylogenetics and distribution of the genus Phrynocephalus ssp. (Sauria: Agamidae) wirthe the paleogeographical changes during Cenozoic era in Tibet plateau. – Zoological Research, 20: 178-185. (in chinesisch)

WHITEMAN, R.S: (1978): Evolutionary history of the lizard genus Phrynocephalus (Lacertilia, Agamidae). – Master Thesis, California State University, Fullerton. VIII + 113 S.

ZENG, X.M., WANG, Y.Z., LIU, Z.J., FANG, Z.L., WU, G.F., PAPENFUSS, T.J. & J.R. MACEY (1997): Karyotypes of nine species in the genus Phrynocephalus, with discussion of karyotypic evolution of Chinese Phrynocephalus. – Acta Zool. Sinica, 43: 399-410. (in chinesisch)

ZHAO, K.T. (1997): Notes on the Chinese toad-headed agamids and its diagnostic characters. – J. Suzhou Railway Teachers College, 14: 27-32. (in chinesisch)

ZHAO, K.T. (1999): Phrynocephalus Kaup, 1825. – In: Zhao, E.M. ZHA, K.T. & K.Y. ZHOU (eds.): Fauna Sinica Vol. 2. Squamata. Lacertilia. Science Press, Beijing: 151-193.

Phrynocephalus ananjevae MELNOKOV et al., (2013)

MELNIKOV, D., MELNIKOVA, E., NAZAROV, R. & M. RAJABIZADEH (2013): Taxonomic revision of Phrynocephalus persicus DE FILIPPI, 1863 complex with description of a new species from Zagros, Southern Iran. - СОВРЕМЕННАЯ ГЕРПЕТОЛОГИЯ, 13 (1-2): 34–46.

Phrynocephalus arabicus ANDERSON, 1894

Arabische Krötenkopfagame / Arabian Toadhead Agama

ABU BAKER, M., QUARQAZ, M., RIFAI, L., HAMIDAN, N., AL OMARI, K., MODRY, D. & Z. AMR (2004): Results of herpetofaunal inventory of Wadi Ramm protected area, with notes on some relict species. – Russian Journal of Herpetology, 11 (1): 1-5.

AL-SIRHAN, A.-R. & G. BROWN (2010): The status of the two Toad-headed Agamas, Phrynocephalus arabicus (Anderson, 1894), and P. maculatus (Anderson, 1872), in Kuwait. – Zoology in the Middle East, 51.

ANDERSON, J. (1894): On two new species of agamoid lizards from the Hadramut, South-Eastern Arabia. – Ann. Mag. Nat. Hist. (6) 14: 377.

ANDERSON, S.C. (1999): Phrynocephalus arabicus Anderson, 1894. - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 84-85.

LONDEI, T. (2015): Arabian sand boa Eryx jayakari (Squamata: Boidae) preying on Arabian toad-headed agama Phrynocephalus arabicus (Squamata: Agamidae): a nocturnal-to-diurnal species interaction - Herpetology Notes 8: 155-156.

MANDAVILLE, J.P. (1968): The Toad-head from Najd and other reptiles. – Saudi Aramco World, 1968 (september-october): 30-33.

MELNIKOV, D., MELNIKOVA, E., NAZAROV, R., RAJABIZADEH, M., AL-JOHANY, A.,& S. ZUHAIR (2014): Taxonomic revision of Phrynocephalus arabicus ANDERSON, 1984 complex with description of a new species from Ahvaz, South-Western Iran. – Russian Journal of Herpetology, 21 (2): 149-159.

ROSS, W. (1995): Tail signalling in populations of Phrynocephalus arabicus, ANDERSON, 1894 (Reptilia: Agamidae). – Zoology in the Middle East, 11: 63-71.
Kurzfassung:
Es werden Beobachtungen zur ventralen Schwanzfürbung in bezug auf das Sexual- und Sozialverhalten der Arabischen Krötenkopfagame Phrynocephalus arabicus im östlichen Saudfi Arabien mitgeteilt.

SCHUSTER, R.K. (2012): A new species of Oochoristica (Cestoda, Linstowiidae) from the Arabian Toad-Headed Agama, Phrynocephalus arabicus (Sauria, Agamidae), from the United Arab Emirates. – Vestnik Zoologii, 46 (3): 29-32.

Eight complete strobilae of Oochoristica phrynocephali Schuster, sp. n. were recovered from small intestines of 4 out of 13 Arabian toad-headed agamas, Phrynocephalus arabicus from Dubai emirate, United Arab Emirates. O. phrynocephali belongs to a group of Oochoristica species possessing circular suckers and fewer than 25 testes in a single cluster. It can be distinguished from O. sobolevi, O. elongata, O. parvogenitalis, O. feliui, O. jonnesi, O. macallisteri, O. lygosomatis, O. junkea and O. novaezealandae by smaller scolices and lesser diameter of suckers.

STARKES, J. & T. ACKERMANN (2021): Ein sandiger und nebliger Weg – zur Haltung und Vermehrung der arabischen Krötenkopfagame Phrynocephalus arabicus (ANDERSON, 1894). – Sauria, Berlin, 43 (4): 1-12.

WITTENBERG, J. (1992): First records of Arabian Toad-head agamid, (Phrynocephalus arabicus) (Sauria: Agamidae) in Jordan. – Zoology in the Middle East, Heidelberg, 7: 59-64.
Kurzfassung:
Für Jordanien wurde erstmals eine Agame des Genus Phrynocephalus nachgewiesen. In Sanddünengebieten mit Büschen beim Wadi Didi (ca. 29°37’N, 35°40’E) im Süden des Landes wurden am 8.4.1988 erst ein Exemplar, dann ca. 4 km südlich am 19.4.1990 zwei Exemplare fotografiert und als Arabische Krötenkopfagame (P. arabicus) nachbestimmt. Die Möglichkeit, daß es sich um P. maculatus gehandelt haben könnte, wird anhand von Merkmalen, Habitat und Verbreitung ausgeschlossen.

Phrynocephalus axillaris BLANFORD, 1875

Krötenkopfagame

AUTUMN, K. & Y.Z. WANG (1988): Preliminary observations on the ecology of Phrynocephalus axillaris and Eremias velox in the Turpan Depression, Xinjiang Uygur Autonomous region, China. – Chinese Herpet. Res., 2 (1): 6-13.

BLANFORD, W.T. (1875): List of Reptilia and Amphibia collected by the late Dr. STOLICZKA in Kashmir, Ladák, Eastern Turkestán, and Wakhán, with descriptions of new species. – Journal of the Asiatic Society of Bengal, Vol. XLIV Part II (Natural History): 191-196.

DUNAYEV, E.A. (2020): History of Study, Taxonomy, Distribution, and Ecology of Phrynocephalus nasatus Golubev et Dunayev, 1995 (Reptilia: Agamidae). – Russian Journal of Herpetology, 27 (2): 87-96.

Herein we provide a historical overview of the study of Phrynocephalus nasatus, a species that has been known by poorly preserved type materials only, collected by A. I. Wilkins in Aksu region, China, in 1883. Information on species ecology and distribution is given for the first time in 130 years. The species range consists of disjunct areas which are divided by river valleys covering 4000 km², at the altitude of 2000 m a.s.l., in the eastern spurs of the Jengish Chokusu mountain. Ph. nasatus prefers clay-gravel biotopes with scattered vegetation. This species differs from Ph. axillaris, with which it was erroneously synonymized lately, by the utricular nasal scales (the trait is absent in all other species of this genera) and at least 23 other traits. The intravital coloration is described.

DUNAYEV, E.A. (2021): Characteristics of the biotopic and trophic ecology of toad-headed agamas (Reptilia: Agamidae: Phrynocephalus spp.) from the Xinjiang Uygur Autonomous Region of China. - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p 105-107. (in Russisch)

Information about the biotopic and trophic preferences is given for the three species of toad-headed agamas (Phrynocephalus). The diet of Ph. axillaris is signiicantly wider than that of Ph. nasatus and Ph. forsythii, which directly correlates with the breadth of its bio-topic niche. Ph. axillaris lives in gravelly-clay deserts with pebbles, saline areas, and sandy massifs, which determines the diversity of vegetation and associated insects. Ph. nasatus occurs in intermountain clay-gravelly plains cut by watercourses with sparse vegetation cover (Dunayev, 2020). Ph. forsythii lives in hilly sands and depressions between barkhan dunes with compressed sand. Beetles and ants form the basis of diet in toad-headed agamas. The presence of shed epithelium in the faeces is likely explained by the need to consume microelements entering the skin from the soil. Nutritional values are given in percentage for prevalence and abundance in the faeces for all three species examined.

LI, J., GUO, X.G. & Y. WANG (2010): Population genetic differentiation of Phrynocephalus axillaris in East of Xinjiang Uygur Autonomous Region based on sequence variation of mitochondrial ND4-tRNALeu gene. – Zool. Res., 31 (4): 361-369. (In Chinese).

A 838 bp fragment of mtDNA ND4-tRNALeu gene was sequenced for 66 individuals from five populations (DB: Dabancheng, TU: Turpan, SS: Shanshan, HL: Liushuquan, HD: East district of Hami) of Phrynocephalus axillaris distributed in east of Xinjiang Uygur Autonomous Region. Seventeen haplotypes were identified from 29 nucleotide polymorphic sites in the aligned 838 bp sequence. Excluding DB, there were relatively high haplotype diversity [(0.600±0.113)≤h≤(0.922±0.012)] and low nucleotide diversity[(0.00082±0.00020)≤π≤(0.00917±0.00037)] in the whole and the rest four populations. The result of phylogenetic analysis showed that the haplotypes from five populations formed clade I (DB+TU+SS) and clade II (HL+HD). Network analysis further revealed the phylogeographical structure of these five populations. Analysis of molecular variance (AMOVA) indicated that the significant genetic divergence (81.67%, P<0.01) existed among populations, which was not obviously correlated with the geographic distances between populations inferred from isolation-with-distance analysis (r=0.5324, P=0.1196). The demography analysis implied that TU and HL might have experienced the recent population expansion; SS and HD might be relatively stable. With extremely low genetic diversity, DB might be influenced by the special environment and be experiencing the bottleneck effect. It followed that the significant genetic differentiation among the five populations of P. axillaris might be the consequence of habitat changes resulting from the geological events and climate oscillation since Pleistocene and genetic.

XIE, H., JIANG, L.-X., ZHAO, W. et al (2023): Genetic diversity analysis of Phrynocephalus axillaris based on nucleair gene and microsatellite. – J. Anhui Agric. Sci., 51 (6): 81-87.

ZHANG, Q., XIA, L., HE, J.B., WU, Y., FU, J. & Q. YANG (2010): Comparison of phylogeographic structure and population history of two Phrynocephalus species in the Tarim Basin and adjacent areas. – Molec. Phylogen. Evol., 57 (3): 1091-1104.

An aridification of the Tarim Basin and adjacent areas since middle Pleistocene has produced significant genetic structuring of the local fauna. We examined the phylogeographic patterns, population structure and history of Phrynocephalus axillaris and Phrynocephalus forsythii using a mitochondrial fragment ND4- tRNALEU. Phylogenetic hypotheses were constructed using maximum parsimony and Bayesian inference, and the divergence times of major lineages were estimated by BEAST. Population structure and history were inferred by nested clade analysis, neutrality tests, mismatch distribution, and isolation by distance analysis. The two species might have experienced different evolutionary history throughout their current distribution. For P. forsythii, a vicariant event, as a consequence of geological isolation and desert expansion, might have produced the significant divergence between the Tarim and the Yanqi populations. For P. axillaris, populations of the Yanqi, Turpan and Hami Basins might have been established through dispersal during demographic expansion. Climatic fluctuations caused alternate expansion and shrinkage of rivers and oases several times, which likely led to habitat fragmentation for both species. Interaction between vicariance, dispersal and habitat fragmentation produced the current distribution and genetic diversity. The observed difference between the two species may be due partially to their different reproductive modes (ovoviviparous vs. oviparous).

Phrynocephalus clarkorum ANDERSON & LEVITON,1967

ANDERSON, S.C. (1999): Phrynocephalus clarkorum Anderson and Leviton, 1967. - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 85-86.

ANDERSON, S.C. & A.E. LEVITON (1967): A new species of Phrynocephalus (Sauria: Agamidae) from Afghanistan, with remarks on Phrynocephalus ornatus Boulenger. - Proc. Cal. Acad. Sci., 35 (11): 227-234.

CLARK, R.J. (1992): Notes on the distribution and ecology of Phrynocephalus clarkorum Anderson & Leviton 1967 and Phrynocephalus ornatus Boulenger 1887 in Afghanistan. – Herpetological Journal, 2: 140-142.v

Phrynocephalus forsythii ANDERSON, 1872

Forsyth´s Toadhead Agama

ANDERSON, J. (1872): Description of Phrynocephalus maculatus and P. forsythii. - In: On some Persian, Himalayan and other Reptiles. - Proceedings of the Zoological Society of London 1872 : 371-404.

CHEN, D., ZHOU, T. & X. GUO (2015): The complete mitochondrial genome of Phrynocephalus forsythii (Reptilia, Squamata, Agamidae), a toad-headed agama endemic to the Taklamakan Desert. – Mitochondria DANN, 2015. DOI: 10.3109/19401736.2014.1003837.

The complete mitochondrial genome was sequenced from a viviparous toad-headed agama, Phrynocephalus forsythii, which is endemic to the Taklamakan Desert. The mitogenome sequence was 17,542 bp in size, containing 13 protein-coding genes, 22 tRNA genes, two rRNA genes and a control region (D-loop). The gene arrangement and composition of P. forsythii is identical to the mitogenome of P. theobaldi in that tRNA-Pro was translocated immediately downstream of tRNA-Phe. The D-loop comprised two parts, one existing between tRNA-Thr and tRNA-Phe, and another containing 12 copies of 36-bp tandem repeats inserting between tRNAPro and 12S rRNA. The complete mitogenome sequence of P. forsythii may provide fundamental data for unveiling the phylogenetic origin and adaptive evolution related to Phrynocephalus viviparity.

DUNAYEV, E.A. (2021): Characteristics of the biotopic and trophic ecology of toad-headed agamas (Reptilia: Agamidae: Phrynocephalus spp.) from the Xinjiang Uygur Autonomous Region of China. - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p 105-107. (in Russisch)

SHENBROT, G.I. & D.V. SEMYONOV (1990): A new species of the genus Phrynocephalus (Reptilia, Agamidae) from Turkmenia. - Zoologicheskii Zhurnal 69 (9): 154-156.

Phrynocephalus guttatus GMELIN, 1789

Gefleckter Krötenkopf /Spotted Toadhead Agama

ANANJEVA, N.B. (1981): Phrynocephalus guttatus (Gmelin 1789) – Gefleckter Krötenkopf. – In: Böhme, W. (Hrsg.): Handbuch der Reptilien und Amphibien Europas. Vol. 1. Echsen (Sauria) I. Akademische Verlagsgesellschaft, Wiesbaden. 180-190.

DUNAYEV, E.A., SOLOVYEVA, E.N. & N.A. POYARKOV (2020): Taxonomy, Phylogeny and Distribution of Phrynocephalus (superspecies guttatus) (Reptilia: Agamidae). - Current Studies in Herpetology, 20 (1-2): 16–34. (In Russisch).

7 qualitative pholidosis characters were revealed, according to which the phylogenetic groups of spotted toad-headed agamas, Phrynocephalus (superspecies guttatus), reliably differ from each other, and a diagnostic key was designed based thereon for distinguishing representatives of the group. Molecular phylogenetic analysis of a fragment of the COI gene of mtDNA confirmed the differentiation of Ph. melanurus in two lineages; also Ph. incertus and Ph. kuschakewitschi stand apart with high supports. The subspecies Ph. g. kalmykus on the obtained dendrogram represents a separate lineage close to the nominative subspecies Ph. g. guttatus, whereas Ph. g. salsatus, by contrast, falls into one clade with Ph. g. guttatus.

GOLUBEV, M.L. (1989): Phrynocephalus guttatus (GMEL. or Ph. versicolor str. (Reptilia, Agamidae) which one inhabits Kazakhstan? – Vestnik zoologii, Kiev, 23 (5): 38-45. (in russisch)

GOLUBEV, M.L., GORELOV, Y.K., DUNAYEV, E.A. & T.I. KOTENKO (1995): On the finding of Phrynocephalus guttatus (Gmel.) (Sauria, Agamidae) in Turkmeniya and its taxonomic status. – Byulleten Moskovskogo Obshchestva Ispytalelei Prirody Otdel Biologicheskii, 100 (3): 31-39. (in Russisch)

GORDEEV, D.A., ZHAKUPOVA, G.A. & S.K. PRILIPKO (2014): Ecology and morphology of the spotted toad agama (Phrynocephalus guttatus (Gmelin, 1789) in the north area in the Volga region (Volgograd region). – In: Symbiosis-Russia 2014: materials of the VII All-Russian Congress of Young Biologists (Ekaterinburg, October 6-10, 2014). - Ekaterinburg: Ural University Publishing House, 2014: 198-200. (In Russian).

Presents data on the distribution, morphology, population dynamics of Phrynocephalus guttatus, and analysis of the geographic variability of the basic metric characteristics.

GORIN, V.A., DUNAYEV, E.A. & B.D. VASSILIEV (2023): Step by step examination of tail movement sequences reveals functional differentiation in signals of spotted Toad-headed Agamas Phrynocephalus guttatus (Gmelin, 1789) (Reptilia: Agamidae). – Russian Journal of Herpetology, 30 (1): 56-64.

Phrynocephalus lizards are well-known for their remarkable tail movements. Possible function of such displays still remains unclear. We present a new approach for studying displays of Phrynocephalus, analyzing them as a sequence of movements. Here, we describe four tail movements of Phrynocephalus guttatus on the basis of observations on the population from the surroundings of Astrakhan, Russia. We found that movement sequences are structured and two main stereotyped patterns for movement sequences are suggested to distinguish depending on function they perform — aggressive or submissive. We also explain differences in preference and structure of movement sequences for lizards of different sex and age groups from the point of their social status.

KIDOV, A.A., KHAYRUTDINOV, I.Z., IVANOV, A.A. & E.A. KIDOVA (2020): Age structure and growth of the spotted toadhead agama, Phrynocephalus guttatus (reptilia, sauria, agamidae) in Terek Sand Massif (Eastern Ciscaucasia). – Nat. Sci. Zool., 3 (31): 56-67.(In Russian).

Background. The spotted toadheaded agama is one of the most widely distributed species of the genus. Until now, attempts have been made to determine the age and growth of this lizard only by selecting of size groups or individual tagging. The purpose of the study is to estimate the age structure of the population and the growth characteristics of the spotted toad-head agama in Terek Sand Massif using the method of skeletochronology. Materials and methods. 81 lizards were caught in the vicinity of the Peschaniy set[1]tlement in Kursk district Stavropol Krai. The body length (SVL) and tail length (LT) were measured. To determine the age with skeletochronology method, the phalange of third finger from hind right limb was used. After these procedures, the animals were released into the nature. Results. Most of the captured lizards were young animals (39,5 %) and adult males (39,5 %). Adult females were 21,0 %. In accordance to the age determined by the method of skeletochronology, there were juveniles (39,5 %), yearlings (39,5 %), two-year-olds (19,8 %) and three-year-olds (1,2 %). In the group of adult males, the absolute majority were yearlings (96,9 %), and there was only one two-year-old lizard (3,1 %). In adult females, the majority of individuals were two years old (88,2 %) and only one specimen was one year (5,9 %) and three years (5,9 %) old. The length of lizard’s body at different age was statistically significantly different. The largest sizes had the oldest animals. Conclusions. Lizards of this species grow intensively throughout their lives, differing well in body length in different age groups. For this reason, it is possible to distinguish individuals of different age groups by size with high confidence.

LIU, Y., SONG, Y. LI, W. & L. SHI (2012): Reproductive Strategy and Cycle of the Toad-headed Agama Phrynocephalus grumgrzimailoi (Agamidae) in Xinjiang, China. - Asian Herpetological Research, 3 (3): 198-204.The reproductive strategy and cycle of Phrynocephalus grumgrzimailoi were studied at a locality close to Urumqi City, Xinjiang, China. The hatchlings of P. grumgrzimailoi need at least two years to reach sexual maturity, with the smallest mature male and female measured 48.02 mm and 47.01 mm snout-vent length (SVL), respectively. Adult females produce a single clutch per breeding season, with the clutch size ranging from 1 to 5. The clutch size and mass are significantly correlated with female SVL. There is no correlation of mean egg size with clutch size and relative fecundity in P. grumgrzimailoi, suggesting that the trade-off is absent between mean egg size and number. Females increase reproductive output mainly through increasing egg numbers. The copulation period lasts from April to June. Females begin vitellogenesis in April and lay eggs from May to July. Our results suggest that toad-headed lizards tend to select different reproductive strategies to adapt themselves to their arid or semi-arid habitats.

LOTIEV, K. & A. BATKHIEV (2024): Biotopic distribution of three sympatric species of lizards (Lacertilia) in the Terek sands. - Principy èkologii, 2024 (1): 61‒68.

The background species of the saurofauna of the southeastern periphery of the Tersk Sands in the early 1980s were Phrynocephalus guttatus (Gmelin, 1789), Eremias velox (Pallas, 1771) and Eremias arguta (Pallas, 1773). Research conducted in 1983–1984 showed that each of them had different preferred biotopes: Phr. guttatus – blown sands, E. arguta – fixed sands, E. velox – scattered shrubbery. At the turn of the 20th and 21st centuries, Phr. guttatus completely disappeared in the study area on the southeastern periphery of the Tersk Sands, and after 2008 - E. velox. Currently, the territory is inhabited by E. arguta and the widespread Lacerta strigata Eichwald, 1831. The rapid changes in the composition of the saurofauna can be explained primarily by the strict biotopic confinement of lizards in conditions of total steppification of the Terek sands. The transformation of landscapes brought stenobiont species, in the conditions of the Eastern Pre-Caucasus psammophile species of Turanian genesis, to the brink of extinction and created favorable conditions for eurybiont and/ or steppe forms. If existing trends continue, we can expect the complete disappearance of such obligate psammophiles as Phr. mystaceus (Pallas, 1776) and Phr. guttatus in the Terek Sands. They may be followed by Trapelus sanguinolentus (Pallas, 1814) and E. velox. In this case, biotopic preferences of a species determine their current and prospective sociological status.

KUBYKIN, R.A. (1977): Ecological observations on tagged Phrynocephalus guttatus in the lower reaches of the River Il´ya southern Pribalkhash. – In: Darevslkij, I.S. (ed.): Fourth all-Union Herpetological Conference. Questions of Herpetology. – Akademiya Nauk SSSR, Zoologicheskij Institut. Izdatel´stvo ´Nauka´. Leningrad. 122-123. (inRussisch)

KUBYKIN, R.A. (1977): Behaviour of the lizard Phrynocephalus guttatus in the lower reaches of the River Ili (southern Pribalkhash´e. – In: Manteifel, B.P. (ed.): Animal group behaviour. – Izdatel´stvo Nauk, Moscow. 204-206. (in Russisch)

MANILO, V.V. & M.L. GOLUBEV (1993): Karyotype information oon some toad agamas of the Phrynocephalus guttatus species group (Sauria, Agamidae) of the former USSR. – Asiatic Herpetological Research, Berkeley, 5: 105-108.

MEZHZHERIN, S. & M.I. GOLUBEV (1993): Allozyme variation and genetic relationships within the Phrynocephalus guttatus species group (Sauria: Agamidae) in the Former USSR. – Asiatic Herpetological Research, 5: 59-64.

NAKARENOK, E.G. (2003): Distribution and ecological features of the Toad-headed Agama (Phrynocephalus guttatus) of the sandy desert of the northern Caspian region. – Volga Ecol. J., 1: 77-81.

The results of our reptile research in the sandy deserts of the Northern Caspian area in 1975 – 1990 are presented. The habitats, biotopic distribution, and ecological peculiarities of Spotted toad-headed agama are discussed.

NEYMARK, L.A. (2021): On the thermal biology of psammophilous lizards of Kalmykia. - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p 195-197. (in Russisch)

OKSHTEIN, I.L. (2016): The thermal biology of the Phrynocephalus guttatus (Gmelin, 1789) in the Astrakhan region. - Proceedings of the 6th congress of the A.M. Nikolsky Herpetological Society: 107-112. (in Russisch)

Th ermophysiological (maximum 41,3 °С and minimum 35,7 °С temperature of the full activity, mode thermostabilization range 38–40 °С, mode is 38,6 °C) and thermoecological (voluntary maximum and minimum, preferred temperatures, night temperatures 10–22 °С) activity indices were determined in Ph. guttatus in May. Four general forms of behavior were observed: warming (22,5–36,1 °C), thermostabilizing behavior (35,7–41,0 °C), voluntary overheat (36,0–41,3 °С) and cooling (37,8–29,3 °C).

QU, Y.F., GAO, J.F., MAO, L.X. & X. JI (2011): Sexual dimorphism and female reproduction in two sympatric toad-headed lizards Phrynocephalus frontalis and P. versicolor (Agamidae). – Anim. Biol., 61: 139-151.

SHENBROT, G.I. & D.V. SEMENOV (1987): Present distribution and taxonomy of Phrynocephalus guttatus (Reptilia, Agamidae). – Russian Journal of Zoology, Moscow, 66 (2): 259-271. (in russisch)

TABACHISHIN, V.G. (2010): Spatial distribution and abundance trends of spotted toad-headed agama, Phrynocephalus guttatus, in its northern habitat in the Volga region. - Abstracts of the Second International Symposium on Agamid Lizards «DeAgamis2». - Current Studies in Herpetology, 10 (3/4): 155-156.

TABACHISHIN, V.G., KAIBELEVA, E.I., ZAVIALOV, E.V. & G.V. SHLY AKHTIN (2005): Ecological features of Phrynocephalus guttatus in the north of the lower Volga region. - Volga Ecol. J., 2005 (2): 180–184.

The northern habitat boundary of Phrynocephalus guttatus in European Russia has been mapped. Outlying settlements have been found in the Volgograd region: the reptiles inhabit an isolated sand massif on the left bank of the Don river between v. Kamyshi (Kalachev district) in south and v. Peskovatka (Gorodishcheno district) in north. The chromosome set of the species contains 12 pairs of macrochromosomes (1 – 12th pairs) and 11 pairs of microchromosomes (13 – 23th pairs), 2n = 46, NF = 46. The first chromosome pair looks bigger than the others.

TABACHISHIN, V.G. & E.V. ZAVIALOV (2005): Present distribution and taxonomic status of the Spotted Toad-headed Agama Phrynocephalus guttatus (GMELIN, 1789) in the Volga region, Russia. [Gegenwärtige Verbreitung und taxonomischer Status der Gefleckten Krötenkopf-Agame Phrynocephalus guttatus (GMELIN, 1789) in der Wolga-Region, Rußland]. – Herpetozoa, Wien, 18 (3/4): 141-146.

Kurzfassung:
Nach Feldbeobachtungen in den Jahren 1996 bis 2003 bevorzugen die Populationen der Gefleckten Krötenkopf-Agame Phrynocephalus guttatus (GMELIN, 1789) aus der Wolga-Region als Lebensraum verfestigte und mäßig verfestigte Sande mit spärlicher Busch- und Grasvegetation. Untersuchungen von russischem und ukrainischem Museumsmaterial ergaben, dass die Populationen auf den Don-Sanden im Volgograd-Gebiet zu Ph. g. guttatus zu stellen sind. Sie zeigen jedoch aufgrund ihrer Randlage hinsichtlich des Gesamtverbreitungsgebietes der Art gewisse Eigentümlichkeiten in bezug auf die Ausprägung einiger meristischer Merkmale.

WITTIG, W. (2002): Haltung und Nachzucht von Phrynocephalus guttatus. –elaphe N.F., Rheinbach, 10 (3): 29-31. (02.872 / 01.931)

Als ich im Oktober 1984 vier im Westen Kasachstans gefangene Krötenkopf-Agamen der Art Phrynocephalus guttatus (Gefleckter Krötenkopf) erhielt, sah ich mich zunächst vor erhebliche Probleme gestellt, denn diese Echsen galten als heikel und schwierig in der Haltung. Aber obwohl die Tiere einen längeren Transport mit Zwischenaufenthalten hinter sich hatten, lebten sie sich schnell im Terrarium ein und erwiesen sich als haltbare und dankbare Pfleglinge, die durch ihre Munterkeit erfreuten und auch bald zur Fortpflanzung schritten. Über die Haltungsbedingungen und die Nachzuchterfolge will ich im Folgenden berichten.

ZAVYALOV, E.V. & V.G.TABACHISHIN (2000): Spotted toad agame (Sauria, Agamidae, Phrynocephalus guttatus) distribution and its taxonomical status in the Lower Volga area. – Curr. Stud. Herp., 1: 40–47. [In Russisch.]

Relying on the analysis of the materials collected at the Zoological museum of the Saratov State University and the data came from literature the Phrynocephalus guttatus population in the Lower Volga area is spread either over hard and loose sands or sands’ modifications northward, up to Peskovatka village in the Gorodischensky region of Volgograd province. The investigation shows that the local populations in the sands along the Don in Volgograd province fall into the nominative subspecies (Ph. g. guttatus Gmel., 1789).

ZHAN, A. & J. FU (2009): Microsatellite DNA markers for three toad-headed lizard species (Phrynocephalus vlangalii, P. przewalskii and P. guttatus). – Molec. Ecol. Res., 9 (2): 535-538.

To assess the impact of natural landscapes on the population structure of lizards, 10 polymorphic microsatellite DNA markers were developed for the Qinghai toad-headed lizard, Phrynocephalus vlangalii. The number of alleles at these informative loci ranged from four to 28. The novel markers and those previously developed for Phrynocephalus przewalskii were cross-tested among three toad-headed lizard species P. vlangalii, P. przewalskii and P. guttatus. A high cross-utility rate of more than 58% was observed among these three species. These markers are expected to be useful tools for taxonomic considerations as well as population genetic analysis and future conservation management.


Phrynocephalus guttatus guttatus (GMELIN, 1789)

BADMAEVA, V.I. & N.N. SCERBAK (1983): Phrynocephalus guttatus kalmykus ssp. n. (Sauria, Agamidae) from Kalmykia. – Vestnik zoologii, Kiev, 17(6): 34-37. (in russisch)

MISHUSTIN, S.S. & G.V. POLYNOVA (2022): Variability of body weight (Eremias arguta deserti Gmelin, 1789) and (Phrynocephalus guttatus guttatus Gmelin, 1789) in the southeastern part of the Lower Volga region. - Journal of Ecology and Life, 30 (2): 189-200.

One of the aspects of studies of micropopulations of (Eremias arguta deserti Gmelin, 1789) and (Phrynocephalus guttatus guttatus Gmelin, 1789) in the conditions of the southeastern part of the Lower Volga region was the observation of inter-seasonal fluctuations in body weight of individuals. Males of Eremias arguta deserti are consistently heavier in the spring than females. In the autumn periods, the results turned out to be contradictory and do not make it possible to conclude which individuals of which sex have the greatest mass. Statistical processing of materials using the Kruskal — Walli’s criterion (H) did not reveal significant differences in Eremias arguta deserti when comparing the mass of all females and males both for all periods and separately in spring and autumn periods. Phrynocephalus g. guttatus also showed no statistical significance when comparing all males and females for the entire period. However, the results obtained when comparing between females and males during the 2018 season, as well as when comparing females of different years, were statistically significant. In addition, the body weight of fingerlings of both groups, between the autumn seasons of 2017 and 2018, revealed statistical significance. Individuals of both Phrynocephalus g. guttatus and Eremias arguta deserti steadily gain body weight, despite the slowdown or complete stop of the growth of individuals in the former and autotomy in the latter. Regeneration of the tail or its absence obviously affects the growth of the trunk of Eremias arguta deserti.

POLYNOVA, G.V. (2014): The dynamics of the Phrynocephalus guttatus guttatus Gmel. population number in Astrakhan semi-deserts. - News of the Samara Scientific Center of the Russian Academy of Sciences, 61 No. 5 (1) 443-444.

The article is devoted to the long-term investigation in the Phrynocephalus guttatus guttatus Gmel. population number. The author considers that the significant fluctuations in the number of this species are related to the short-living cycle of the population: almost complete replacement most likely occurs in two years.

POLYNOVA, G.V. & A.V. BAZHINOVA (2012): Assessment of the Phrynocephalus guttatus guttatus population number with regard to daily activity of animals. – Zool. Zh., 91 (11): 1411-1414.

The detailed investigation of the activity of a Phrynocephalus guttatus guttatus population in Astrakhan semi_ deserts showed that almost all resident lizards had some rest periods (1–4 days), when they do not emerge on the surface. The account of the lizards’ activity allows proposing some correcting coefficient. This coefficient should be used to obtain more precise data on the population number.

POLYNOVA, G.V., BAZHINOVA, A.V. & E.V. MUSHROOM (2012): Materials on the spatial structure of population Phrynocephalus guttatus guttatus in semi-deserts of the Astrakhan region. - Proceedings of the international conference “Terrestrial vertebrates of arid ecosystems”, dedicated to the memory of N.A. Zarudny, Tashkent-Uzbekistan, October 24-27, 2012. (in Russisch)

POLYNOVA, G.V., BAZHINOVA, A.V., OBUKHOVSKAYA, A.A. & A.V. KRINICHENKOVA (2014): Some characteristics of the activity and thermobiology of Phrynocephalus guttatus guttatus in the semi-deserts of the Astrakhan region. Peoples' Friendship University of Russia, Moscow: 94-97.In the first decade of may in sandy semi-deserts of the Astrakhan region Phrynocephalus guttatus guttatus (Gmel., 1789) has two periods of daily activity, morning and evening, temporal boundaries of which are associated with the temperature of habitats. Temperature features of the microrelief of the territory determine centers of activity of animals.

POLYNOVA, G.V., BAZHINOVA, A.V. & O.E. POLYNOVA (2014): The dynamics of the demographic structure of Phrynocephalus guttatus guttatus population in Astrakhan semi-deserts. – RUDN Journal of Ecology and Life Safety, 2014 (4): 11-24.

The spring settlement of Phrynocephalus guttatus guttatus Gmel. reliably splits into several age groups: 1—2 group of young, 3—4 groups of females and 2– 4 group of males. Senior group of young and two younger groups of mature individuals are aged for about a year and most likely belong to different clutches of eggs of the previous season. The overall ratio sex groups is close to unity, but that in the group of individuals of the first clutch of the year, as a rule, is dominated by females, and from the second — by males. Two older groups of mature animals have age not less than 2 years. The largest and oldest age group includes mostly females. Almost complete replacement of the population in this species most likely occurs in two years.

POLYNOVA, G.V. & S.S. MISHUSTIN (2017): Seasonal changes in the Phrynocephalus guttatus guttatus Gmel. population age and sex structure. - RUDN Journal of Ecology and Life Safety, 25 (3), 431—441. (In Russian).

The autumn studies of the Phrynocephalus guttatus guttatus Gmel. population in the Astrakhan semideserts (August 2011 and August — September 2016) revealed the following features of its age and sex structure. The autumn settlement of the species reliably splits into several age groups: 1—2 group of young, 2—4 groups of females and 2—5 group of males. In the fall season the settlement is dominated by males, especially great was the numerical superiority of the males in the fall season of 2016.The lack of immature individuals, the significant prevalence of older age groups and the predominance of males among adult animals in the autumn 2016 create the picture of the population depression. The unsustainable type of population dynamics, typical for short-living species, may be one of the characteristics that stimulates the processes of population depression. Almost complete replacement of the population is likely to occur both in two or 3—4 years.

POLYNOVA, G.V. MISHUSTIN, S.S. & O.E. POLYNOVA (2022): Spatial structure of the Spotted Toad Agama (Phrynocephalus guttatus guttatus, Agamidae) population in the conditions of overgrowth of semi-deserts in the Astrakhan Region. - Principy Èkologii 2022 (1): 92-110.

The decrease in the number of many reptile species as a result of overgrowth of sandy deserts and semi-deserts is a widespread phenomenon in modern conditions. To understand the prospects of this process, it is necessary to study the mechanisms of population adaptation to changes in the biotope. Such a mobile and effective mechanism is, first of all, a change in the spatial structure of the population, the study of which does not cease to be relevant. The purpose of this study was to clarify the changes occurring in the spatial structure of the spotted toad agama, Phrynocephalus guttatus guttatus (Gmelin, 1789), and their relationship with the developing process of overgrowth of sandy semi-deserts in the Astrakhan region. The data were collected during the 2010–2014 and 2017–2019 field seasons. The place of research was a section of sandy semi-desert near the village of Dosang (46°54'08.7264"N, 47° 54'52.5312"E). In the work, standard methods were used: marking, sex and age determination, mapping of meetings and movements, identification of sedentary and migrating individuals. Investigation of overgrowth processes was based on the description of geobotanical sites. Statistical processing was carried out based on the Mann-Whitney U-test. Software consisted of a set of standard programs, Adobe illustrator and Yandex Map Constructor. The analysis of long-term studies of the spatial structure of the grouping allowed us to build a model of adaptation of the population to the reduction of characteristic habitats. At the initial stages of overgrowth of semi-fixed sands, there was a general increase in the mobility of animals, expressed in an increase in the number of individuals migrating in search of a suitable biotope. The reduction of the biotope caused the re-consolidation of the sedentary population of the grouping. This process occurred during the third and fourth seasons of research. Over-compaction was followed by a decrease in the rate of reproduction, manifested in a reduction in the number of immature individuals. As a result, the combination of factors led to a further decrease in the number of individuals (the fifth year of observations). The ongoing process of overgrowth and the reduction in the number for the tenth year of research resulted in the disintegration of the grouping into parts and a decrease in the population as a whole. The general decrease in the population was confirmed by observations in the adjacent territory and the almost complete disappearance of migrants in the grouping.

RIABOVA, E.D., VETO, D.V., OKSHTEIN, I.L., TISCHENKO, M.A. & A.Y. FRAERMAN (2021): The social contacts of Phrynocephalus guttatus guttatus (Gmelin, 1789) and Phrynocephalus mystaceus mystaceus (Pallas, 1776). - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p. 229-231. (in Russisch)


Phrynocephalus guttatus alpherakii BEDRIAGA,1906


Phrynocephalus guttatus melanurus EICHWALD,1831

DUNYAEV, E.A. & D.V. SEMENOV (1988): The lizard species Phrynocephalus melanurus. - Priroda (Moscow), 1988 (4): 54-55. (In Russian).

SHUANG, L., LIU, L.J. & S. SONG (2014): The complete mitochondrial genome of Grumgzimailo’s toad-headed agama, Phrynocephalus grumgrizimailoi (Reptilia, Squamata, Agamidae). – Mitochondrial DNA, 2014. DOI: 10.3109/19401736.2014.958678

Phrynocephalus helioscopus PALLAS, 1773

Sonnengucker / Sunwatcher Toadhead Agama

ANANJEVA, N.B. (1981): Phrynocephalus heliocopus (Pallas 1771) – Sonnengucker. – In: Böhme, W. (Hrsg.): Handbuch der Reptilien und Amphibien Europas. Vol. 1. Echsen (Sauria) I. Akademische Verlagsgesellschaft, Wiesbaden. 191-202.

ANDERSON, S.C. (1999): Phrynocephalus helioscopus (Pallas, 1771). - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 87-88.

BARAN, I., KASPAREK, M. & M. ÖZ (1989): On the distribution of four species of agama (Agamidae) in Turkey. – Zoology in the Middle East, Heidelberg, 3: 37-46.
Kurzfassung:
Die Verbreitung des Harduns, Agama stellio, der Kaukasischen Agame, A. caucasia, der Ruinenagame, A. ruderata, und des Sonnenguckers, Phrynocephalus helioscopus, in der Türkei wird durch Punktkarten dargestellt. A. stellio und A. caucasia schließen sich gegenseitig horizontal und vertikal aus. Das Areal von A. stellio wird durch die März-Isotherme von 8°C und die Juli- und August-Isothermen von 24°C definiert. A. ruderata kommt zwar in den großen, ursprünglichen Steppengebieten Zentral- und Südost-Anatoliens vor, fehlt aber in den Steppen Ost-Anatoliens.

BEREZOVIKOV, N.N. (2007): New findings of Alsophylax pipiens and Phrynocephalus helioscopus in the south-western foothills of Southern Altay. – Selevinia, 2006: 213. (In Russisch).

BONDARENKO, B.M. & D.G. ZAMOLODCHIKOV (1991): Efficiency of different population estimation methods as applied to a model population of Phrynocephalus helioscopus. – Vestn. Zool., 1991 (5): 78-81. (In Russian)

GRAVENHORST, J.L.C. (1833): Über Phrynosoma orbicularis, Trapelus hispidus, Phrynocephalus helioscopus, Corythophanes cristatus und Chamaeleopsis hernandesii. - Acta Acad. Caes. Leop. Carol. Nat. Cur. (?) 16 (2): 909-958.

KRYMOV, N.G. (2017): On possible estivation of Phrynocephalus helioscopus (PALLAS, 1771) and Eremias arguta (PALLAS, 1773) in the Altai region. - Curr. Stud. Herp., 17 (1-2): 66-70.

Summer counts of Phrynocephalus helioscopus (Pallas, 1771) and Eremias arguta (Pallas, 1773) were conducted in the Altai Region during April – September, 2016. A sharp decline in the activity of lizards in June and July was noted, due to the high temperatures and dry weather. As a result of excavation of some burrows, lizards in a non-active state were found. The possibility of summer hibernation (estivation) of the species is considered.

LIANG, T. & L. SHI (2017): Sexual dimorphism and morphological variation of three populations of Phrynocephalus helioscopus: Test of Bergmann’s Rule, Allen’s Rule and Rensch’s Rule. – Sichuan Journal of Zoology, 36 (3): 249-257. (in Chinesisch)

MELNIKOV, D., ANANJEVA, N.B., RAJABIZADEH, M. & K. MILTO (2009): On Systematics and phylogeography of sun wtcher toad agamas Phrynocephalus helioscopus (PALLAS, 1771). - Abstracts of presentations hold on DeAGAMIS the 1st International Symposium on Agamid Lizards. Bonner Zoologische Beiträge, Bonn, 56 (4): 303.

MEYER, F. & O. ZINKE (1992): Zur Ökologie von Phrynocephalus helioscopus (Pallas, 1771) in der Dschungarischen Gobi (Reptilia; Agamidae). – Bonn. zool. Beitr. 43 (1): 131-144.

PALLAS, P.S. (1771): Description of Phrynocephalus helioscopus – In: Reise durch verschiedene Provinzen des Rußischen Reichs, erster Theil. - St. Petersburg.

PETERS, G., SEMENOV, D.V. & L.J. BORKIN (1990): On the Sunwatcher, Phrynocephalus helioscopus (Pallas, 1771) in Mongolia. – Trudy Zool. Inst. Acad. Nauk SSSR, 207: 224-229.

SCHLEICH, H.-H. (1976): Über Phrynocephalus helioscopus aus Persien (Reptilia, Sauria, Agamidae). – Salamandra, Frankfurt/Main, 12 (4): 189-193. (00.052)
Zusammenfassung:
Phrynocephalus helioscopus ist in seinem Biotop nicht gleichmäßig verteilt. Gruppen von etwa 12-20 Individuen bewohnen Areale von etwa 50 m². Die Hauptnahrung freiolebender Tiere besteht aus Ameisen (bis 1,5 cm Länge). AM Schädelskelett fallen die geflügelten Präfrontalia auf. Weder drohendes noch aggressives Verhalten war zu bemerken. Beim Besprühen stoillten die Tiere ihren Durst. Bei weiteren Haltungsversuchen wäre Besprühen mit destilliertem beziehungsweise kalkarmem Wasser anzuraten.

SCHWENK, K. & H.W. GREENE (1987): Water Collection and Drinking in Phrynocephalus helioscopus: A Possible Condensation Mechanism. - Journal of Herpetology, 21 (2): 134-139.

The drinking behavior of nine captive Phrynocephalus helioscopus was observed. When wetted, animals assumed a stereotyped posture in which the head was depressed to within several mm of the substrate, the limbs splayed, and the hindquarters and tail elevated. Drinking involved repeated, slight tongue protrusion, but without lapping from the substrate. This behavior was elicited in no other context. Simple experiments showed that the interscalar channels transport water across the skin to the mouth by capillary action. This mechanism is compared with that reported for another agamid, Moloch horridus. We hypothesize that the drinking posture 1) facilitates drinking rain water and/or 2) facilitates drinking water condensed on the skin and moved to the mouth by capillary action. We conclude that derived features of water collection and transport shared by Phrynocephalus and Moloch are convergent.

SHENBROT, G.I. & G.S. ANTONOVA (1981): Comparative ecology of Phrynocephalus helioscopus and P. reticulatus in the conditions of their common habitat, south of Bukhara region (Uzbek SSR). - In: Darevsky, I.S., Ananjeva, N.B., Barkagan, Z.S., Borkin, L.Y., Sokolova, T.M. & N.N. Szczerbak (eds.): The problems of herpetology: abstracts. Nauka, Leningrad: 156-157. (in Russian).

SOKOLOV, V.E., DAREVSKY, I.S., KOTOVA, E.L. & O.F. CHERNOVA (1997): Specialized skin organs of Phrynocephalus helioscopus (Reptilia, Squamata, Agamidae). – Zool. Zhurn., 76 (4): 466-472. (In Russian)

SOLOVYEVA, E.N. (2010): Molecular differentiation and distribution within the species complex of Phrynocephalus helioscopus (Reptilia: Agamidae). - Abstracts of the Second International Symposium on Agamid Lizards «DeAgamis2». - Current Studies in Herpetology, 10 (3/4): 154-155.

SOLOVYEVA, E.N., POYARKLOV, N.A., DUNAEV, E.A., DUYSEBAYEVA, T.N. & A.A. BANNIKOVA (2011): Molecular Differentiation and Taxonomy of the Sunwatcher Toad-[1]Headed Agama Species Complex Phrynocephalus Superspecies helioscopus (Pallas 1771) (Reptilia: Agamidae). - Russian Journal of Genetics, 47(7): 842–856.

Lizards of the sunwatcher toad[1]headed agama species complex Phrynocephalus superspecies helioscopus, mostly distributed in Central Asia and Middle East, were examined using analysis of variation at the mitochondrial cytochrome oxidase c subunit I gene fragment and fingerprint analysis of nuclear DNA (inter[1]SINE PCR technique). A total of 86 individual tissue samples from 53 populations, to the full extent representing different parts of the species complex range, were subjected to molecular genetic examination, and surprisingly deep differentiation was revealed. The populations analyzed split into 12 isolated phylo[1]groups, many of which were characterized by a narrow range and genetic isolation. Monophyly of sunwatcher (Ph. helioscopus) and Persian (Ph. persicus) toad[1]headed agamas was confirmed. However, both of these spe[1]cies probably represent the species complexes. Zoogeography of Central Asiais discussed.

SOLOVYEVA, E.N., DUNAYEV, E.A. & N.A. POYARKOV (2012): Interspecific taxonomy of sunwatcher toadhead agama species complex (Phrynocephalus helioscopus, Squamata). – Zool. Zhurn., 91 (11): 1377-1396. (in Russisch)

The taxonomic revision of Ph. helioscopus as a species with 7 valid subspecies (Ph. h. helioscopus (Pallas 1771), Ph. h. varius Eichwald 1831, Ph. h. cameranoi Bedriaga 1907, Ph. h. saidalievi Sattorov 1981, Ph. h. sergeevi ssp. n. (Uzbekistan: northeastern Bukhara and Navoi regions), Ph. meridionalis ssp. n. (Uzbekistan: southern Bukhara and Surkhadarya region; Turkmenistan: Kugitang foothills in Lebap veloyat; probably, northern Afghanistan), Ph. h. turcomanus ssp. n. (southeastern Turkmenistan, northeastern Iran; probably, western Afghanistan) is given. The new taxa are described and diagnostic keys for all subspecies are given. An insignificant divergence of morphometric parameters and overlap of standart folidosis parameters are shown. The coloration of the ventral side of the tail and some other new folidosis traits allowed dividing all subspecies in phyletic groups. These groups were supported by analysis of COI of mtDNA and fingerprint analysis of nuDNA. Toad-headed agamas from Karatau region still have unclear taxonomic position. The level of taxonomic differentiation of different forms is also discussed.


Phrynocephalus helioscopus helioscopus PALLAS, 1773

RUSTAMOV, A.K. & S. SHAMMAKOV (1967): Ecology of Phrynocephalus helioscopus helioscopus Pallas in Turkmenia. – Zool. Zh., 46: 741-748. (in Russisch)


Phrynocephalus helioscopus cameranoi BEDRIAGA, 1907


Phrynocephalus helioscopus meridionalis DUNAYEV, SOLOVYEVA & POYARKOV, 2012


Phrynocephalus helioscopus saidalievi SATTOROV, 1981

MANILO, V.V., GOLUBEV, M.L. & T. SATTAROV (1991): The karyotype of Phrynocephalus helioscopus saidalievi (Sauria, Agamidae) of the Ferghana Valley. (In Russisch) – Vestn. Zool., 1991 (2): 79-81. (in Russisch)


SATTOROV, T. (1981): Phrynocephalus helioscopus saidalievi ssp. n. (Sauria, Reptilia) - a new subspecies of Ph. helioscopus from the Ferghana Valley. - Vestnik Zoologii 1981 (1): 73-75. (in Russisch).


Phrynocephalus helioscopus sergeevi DUNAYEV, SOLOVYEVA & POYARKOV, 2012


Phrynocephalus helioscopus turcomanus DUNAYEV, SOLOVYEVA & POYARKOV, 2012


Phrynocephalus helioscopus varius EICHWALD, 1831 

EICHWALD, E. (1831): Description of Phrynocephalus helioscopus varius; Ph. reticulatus in “Zoologia specialis, quam expositis animalibus tum vivis, tum fossilibus potissimuni rossiae in universum, et poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi”. Zawadski, Vilnae.

Lichtenstein´s Toadhead Agama

ANDERSON, S.C. (1999): Phrynocephalus interscapularis  Lichtenstein, 1856. - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 88-89.

DUJSEBAYEVA, T.N. (1994): The topography and numerical distribution of skin sense organs in the skin of Phrynocephalus interscapularis (Lacertilia: Agamidae). – Sedlevinia, 2 (4): 10-14. (in Russisch)

ULMASOV, K., ZATSEPINA, O., MOLODTSOV, V. & M. EVGEN’EV (1999): Natural body temperature and kinetics of heat-shock protein synthesis in the toad-headed agamid lizard Phrynocephalus interscapularis. – Amphibia-Reptilia, Leiden, 20 (1): 1-9.


Phrynocephalus interscapularis interscapularis LICHTENSTEIN, 1856

Phrynocephalus luteoguttatus BOULENGER, 1887

Yellow-speckled Toad-headed Agama

ANDERSON, S.C. (1999): Phrynocephalus luteoguttatus Boulenger, 1887. - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 89-90.

Phrynocephalus maculatus ANDERSON, 1872

Gefleckte Krötenkopfagame / Blacktail Toadhead Agama

ABU BAKER, M., ŠIROKÝ, P., AMR, Z. & D. MODRÝ (2005): Discovery of a population of Phrynocephalus maculatus ANDERSON, 1872 in the Hashemite Kingdom of Jordan. [Erstnachweis von Phrynocephalus maculatus ANDERSON, 1872 für Jordanien]. – Herpetozoa, Wien, 18 (3/4): 107-113.

Kurzfassung:
Die vorliegende Arbeit beinhaltet den Erstnachweis von Phrynocephalus maculatus ANDERSON, 1872 in Jordanien. Diagnostisch relevante morphologische Daten werden angegeben und der Lebensraum wird beschrieben. Taxonomie und Variabilität innerhalb P. maculatus werden diskutiert und mit dem Untersuchungsmaterial verglichen. Trotz einiger geringfügiger Unterschiede zur Originalbeschreibung werden die aufgesammelten Exemplare der Unterart P. m. longicaudatus HAAS, 1957 zugeordnet.

AL-BARAZENGY, A.N. (2015): First observations on Phrynocephalus maculatus longicaudatus Haas, 1957 (Squamata: Sauria: Agamidae) in Iraq. - Bull. Iraq nat. Hist. Mus, 13 (3): 1-7.

The present paper confirmed the presence of Phrynocephalus maculatus longicaudatus Haas, 1957 in Iraq and recorded the first observations of this taxon in Al-Muthanna province southwestern of Iraq. The existence of the species is yet uncertain in Iraq. The habitat and morphological characteristics of this species were reviewed.

AL-SIRHAN, A.-R. & G. BROWN (2010): The status of the two Toad-headed Agamas, Phrynocephalus arabicus (Anderson, 1894), and P. maculatus (Anderson, 1872), in Kuwait. – Zoology in the Middle East, 51.

ANDERSON, J. (1872): Description of Phrynocephalus maculatus and P. forsythii. - In: On some Persian, Himalayan and other Reptiles. - Proceedings of the Zoological Society of London 1872 : 371-404.

ANDERSON, S.C. (1999): Phrynocephalus maculatus Anderson, 1872. - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 90-92.

BOGDANOV, O.P., ATAEV, C. & S. SHAMMAKOV (1974): The finding of Phrynocephalus maculatus in the USSR. – Zoologicheskij Zh., 53 (2): 304-305. (in Russisch)

MALEKMOHAMMADI-KALAHROUDI, M. & V. HOJATI (2014): The Female Reproductive Cycle of the Spotted Toad-headed Agama, Phrynocephalus maculatus (Sauria: Agamidae) in Iran. - Iranian Journal of Animal Biosystematics (IJAB) 10(2): 185-194.

The Spotted Toad-headed Agama, Phrynocephalus maculatus, is a member of the Agamidae family distributed in the central and south-eastern deserts of Iran. Iranian specimens are rare in collections. In this research, the female reproductive cycle of this species was studied from April 5 to August 5, 2013. Totally, 15 adult females were collected by hand at midday from southern parts of Damghan County, located in Semnan Province of Iran. Ovaries were removed and processed for histological and morphometric studies. The oogenic cycle begins from early April, mating occurs at the beginning of May, with oviposition occurring from late May to mid July. Females lay 2-3 eggs per clutch with the possibility of producing a secondary clutch later in the season. Maximum reproductive activity occurs in May and early June and reduces from early July and ends in August. There was no significant difference between the right and left side of the reproductive system. Hence, oogenesis occurs from April through July, P. maculatus follows an associated reproductive cycle typical for temperate species.

RAHMANI, S. & V. HOJATI (2015): The Male Reproductive Cycle of the Spotted Toad-Headed Agama,Phrynocephalus maculatus, in Iran. – Int. J. Zool. 2015: 1-5. http://dx.doi.org/10.1155/2015/521406

Thespotted toad-headed agama, Phrynocephalus maculatus, distributed in the central and southeastern arid regions of Iran, belongs to the Agamidae family. In this research, the male reproductive cycle of this species was studied from 5 April to 5 August, 2013. Totally, 40 adult males were collected at midday from southern deserts of Damghan County, located in Semnan Province of Iran. Testes were removed and processed for morphometric and histological studies. The spermatogenic cycle begins from early April, mating occurs in mid-May, and it ends in August. Maximum reproductive activity occurs in early June and reduces from early July and ends in August. The numbers of seminal vesicles were 33–127 and their diameter varied between 69.0 and 258.3 microns. The diameter of tunica albuginea varied between 3.0 and 8.1 microns. The diameter of germinal layer varied between 10.0 and 110.0 microns.There were significant differences in macroscopic and microscopic testicular characters betweenmonths. Also, there were no significant differences in the testicular and hemipenal characters between the left side and the right side of body. Since spermatogenesis occurs from April through August, P. maculatus follows an associated reproductive cycle typical for temperate species.

ROSS, W. (1989): Notes on ecology and behaviour with special reference to tail signalling in Phrynocephalus maculatus (Reptilia: Agamidae). – Fauna of Saudi Arabia, Jeddah, 10: 417-422.

RUSTAMOV, A.K. & S. SHAMMAKOV (1977): Ecology of Phrynocephalus maculatus. – Zoologicheskij Zh., 56 (9): 1351-1356. (In Russisch)

SCHOLZ, S., SIEGENTHALER, F. & T.M. WILMS (2013): A new locality record of Phrynocephalus maculatus ANDERSON, 1872, from Jordan. – Herpetozoa, Wien, 25 (3/4): 174-179.


Phrynocephalus maculatus maculatus ANDERSON, 1872

HOJATI, V., MALEKMOHAMMADI, M. & S. RAHMANI (2014): A preliminary study on the biology of the Black-tailed Toad Agama, Phrynocephalus maculatus maculatus in Iran. - CIBTech J. Zool., 3 (3): 60-67.

The understudied Black-tailed toad agama, Phrynocephalus maculatus maculatus, (Anderson, 1872) belongs to the Agamidae family. Iranian specimens are rare in collections and are distributed in the central and south-eastern deserts of Iran. In this research, biological studies including food habits, morphology, behaviors and habitats of these species were performed from April to September, 2013. A total of 30 adult specimens including 15 adult males and 15 adult females were collected by hand at midday from southern parts of Damghan County, located in Semnan Province of Iran. Results show that the animal is active from early April to September, and that it hibernates from October to March. They are sit-and-wait predators. They are insectivores and their main food items belong to seven insect families including: Formicidae, Tenebrionidae, Acrididae, Noctuidae, Termitidae, Muscidae and Ixodidae. No plant consumption was observed in this species. This agama inhabits the desert, especially harder sandy surfaces. Ph. maculatus maculatus is a fairly understudied subspecies in the Middle East region and this study presents some useful information regarding this poorly known animal.


Phrynocephalus maculatus longicaudatus HAAS, 1957

HAAS, G. (1957): Some amphibians and reptiles from Arabia. - Proc. Cal. Acad. Sci. 29 (3): 47-86.

Phrynocephalus mystaceus (PALLAS, 1776)

Bärtiger Krötenkopf / Secret Toadhead Agama

AKHMEDENOV, K.M. & A.G. BAKIEV (2023): Finds of the eared roundhead Phrynocephalus mystaceus (Pallas, 1776) in western Kazakhstan regions of the Republic of Kazakhstan. – In: Davygora, A.V. (Ed.): Terrestrial vertebrates of arid and subarid ecosystems of the Aral-Caspian region. Materials of the III International Conference, dedicated to the memory of the outstanding ornithologist, naturalist and traveler Nikolai Alekseevich Zarudny, Orenburg, April 24–28, 2023: 10-14.

ANANJEVA, N.B. (1981): Phrynocephalus mystaceus (Pallas 1776) – Bärtiger Krötenkopf. – In: Böhme, W. (Hrsg.): Handbuch der Reptilien und Amphibien Europas. Vol. 1. Echsen (Sauria) I. Akademische Verlagsgesellschaft, Wiesbaden. 203-216.

ANANJEVA, N.B. (1986): On the validity of Megalochilus mystaceus (Pallas 1776). – Proceedings of the Zoological Institute, 157, USSR Academy of Science, Leningrad. 4-13. (in russisch)

ANDERSON, S.C. (1999): Phrynocephalus mystaceus (Pallas, 1776) - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 92-93.

BADMAEVA, V.I., LEBEDENKO, N.A. & N.A. SAVINA (1981): Diel activity pattern of Phrynocephalus mystaceus in Kalmykia]. - In: Darevsky, I.S., Ananjeva, N.B., Barkagan, Z.S., Borkin, L.Y., Sokolova, T.M. & N.N. Szczerbak (eds.): The problems of herpetology: abstracts. Nauka, Leningrad: 11-12. (in Russian).

BELYALOV, O.V. (2000): About attack of toad-headed agama (Phrynocephalus mystaceus Pall.) on fledglings of desert warbler (Sylvia nana Hemprich et Ehrenberg). – Selevinia, 2000 (1-4): 220. (In Russian).

CHERLIN, V., OKSHTEIN, I., ALIEVA, S. & A. MAGOMEDOVA (2022): Estimation of the number of the toad-headed agama (Phrynocephalus mystaceus) and the stepperunner (Eremias arguta) on the Sarykum dunes and their surroundings (Dagestan Republic, Russian Federation). [in Russ. with Engl. summ.]. - Principy èkologii. 11 (4): 3-21.

Variants of estimating the number of different lizard species in natural populations are proposed. In one sense or another, we are talking about an accounting site of a known area. But depending on the biological features of different lizard species, the accounting indicators may be different: either it is a fixation of the total number of lizards living on the accounting site (which is possible for the toad-headed agama Phrynocephalus mystaceus), or it is a fixation of the number of lizards encounters on accounting routes of a certain length and width, i.e. a certain area (which is possible for the steppe runner Eremias arguta). Correction coefficients should be introduced into the accounting results, which should take into account that: 1) the distribution of lizards across the territory is uneven due to microbiotopic differences (K1), 2) not all lizards that live in the this territory may be active on the surface every day (K2), 3) when using different registration methods, primary accounting data allow registering a different proportions of lizards from their total number inhabiting this site (K3). After such an adjustment of the accounting results, it is possible to calculate the density of lizard settlements and the absolute number of lizards for any separate sections of entire territories. As a result of our work, we have so far determined the composition of these coefficients. Subsequent studies should lead to the development of standardized methods for determining these coefficients. According to our calculations, the density of the population of toad-headed agamas in the area of their most compact habitat on the large Sarykum dune can be up to 125 individuals/ha, and their total number can be estimated at about 7000-7500 individuals. For steppe runners, their population density in places of compact habitat in the vicinity of the small Sarykum dune can be 18.0-23.4 ind./ha, and their total number in the area of about 9 ha, where we carried out our research work, could be 180-220 individuals. By standardizing the methods of evaluating different species of lizards, it is possible to organize correct long-term monitoring of the state of their populations.

DATHE, F. (1986): Phrynocephalus mystaceus (Pallas, 1776) Bärtiger Krötenkopf (Familie: Agamidae, Agamen). – Aquarien Terrarien, Leipzig, 33 (8): 288.

DIECKMANN, M. (2011): Bemerkungen zur Haltung des Bärtigen Krötenkopf Phrynocephalus mystaceus (PALLAS, 1776). – Iguana, 24 (1): 21-26.

DUJSEBAYEVA, T.N. & D.V. MALAKHOV (2006): Postembryonic growth of the steppe agama, Trapelus sanguinolentus (Pall., 1814) and the toad-headed agama, Phrynocephalus mystaceus (Pall., 1776) in terrarium. – Selevinia, 2005: 143-146. (In Russisch).

GOLUBEV, M.L. & T.S. SATTOROV (1992): On intraspecific structure and intraspecific relations of the ear-folded toad agama Phrynocephalus mystaceus (Reptilia, Agamidae). – Vestnik Zoologii, 1992 (3): 26-32. (in Russisch)

KHONYAKINA, Z.P. (1962): Population dynamics of Phrynocephalus mystaceus Pall. in Kumtorkali region of Daghestan ASSR. – Zool. Zh., 41: 778-780.

KRASSOWSKY, D.B. (1932): Beitrag zur Systematik von Phrynocephalus mystaceus (Pall.). - Zoologischer Anzeiger, Leipzig 97(7/8): 225-228.

MERTENS, R. (1952): Der „Bärtige Krötenkopf“ und seine Warnstellung (Phrynocephalus mystaceus). – Natur und Volk, 82: 15-19.

MOLAVI, F., MOHAMMADIAN-KALAT, T., KIABI, B.H., KAMI, H.G. & Y. MORTEZA (2014): New distribution record of Phrynocephalus mystaceus Pallas 1776 (Sauria: Agamidae) from Iran. - Check List, 10 (2): 450-452.

The Toad-headed agama, Phrynocephalus mystaceus Pallas 1776, was described in 1999 from eastern Khorasan by Anderson. Seven specimen of The Toad-headed agama were collected in Khar Turan National Park during fieldwork from June 2008 to June 2009. The new locality of the species is situated about 900 km west of the type locality. This record indicates a wider distribution of Phrynocephalus mystaceus on the Iranian plateau than previously thought. Information on morphological characters and habitat is presented.

MÜLLER, H.D. (2002): Der Bärtige Krötenkopf Phrynocephalus mystaceus (PALLAS, 1776) im Terrarium. – Draco, Münster, 3 (2): 50-57. (02.931)

NEYMARK, L.A. (2021): On the thermal biology of psammophilous lizards of Kalmykia. - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p 195-197. (in Russisch)

PALLAS, P.S. (1776): Description of Phrynocephalus mystaceus – In: Reise durch verschiedene Provinzen des Russischen Reichs, Dritter Theil. St. Petersburg.

PANOV, E.N., TSELLARIUS, A.Y. & V.A. NEPOMNYASHCHIKH (2004): Motor coordinations in behavior of the toad-headed agama (Phrynocephalus mystaceus, Reptilia, Agamidae): Signal functions and endogenous rhythms. - Zoologicheskii Zhurnal 83(8): 971-982. (In Russisch).

PANOV, E.N., TSELLARIUS, A.Y. & V.A. NEPOMNYASHCHIKH (2004): Motor coordinations in behavior of the toad-headed agama (Phrynocephalus mystaceus, Reptilia, Agamidae): Signal functions and endogenous rhythms. – Entomological Review, 84 (2): 185-194.

Long-term observations and videotape recording of the behavior of territorial animals were used to study its temporal organization in male toad-headed agamas. The total recording time was 10.5 h, including nine continuous recordings of six mature males totaling more than 6 h. An analysis was made of the temporal pattern of complex tail movements, which proved to be a standard component of male behavior even outside of a social context. The basic element of this behavior is a set of stereotyped tail movements (a cycle) performed in a rhythmic sequence, designated a series. One cycle is approximately 7 s long, and the number of cycles per series varies in a broad range (1–20). The intensity of tail movements comprising the cycle gradually decreases within each series and in consecutive series; the series themselves gradually become shorter, whereas intervals between them increase. The cyclicity of series performance under stationary conditions indicates that this behavior is probably under constant endogenous control, which casts doubt on the concept that the stereotyped tail movements in male toad-headed agamas have primarily a communicative function.

POLYNOVA, G.V. & O.E. POLYNOVA (2022): Toad-Headed Agama (Phrynocephalus mystaceus Pallas): The State of the Population on the Sarykum Sand Massif (Dagestan, Russia). – Proc. Kazan Univ. Nat. Sci. Ser., 164 (2): 328-341.(In Russian).

This article provides a detailed description and analysis of two settlements of toad-headed agamas (Phrynocephalus mystaceus Pall.) on the Sarykum sand massif based on the data collected during the first ten days of May 2019 (105 individuals – 31 males, 49 females, and 25 immature individuals) and 2021 (115 individuals – 26 males, 22 females, and 67 immature individuals), respectively. The age and sex structure of both settlements was studied to assess the state of the species population. All individuals had their body length measured and were labelled, either permanently or temporarily. The statistical significance of the results obtained was determined using the nonparametric Mann-Whitney test. It was revealed that both settlements had the following sex and age groups: immature individuals and mature males and females. Mature individuals of both sexes were represented by two age stages: those aged two years, as well as three years and older. The core of the population consisted of young individuals and males and females aged two years. In the first settlement, mature individuals were more abundant, and females dominated among them. In the second settlement, immature individuals prevailed, and males dominated among mature individuals. The local density of the second settlement inaccessible to tourists was 2.6 times higher. The configurations of the sex and age pyramids showed that the first set-tlement affected by the recreational load was in the process of reduction, while the second settlement with no anthropogenic input turned out to be stable. It was concluded that the recreational load and habitat overgrowth speak for the need to monitor the toad-headed agama population of the Sarykum sand massif.

POLYNOVA, G.V. & O.E. POLYNOVA (2023): Foster’s or island rule in populations of Phrynocephalus mystaceus and Eremias velox (Reptilia, Lacertilia) on the sandy massif Sarykum. - Curr. Stud. Herpetol., 23 (3-4): 154-159.

The sandy massif Sarykum, whose age is about 100 thousand years, is an island habitat for psammophilic species of terrestrial vertebrates. The paper presents new morphometric data on the populations of two species of psammophilous lizards living in this area. These are the nominative subspecies of the Secret Toadheaded Agama (Phrynocephalus mystaceus mystaceus Pallas, 1776) and the Caucasian Central Asian Racerunner (Eremias velox caucasica Lantz, 1928). Body length of sexually mature males of the Secret Toadheaded Agama averages 76.5±3.7 mm (n = 30), and adult females – 68.9±4.2 mm (n = 29). Comparison of the obtained materials with similar parameters of the Kazakhstan population of the subspecies shows that mature individuals of the Sarykum population are significantly smaller: for males td = 1.33 ≥ tst with a confidence level α = 0.80, and for females td = 2.07 ≥ tst with α = 0.95. It is known from the literature that all the pre-Caucasian populations of this species are isolated. Perhaps the small size of mature individuals in them also serve as an example of the manifestation of Foster's rule. The data of the presented study indicate a similar feature of the Sarykum population of the Caucasian Central Asian Racerunner. The body length of mature males at Sarykum is 63.6±2.9 mm (n = 9), and that of females is 58.4±3.0 mm (n = 17). The calculation of the reliability of differences by the Student coefficient shows that the length of the trunk of males (td = 2) and females (td = 0.61) of the Sarykum population does not statistically differ from the averaged materials for the region. At the same time, mature individuals of the Sarykum population are significantly smaller than the nominative subspecies from Kazakhstan: for males td = 1.40 ≥ tst with a confidence probability α = 0.80, and for females td = 2.20 ≥ tst with α = 0.95. It is obvious that Foster's rule does not manifest itself in the subspecies population living on the Sarykum sandy massif, and the conspicuous small size of mature individuals is determined by comparison with the size of the nominative subspecies. An interesting fact is that immature foot-and-mouth of both species do not differ in size from individuals of the same age of other populations. Probably, at this stage of ontogenesis, the overall physiologically optimal size for the species is preserved.

RAIMBEKOVA, E.E. (2011): Diurnal activity of Phrynocephalus mystaceus and its relation to food supply. – Selevinia, 2010: 180-181. (In Russian).

SEMENOV, D.V. & G.I. SHENBROT (1990): Species of the genus Phrynocephalus of the USSR fauna. Description of a new subspecies with remarks on the taxonomic status of Phrynocephalus mystaceus (Reptilia, Agamidae). – Zool. Zh. (USSR), 69 (5): 76-81.

SHAMMAKOV, S. & G. MARKOV (1970): Occurrence of a nematode Abbreviata turcomanica on Phrynocephalus mystaceus in central Karakum. – Izv. Akad. Nauk turkmen. SSR (Biol.), 1970 (6): 76-78. (in Russisch)

SHAMMAKOV, S. & K. NIZAMUTDINOVA (1970): On ecology of Phrynocephalus mystaceus Pallas in Central Karkum. – Izv. Akad. Nauk turkmen. SSR (Biol.), 1970: 66-70. (in Russisch)

SHAW, G. & E. NODDER (1803): Lacerta aurita – The Lobe-Cheeked Lizard. -  The Naturalist’s Miscellany 15 (unnumbered pages).

SHIBANOV, N. (1941): On the age-variability and geographical forms of Phrynocephalus mystaceus (Pall.). – Arch. Mus. zool. Moscou, 6: 201-210. (in Russisch)

SOLOVYEVA, E.N., DUNAYEV, E.N., NAZAROV, R.A., RADJABIZADEH, M. & N.A. POYARKOV (2018): Molecular and morphological differentiation of Secret Toad-headed agama, Phrynocephalus mystaceus, with the description of a new subspecies from Iran (Reptilia, Agamidae). – ZooKeys, 748: 97-129.

The morphological and genetic variation of a wide-ranging Secret Toad-headed agama, Phrynocephalus mystaceus that inhabits sand deserts of south-eastern Europe, Middle East, Middle Asia, and western China is reviewed. Based on the morphological differences and high divergence in COI (mtDNA) gene sequences a new subspecies of Ph. mystaceus is described from Khorasan Razavi Province in Iran. Partial sequences of COI mtDNA gene of 31 specimens of Ph. mystaceus from 17 localities from all major parts of species range were analyzed. Genetic distances show a deep divergence between Ph. mystaceus khorasanus ssp. n. from Khorasan Razavi Province and all other populations of Ph. mystaceus. The new subspecies can be distinguished from other populations of Ph. mystaceus by a combination of several morphological features. Molecular and morphological analyses do not support the validity of other Ph. mystaceus subspecies described from Middle Asia and Caspian basin. Geographic variations in the Ph. mystaceus species complex and the status of previously described subspecies were discussed.

TABACHISIN, V. (2002): Long-eared roundhead. – Aquarium, 2002 (5): 38-39. (in Russisch).

TABACHISIN, V. (2003): Long-eared roundhead (keeping). – Aquarium, 2003 (2): 36-37. (in Russisch).

VELDRE, S.R. (1964): On the reality of subspecies division for the lizard Phrynocephalus mystaceus (Pall.). – Vestn. leningr. Univ. (Biol. Ser.), 1964 (3): 34-40. (in Russisch)

WHITING, M.J., NOBLE, D.W.A. & Y. QI (2022): A potential deimatic display revealed in a lizard. – Bio. J. Linn. Soc., 136: 455-465.

Conspicuously coloured signals may evolve via sexual selection to be ornaments or armaments, thereby conferring a fitness advantage to their bearer. Conversely, conspicuous colours may also evolve under natural selection as either aposematic signals or deimatic displays that deter attacks from predators. While conspicuous colour patches may evolve for one purpose (e.g. quality indicators), they may later be co-opted for another (e.g. anti-predator defence). Phrynocephalus mystaceus is a cryptic agamid lizard with flaps in both sexes that when folded against the head are inconspicuous, but when deployed are predicted to be highly conspicuous and to increase the appearance of body size. We tested whether head flaps play a role in social signalling via courtship or as status signals during contests in both sexes. We also tested whether the head flaps have an anti-predator function by simulating predatory encounters. Head flaps were never deployed in courtship or during contests and, therefore, are unlikely to be under sexual selection. However, head flaps and their deployment during simulated predatory encounters were consistent with the predictions associated with deimatic display theory. First, head flaps were similar in form and function between sexes. Second, they were highly conspicuous to both avian and snake predators. Third, there was a rapid transition from crypsis to conspicuousness when they deployed their head flaps during a late stage of predation, the subjugation phase, consistent with an ambush. Confirmation of the deimatic display hypothesis will require future testing of receiver responses.


Phrynocephalus mystaceus mystaceus (PALLAS, 1776)

Bärtiger Krötenkopf / Secret Toadhead Agama

OKSHTEIN, I.L., CHERLIN, V.A., VETO, D.V., KOZHARA, V.V., KORNEEVA, A.A., LOZHKINA, A.V., MIRONENKO-MARENKOV, A.D., RIABOVA, E.D., TISCHENKO, M.A., TUROVA, M.A., FRAERMAN, A.Y. & A.A. YATSUK (2021): Twilight activity in secret toadhead agamas (Phrynocephalus mystaceus mystaceus) on the Sarykum Barkhan Dune in Dagestan. - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p 199-201. (in Russisch)

POLYNOVA, G.V. (2016): An account of the activity of the secret toadhead agama (Phrynocephalus mystaceus mystaceus Pall. 1776) during assessments of its population Proceedings of the 6th congress of the A.M. Nikolsky Herpetological Society: 120-125. (in Russisch)

The detailed study of the activity of the Phrynocephalus mystaceus mystaceus population in Astrakhan semi-deserts showed that almost all resident lizards had some rest periods (1–4 days), when they do not emerge on the surface. The account of the lizards’ activity allows proposing some correcting coefficient, which is 1.88. Th is coeffi cient should be used to obtain more precise data on the population number based on one-day registrations.

POLYNOVA, G.V. (2016): Consideration of activity features in assessment population size of the Eared Roundhead (Phrynocephalus mystaceus mystaceus Pall. 1776). – Bull. Univ.St-Petersbourg. Series 3. Biologie, 2016 (3): 120–125.

The detailed study of the activity of the Phrynocephalus mystaceus mystaceus population in Astrakhan semi-deserts showed that almost all resident lizards had some rest periods (1–4 days), when they do not emerge on the surface. The account of the lizards’ activity allows proposing some correcting coeffi cient, which is 1.88. This coeffi cient should be used to obtain more precise data on the population number based on one-day registrations.

POLYNOVA, G.V. & O.E. POLYNOVA (2021): Assessment of the status of the Phrynocephalus mystaceus mystaceus population on the Sarykum Dune, Dagestan State Nature Reserve. - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p. 212-213. (in Russisch)

Five-year observations of the intra-population grouping of the toadheaded agama Phrynocephalus mystaceus mystaceus (Pallas, 1776) were carried out in semi-deserts of the Astrakhan region. It showed a reduction in its number up to complete extinction in this territory. The main reason for the degradation of the grouping was the overgrowth of weakly fixed and semi-fixed sandy areas - a typical biotope for the species. The process of overgrowth is shown on the materials of the description of geobotanical sites, the projective cover of which significantly increased during the observation period. The nucleus of the intrapopulation group was composed of sedentary sexually mature individuals, and its stability was determined by attachment to the site of females that repeatedly met in the study area in successive seasons. Sedentary males and females disappeared from the territory at the same time. The experience of significant changes by the species grouping in the characteristic biotope for a lot of years was due to the influx of migrants passing through the studied territory. This mechanism is well known in mammals and insufficiently studied in reptiles. The flow of migrants mainly consisted of immature individuals, an age group with increased mobility, which usually serves the purpose of resettlement. The decrease in the influx of nomadic individuals, apparently related to the general decline in the number of species in the surrounding area, led, in the end, to the disappearance of the group.

POLYNOVA, G.V. & O.E. POLYNOVA (2021): On the extinction of the intrapopulation group of the long-eared roundhead Phrynocephalus mystaceus mystaceus (Pallas, 1776) in the overgrown semi-deserts of the Astrakhan region – Principles of ecology, 2021 (1): 43-51.(In Russian).

Five-year observations of the intra-population grouping of the toadheaded agama Phrynocephalus mystaceus mystaceus (Pallas, 1776) were carried out in semi-deserts of the Astrakhan region. It showed a reduction in its number up to complete extinction in this territory. The main reason for the degradation of the grouping was the overgrowth of weakly fixed and semi-fixed sandy areas - a typical biotope for the species. The process of overgrowth is shown on the materials of the description of geobotanical sites, the projective cover of which significantly increased during the observation period. The nucleus of the intrapopulation group was composed of sedentary sexually mature individuals, and its stability was determined by attachment to the site of females that repeatedly met in the study area in successive seasons. Sedentary males and females disappeared from the territory at the same time. The experience of significant changes by the species grouping in the characteristic biotope for a lot of years was due to the influx of migrants passing through the studied territory. This mechanism is well known in mammals and insufficiently studied in reptiles. The flow of migrants mainly consisted of immature individuals, an age group with increased mobility, which usually serves the purpose of resettlement. The decrease in the influx of nomadic individuals, apparently related to the general decline in the number of species in the surrounding area, led, in the end, to the disappearance of the group.

POLYNOVA, G.V. & O.E. POLYNOVA (2024): Specific features of the size structure of marginal populations of the nominative subspecies of the secret toad-headed agama (Phrynocephalus mystaceus mystaceus) (Agamidae, Reptilia). - Povolzhskiy Journal of Ecology, 2024 (1): 52-63. (In Russian, English summary).

PORCU, D. (2019): Skurril und agil: Bärtige Krötenkopfagamen im Terrarium. – Reptilia, 24 (5): 34-36.

RIABOVA, E.D., VETO, D.V., OKSHTEIN, I.L., TISCHENKO, M.A. & A.Y. FRAERMAN (2021): The social contacts of Phrynocephalus guttatus guttatus (Gmelin, 1789) and Phrynocephalus mystaceus mystaceus (Pallas, 1776). - In: Dunayev, E.A. & N.A. Poyarkov et al. (ed.): Problems of Herpetology Program and abstracts of the VIII congress of the A.M. Nikolsky Herpetological Society (NHS) of the Russian Academy of Sciences “Current herpetological research in Eurasia” October 3-9. 2021. Moscow. p. 229-231. (in Russisch)

SEMENOV, D.V. & G.I. SHENBROT (1990): Species of the genus Phrynocephalus of the USSR fauna. Description of a new subspecies with remarks on the taxonomic status of Phrynocephalus mystaceus (Reptilia, Agamidae). – Zool. Zh. (USSR), 69 (5): 76-81.

TADEVOSYAN, T.L. (2007): The role of vegetation in microhabitat selection of syntopic lizards, Phrynocephalus persicus, Eremias pleskei, and Eremias strauchi from Armenia. – Amphibia-Reptilia, 28 (3): 444-448.

Bärtiger Krötenkopf / Secret Toadhead Agama

Phrynocephalus ornatus BOULENGER, 1887

Krötenkopfagame

ANDERSON, S.C. (1999): Phrynocephalus ornatus Boulenger, 1887 - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 93-94.

ANDERSON, S.C. & A.E. LEVITON (1967): A new species of Phrynocephalus (Sauria: Agamidae) from Afghanistan, with remarks on Phrynocephalus ornatus Boulenger. – Proceedings of the California Academy of Sciences, ser. 4, 35 (11): 227-234.

CLARK, R.J. (1992): Notes on the distribution and ecology of Phrynocephalus clarkorum Anderson & Leviton 1967 and Phrynocephalus ornatus Boulenger 1887 in Afghanistan. – Herpetological Journal, 2: 140-142.


Phrynocephalus ornatus ornatus BOULENGER, 1887


Phrynocephalus ornatus vindumi GOLUBEV,1998

GOLUBEV, M.L. (1998): A new subspecies of Phrynocephalus ornatus Boulenger from eastern Iran, with a key to South-Westernand Middle Asian microphrynocephalids. - Hamadryad, 23 (2): 162–168.

Phrynocephalus przewalskii STRAUCH, 1876

Przewalski´s Toadhead Agama

CHANG CHENG (1995): Histological changes in the epidermis of Phrynocephalus prezewalskii during the sloughing cycle. – Journal of Lanzhou University Natural Sciences, 31 (1): 55-60. (in Chinesisch)

CHANG CHENG & WANG ZIREN (1996): Morphological studies of the skin receptor of Phrynocephalus prezewalskii. – Journal of Lanzhou University Natural Sciences, 32 (1): 92-97. (in Chinesisch)

DUNAYEV, E.A., SOLOVYEVA, E.N. & N.A. POYARKOV (2021): Systematics, Phylogeny, and Evolution of Phrynocephalus (Superspecies przewalskii) (Reptilia: Agamidae). – Russian Journal of Herpetology, 28 (1): 43-59.The superspecies przewalskii group of Phrynocephalus includes several taxa with unclear taxonomic status. We analyze a fragment of the mitochondrial DNA gene COI and the body patterns of 275 specimens including type specimens. The results resolve the taxonomy and phylogeny of the group and we provide a diagnostic key for the specie

FU, J. (2010): Male-mediated gene flow in the toad-headed lizards Phrynocephalus przewalskii. - Abstracts of the Second International Symposium on Agamid Lizards «DeAgamis2». - Current Studies in Herpetology, 10 (3/4): 144.

GOZDZIK, A. & J. FU (2009): Are Toad-Headed Lizards Phrynocephalus przewalskii and P. frontalis (Family Agamidae) the Same Species? Defining Species Boundaries with Morphological and Molecular Data. - Russ. J. Herpetol. 16 (2): 107-118.

The paper deals with the relationship between the body temperatures of Phrynocephalus przewalskii (Sthauch) and Eremias multiocellata (Guenther) and the environmental temperatures,their selections of environmental temperatures and their resistance against low and high temperatures. The body temperatures of przewalskii and multiocellata were negatively interrelated to environmental temperatures (P<0.001). Under the same temperatures. The body temperatures of przewalskii were 3 °C higher than that of multiocellata. The environmental temperatures selected by the former were 38-40 °C,whereas the latter required only 35-37 °C. The hot and dead temperatures of przewalskii were higher than those of multiocellata. The threshhold of the hot and dead temperatures in przewalskii varied from 44 °C to 48 °C and its highest dead temperature (TL[50]) was up to 48 °C,and that in multiocellata was 42-46 °C and its highest hot temperature (TL[50]) was 46 °C. The ability that two species can resist against low temperatures was equal. The cold and dead temperatures varied from 0 °C to -3 °C. The cold and dead temperatures (TL[50]) of przewalskii were -2.3 °C, but those of multiocellata were -2.5 °C. These significant differences between the two species are concerned with the characteristics of each, habitats and sizes of the bodies.

LI, S.-R., HAO, X., BI, J.-H. & W.-G. DU (2017): Why do female desert lizards construct burrows to lay eggs? – Behaviour, 154 (9-10): 1051-1068.

Many oviparous animals construct well-designed nests to provide relatively favourable conditions for their eggs and hatchlings, but the direct evidence that nest structure can determine their reproductive success is insufficient. In the present study, we explored the structure of nests and ist effect on nest environments and reproductive success in the toad-headed agama (Phrynocephalus przewalskii). We observed that female P. przewalskii constructed burrow nest consisting of an inclined tunnel and an expanded chamber. We constructed artificial nests with or without the burrow to determine how burrows influence nest environments, egg survival and successful emergence of hatchlings. Our results indicated that burrow nests had higher and more stable humidity than non-burrow nests. More importantly, egg survival and the emergence success of hatchlings were significantly higher for burrow nests than for non-burrow nests. Therefore, our manipulation experiments provide direct evidence that maternal nest construction behaviour could determine parental reproductive success in reptile.

LI, S.-R., WANG, Y., MA, L., ZENG, Z.-G., BI, J.-H. & W.-G. DU (2017): Thermal ecology of three coexistent desert lizards: Implications for habitat divergence and thermal vulnerability. – Journal of Comparative Physiology B, 2017. doi:10.1007/s00360-017-1087-4

How ectotherms exploit thermal resources has important implications for their habitat utilization and thermal vulnerability to climate warming. To address this issue, we investigated thermal relations of three sympatric lizard species (Eremias argus, Eremias multiocellata, and Phrynocephalus przewalskii) in the desert steppe of Inner Mongolia, China. We determined the thermoregulatory behavior, body temperature (T_b), operative temperature (T_e), selected body temperature (T_sel), and critical thermal maximum (CT_max) of adult lizards. Based on these physiological parameters, we quantified the accuracy and effectiveness of thermoregulation as well as thermal-safety margin for these species. The three species were accurate and effective thermoregulators. The P. przewalskii preferred open habitats, and had a higher T_b than the two Eremias lizards, which preferred shade habitats and shuttled more frequently between the shade and sun. This indicated that the three sympatric lizards have different thermoregulatory behavior and thermal physiology, which might facilitate their coexistence in the desert steppe ecosystem. In addition, the P. przewalskii had higher T_sel and CT_max, and a wider thermal-safety margin than the two Eremias lizards, suggesting that the two Eremias lizards would be more vulnerable to climate warming than P. przewalskii.

LI, X., WANG, Y., LI, M., MEN, S., PU, P., TANG, X. & Q. CHEN (2017): Comparison of cold hardiness of two Toad-headed lizards from different altitudes. – Sichuan Journal of Zoology, 36 (3): 300-305. (in Chinesisch)

LI, T., ZHAO, B., ZHOU, Y.K., HU, R. & W.G. DU (2014): Thermoregulatory behavior is widespread in the embryos of reptiles and birds. – The American Naturalist, 183 (3): 443-451.

Recent studies have demonstrated that thermoregulatory behavior occurs not only in posthatching turtles but also in turtles prior to hatching. Does thermoregulatory behavior also occur in the embryos of other reptile and bird species? Our experiments show that such behavior is widespread but not universal in reptile and bird embryos. We recorded repositioning within the egg, in response to thermal gradients, in the embryos of three species of snakes (Xenochrophis piscator, Elaphe bimaculata, and Zaocys dhumnades), two turtles (Chelydra serpentina and Ocadia sinensis), one crocodile (Alligator sinensis), and four birds (Coturnix coturnix, Gallus gallus domesticus, Columba livia domestica, and Anas platyrhynchos domestica). However, we detected no significant thermoregulation by the embryos of two lizard species (Takydromus septentrionalis and Phrynocephalus frontalis). Overall, embryonic thermoregulatory behavior is widespread in reptile as well as bird species but may be unimportant in the small eggs laid by most lizards.

LIU, C.-J., ZHAO, W., ZHOU, R. & N.-F. LIU (2012): Phylogeny and genetic diversity of Phrynocephalus przewalskii. – Zool. Res., 33 (2): 127-132.

Phrynocephalus przewalskii is one kind of lizards inhabiting inclusively in the desert that has controversial viewpoints on its phylogeny. Based on mitochondrial ND2 gene of 119 samples from 12 geographic populations, we analyzed the effects of environmental factors on the variation of genetic diversity, as well as ist relationship to P. versicolor. The results showed that these populations clustered into three major lineages, with P. versicolor embedded within one lineage. The twelve populations had great genetic diversity variation, which was tightly linked with local altitude, annual precipitation, and variation of annual precipitation. High latitudes, increased annual precipitation and great variation in annual precipitations may all have resulted in the decrease of genetic diversity. It thus assumed that altitude can change the genetic diversity of different geographic populations of P. przewalskii resulting from the effects of different local annual precipitation.

LIU, N., LI, R. & H. SUN (1993): The effects of environmental factors on population density of Phrynocephalus przewalskii. – Zool. Res., 14 (4): 319-325. (in chinesisch)

LIU, N.F., LIU, C.J., ZHAO, W. & R. ZHOU (2012): Phylogeny and genetic diversity of Phrynocephalus przewalskii. – Zool. Res., 33 (2): 127-132. (In Chinese).

Phrynocephalus przewalskii is one kind of lizards inhabiting inclusively in the desert that has controversial viewpoints on its phylogeny. Based on mitochondrial ND2 gene of 119 samples from 12 geographic populations, we analyzed the effects of environmental factors on the variation of genetic diversity, as well as ist relationship to P. versicolor. The results showed that these populations clustered into three major lineages, with P. versicolor embedded within one lineage. The twelve populations had great genetic diversity variation, which was tightly linked with local altitude, annual precipitation, and variation of annual precipitation. High latitudes, increased annual precipitation and great variation in annual precipitations may all have resulted in the decrease of genetic diversity. It thus assumed that altitude can change the genetic diversity of different geographic populations of P. przewalskii resulting from the effects of different local annual precipitation.

MA, L., SUN, B.-J., LI, S.-R., HAO, X., BI, J.-H. & W.-G. DU (2018): The vulnerability of developing embryos to simulated climate warming differs between sympatric desert lizards. – JEZ-A Ecological and Integrative Physiology, 2018. https://doi.org/10.1002/jez.2179

The vulnerability of species to climate warming varies along latitudinal and elevational clines, but how sympatric species vary in vulnerability to climate warming remains largely unknown. We experimentally simulated nest temperatures of two sympatric lizards with divergent microhabitat preferences (Phrynocephalus przewalskii and Eremias argus), under climate warming senarios, to determine the response of embryos to increased mean temperatures and heat waves. Our study demonstrated that simulated climate warming reduced hatching success and hatchling size and growth in E. argus (that prefers closed microhabitats), but had less effect in P. przewalskii (that occupies open microhabitats). The reduced growth rate of E. argus hatchlings was associated with a decrease in metabolic rate, which was more evident in hatchling E. argus than in P. przewalskii. Our results suggest lizards that prefer closed microhabitats may be more vulnerable to climate warming than those that prefer open microhabitats; further studies are needed to test this hypothesis. More generally, the divergent responses of sympatric species to climate warming highlights the importance of distinguishing the thermal sensitivity of behavior and physiology for each species of a community, in order to make predictions about the impacts of climate warming at regional scales.

SONG, Z., CHEN, L., CHEN, Q. & S. CHEN (1987): Studies on the breeding habit of Phrynocephalus przewalskii. - Acta Herpetol. Sin., 6 (1): 12-17. (In Chinese, English summary).

TANG, X., YUE, F., MA, M., WANG, N.B., HE, J. & Q. CHEN (2012): Effects of thermal and hydric conditions on egg incubation and hatchling phenotypes in two Phrynocephalus lizards. – As. Herpetol. Res., 3 (3): 184-191.

Flexible-shelled eggs of the lizards Phrynocephalus przewalskii and P. versicolor were incubated under different thermal and hydric conditions to elicit the effects of incubation environment on hatching success, embryonic development and duration as well as hatchling phenotypes. Embryogenesis of the two species was not sensitive to changes in the hydric environment except P. przewalskii incubated in 30°C group. Temperature significantly altered the duration of embryogenesis, with cooler temperatures leading to a longer incubation period. Hatching success was greater at 26 and 30°C than at 34°C. The hatchlings incubated at 26 and 30°C had longer snout-vent length, larger body mass, and better locomotor performance than those incubated at 34°C. Compared to P. przewalskii, P. versicolor had a shorter incubation period and yielded smaller hatchlings, which then had a higher survival rate in cooler and drier habitats. We conclude that an incubation temperature of 30°C would produce the best balance among developmental rate, hatching success, and post-hatching performance. We speculate that the upper temperature limit for incubation of P. versicolor eggs may be slightly higher than 34°C.

URQUHART, J., WANG, Y. & J. FU (2009): Historical vicariance and male-mediated gene flow in the toad-headed lizards Phrynocephalus przewalskii. – Molec. Ecol., 18 (17): 3714-3729.

Using mitochondrial and microsatellite DNA data and a population genetic approach, we tested male-mediated gene flow in the toad-headed lizards Phrynocephalus przewalskii. The mitochondrial DNA (ND2 gene), on the one hand, revealed two major lineages and a strong population genetic structure (FST = 0.692; FST￠ = 0.995). The pairwise differences between the two lineages ranged from 2.1% to 6.4% and the geographical division of the two lineages coincided with a mountain chain consisting of the Helan and Yin Mountains, suggesting a historical vicariant pattern. On the other hand, the nuclear microsatellite DNA revealed a significant but small population genetic structure (FST = 0.017; FST￠ = 0.372). The pairwise FST among the nine populations examined with seven microsatellite DNA loci ranged from 0.0062 to 0.0266; the assignment test failed to detect any naturally occurring population clusters. Furthermore, the populations demonstrated a weak isolation by distance and a northeast to southwest clinal variation, rather than a vicariant pattern. A historical vicariant event followed by male-mediated gene flow appears to be the best explanation for the data. Approximately 2–5 Ma, climatic change may have created an uninhabitable zone along the Helan-Yin mountain chain and initiated the divergence between the two mitochondrial lineages. With further climatic changes, males were able to disperse across the mountain chain, causing sufficient gene flow that eventually erased the vicariant pattern and drastically reduced the population genetic structure, while females remained philopatric and maintained the mitochondrial DNA (mtDNA) divergence. Although polygyny mating system and female philopatry may partially contribute to the reduced movement of females, other hypotheses, such as female intrasexual aggression, should also be explored.

WANG, X., LI, S., LI, L., ZHANG, F., HAN, X., BI, J. & B. SUN (2019): Thermal-physiological Strategies Underlying the Sympatric Occurrence of Three Desert Lizard Species. - Asian Herpetological Research, 10 (3): 190-196.

Sympatric reptiles are the ideal system for investigating temperature-driven coexistence. Understanding thermally physiological responses of sympatric lizards is necessary to reveal the physiological mechanisms that underpin the sympatric occurrence of reptiles. In this study, we used three lizard species, Eremias argus, E. multiocellata, and Phrynocephalus przewalskii, which are sympatric in the Inner Mongolia desert steppe, as a study system. By comparing their resting metabolic rates (RMR) and locomotion at different body temperatures, we aimed to better understand their physiological responses to thermal environments, which may explain the sympatric occurrence of these lizards. Our results showed that E. argus had significantly higher RMR and sprint speed than E. multiocellata, and higher RMR than P. przewalskii. In addition, the optimal temperature that maximized metabolic rates and locomotion for E. argus and E. multiocellata was 36°C, whereas for P. przewalskii it was 39°C. Our study revealed the physiological responses to temperatures that justify the sympatric occurrence of these lizards with different thermal and microhabitat preferences and active body temperatures. Eremias argus and E. multiocellata, which have lower body temperatures than P. przewalskii, depend on higher RMR and locomotion to compensate for their lower body temperatures in field conditions. Our study also highlights the importance of using an integrative approach, combining behavior and physiology, to explore the basis of sympatric occurrence in ectothermic species.

WANG, Y. & H. WANG (1993): Geographic variation and diversity in three species of Phrynocephalus in the Tengger Desert, Western China. - Asiatic Herpetological Research 5: 65-73.

WANG, Y., JI, W., ZHAO, W., YU, N. & N. LIU (2011): Geographic Variation in Clutch and Egg Size for the Lizard Phrynocephalus przewalskii (Squamata: Agamidae). - Asian Herpetological Research 2 (2): 97-102.

In order to tease apart proximate vs. ultimate sources of variation in reproductive strategy, studies have increasingly focused on populations rather than species as the unit of interest. The reproductive parameters of Phrynocephalus przewalskii (Agamidae) in different populations within the same phylogenetic clade were compared in this study. Female SVL, clutch size, egg volume and clutch volume varied significantly among populations. With increase in latitude, clutch size increased, while egg size decreased. Relatively fewer but larger eggs were produced with increasing of population density. Food availability had positive effects on clutch size, but no effect on egg size. Our result indicated that latitude, food availability and population density may be the proximate factors affecting the reproductive parameters of P. przewalskii.

XU, H. & F. YANG (1993): Age classification and growth model of Phrynocephalus przewalskii. – Ecol. Modell., 70 (1-2): 127-135.

ZENG, Z.G., ZHAO, J.M. & B. SUN (2013): Life history variation among geographically close populations of the toad-headed lizard (Phrynocephalus przewalskii): exploring environmental and physiological associations. - Acta Oecol., 51: 28-33.

Geographic variation in life history traits has been extensively studied along latitudinal and altitudinal clines, but life history variation among geographically close populations has received much less attention. We collected gravid female toad-headed lizards (Phrynocephalus przewalskii) and environmental data from three localities (Alxa Zuoqi, Alxa Youqi, and Shandan) across the Gobi desert in China, to examine among-population differences in reproductive strategies. The precipitation was significantly lower in Alxa Youqi than Alxa Zouqi and Shandan. Food availability was highest in Shandan, lowest in Alxa Zuoqi, with Alxa Youqi in between. Females from Shandan population were larger and produced more and larger eggs than their counterparts from the other two populations. Incubation period also differed among the populations, with the lowest incubation period in Alxa Youqi population, and the longest incubation period in Alxa Zuoqi population. Our data on the physiological mechanisms of incubation period indicated that the shortened incubation period in Alxa Youqi population was due to advanced embryogenesis completed prior to oviposition rather than higher embryonic heart rates during incubation. Therefore, our data support the hypothesis that geographically close populations can show different reproductive strategies if environmental factors vary among these populations.

ZHAN, A. & J. FU (2009): Microsatellite DNA markers for three toad-headed lizard species (Phrynocephalus vlangalii, P. przewalskii and P. guttatus). – Molec. Ecol. Res., 9 (2): 535-538.

Animals depend on their habitats to complete a life cycle and habitat selection will affect all their other selections. Parameters of 173 selected and 227 controlled habitats of Phrynocephalus przewalskii from four populations around the Tengger desert were compared using principal component analysis, ANOVA and Mann-Whitney U-test. It was found that abitat selection was similar between males and females, but differed in seasons or among the populations. Before breeding, P. przewalskii selected sites with a lower cover of vegetation to achieve optimal body temperature. In order to recruit the energy used in reproduction and accumulate energy for passing the winter, they would favor sites with a high density and cover of grass after breeding. No significant seasonal difference was found in the controlled habitats; so the variations in the selected habitats were mainly caused by active selection. Geographic variation patterns in selected and controlled habitats were similar. Thus the geographic variations in P. przewalskii habitat selection might have mainly contributed to the environment variations. Moreover, P. przewalskii would weigh the benefit between foraging and avoiding predation, and make different decisions in different conditions. For example, P. przewalskii would favor bathing and foraging when the vegetation coverage was high, but mainly favored foraging when the food was short.

ZHAO, W. & N. LIU (2013): Sexual Dimorphism of Head Size in Phrynocephalus przewalskii:Testing the Food Niche Divergence Hypothesis. - Asian Herpetological Research  4 (4): 242–247.

Sexual size dimorphism (SSD) is a general phenomenon in lizards, and can evolve through sexual selection or natural selection. But natural selection, which was thought to operate mainly through reducing the competition between the two sexes (niche divergence hypothesis), gave rise to a lot of controversy. We tested the niche divergence hypothesis in the toad-headed lizard Phrynocephalus przewalskii by comparing diet composition and prey sizes between males and females. The species was found to be sexual dimorphic, with males having relatively larger snout-vent length, head width, head length, and tail length, while females have relatively larger abdomen length. Based on analysis of 93 studied stomachs, a total of 1359 prey items were identified. The most common prey items were formicid, lygaeid and tenebrionid. The two sexes did not differ in the relative proportions of prey size categories they consumed and the dietary overlap based on prey species was high (O = 0.989). In addition, the meal size, the volume or any maximal dimension of the largest prey item in the stomach was not explained by the sexes. According to our results, food niche divergence might not play an important role in the SSD evolution of P. przewalskii.

ZHAO, W. & N. LIU (2014): The proximate causes of sexual size dimorphism in Phrynocephalus przewalskii. - PLoS ONE, 9 (1): e85963.

Sexual size dimorphism (SSD) is a common phenomenon and is a central topic in evolutionary biology. Recently, the importance of pursuing an ontogenetic perspective of SSD has been emphasized, to elucidate the proximate physiological mechanisms leading to its evolution. However, such research has seldom focused on the critical periods when males and females diverge. Using mark-recapture data, we investigated the development of SSD, sex-specific survivorship, and growth rates in Phrynocephalus przewalskii (Agamidae). We demonstrated that both male and female lizards are reproductively mature at age 10–11 months (including 5 months hibernation). Male-biased SSD in snout-vent length (SVL) was only found in adults and was fully expressed at age 11 months (June of the first full season of activity), just after sexual maturation. However, male-biased SSD in tail length (TL), hind-limb length (LL), and head width (HW) were fully expressed at age 9–10 months, just before sexual maturation. Analysis of age-specific linear growth rates identified sexually dimorphic growth during the fifth growth month (age 10–11 months) as the proximate cause of SSD in SVL. The males experienced higher mortality than females in the first 2 years and only survived better than females after SSD was well developed. This suggests that the critical period of divergence in the sizes of male and female P. przewalskii occurs between 10 and 11 months of age (May to June during the first full season of activity), and that the sexual difference in growth during this period is the proximate cause. However, the sexual difference in survivorship cannot explain the male-biased SSD in SVL. Our results indicate that performance-related characteristics, such as TL, HW, and LL diverged earlier than SVL. The physiological mechanisms underlying the different growth patterns of males and females may reflect different energy allocations associated with their different reproductive statuses.

ZHAO, W., YU, N., WANG, Y., JI, W. & N.F. LIU (2011): Female reproductive cycles of Phrynocephalus przewalskii (Lacertilia: Agamidae) in the Tengger Desert, China. - As. Herpetol. Res., 2 (1): 30-35.

The timing of reproduction can significantly affect an offspring’s fitness, thereby also influencing the fitness of the parents, especially in species inhabiting extreme environments, such as deserts. Female reproductive cycles in Phrynocephalus przewalskii were studied from April to September 2008. Significant cycles of gonadal volume were found in all studied populations and the cycles were similar among the various populations. Females began vitellogenesis in April and contained oviductal eggs form May to June. Gonad volume decreased significantly in July and reached minimum volume from August to September. The follicular growth was negatively correlated with increasing precipitation and temperature in all populations. Hatching occurs during summer and early fall, when most of the annual rainfall occurs. Mean clutch size based on all populations was 2.7 ± 0.9 SE (n = 71).

Phrynocephalus putajatai

JI, X.,  WANG, Y.-Z. & & Z. WANG (2009): New species of Phrynocephalus (Squamata, Agamidae) from Qinghai, Northwest China. - Zootaxa 1988: 61-68.

A new viviparous species of Phrynocephalus from Guinan, Qinghai, China, is described. Phrynocephalus guinanensis sp. nov., differs from all congeners in the following combination of characters: body large and relatively robust; dorsal ground color of head, neck, trunk, limbs and tail brown with weak light brown mottling; lateral ground color of head, neck, trunk and tail light black with weak white-gray mottling in adult males, and green with weak white-gray mottling in adult females; ventral ground color of tail white-gray to black in the distal part of the tail in adult males, and totally white-gray in adult females; ventral surfaces of hind-limbs white-gray; ventral surfaces of fore-limbs brick-red in adult males, and white-gray in adult females; ventral ground color of trunk and head black in the center but, in the periphery, brick-red in adult males and white-gray in adult females. Phrynocephalus guinanensis sp. nov. typically uses desert habitats, whereas P. vlangalii, a species closely related to the new form, uses a variety of arid and semi-arid habitats. It is the nineteenth species of Phrynocephalus recorded from China.

JIN, Y., YANG, Z., BROWN, R.P., LIAO, P. & N.F. LIU (2014): Intraspecific lineages of the lizard Phrynocephalus putjatia from the Qinghai-Tibetan Plateau: impact of physical events on divergence and discordance between morphology and molecular markers. – Molec. Phylogen. Evol, 71: 288-297.

The Northeastern Qinghai-Tibetan Plateau (NQTP) contains many physical features that may have contributed to its rich biological diversity. We explored the systematics and genetic structures of the sand lizards Phrynocephalus putjatia, distributed across semi-deserts or rocky steppe habitats, and Phrynocephalus guinanensis, found only in sand dune habitats, from the NQTP using mitochondrial (ND2 and tRNAs) and nuclear (RAG-1) markers. Bayesian analyses revealed two main monophyletic mtDNA groups that separate populations of the Qinghai Lake Basin (QLB) from populations to the southeast of this basin (SEQL), but these did not correspond to the two morphological species. The QLB populations are divided into western and eastern groups. Two major groups with similar geographical structuring were also detected for the nuclear RAG-1 marker, but with some geographical discordance. A Bayesian species delimitation analysis did not support division of P. putjatia/P. guinanensis into separate species. Bayesian dating of mtDNA suggests that the earliest divergence within this group occurred less than 2 Ma, which seems to be explained by mountain uplift between the QLB and the SEQL regions. Other Pleistocene events may explain further genetic structuring. Overall, we do not detect reciprocal monophyly of markers between morphological species but note that they do appear to represent ecological forms.

LI, J., TONG, H., ZHANG, K., YANG, K., LUO, Y. & Y. JIN (2016): Chromosomal karyotype characteristics and systematic cluster analysis of Phrynocephalus guinanensis. – Sichuan Journal of Zoology, 35 (3): 391-394. (in Chinesisch)

TONG, H. & Y. JIN (2014): The complete mitochondrial genome of an agama, Phrynocephalus putjatia (Reptilia, Squamata, Agamidae). – Mitochondrial DNA, 2014. DOI: 10.3109/19401736.2014.926538

The complete mitochondrial genome was sequenced from the toad-headed viviparous lizard, Phrynocephalus putjatia. The mitogenome was 16,283 bp in length; it contained 13 protein coding, 22 tRNA, 2 rRNA genes and 2 control regions. The gene order and compositions were identical with all published congeneric mitogenomes for the fragment between 12 s RNA and tRNA-Thr, but with some differences for the remaining sequences including CR, tRNA-Pro and tRNA-Phe. The characteristics of the mitogenome was analyzed and discussed in detail.

Phrynocephalus raddei BOETTGER, 1888

ANDERSON, S.C. (1999): Phrynocephalus raddei Boettger, 1890 - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 96-97.

BOETTGER, O. (1888): Über die Reptilien und Batrachier Transcaspiens. - Zool. Anz. 11: 259-263.

SHAMMAKOV, S.S., ATAEV, C.A. & Z.Y. KAMALOVA (1973): The ecology of Phrynocephalus raddei in Turkmenia. – Ekologiya, 1973 (6): 80-83.


Phrynocephalus raddei raddei BOETTGER, 1888


Phrynocephalus raddei boettgeri BEDRIAGA, 1906

Reticulated Toad-headed Agama

EICHWALD, E. (1831): Description of Phrynocephalus helioscopus varius; Ph. reticulatus in “Zoologia specialis, quam expositis animalibus tum vivis, tum fossilibus potissimuni rossiae in universum, et poloniae in specie, in usum lectionum publicarum in Universitate Caesarea Vilnensi”. Zawadski, Vilnae.

MEZHZHERIN, S. & M.L. GOLUBEV (1998): Allozyme variation and genetic differentiation of toad agama species group Phrynocephalus ocellatus (=reticulatus) (Reptilia: Agamidae). - Byull. Mosk. Obshch. Isp. Prir. Otd. Biol., 103 (6): 9-16. (In Russian, English summary).

SHENBROT, G.I. & G.S. ANTONOVA (1981): Comparative ecology of Phrynocephalus helioscopus and P. reticulatus in the conditions of their common habitat, south of Bukhara region (Uzbek SSR). - In: Darevsky, I.S., Ananjeva, N.B., Barkagan, Z.S., Borkin, L.Y., Sokolova, T.M. & N.N. Szczerbak (eds.): The problems of herpetology: abstracts. Nauka, Leningrad: 156-157. (in Russian).


Phrynocephalus reticulatus reticulatus EICHWALD, 1831


Phrynocephalus reticulatus bannikovi DAREVSKY, RUSTAMOV & SHAMMAKOV, 1976

OVEZMUKHAMMEDOV, A. (1977): A new Coccidia Isospora rustamovi, sp. n. from the lizard (Phrynocephalus reticulatus bannicovi) in Turkmenistan. – Izvestiya Akad. Nauk Turkmen. SSR (Biol.), 1977 (6): 67-78. (in Russisch)

SHAMMAKOV, S. (1977): On the ecology of Phrynocephalus reticulatus bannicovi. – In: Darevskij, I.S. (ed.): Fourth all-Union Herpetological Conference. Questions of Herpetology. – Akademiya Nauk SSSR, Zoologicheskij Institut. Izdatel´stvo ´Nauka´. Leningrad. 230.

Phrynocephalus roborowskii BEDRIAGA, 1907

Roborowski´s Toadhead Agama

BEDRIAGA, J. von (1905) Verzeichnis der von der Central-Asiatischen Expedition unter Stabs-Kapitän W. Roborowski in den Jahren 1893-1895 gesammelten Reptilien. - Annuaire du Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg (1905) 10 (3-4) : 159-200.

Phrynocephalus rossikowi NIKOLSKY, 1898

Uzbekistan Toadhead Agama

GOLUBEV, M.L., MANILO,V.V. & A.A. TOKAR (1994): Geographic variability of Phrynocephalus rossikowi Nik. (Reptilia: Agamidae) in Turkmenistan and adjacent regions [including karyotype]. In: Fet.,V. & K.I. Atamuradov (eds.): Biogeography and Ecology of Turkmenistan. Kluwer Academic Publishers, The Netherlands: 351-364.

We studied karyotypes and morphology of five samples of Ph. rossikowi from different parts of the species range. Revealed variability shows that peripheral populations of this species undergo intensive speciation; for further taxonomic conclusions, additional detailed studies of populations from Chardzhou and Lebab are required. Analysis of variability shows that the Amudarya and Syrdarya Rivers played an important role in the formation of modern range of Ph. rossikowi. We consider central populations as a relatively stable core as compared to peripheral ones. It is suggested that Ph. rossikowi originated through adaptation of ancestral lizards to the local stony semi-desert environment in the West Kizylkum. The most ancient part of the range of Ph. rossikowi probably lies within the alluvial fan submontane plain of the Sultanuizdagh Range and the Pitnyak elevation in East Turkmenistan, adjacent to the south. The most ancient part of the submontane plain of the Sultanuizdagh has an early Pleistocene age (Kogai 1957). Therefore, our opinion differs from that of Ananjeva and Tunijev (1992), who suggested that Ph. rossikowi is a Pliocene species whose range shifted during the Pleistocene from the middle portion of the Amudarya River to its lower part.

NIKOLSKY, A.M. (1898): Dwa nowych wida jasceric iz Rossii [On two new lizards from Russia; in Russian and Latin]. -Annuaire Musée Zoologique de l’Académie Impériale des Sciences de St.-Pétersbourg 3: 284-288.


Phrynocephalus rossikowi rossikowi NIKOLSKY, 1898



Phrynocephalus rossikowi shammakowi SZCZERBAK & GOLUBEV, 1979

SZCZERBAK, N.N. & M.L. GOLUBEV (1979): The new subspecies of the toad-headed agama — Phrynocephalus rossikowi shammakowi ssp. n. (Reptilia, Sauria, Agamidae) from the central region of the Kara Kum Desert. [in Russian with English summary]. - Vestnik Zoologii, Kiev 6: 81–82.

MANILO, V.V. & M.L. GOBULEV (1993): Description of karyotype of Phrynocephalus rossikowi shammakowi (Reptilia, Agamidae). - Vestnik Zoologii, 1993 (2): 74-76.

Phrynocephalus scutellatus OLIVIER, 1807

Gray Toadhead Agama

ANDERSON, S.C. (1999): Phrynocephalus scutellatus Olivier, 1807  - In: Lizards of Iran. Society for the Study of Amphibians and Reptiles. Oxford, Ohio: 97-99.

OLIVIER, G.A. (1807): Description of Phrynocephalus scutellatus. – In: “Voyage dans l’Empire Othoman, l’Egypte et la Perse”. Vol. 3. Agasse, Paris.

RAHIMIAN, H., SHAFIEI, S., RASTEGAR POUYANI, N. & E. RASTEGAR POUYANI (2015): Phylogenetic relationships of the gray-toad agama, Phrynocephalus scutellatus (Olivier, 1807), species complex from Iran. – Zootaxa 3990 (3): 369-380.

The gray toad agama, Phrynocephalus scutellatus (Olivier, 1807) species complex is confined to the Iranian plateau, and forms one of the most widespread, but rarely studied species of the family Agamidae. It represents a complex with many local populations inhabiting a variety of habitats, and exhibiting considerable morphological, genetic and ecological vari-ations. We analyzed sequences of the mitochondrial ND2 gene and tRNA-Trp and tRNA-Ala derived from 89 geograph-ically distant populations. The sequences data strongly support a basal separation of the populations of southeastern— south-central Iran from those occurring in the North. The subsequent radiation, fragmentation, and evolution of these ma-jor assemblages have led to four discernible geographical lineages in Iran: southeastern—south-central, west-central, east—northeastern and Khaf. The southeastern—south-central radiation is the earliest lineage and Khaf lineage is proba-bly related to the Afghan plateau. Separation of northern clades from each other can be explained by the presence of large deserts in central Iran. Due to the lack of sufficient geological information, the divergence between the northern and southern clades cannot be explained by the present data.

Phrynocephalus strauchi NIKOLSKY, 1899

KAMALOVA, Z.Y. (1970): Feeding habits of Phrynocephalus reticulates strauchi Nik. in the sands of the Fergana Valley during the summer season. – Ekologiya, 1970 (5): 102-103. (in Russisch)

DUNAYEV, E.A. (1995): Reviewed description of the types of Phrynocephalus strauchi Nikolsky, 1899 (Squamata, Agamidae) and materials on the history of its study, distribution, and variability. – Russian Journal of Herpetology, 2 (2): 87-94.

Phrynocephalus theobaldi BLYTH, 1863

Theobald´s Toad-headed Agama / Toad Mounted Lizard / Snow Lizard

BLYTH, E. (1863): Description of Phrynocephalus theobaldi. - In: Report of the Curator, Zoological Department. Journal of the Asiatic Society of Bengal. 32 (1) : 73-90.

JIN, Y. & P. LIAO (2015): An elevational trend of body size variation in a cold-climate agamid lizard, Phrynocephalus theobaldi. - Current Zoology 61 (3): 444–453.

The pattern that many ectotherms have smaller body sizes in cold environments follows the converse to Bergmann’s rule and is most frequently found in lizards. Allen’s rule predicts animals from warm climates usually have longer tails and limbs, while these traits tend to be shorter in individuals from cold climates. We examined body size variation in an endemic Chinese lizard Phrynocephalus theobaldi along a broad elevational gradient (3,600–5,000 m on the Qinghai-Tibetan Plateau). Female body size showed a U-shaped cline, decreasing with increased elevation within the range 3,600–4,200 m, but increasing at elevations > 4200 m. Male body size continued to increase with increasing elevations. Both sexes showed an increased pattern of extremity length with elevation that does not conform to Allen’s rule. Limb length and tail length increased along the elevational gradients. In terms of color pattern, an abdominal black speckled area appears at elevations >4,200 m. This trait increases in size with increased elevation. Unlike most studies, our results indicated that annual sunshine hours corresponding to the activity period of the lizards could play an important role on the positive body size cline in environments at very high elevations > 4200 m

WANG, Y., ZENG, X., FANG, Z., LIU, Z., WU, G., PAPENFUSS, T.J. & R.J. MACEY (1996): A new species of the genus Phrynocephalus – Phrynocephalus zetangensis sp. nov. – Zoological Research, 17 (1): 27-29. (in Chinesisch)

WANG, Y.-z., ZENG, X.-m., WU, G.-f., LIU, Z.-j., FANG, Z.-I., PAPENFUSS, T.J. & R.J. MACEY (1996): A newly recorded species of Phrynocephalus from China. – Acta Zootaxonomica Sinica, 21 (1): 9. (in Chinesisch)
Mongolischer Krötenkopf

DUNAYEV, E.A. & N.A. POYARKOV (2010): Phylogeny, phylogeography and

Phrynocephalus versicolor STRAUCH, 1876

Mongolischer Krötenkopf

DUNAYEV, E.A. & N.A. POYARKOV (2010): Phylogeny, phylogeography and identification of Asian toad-headed agamas Phrynocephalus (superspecies versicolor). - Abstracts of the Second International Symposium on Agamid Lizards «DeAgamis2». - Current Studies in Herpetology, 10 (3/4): 142–143.

GOLUBEV, M.L. (1989): Phrynocephalus guttatus (GMEL. or Ph. versicolor str. (Reptilia, Agamidae) which one inhabits Kazakhstan? – Vestnik zoologii, Kiev, 23 (5): 38-45. (in russisch)

GOLUBEV, M.L. (1992): Variegated Toad Agama Phrynocephalus versicolor (Reptilia, Agamidae) of the Djungar Gate (East Kazakhstan) wit Notes on Systematics of the Species. – Vestnik Zoologii 1992 (2): 31-38. (in Russisch)

GOLUBEV, M.L. (1993): The variegated toad agama in Djungar Gate (Eastern Kazakhstan) with notes on certain systematic problems of Phrynocephalus versicolor str. (Reptilia: Agamidae). - Asiatic Herpetological Research, 5: 51-58. 9528

KIRMSE, W. & V. SINZ (1968): Freileben und Terrarienhaltung des Mongolischen Krötenkopfes, Phrynocephalus versicolor. - Aquarien Terrarien, Leipzig, 15 (12): 412-415. (1127)

KÖHLER, D. (1982): Begegnungen mit Krötenkopfagamen. – Aquarien Terrarien, Leipzig, 29 (9): 317. (00.171)

KUBYKIN, R.A. & Z.K. BRUSHKO (1990): Important representatives of rare species. Phrynocephalus versicolor. - In: Kovshar, A.F. (ed.): Rare animals of desert regions: problems of protecting Kazakhstan vertebrates genofond [sic]. Nauka Kazakhstan, Alma Ata: 217-229. (In Russisch).

QU, Y.F., GAO, J.F., MAO, L.X. & X. JI (2011): Sexual dimorphism and female reproduction in two sympatric toad-headed lizards Phrynocephalus frontalis and P. versicolor (Agamidae). – Anim. Biol., 61: 139-151.

SONG, S., LI, D., ZHANG, C., JIANG, K., ZHANG, D. & C. CHANG (2014): The complete mitochondrial genome of the color changeable toad-headed agama, Phrynocephalus versicolor (Reptilia, Squamata, Agamidae). – Mitochondrial DNA 2014. DOI: 10.3109/19401736.2014.933329.

The complete mitochondrial genome sequence of color changeable toad-headed agama, Phrynocephalus versicolor, was determined using polymerase chain reaction (PCR), longand- accurate PCR and directly sequencing by primer walking. The entire mitochondrial genome of P. versicolor was 16,429 bp in length, the accession was KJ749841 and the content of A, T, C, and G were 36.1%, 26.5%, 24.9% and 12.5%, respectively, which was similar to most vertebrate. The complete mitochondrial genome of P. versicolor contain 13 protein-coding genes, 2 rRNA genes, 23 tRNA genes, plus one control region and was similar to those of other Phrynocephalus sand lizards in gene arrangement and composition, except that tRNA-Phe and tRNA-Pro were exchanged and tRNA-Phe had two copies. The control region comprised three parts, one between tRNA-Thr and tRNA-Phe, a second between tRNA-Pro and tRNA-Phe, and a third between tRNA-Phe and 12S RNA. The complete mitochondrial genome of P. versicolor provided fundamental data for resolving phylogenetic relationship problems related to Agaimidae and genus Phrynocephalus.

STRAUCH, A. (1876): Description of Phrynocephalus versicolor and P. vlangalii. - 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). (in Russisch)

TANG, X., YUE, F., MA, M., WANG, N.B., HE, J. & Q. CHEN (2012): Effects of thermal and hydric conditions on egg incubation and hatchling phenotypes in two Phrynocephalus lizards. – As. Herpetol. Res., 3 (3): 184-191.

Using mtDNA sequence data and a phylogenetic approach, we examined the divergence and evolutionary history of the Toad-Headed Lizards of the Phrynocephalus versicolor species complex, which are currently assigned to three species, P. versicolor, Phrynocephalus frontalis, and Phrynocephalus przewalskii. Both parsimony and likelihood-based analyses resulted in three clades, which do not conform to the current species designation. The three species did not form mutually exclusive monophyletic groups. Instead, the populations from Tengger Desert, Ordos Highland, and further east, which include P. frontalis, P. przewalskii, and several populations of P. versicolor, formed a clade and we recommend recognizing them as a single species under the name P. przewalskii. The populations from the northwestern part of the complex’s distribution also formed a monophyletic group, for which we recommend retaining the name P. versicolor. Finally, the southwestern populations share a unique evolutionary history and should be recognized as a new species, Phrynocephalus sp. The resulting phylogeny suggests that the species diversity and distribution patterns of P. przewalskii, P. versicolor, and Phrynocephalus sp. can be better explained by vicariance, rather than the traditional west-to-east dispersal hypothesis.

WANG, Y. & H. WANG (1993): Geographic variation and diversity in three species of Phrynocephalus in the Tengger Desert, Western China. - Asiatic Herpetological Research 5: 65-73.


Phrynocephalus versicolor versicolor STRAUCH,1876


Phrynocephalus versicolor doriai BEDRIAGA, 1909


Phrynocephalus versicolor hispidus BEDRIAGA,1909


Phrynocephalus versicolor kulagini BEDRIAGA, 1909

Phrynocephalus vlangalii STRAUCH, 1876

Ching Hai Toadhead Agama

ANONYMOUS (2017): Krötenkopfagamen und eine tiergeografische Regel. – TERRARIA/elaphe, 4/2017: 71.

GUO, X., LIU, L. & Y. WANG (2012): Phylogeography of the Phrynocephalus vlangalii Species Complex in the Upper Reaches of the Yellow River Inferred from mtDNA ND4-tRNALEU Segments. -  Asian Herpetological Research 3 (1): 52-68.

The Ching Hai Toad-headed Agama (Phrynocephalus vlangalii) complex is a small toad-headed viviparous lizard that is endemic to the Qinghai-Tibetan Plateau. A fragment of mtDNA ND4-tRNALEU from 189 samples in 26 populations was used to infer the phylogeographic history of this species complex in the upper reaches of the Yellow River. Phylogenetic analyses revealed that P. vlangalii and another proposed species (P. putjatia) do not form a monophyletic mtDNA clade, which in contrast with a previous study, includes P. theobaldi and P. forsythii. Lineage diversification occurred in the Middle Pleistocene for P. vlangali (ca. 0.95 Ma) and in the Early Pleistocene for P. putjatia (ca. 1.78 Ma). The uplift of the A’nyemaqen Mountains and glaciations since the mid-late Pleistocene, especially during the Kunlun Glaciation, are considered to have promoted the allopartric divergence of P. vlangalii. The diversification of P. putjatia may be triggered by the tectonic movement in the Huangshui River valley during the C phase of Qingzang Movement. Subsequently, the glacial climate throughout the Pleistocene may have continued to impede the gene flow of P. putjatia, eventually resulting in the genetic divergence of P. putjatia in the allopatric regions. Demographic estimates revealed weak population expansion in one lineage of P. vlangalii (A2, the Qaidam Basin lineage) and one lineage of P. putjatia (B2, the north Qinghai Lake lineage) after approximately 42 000 years before present. However, constant population size through time was inferred for two lineages (A1 and B1), the source of Yellow River lineage of P. vlangalii and the southeast of Qinghai Lake lineage of P. putjatia, possibly due to stable populations persisting in areas unaffected by glacial advances. Our results also suggest: 1) at least four differentiated lineages of P. vlangalii complex may have evolved allopatrically in different regions during the Pleistocene glaciation events; 2) in support of several recent studies, P. putjatia is a valid species, having a more wide distribution than previously considered; and 3) a hypothesis referring to P. v. hongyuanensis, inhabiting in the source region of the Yellow River, being synonymous with P. v. pylozwi is supported.

HUANG, Y., WU, X.; LI, Y.H. & W.B.LIAO (2014): Global warming, body size and conservation in a Qinghai-Tibet Plateau lizard. The Herpetological Journal 24 (3): 161-166.

Global mean temperatures have increased by 0.3–0.6°C since the late 19th century, affecting the physiology, distributions, phenology and adaptations of plants and animals. In the Qinghai-Tibet Plateau, average annual temperatures increased by an average of 0.25°C per decade from the 1970s to the 1990s, and by an average of 0.34°C per decade thereafter. Using museum collections from the 1950s to the 2000s and published references, we tested the hypothesis that body size of the toad-headed lizard Phrynocephalus vlangalii in the Qinghai-Tibet Plateau declined between 1954 and 2008 as a response to global warming. However, body size of males and females did not vary significantly between 1954 and 2008, probably due to the reciprocity between higher food availability and earlier age at sexual maturity. We suggest that human activity might result in declining population sizes in the future despite the lack of an apparent current response to changing climates.

JIN, Y.-T., BROWN, R.P. & N. LIU (2008): Cladogenesis and phylogeography of the lizard Phrynocephalus vlangalii (Agamidae) on the Tibetan plateau. - Molecular Ecology 17: 1971–1982.

Phrynocephalus vlangalii is restricted to dry sand or Gobi desert highlands between major mountain ranges in the Qinghai (Tibetan) Plateau. Mitochondrial DNA (mtDNA) sequence (partial ND2, tRNATrp and partial tRNAAla) was obtained from 293 Phrynocephalus sampled from 34 sites across the plateau. Partitioned Bayesian and maximum parsimony phylogenetic analyses revealed that P. vlangalii and two other proposed species (P. erythrus and P. putjatia) together form a monophyletic mtDNA clade which, in contrast with previous studies, does not include P. theobaldi and P. zetangensis. The main P. vlangalli clade comprises seven well-supported lineages that correspond to distinct geographical areas with little or no overlap, and share a most recent common ancestor at 5.06 ± 0.68 million years ago (mya). This is much older than intraspecific lineages in other Tibetan animal groups. Analyses of molecular variance indicated that most of the observed genetic variation occurred among populations/regions implying long-term interruption of maternal gene flow. A combined approach based on tests of population expansion, estimation of node dates, and significance tests on clade areas indicated that phylogeographical structuring has been primarily shaped by three main periods of plateau uplift during the Pliocene and Pleistocene, specifically 3.4 mya, 2.5 mya and 1.7 mya. These periods corresponded to the appearance of several mountain ranges that formed physical barriers between lineages. Populations from the Qaidam Basin are shown to have undergone major demographic and range expansions in the early Pleistocene, consistent with colonization of areas previously covered by the huge Qaidam palaeolake, which desiccated at this time. The study represents one of the most detailed phylogeographical analyses of the Qinghai Plateau to date and shows how geological events have shaped current patterns of diversity.

JIN, Y., LIU, N. & J. LI (2007): Elevational variation in body size of Phrynocephalus vlangalii in the North Qinghai-Xizang (Tibetan) Plateau. -Belg. J. Zool. 137 (2): 197-202.

We examined elevational and environmental aspects of body size variation in the Qinghai toad-headed lizard, Phrynocephalus vlangalii, using principal component analysis (PCA) of 9 morphological traits taken from 565 lizards from 17 populations. The first principal component (PC1) accounted for 67% of the size variation in males and 62% in females. For both males and females, PC1 decreased with increasing elevation. When analyzed in relation with respect to environmental variables, body size showed positive relationship with temperature, air pressure, and activity season length, but showed weaker or inconsistent relationships with rainfall and humidity. The described pattern is the converse of Bergmann’s rule for this lizard species and suggests that this body size pattern is driven by temperature, air pressure or length of the activity season.

LI, J., ZHOU, R. & N. LIU (2014): Life-history variation among three populations of the toad-headed lizard Phrynocephalus vlangalii along an elevation gradient on the northeastern Tibetan Plateau. – Herp. J., 24 (1): 17-23.

Elevation and related environmental factors is a main factor for life-history variation in ectothermic species. To test if altitude can explain variation in life-history traits in the toad-headed lizard Phrynocephalus vlangalii, we sampled gravid females from three localities (Delingha: elevation 2910m, Daotanghe: 3367m; Maduo: 4257m) and measured parturition time, snout-vent length (SVL) at sexual maturity, female body size and postpartum body condition, litter size, litter mass, relative litter mass (RLM), size and mass of newborn offspring. Females gave birth between 25 July and 28 August, and females from lower elevations were earlier than those from higher elevations. Female SVL at sexual maturity and mean SVL, litter size, litter mass, relative litter mass and offspring size varied among populations, whereas female post-parturition body mass and offspring mass remained invariant. Females from the highest elevation had a greater SVL at sexual maturity, mean SVL and abdominal length. Both litter size and litter mass were positively correlated with female SVL. Females from the higher elevation localities (Maduo and Daotanghe) had a lower RLM than those from the lowest elevation (Delingha). Furthermore, females from the highest elevation produced fewer and larger (SVL but not mass) offspring than those from the other localities. Trade-offs between offspring size and number were detected in Delingha and Daotanghe, respectively, but not in Maduo. At a given level of litter size, offspring size was similar between Delingha and Daotanghe.

LI, W., LIANG, S., WANG, H., XIN, Y., LU, S., TANG, X. & Q. CHEN (2016): The Effects of Chronic Hypoxia on Thermoregulation and Metabolism in Phrynocephalus vlangalii. - Asian Herpetological Research 7 (2): 103-111.Phrynocephalus vlangalii are widely distributed on Tibetan plateau spanning diverse altitudes and habitats. In the present study, P. vlangalii were exposed to 8% oxygen concentration in a hypoxic chamber for 6 weeks. Then the body temperature (Tb), standard metabolic rate (SMR), heart rate and metabolic enzyme activities of the lizards were measured at 20°C and 30°C. The results indicated that hypoxia exposure decreased Tb, SMR and heart rate. Lactate dehydrogenase (LDH) activity of 8% O2 group became significant elevated in liver and skeletal muscle compared with control group at 20°C, but descended significantly in heart. Using electrophoresis we found that LDH contains five isozymes (LDH1, LDH2, LDH3, LDH4 and LDH5) and are expressed specifically in liver, skeletal muscle and heart. Citrate synthase (CS) activity in the liver also decreased at 20°C and 30°C. No significant difference of CS activity was found between the two groups in skeletal muscle and heart.

LIU, L., GUO, X. & Y. WANG (2010): Phylogeography of Phrynocephalus vlangalii complex on the upper reaches of the Yellow River inferred from mtDNA ND4-tRNAleu sequences. - Abstracts of the Second International Symposium on Agamid Lizards «DeAgamis2». - Current Studies in Herpetology, 10 (3/4): 147–148.

LU, H.-L., JIANG, C.-Q. & X. JI (2015): Locomotor costs of pregnancy in a viviparous toad-headed lizard, Phrynocephalus vlangalii (Agamidae). – Herp. J., 25 (3): 149-154.

Locomotor impairment during pregnancy can be attributed to physical burden or physiological changes associated with pregnancy. However, the degree to which physical and physiological changes affect reproductive costs likely varies between species. Here, we used the Qinghai toad-headed lizard (Phrynocephalus vlangalii) as a model to assess locomotor costs during pregnancy and the relative impact of physical and physiological effects in pregnant viviparous lizards. The locomotor costs of pregnancy were pronounced: sprint speed decreased gradually throughout pregnancy, reached a minimum at parturition and increased slowly thereafter. The reduced speed in pregnant females was not related to relative litter mass. Compared with the locomotion of non-reproductive females or males, pregnant females exhibited lower speeds and shorter stride lengths. These results suggest that, despite having a physical effect on locomotor performance, physiological changes associated with pregnancy likely play a major role in locomotor impairment in pregnant P. vlangalii.

NOBLE, D.W.A., QI, Y. & J. FU (2010): Species delineation using Bayesian model-based assignment tests: a case study using Chinese toad-headed agamas (genus Phrynocephalus). - BMC Evolutionary Biology 10: 197.

Background: Species are fundamental units in biology, yet much debate exists surrounding how we should delineate species in nature. Species discovery now requires the use of separate, corroborating datasets to quantify independently evolving lineages and test species criteria. However, the complexity of the speciation process has ushered in a need to infuse studies with new tools capable of aiding in species delineation. We suggest that modelbased assignment tests are one such tool. This method circumvents constraints with traditional population genetic analyses and provides a novel means of describing cryptic and complex diversity in natural systems. Using toadheaded agamas of the Phrynocephalus vlangalii complex as a case study, we apply model-based assignment tests to microsatellite DNA data to test whether P. putjatia, a controversial species that closely resembles P. vlangalii morphologically, represents a valid species. Mitochondrial DNA and geographic data are also included to corroborate the assignment test results.
Results: Assignment tests revealed two distinct nuclear DNA clusters with 95% (230/243) of the individuals being assigned to one of the clusters with > 90% probability. The nuclear genomes of the two clusters remained distinct in sympatry, particularly at three syntopic sites, suggesting the existence of reproductive isolation between the identified clusters. In addition, a mitochondrial ND2 gene tree revealed two deeply diverged clades, which were largely congruent with the two nuclear DNA clusters, with a few exceptions. Historical mitochondrial introgression events between the two groups might explain the disagreement between the mitochondrial and nuclear DNA data. The nuclear DNA clusters and mitochondrial clades corresponded nicely to the hypothesized distributions of P. vlangalii and P. putjatia.
Conclusions: These results demonstrate that assignment tests based on microsatellite DNA data can be powerful tools for distinguishing closely related species and support the validity of P. putjatia. Assignment tests have the potential to play a significant role in elucidating biodiversity in the era of DNA data. Nonetheless, important limitations do exist and multiple independent datasets should be used to corroborate results from assignment programs.

QI, Y., LI, S., SUO, L., LI, H. & Y. WANG (2011): An Ethogram of the Toad-headed Lizard Phrynocephalus vlangalii during the Breeding Season. - Asian Herpetological Research 2 (2): 110-116.

In this study, we intended to construct an ethogram of the toad-headed lizard Phrynocephalus vlangalii by classifying and describing its behaviors using the Posture-act-environment (PAE) method. Through direct field observation and video recording, we obtained eight postures, sixty-two acts and ten environments, and thus we formed an ethogram consisting of eighty-three behaviors encoded with PAE. A complete ethogram for a species could provide systematic behavioral information which helps to understand the connections among related behaviors and provides background information for specific behavioral study, hence the present ethogram will be greatly useful for future behavioral studies on P. vlangalii as well as other Phrynocephalus species.

SHAO, H., FAN, L., XU, X.J., XU, W.Q., LIU, B.F., WANG, J.L., LIU, N.F. & S.T. ZHAO (2012): Characterization of adult neurogenesis in lizard Phrynocephalus vlangalii (Agamidae: Reptilia). – Ital. J. Zool., 79 (4): 547-558.

Adult neurogenesis, a process of giving rise to neurons and glia from progenitors residing in restricted regions of the adult central nervous system (CNS) throughout life, varies considerably across species. The role of adult neurogenesis in reptiles, the amniotic vertebrates that possess spontaneous regenerative capacity is less known. In the present study, we used Phrynocephalus vlangalii – a typical reptile in northwest of China to investigate the cell proliferation and general pattern of glial fibrillary acidic protein (GFAP)-immunoreactivity in the brain of lizard. Compared with mammals, the brains of P. vlangaliiowned have simpler cytoarchitectonical patterns but a more widespread neurogenesis. Bromodeoxyuridine (BrdU)-labeled cells were mostly distributed in the ventricular zone of lateral ventricle with cell proliferation highest in the region referred to as anterior olfactory nucleus (AON). In addition, GFAP-immunostaining demonstrated differences in element, abundance and distribution. GFAP-positive cells were fundamentally represented by radial glial structures in telencephalon, diencephalon and mesencephalon. Furthermore, an intriguing finding was the emergence of GFAP-positive, star-shaped cells in the mesencephalon. We conclude that the cell proliferation and corresponding functions of glial lineage cells in lizards are closely implicated with phylogenetic development. Hence, reptiles provide excellent access for extracting core mechanisms of adult neurogenesis, which lays a good foundation to study the effect of different stimuli and exposure on the adult neurogenesis of reptiles.

STRAUCH, A. (1876): Description of Phrynocephalus versicolor and P. vlangalii. - 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). (in Russisch)

WANG, Y.Z. & Y.M. JIANG (1992): Study on the taxonomy of Phrynocephalus vlangalii hongyuanensis: a new species of the genus Phrynocephalus (Lacertilia: Agamidae). – In: Jiang, Y.M. (ed.): A collection of papers on Herpetology. Sichuan Science and Technology Press, Chendu, China. Pp. 110-115.

ZHAN, A. & J. FU (2009): Microsatellite DNA markers for three toad-headed lizard species (Phrynocephalus vlangalii, P. przewalskii and P. guttatus). – Molec. Ecol. Res., 9 (2): 535-538.

According to the distribution of Phrynocephalus vlangalii hongyuanensis in Zoige Wetland，three geographic units: Zoige Xiaman (XM)，Hongyuan (HY)，both in Sichuan Province and Maqu (MQ) in Gansu Province were defined. We used molecular methods to reveal these unit’s genetic variation and diversity. A 785bp fragment of the mtDNA ND4-tRNAleu was determined from 72 samp1es in seven populations of P. vlangalii hongyuanensis. Seven variable nucleotide sites and nine haplotypes were identified in the 785bp fragments. As a whole，the haplotype diversity was high (0.806±0.024)，but the nucleotide diversity was low (0.00231±0.00016). In a single population，MQa，MQb and XMb had very low genetic diversities，and XMc had a much higher one. The Kimura 2-parameter distances among all the populations were small (0.001-0.005)，and the distance between MQa and XMa was the greatest. Analysis of molecular variance (AMOVA) showed that the three units were distinctly different (P<0.01)，and 62.61% of the total genetic diversity was attributable to variation among units. There were 3 haplotypes shared among XM and HY，and no geographic clustering was observed except MQ from the TCS network. The results from the mismatch distribution analysis and Fu’s Fs test (Fs=－2.21937) implied that there might be a recent population expansion in the XM unit，and this may be the reason why XM had a high haplotype diversity but a low nucleotide diversity. We estimate that the MQ and XMb have lower diversities because of some very recent geographic events，such as the formation of the Yellow river’s upriver and the Zoige Wetland. Although they are distinctly different，not enough time has passed for them to have diverged a great genetic distance.

zurück / back