PHRYNOCEPHALUS PHYLOGENY 



Ecological analogues of Phrynocephalus 



Small diurnal lizards, that are sit-and-wait foragers, have high body 

 temperatures when active and in many cases signal with their tails, 

 are found in several desert systems. Apart from Phrynocephalus, 

 they include the agamids Ctenophorus and Tympanocryptis in Aus- 

 tralia, the phrynosomatid sand lizards in North America (Uma, 

 Callisaurus, Holbrookia and Cophosaurus), tropidurines in south 

 America (Leiolaemus), geckoes in southern Arabia and Somalia 

 (Pristurus) and lacertids in Southwest Africa (Meroles anchietae). 

 However, although they show significant parallels in morphology 

 and behaviour, these derived features are not necessarily assembled 

 in the same order (Arnold, 1994). 



Nomenclature 



As presently understood, Phrynocephalus is a well-defined clade 

 defined by six synapomorphies not found in closely related agamids 

 (numbers 1.1, 12.1, 23, 35, 37.1 and 46 in the present data set). 

 Besides lacking these, Bufoniceps, the sister taxon of 

 Phrynocephalus, possesses at least one apomorphy not found in the 

 latter genus, namely a very short tail. Golubev & Dunayev (1997) 

 suggested that Bufoniceps should be expanded to include P. 

 mystaceus, P. maculatus, P. arabicus, P. ornatus, P. clarkorum, P. 

 luteoguttatus, P. euptilopus, P. interscapularis and P. sogclianus. 

 These are all basal members of Phrynocephalus and their inclusion 

 in Bufoniceps would create a new grouping that is clearly paraphy letic 

 and reduce Phrynocephalus to a smaller and less well defined clade. 

 The suggestion should consequently be rejected. 



Acknowledgements. I am grateful to Jens Vindum (California Acad- 

 emy of Sciences for the loan of material of Phynocephalus roborowskih, 

 P. rossikowi and P. strauchi. Ed Wade drew Figures 7-11. 



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