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© Rune Midtgaard


Taxonomy of the family Agamidae
Bibliography of the genus Phrynocephalus
Biodiversity of the family Agamidae








Genus
Phrynocephalus

Toad-headed Agamas, Toad Agamas

Krötenkopfagamen

Tudsehovedagamer

1825 Phrynocephalus Kaup (type species: Lacerta caudivolvula Pallas 1814)
1831 Megalochilus Eichwald (type species: Lacerta aurita Gmelin 1789; syn. Wermuth 1967; Szczerbak 1994; Barabanov & Ananjeva 2007)
1843 Helioscopus Fitzinger (type species: Phrynocephalus helioscopus Wagler; syn. Wermuth 1967)
1843 Phrynosaurus Fitzinger (type species: Phrynocephalus olivieri Dumeril & Bibron 1837; syn. Wermuth 1967)
1843 Saccostoma Fitzinger (type species: Phrynocephalus auritus Fitzinger [=Lacerta aurita Gmelin 1789]; syn. Wermuth 1967)
2007 Oreosaura Barabanov & Ananjeva (type species: Phrynocephalus vlangalii Strauch 1876)
Contents: 44 species, of which 17 (38.6%) are endemic.
Endemism: 0% 100%
Remarks: Previously included Bufoniceps laungwalaensis. Taxonomy of Phrynocephalus is still far from clear. Status of some forms is a matter of controversy, and various authors have disagreed regarding validity of forms and synonymies. The species list below follows Barabanov & Ananjeva (2007), with some adjustments based on subsequent works.
Distribution: Middle East, C. and E. Asia
Reported from: Afghanistan, Armenia, Azerbaijan, China (Gansu, Hebei, Inner Mongolia, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Tibet, Xinjiang), India (Jammu & Kashmir), Iran, Iraq, Jordan, Kazakhstan, Kuwait, Kyrgyzstan, Mongolia, Nepal, Oman, Pakistan, Qatar, Russia, Saudi Arabia, Tajikistan, Turkey, Turkmenistan, United Arab Emirates, Uzbekistan, Yemen.


Phrynocephalus sp.
© Rune Midtgaard

Phrynocephalus ahvazicus

Ahvaz Toad-headed Agama

Ahvaz-Krötenkopfagame

Ahvaz-tudsehovedagam

2014 Phrynocephalus ahvazicus Melnikov, Melnikova, Nazarov, Rajabizadeh, Al-Johany, Amr & Ananjeva

Remarks: Previously confused with arabicus (see remarks under that species). Records of arabicus from Kuwait (e.g., Al-Sirhan & Brown 2010), southern Iraq (Mohammed, Rhadi, Rastegar-Pouyani, Rastegar-Pouyani & Yousefkhani 2017), and southwestern Iran (e.g., Kamali 2020) are doubtful and provisionally assigned to ahvazicus, the species of the arabicus group occurring most closely to this region.
Distribution: Iran, Iraq, Kuwait.


Phrynocephalus ananjevae

Zagros Toad-headed Agama

Zagros-Krötenkopfagame

Zagros-tudsehovedagam

2013 Phrynocephalus ananjevae Melnikov, Melnikova, Nazarov & Rajabizadeh

Distribution: Iran.


Phrynocephalus arabicus

Arabian Toad-headed Agama

Arabische Krötenkopfagame

Arabisk Tudsehovedagam

1894 Phrynocephalus arabicus Anderson

Remarks: Now restricted to southern Arabia. Previously included macropeltis, nejdensis, and the population now assigned to ahvazicus. Records of arabicus from Kuwait (e.g., Al-Sirhan & Brown 2010), southern Iraq (Mohammed, Rhadi, Rastegar-Pouyani, Rastegar-Pouyani & Yousefkhani 2017), and southwestern Iran (e.g., Kamali 2020) are doubtful and provisionally assigned to ahvazicus, the species of the arabicus group occurring most closely to this region. Nevertheless, Kamali (2020) listed both ahvazicus and arabicus from Iran. A record from Qatar (Gardner 2013) is provisionally assigned to macropeltis, the species of the arabicus group occurring most closely to Qatar.
Distribution: Oman, Saudi Arabia, Yemen.


Phrynocephalus axillaris

East Turkestan Toad-headed Agama

Ostturkestanische Krötenkopfagame

Østturkestansk Tudsehovedagam

1875 Phrynocephalus axillaris Blanford
1906 Phrynocephalus axillaris var. ericae Bedriaga "1905" (Barabanov & Ananjeva 2007)
1906 Phrynocephalus axillaris var. idae Bedriaga "1905" (Barabanov & Ananjeva 2007)
1906 Phrynocephalus axillaris var. murielis Bedriaga "1905" (Barabanov & Ananjeva 2007)
1906 Phrynocephalus koslowi Bedriaga "1905" (Barabanov & Ananjeva 2007)
1907 Phrynocephalus axillaris var. klemenzi Bedriaga (Barabanov & Ananjeva 2007)
1907 Phrynocephalus axillaris var. maculata Bedriaga [not Phrynocephalus maculata Anderson 1872] (Barabanov & Ananjeva 2007)
1910 Phrynocephalus Ludovici Mocquard (Barabanov & Ananjeva 2007)

Remarks: Previously included nasatus.
Distribution: China (Gansu, Xinjiang).


Phrynocephalus bannikovi

Krasnovodsky Toad-headed Agama

Krasnovodsky-Krötenkopfagame

Krasnovodsky-tudsehovedagam

1976 Phrynocephalus reticulatus bannikovi Darevsky, Rustamov & Shammakov
Phrynocephalus bannikovi Lichnova 1992
Phrynocephalus ocellatus bannikovi Szczerbak 1994

Remarks: Previously treated as a subspecies of reticulatus. Lichnova (1992) raised bannikovi to status as a separate species. Validity confirmed by Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss (2018).
Distribution: Turkmenistan.





Phrynocephalus clarkorum

Afghan Toad-headed Agama

Afghanische Krötenkopfagame

Afghansk Tudsehovedagam

1967 Phrynocephalus clarkorum Anderson & Leviton

Distribution: Afghanistan, Pakistan.


Phrynocephalus erythrurus

Northwest Tibetan Toad-headed Agama

Nordwesttibetanische Krötenkopfagame

Nordvesttibetansk Tudsehovedagam

1909 Phrynocephalus erythrurus Zugmayer
1909 Phrynocephalus vlangalii parva Bedriaga
Phrynocephalus parvus Tsarevsky 1927
Phrynocephalus erythrurus parva Jin & Liu 2010

Remarks: Regarded as a synonym of theobaldi by Barabanov & Ananjeva (2007), but as a valid species by Jin & Liu (2010). The taxon parva was previously regarded as a synonym of vlangalii (Barabanov & Ananjeva 2007). This species occupies the highest regions of any reptile on earth (Jin & Liu 2010).
Distribution: China (Qinghai, Tibet, Xinjiang).


Phrynocephalus euptilopus

Darband Toad-headed Agama, Sand Toad Agama, (Spotted Toad-headed Agama)

Darband-Krötenkopfagame

Darband-tudsehovedagam

1897 Phrynocephalus euptilopus Alcock & Finn "1896"

Remarks: Although reported from India by several authors (e.g., Daniel 1983; Tikader & Sharma 1992; Das 1997; Daniel 2002), there seem to be no verified records from the country (Das 1996; Barabanov & Ananjeva 2007; Venugopal 2010).
Distribution: Afghanistan, Pakistan.


Phrynocephalus forsythii

Yarkand Toad-headed Agama

Yarkand-Krötenkopfagame

Yarkand-tudsehovedagam

1872 Phrynocephalus forsythii Anderson
Phrynocephalus theobaldi forsythii Boulenger 1885

Distribution: China (Xinjiang).


Phrynocephalus frontalis

Shanxi Toad-headed Agama

Shanxi-Krötenkopfagame

Shanxi-tudsehovedagam

1876 Phrynocephalus frontalis Strauch

Remarks: Previously regarded as a synonym of przewalskii (e.g., Barabanov & Ananjeva 2007). Retained herein as a valid species, following Melville, Hale, Mantziou, Ananjeva, Milto & Clemann (2009) and Dunayev, Solovyeva & Poyarkov (2021).
Distribution: China (Beijing, Gansu, Hebei, Inner Mongolia, Ningxia, Shaanxi, Shanxi).





Phrynocephalus golubewii

Bami Toad-headed Agama, (Golubev's Toad-headed Agama)

Bami-Krötenkopfagame

Bami-tudsehovedagam

1990 Phrynocephalus golubewii Shenbrot & Semyonov

Distribution: Turkmenistan.


Phrynocephalus guinanensis

Guinan Toad-headed Agama

Guinan-Krötenkopfagame

Guinan-tudsehovedagam

2009 Phrynocephalus guinanensis Ji, Wang & Wang

Remarks: Jin & Brown (2019) suggested that guinanensis might be a synonym of of putjatai. Not listed for China by Wang, Ren, Chen, Lyu, Guo, Jiang, Chen, Li, Guo, Wang & Che (2020).
Distribution: China (Qinghai).


Phrynocephalus guttatus

Spotted Toad-headed Agama, Tail Roller

Gefleckter Krötenkopf, Schwanzroller

Plettet Tudsehovedagam

1789 Lacerta guttata Gmelin
Agama guttata Daudin 1802
Phrynocephalus guttatus Kaup 1827
1814 Lacerta caudivolvula Pallas (Barabanov & Ananjeva 2007)
Agama caudivolvula Lichtenstein 1823
Phrynocephalus caudivolvulus Fitzinger 1826
1831 Phrynocephalus melanurus Eichwald (Barabanov & Ananjeva 2007)
Phrynocephalus guttatus melanurus Melville, Hale, Mantziou, Ananjeva, Milto & Clemann 2009
1831 Phrynocephalus Pallasii Gray (Barabanov & Ananjeva 2007)
1905 Phrynocephalus alpherakii Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)
Phrynocephalus caudivolvulus alpherakii Tsarevsky 1926
Phrynocephalus guttatus alpherakii Golubev 1992
1905 Phrynocephalus caudivolvolus var. incerta Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)
Phrynocephalus guttatus incerta Melville, Hale, Mantziou, Ananjeva, Milto & Clemann 2009
1905 Phrynocephalus kuschakewitschi Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)
Phrynocephalus caudivolvulus kuschakewitschi Tsarevsky 1926
Phrynocephalus guttatus kuschakewitschi Terentjev & Chernov 1949
1905 Phrynocephalus suschkini Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)
1906 Phrynocephalus bergi Elpatjevsky "1907" (Barabanov & Ananjeva 2007)
1906 Phrynocephalus haeckeli Bedriaga "1905" [nomen nudum] (Barabanov & Ananjeva 2007)
1906 Phrynocephalus isseli Bedriaga "1905" [nomen nudum] (Barabanov & Ananjeva 2007)
1906 Phrynocephalus salenskii Bedriaga "1905" [nomen nudum] (Barabanov & Ananjeva 2007)
1907 Phrynocephalus acutirostris Bedriaga (Barabanov & Ananjeva 2007)
1907 Phrynocephalus Arcellazzii Bedriaga (Barabanov & Ananjeva 2007)
1907 Phrynocephalus Haeckeli Bedriaga (Barabanov & Ananjeva 2007)
1907 Phrynocephalus Isseli Bedriaga (Barabanov & Ananjeva 2007)
1907 Phrynocephalus Salenskyi Bedriaga (Barabanov & Ananjeva 2007)
Phrynocephalus guttatus salenskyi Szczerbak 2003
1909 Phrynocephalus Grum-Grzimailoi [grumgrzimaloi] Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus helioscopus levis Kascenko (Barabanov & Ananjeva 2007)
Phrynocephalus helioscopus laevis Nikolski 1915 [substitute name for Phrynocephalus helioscopus var. levis Kascenko 1909]
1913 Phrynocephalus caudivolvolus var. moltschanovi Nikolski (Barabanov & Ananjeva 2007)
Phrynocephalus moltschanovi Semenov & Shenbrot 1982
1926 Phrynocephalus bedrjagai Zarevsky (Barabanov & Ananjeva 2007)
1979 Phrynocephalus albo-lineatus Zhao (Barabanov & Ananjeva 2007)
1983 Phrynocephalus guttatus kalmykus Badmaeva & Shcherbak (Barabanov & Ananjeva 2007)
1987 Phrynocephalus versicolor paraskiwi Semenov, Brushko, Shenbrot & Kubykin (Barabanov & Ananjeva 2007)
1995 Phrynocephalus guttatus salsatus Gorelov, Dunayev & Kotenko in Golubev, Gorelov, Dunayev & Kotenko (Barabanov & Ananjeva 2007)

Other common names:
moltschanovi: Molchanov's Toad-headed Agama
Remarks: See under reticulatus. Ananjeva, Guo & Wang (2011) listed albolineatus as a separate species. Liu, Song, Li & Shi (2012) treated grumgrzimaloi as a separate species. Melville, Hale, Mantziou, Ananjeva, Milto & Clemann (2009) suggested incerta, kuschakewitschi, melanurus, and alpherakii deserve recognition as separate species, but this was regarded as premature by Sindaco, Venchi & Grieco (2013).
Distribution: China (Xinjiang), Kazakhstan, Russia, Turkmenistan, Uzbekistan.


Phrynocephalus helioscopus

Central Asian Sunwatcher

Zentralasiatische Sonnengucker

Centralasiatisk Solkigger

1771 Lacerta helioscopa Pallas
Agama helioscopus Lichtenstein 1823
Phrynocephalus helioscopus Kaup 1827
1789 Lacerta uralensis Gmelin (Barabanov & Ananjeva 2007)
1831 Phrynocephalus nigricans Eichwald (Barabanov & Ananjeva 2007)
1831 Phrynocephalus varius Eichwald (Barabanov & Ananjeva 2007)
Phrynocephalus helioscopus varius Barabanov & Ananjeva 2007
1841 Lacerta gibba Eichwald [unavailable name] (Barabanov & Ananjeva 2007)
1907 Phrynocephalus helioscopus var. Cameranoi Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus helioscopus saposhnikovi Kascenko (Barabanov & Ananjeva 2007)
1912 Phrynocephalus helioscopus orientalis Bedriaga (Barabanov & Ananjeva 2007)
1981 Phrynocephalus helioscopus saidalievi Sattorov (Barabanov & Ananjeva 2007)
Phrynocephalus saidalievi Eremchenko & Panfilov 1999
2012 Phrynocephalus helioscopus meridionalis Solovyeva, Dunayev & Poyarkov
2012 Phrynocephalus helioscopus sergeevi Solovyeva, Dunayev & Poyarkov

Remarks: Previously included horvathi, persicus, and turcomanus.
Distribution: China (Inner Mongolia, Xinjiang), Iran, Kazakhstan, Kyrgyzstan, Mongolia, Russia, Tajikistan, Turkmenistan, Uzbekistan.


Phrynocephalus hispidus

Dzungarian Toad-headed Agama, (Spotted Toad-headed Agama)

Dsungarischer Krötenkopfagame

Dzungarisk Tudsehovedagam

1909 Phrynocephalus versicolor var. hispida Bedriaga
Phrynocephalus hispidus Buehler, Zoljargal, Purvee, Munkhbayar, Munkhbaatar, Batsaikhan, Ananjeva, Orlov, Papenfuss, Roldán-Pina, Douchindorj, Grismer, Oaks, Brown & Grismer 2021

Remarks: Previously regarded as a synonym of versicolor (e.g., Barabanov & Ananjeva 2007). Regarded as a separate species herein, following Buehler, Zoljargal, Purvee, Munkhbayar, Munkhbaatar, Batsaikhan, Ananjeva, Orlov, Papenfuss, Roldán-Pina, Douchindorj, Grismer, Oaks, Brown & Grismer (2021) and Dunayev, Solovyeva & Poyarkov (2021).
Distribution: China (Gansu, Xinjiang), Mongolia.





Phrynocephalus hongyuanensis

Sichuan Toad-headed Agama

Sichuan-Krötenkopfagame

Sichuan-tudsehovedagam

1980 Phrynocephalus vlangalii hongyuanensis Zhao, Jiang & Huang in Jiang, Huang & Zhao
Phrynocephalus hongyuanensis Wang & Jiang 1992

Remarks: Previously treated as a subspecies of vlangalii. Wang & Jiang (1992) raised hongyuanensis to status as a separate species. Validity confirmed by Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss (2018). Not listed for China by Wang, Ren, Chen, Lyu, Guo, Jiang, Chen, Li, Guo, Wang & Che (2020).
Distribution: China (Sichuan).





Phrynocephalus horvathi

Turkish Toad-headed Agama, (Horvath's Toad-headed Agama)

Türkische Krötenkopfagame

Tyrkisk Tudsehovedagam

1894 Phrynocephalus helioscopus var. horvathi Mehely
Phrynocephalus horvathi Melnikov, Melnikova, Nazarov & Rajabizadeh 2013

Remarks: Previously regarded as a subspecies or a synonym of helioscopus (e.g., Mehely 1894) or of persicus (e.g., Anderson 1999; Barabanov & Ananjeva 2007). Arakelyan, Danielyan, Corti, Sindaco & Leviton (2011) suggested that horvathi deserved recognition as a separate species. It was formally revalidated by Melnikov, Melnikova, Nazarov & Rajabizadeh (2013).
Distribution: Armenia, Azerbaijan, Iran, Turkey.


Phrynocephalus interscapularis

Rose-shouldered Toad-headed Agama, Spiny Toad-headed Agama

Rosaschulter-Krötenkopfagame, Stachelkopf

Rosenskuldret Tudsehovedagam

1856 Phrynocephalus interscapularis Lichtenstein

Remarks: Previously included sogdianus. Listed for Iran by Kamali & Anderson (2015) and (Nasrabadi, Rastegar-Pouyani, Rastegar-Pouyani & Gharzi 2017), but omitted(?) by Kamali (2020).
Distribution: Afghanistan, Iran, Kazakhstan, Turkmenistan, Uzbekistan.


Phrynocephalus kulagini

Tuva Toad-headed Agama

Tuva-Krötenkopfagame

Tuva-tudsehovedagam

1909 Phrynocephalus versicolor var. Kulagini Bedriaga
Phrynocephalus kulagini Buehler, Zoljargal, Purvee, Munkhbayar, Munkhbaatar, Batsaikhan, Ananjeva, Orlov, Papenfuss, Roldán-Pina, Douchindorj, Grismer, Oaks, Brown & Grismer 2021

Remarks: Previously regarded as a synonym of versicolor (e.g., Barabanov & Ananjeva 2007). Regarded as a separate species herein, following Buehler, Zoljargal, Purvee, Munkhbayar, Munkhbaatar, Batsaikhan, Ananjeva, Orlov, Papenfuss, Roldán-Pina, Douchindorj, Grismer, Oaks, Brown & Grismer (2021) and Dunayev, Solovyeva & Poyarkov (2021).
Distribution: Mongolia, Russia.





Phrynocephalus longicaudatus

Long-tailed Toad-headed Agama

Langschwanz-Krötenkopfagame

Langhalet Tudsehovedagam

1957 Phrynocephalus maculatus longicaudatus Haas
Phrynocephalus longicaudatus Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss 2018

Remarks: Not mentioned for Saudi Arabia by Aloufi, Amr, Baker & Hamidan (2019).
Distribution: Iraq, Jordan, Kuwait, Oman, Saudi Arabia, United Arab Emirates, Yemen.





Phrynocephalus lutensis

Lut Toad-headed Agama

Lut-Krötenkopfagame

Lut-tudsehovedagam

2015 Phrynocephalus lutensis Kamali & Anderson

Distribution: Iran.


Phrynocephalus luteoguttatus

Yellow-speckled Toad-headed Agama, Spiny-headed Toad-headed Agama

Gelbfleckiger Krötenkopfagame

Gulprikket Tudsehovedagam

1887 Phrynocephalus luteoguttatus Boulenger

Remarks: Although reported from India by Das (1997), there are no verified records from the country (Venugopal 2010).
Distribution: Afghanistan, Pakistan.


Phrynocephalus macropeltis

Dhahran Toad-headed Agama

Dhahran-Krötenkopfagame

Dhahran-tudsehovedagam

1957 Phrynocephalus nejdensis macropeltis Haas
Phrynocephalus macropeltis Melnikov, Melnikova, Nazarov, Rajabizadeh, Al-Johany, Amr & Ananjeva 2014

Remarks: Previously regarded as a synonym of arabicus (e.g., Barabanov & Ananjeva 2007). Revalidated as a separate species by Melnikov, Melnikova, Nazarov, Rajabizadeh, Al-Johany, Amr & Ananjeva (2014). A record from Qatar of arabicus (Gardner 2013) is provisionally assigned to macropeltis, the species of the arabicus group occurring most closely to Qatar.
Distribution: Qatar, Saudi Arabia, United Arab Emirates.


Phrynocephalus maculatus

Black-tailed Toad-headed Agama, Whip-tailed Toad-headed Agama, (Spotted Toad-headed Agama)

Schwarzschwanz-Krötenkopfagame

Sorthalet Tudsehovedagam

1872 Phrynocephalus maculatus Anderson
1896 Phrynocephalus spiniventris Nikolsky (Barabanov & Ananjeva 2007)
Phrynocephalus maculatus spiniventris Nikolsky 1900

Remarks: Previously included longicaudatus. Records from Turkmenistan (e.g., Szczerbak 1994) refer to golubewii (Shenbrot & Semenov 1990; Barabanov & Ananjeva 2007).
Distribution: Afghanistan, Iran, Pakistan.


Phrynocephalus mystaceus

Bearded Toad-headed Agama

Bärtiger Krötenkopfagame

Skægget Tudsehovedagam

1776 Lacerta mystacea Pallas
Agama mystacea Merrem 1820
Phrynocephalus mystacia [sic] Kaup 1827
Megalochilus mystaceus Ananjeva 1986
1776 Lacerta aurita Pallas (Barabanov & Ananjeva 2007)
Gekko auritus Latreille 1801
Agama aurita Daudin 1802
Phrynocephalus auritus Fitzinger 1826
Megalochilus auritus Eichwald 1831
Saccostoma auritum Fitzinger 1843
1802 Lacerta lobata Shaw [substitute name for Lacerta aurita Gmelin 1789] (Barabanov & Ananjeva 2007)
1932 Phrynocephalus mystaceus galli Krassowsky (Barabanov & Ananjeva 2007)
1932 Phrynocephalus mystaceus mystaceus natio Dagestanica Krassowsky (unavailable name; Barabanov & Ananjeva 2007)
Megalochilus mystaceus dagestanica Ananjeva 1987 "1986" (Barabanov & Ananjeva 2007)
1990 Phrynocephalus mystaceus aurantiacocaudatus Semyonov & Shenbrot
2018 Phrynocephalus mystaceus khorasanus Solovyeva, Dunayev, Nazarov, Radjabizadeh & Poyarkov

Introduced to: Azerbaijan.
Distribution: Afghanistan, China (Xinjiang), Iran, Kazakhstan, Russia, Tajikistan, Turkmenistan, Uzbekistan.


Phrynocephalus nasatus

Aksu Toad-headed Agama

Aksu-Krötenkopfagame

Aksu-tudsehovedagam

1995 Phrynocephalus nasatus Golubev & Dunayev

Remarks: Previously regarded as a synonym of axillaris (e.g., Barabanov & Ananjeva 2007). Revalidated as a separate species by Dunayev (2020).
Distribution: China (Xinjiang).


Phrynocephalus nejdensis

Nejd Toad-headed Agama

Nejd-Krötenkopfagame

Nejd-tudsehovedagam

1957 Phrynocephalus nejdensis Haas

Remarks: Previously regarded as a synonym of arabicus (e.g., Barabanov & Ananjeva 2007). Revalidated as a separate species by Melnikov, Melnikova, Nazarov, Rajabizadeh, Al-Johany, Amr & Ananjeva (2014).
Distribution: Jordan, Saudi Arabia.


Phrynocephalus ornatus

Southern Striped Toad-headed Agama, Black-and-White Toad-headed Agama

Südliche Gestreifte Krötenkopfagame

Sydlig Stribet Tudsehovedagam

1887 Phrynocephalus ornatus Boulenger

Remarks: Previously included vindumi.
Distribution: Afghanistan, Pakistan.


Phrynocephalus persicus

Persian Toad-headed Agama

Persische Krötenkopfagame

Persisk Tudsehovedagam

1863 Phrynocephalus persicus De Filippi
Phrynocephalus helioscopus persicus Terentjev & Chernov 1949

Remarks: Previously regarded as a subspecies of helioscopus. Revalidated as a separate species by Anderson (1999). Previously included horvathi.
Distribution: Armenia, Azerbaijan, Iran.


Phrynocephalus przewalskii

Alashan Toad-headed Agama

Alasha-Krötenkopfagame

Alasha-tudsehovedagam

1876 Phrynocephalus Przewalskii Strauch
1876 Phrynocephalus affinis Strauch (Barabanov & Ananjeva 2007)
1906 Phrynocephalus pewzowi Bedriaga "1905" [nomen nudum] (Barabanov & Ananjeva 2007)
1907 Phrynocephalus Pewzowi Bedriaga (Barabanov & Ananjeva 2007)
1907 Phrynocephalus steindachneri Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus carinilabris Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus ciliaris Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus potanini Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus Reldæ Bedriaga (Barabanov & Ananjeva 2007)
1927 Phrynocephalus birulai Zarevsky (Barabanov & Ananjeva 2007)
1927 Phrynocephalus carinatus Zarevsky (Barabanov & Ananjeva 2007)
1927 Phrynocephalus elegans Zarevsky (Barabanov & Ananjeva 2007)
1927 Phrynocephalus parvulus Zarevsky (Barabanov & Ananjeva 2007)
1927 Phrynocephalus suschkinianus Zarevsky (Barabanov & Ananjeva 2007)
1984 Phrynocephalus werneri Peters [unavailable name] (Barabanov & Ananjeva 2007)

Remarks: Previously included frontalis.
Distribution: China (Beijing, Gansu, Inner Mongolia, Ningxia).


Phrynocephalus putjatai

Guide Toad-headed Agama

Guide-Krötenkopfagame

Guide-tudsehovedagam

1909 Phrynocephalus Putjatai Bedriaga

Remarks: Zhao & Adler (1993) regarded the status of this species as uncertain. Barabanov & Ananjeva (2007) treated it as a valid species. Validity supported by Guo & Wang (2007) and Noble, Qi & Fu (2010). Specific epithet occasionally misspelled putjatia (e.g., Noble, Qi & Fu 2010).
Distribution: China (Gansu, Qinghai).


Phrynocephalus raddei

Trans-Caspian Toad-headed Agama

Trans-Kaspische Krötenkopfagame

Transkaspisk Tudsehovedagam

1888 Phrynocephalus raddei Boettger
1905 Phrynocephalus raddei var. boettgeri Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)
Phrynocephalus reticulatus boettgeri Tsarevsky 1926
1915 Phrynocephalus raddei var. bilkewitschi Nikolski (Barabanov & Ananjeva 2007)
1915 Phrynocephalus raddei var. zarudnyi Nikolski (Barabanov & Ananjeva 2007)

Remarks: Previously regarded as a synonym of reticulatus (e.g., Terentjev & Chernov 1949; Wermuth 1967), but as a valid species by Szczerbak (1994) and Dunayev (1996). Although presence in Iran has previously been questioned (e.g., Anderson 1999; Safaei-Mahroo & al. 2015), raddei was recently confirmed for the country (Khosravani, Khani, Oraie & Rasyegar-Pouyani 2014), although not listed for Iran by Kamali (2020).
Distribution: Afghanistan, Iran, Tajikistan, Turkmenistan, Uzbekistan.


Phrynocephalus reticulatus

Reticulated Toad-headed Agama, Netted Toad-headed Agama, (Ladakh Toad-headed Agama)

Netz-Krötenkopfagame

Retikuleret Tudsehovedagam

1823 Agama ocellata Lichtenstein in Eversmann (Barabanov & Ananjeva 2007; see remarks)
Phrynocephalus ocellatus Golubev 1991
1831 Phrynocephalus reticulatus Eichwald
Phrynocephalus caudivolvolus reticulatus Nikolsky 1915
Phrynocephalus theobaldi reticulatus Das 2003
1905 Phrynocephalus boulengeri Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)
Phrynocephalus nikolskii boulengeri Nikolsky 1915
1905 Phrynocephalus nikolskii Bedriaga in Nikolsky (Barabanov & Ananjeva 2007)

Remarks: Previously included bannikovi. Agama ocellata Lichtenstein in Eversmann 1823 was formerly regarded as a synonym of caudivolvolus (e.g., Boulenger 1885) or of guttatus (e.g., Lantz 1924; Wermuth 1967). Golubev (1991) demonstrated that ocellata is a senior synonym of reticulatus and referred to the species as ocellatus. In the interest of taxonomic stability, the species is here referred to as reticulatus, following Barabanov & Ananjeva (2007), since this name has been used many times for more than a century. P. raddei was previously confused with reticulatus (e.g., Wermuth 1967), and older records may refer to either species. Records from Afghanistan (e.g., Leviton & Anderson 1970 [as reticulatus boettgeri]) refer to raddei (Clark 1990).
Distribution: Turkmenistan, Uzbekistan.


Phrynocephalus roborowskii

Zaidam Toad-headed Agama

Zaidam-Krötenkopfagame

Zaidam-tudsehovedagam

1906 Phrynocephalus vlangalii var. roborowskii Bedriaga "1905"
Phrynocephalus roborowskii Peters 1984

Remarks: Regarded as a synonym of vlangalii by Zhao & Adler (1993), but as a valid species by Barabanov & Ananjeva (2007). Not listed for China by Wang, Ren, Chen, Lyu, Guo, Jiang, Chen, Li, Guo, Wang & Che (2020).
Distribution: China (Qinghai).


Phrynocephalus rossikowi

Khentau Toad-headed Agama

Khentau-Krötenkopfagame

Khentau-tudsehovedagam

1899 Phrynocephalus rossikowi Nikolsky
1915 Phrynocephalus rossikowi relictus Nikolski (Barabanov & Ananjeva 2007)
1979 Phrynocephalus rossikowi shammakowi Shcherbak & Golubev (Barabanov & Ananjeva 2007)

Distribution: Turkmenistan, Uzbekistan.


Phrynocephalus sakoi

Sharqiyah Toad-headed Agama

Sharqiyah-Krötenkopfagame

Sharqiyah-tudsehovedagam

2015 Phrynocephalus sakoi Melnikov, Melnikova, Nazarov, Al-Johany & Ananjeva

Distribution: Oman.


Phrynocephalus scutellatus

Grey Toad-headed Agama, Banded Toad-headed Agama

Graue Krötenkopfagame

Grå Tudsehovedagam

1807 Agame scutellata [incorrect original spelling] Olivier
Phrynocephalus scutellatus Moquard 1910
1837 Phrynocephalus olivieri Duméril & Bibron (Barabanov & Ananjeva 2007)
1845 Phrynocephalus tickelii Gray (Barabanov & Ananjeva 2007)
1907 Phrynocephalus olivieri var. brevipes Nikolski (Barabanov & Ananjeva 2007)
1907 Phrynocephalus olivieri var. carinipes Nikolski (Barabanov & Ananjeva 2007)

Distribution: Afghanistan, Iran, Pakistan.


Phrynocephalus sogdianus

Tajik Toad-headed Agama

Tadschikische Krötenkopfagame

Tadsjikisk Tudsehovedagam

1948 Phrynocephalus interscapularis sogdianus Chernov
Phrynocephalus sogdianus Sokolovsky 1975

Remarks: Previously treated as a subspecies of interscapularis. Sokolovsky (1975) raised sogdianus to status as a separate species. Validity confirmed by Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss (2018).
Distribution: Afghanistan, Tajikistan, Uzbekistan.





Phrynocephalus strauchi

Fergana Valley Toad-headed Agama, (Strauch's Toad-headed Agama)

Fergana-Krötenkopfagame

Fergana-tudsehovedagam

1899 Phrynocephalus strauchi Nikolsky
Phrynocephalus reticulatus strauchi Terentjev & Chernov 1949

Remarks: Previously treated as a subspecies of reticulatus. Mesherin & Golubev (1989) revalidated strauchi to status as a separate species. Validity confirmed by Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss (2018).
Distribution: Kyrgyzstan, Tajikistan, Uzbekistan.


Phrynocephalus theobaldi

Kashmir Toad Agama, (Theobald's Toad Agama)

Himalaya-Krötenkopfagame

Kashmir-tudsehovedagam

1863 Phrynocephalus theobaldi Blyth
1867 Phrynocephalus stoliczkai Steindachner (Barabanov & Ananjeva 2007)
1909 Phrynocephalus vlangalii var. lidskii Bedriaga (Barabanov & Ananjeva 2007)
Phrynocephalus lidskii Tsarevsky 1927
1984 Phrynocephalus alticola Peters (Barabanov & Ananjeva 2007)
1996 Phrynocephalus zetangensis Wang, Zeng, Fang, Liu, Wu, Papenfuss & Macey (the authors also used the spelling zetanensis in the original decription) (Barabanov & Ananjeva 2007; Guo & Wang 2007)
1999 Phrynocephalus theobaldi orientalis Wang, Papenfuss & Zeng in Wang, Zheng, Fang, Wu, Papenfuss & Macey [not Phrynocephalus helioscopus var. orientalis Bedriaga 1912] (Barabanov & Ananjeva 2007)
2002 Phrynocephalus theobaldi lhasaensis Barabanov, Ananjeva, Papenfuss & Wang [replacement name for Phrynocephalus theobaldi orientalis Wang, Papenfuss & Zeng in Wang, Zheng, Fang, Wu, Papenfuss & Macey 1999] (Barabanov & Ananjeva 2007)

Remarks: Previously included erythrurus. Ananjeva, Guo & Wang (2011) questioned the presence of theobaldi in Xinjiang, referring to Zhao (1979).
Distribution: China (Tibet, Xinjiang), India (Jammu & Kashmir), Nepal.


Phrynocephalus turcomanus

Turkmen Sunwatcher

Turkmenische Sonnengucker

Turkmensk Solkigger

2012 Phrynocephalus helioscopus turcomanus Solovyeva, Dunayev & Poyarkov
Phrynocephalus turcomanus Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss 2018

Distribution: Iran, Turkmenistan.





Phrynocephalus versicolor

Piebald Toad-headed Agama

Mongolischer Krötenkopfagame

Mongolsk Tudsehovedagam

1876 Phrynocephalus versicolor Strauch
1906 Phrynocephalus guentheri Bedriaga "1905" (Barabanov & Ananjeva 2007)
1906 Phrynocephalus versicolor var. doriai Bedriaga "1905" [nomen nudum] (Barabanov & Ananjeva 2007)
1909 Phrynocephalus Blanfordi Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus versicolor var. Bogdanowi Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus versicolor var. Doriai Bedriaga (Barabanov & Ananjeva 2007)
1909 Phrynocephalus versicolor var. Siebenrocki Bedriaga (Barabanov & Ananjeva 2007)
1930 Phrynocephalus rostralis Zarevsky (Barabanov & Ananjeva 2007)
1958 Phrynocephalus stejnegeri Bannikov [nomen nudum] (Barabanov & Ananjeva 2007)
1984 Phrynocephalus steynegeri Peters [unavailable name] (Barabanov & Ananjeva 2007)
1995 Phrynocephalus immaculatus Zhao (Barabanov & Ananjeva 2007)

Remarks: Previously included hispidus and kulagini. Records from Kazakhstan refer to the former subspecies paraskiwi, now considered a synonym of guttatus (Golubev 1989; Szczerbak 2003; Barabanov & Ananjeva 2007).
Introduced to: Philippines (Luzon).
Distribution: China (Gansu, Inner Mongolia, Ningxia, Xinjiang), Mongolia.


Phrynocephalus vindumi

Northern Striped Toad-headed Agama

Nördliche Gestreifte Krötenkopfagame

Nordlig Stribet Tudsehovedagam

1998 Phrynocephalus ornatus vindumi Golubev
Phrynocephalus vindumi Macey, Schulte, Ananjeva, Dyke, Wang, Orlov, Shafiei, Robinson, Dujsebayeva, Freund, Fischer, Liu & Papenfuss 2018

Distribution: Iran.





Phrynocephalus vlangalii

Quinghai Toad-headed Agama

Quinghai-Krötenkopfagame

Quinghai-tudsehovedagam

1876 Phrynocephalus Vlangalii Strauch
1906 Phrynocephalus vlangalii var. nanschanica Bedriaga "1905" (Barabanov & Ananjeva 2007)
Phrynocephalus nanschanicus Tsarevsky 1927
1909 Phrynocephalus Vlangalii var. geckoides Bedriaga (Barabanov & Ananjeva 2007)
Phrynocephalus geckoides Tsarevsky 1927
1909 Phrynocephalus Vlangalii var. Grombtschewskii Bedriaga (Barabanov & Ananjeva 2007)
Phrynocephalus grombtschewskii Tsarevsky 1927
1909 Phrynocephalus Vlangalii var. Pylzowi Bedriaga (Barabanov & Ananjeva 2007)
Phrynocephalus pylzowi Tsarevsky 1927
Phrynocephalus parvus pylzowi Tsarevsky 1927
1984 Phrynocephalus occidentalis Peters [unavailable name] (Barabanov & Ananjeva 2007)

Remarks: Previously included hongyuanensis, now a valid species, and parva, now a synonym of erythrurus.
Distribution: China (Gansu, Qinghai, Sichuan, Xinjiang).