Macey, Robert, Schulte, James A., Ananjeva, Natalia B., Van, Erik T., Wang, Yuezhao, Orlov, Nikolai, Shafiei, Soheila, Robinson,, 2018, A molecular phylogenetic hypothesis for the Asian agamid lizard genus Phrynocephalus reveals discrete biogeographic clades implicated by plate tectonics, Zootaxa 4467 (1), pp. 1-81: 51

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F. Phrynocephalus  Species Diversity as Related to Geology, Habitat and Size

Two regions show high species numbers in close proximity. Six species are found on the northern side of the Kopet-Dagh (mountains) along the southern margin of the Caspian Basin in Turkmenistan. Species that inhabit hard substrates are P. rossikowi  , P. raddei  , P. bannikovi  , P. golubewii  , and P. turcomanus  , with P. interscapularis  and P. mystaceus  occurring in soft substrates ( Table 5, Fig. 16View FIGURE 16). Among these species there are major size differences ( Table 6).

Six species are also found at the juncture of the Helmand Basin of southern Afghanistan with the Baluchistan Plateau including the base of the Sulaiman Range of southern Pakistan. Species that inhabit hard substrates are P. maculatus  and P. scutellatus  ( Table 5, Fig. 16View FIGURE 16). Smaller species that inhabit soft substrates are P. clarkorum  , P. ornatus  , and P. luteoguttatus  (Clade C in Figs. 4View FIGURE 4 and 8View FIGURE 8). A larger species inhabiting soft substrates in large floating sand dunes is P. euptilopus  , which was not sampled in this study even though repeated efforts were made in both Afghanistan and Pakistan by the last author. Among these species there are major size differences ( Table 6).

Both of these examples share a plate tectonic past around the northern migration of the Arabian Plate and its impact on ancient tectonic plates to the north in Eurasia. The Kopet-Dagh (mountains), on the northern margin of the Iranian Plateau. descends into the Caspian Basin. This region of the Caspian Basin experienced the most dramatic changes via tectonic uplift, erosion, and sea floor drying of the Paratethys Sea creating unique and diversified habitats (see Figs. 3View FIGURE 3, 9–14View FIGURE 9View FIGURE 10View FIGURE 11View FIGURE 12View FIGURE 13View FIGURE 14; Table 4). Additionally, the Helmand Block of southern Afghanistan is being pushed south by impinging Arabia and further northern invasion of the Indian Plate ( Fig. 9View FIGURE 9). This has created uplift in the Gulf of Oman creating the push-back of the Makron in southern Pakistan ( Figs. 9–12View FIGURE 9View FIGURE 10View FIGURE 11View FIGURE 12).

It is dynamic tectonics of this nature that created a series of habitats leading to speciation among Phrynocephalus  . As adaptive niches were created, size diversity, morphological specializations, background matching, and defensive mimicry developed ( Plates IView PLATE I – VIII). Future work on Phrynocephalus  will need to take into account these aspects with particular attention to plate tectonic induced events in order to investigate phyletic patterns, taxonomy, and evolutionary diversity among one of the world’s most interesting and confusing lizard genera.