Volume 1, Number 1, January 1994 Russian Journal of Herpetology TOBUSHING COMPANY 1 rOLIUM Folium Publishing Co., Moscow, Russia Russian Journal of Herpetology Vol 1, No. 1, 1994, pp. 42 — 52 SOME ASPECTS OF HISTORICAL BIOGEOGRAPHY OF ASIAN ROCK AGAMIDS N. B. Anan’eva1 and B. S. Tuniev2 Oi igiiial article submitted October 21,1993 The genesis of areals of Asian rock agamids belonging to the genus Laudakia Gray, 1845 is considered. The hypothesis about two routes of radiation of these lizards endemic for arid mountain regions of Palearctic is offered. The touthefn route from the hypothetical center of origin in the area of mountain systems of Hindukush and Himalayas passed in the conditions of southern subtropics with well developed belt of sclerophilous forests. This evolutional line with “caucasia” complex remains the relic areas in forest regions. Northern group of species with “himalayam” complex had its speciation in northern subtropic area with more severe climatic situation determined by strong continental and arid conditions. Altitudinal and biotopic differences of Laudakia species under their sympatiic distribution are specially discussed. Key words: Reptilia, Sauna Agamidae, Laudakia, Palearctic Asia, systematics, biogeography, distribu­ tion, evolution The Asian rock agamids of the genus Laudakia phylogenetic scheme is impeded by the difficulty ob­ Gray, 1845 includes 16 species distributed in moun­ taining material for genetic analysis for several groups tain rock landscapes of arid zone from Greece and of species and by the obvious existence of parallel Nile river delta on the west through Middle East and trends in different developmental lines within Central Asia to Gobi Altai on the north-east and Laudakia genus. Bramaputra river on the east. The mountain ring­ We collected data on geographic and biotopic dis­ tailed agamas belonging to this genus have been con­ tribution of rock agamas for their discussion in con­ sidered for a long time as a part of the genus Agama text of notions on the climate and genesis of land­ Daudin, 1802 (Boulenger 1885; Wermuth 1967). Af­ scapes and vegetation. A comparison of recent are- ter Moody’s revision (1980) they were referred to a alographic patterns of the studied species with known distinct genus Stellio Laurenti, 1768. The subsequent data un paleogeography and paleobotany of the region study of the whole complex of mountain ring-tailed of their distribution may be used as additional method agamas showed that it is a paraphyletic group (Joger of research for understanding the trends of reptiles ra­ and Arano 1987; Ananjeva et al. 1990; Joger 1991). diation (Ananjeva and Tuni^ev 1992). It might be es- Taking into consideration these data and some nomen­ pecially fruitful for Laudakia because of the unique clature remarks some authors (Leviton et al. 1992) of­ chorological isolation of this genus (Anan’eva and Pe­ fer to use the generic name Laudakia Gray, 1845 for ters 1990). Asian rock agamids and for Afro-Arabian group of We gathered data on geographic and biotopic dis­ species - -th e generic nameAcanthocercus Fitzinger, tribution of Laudakia caucasia, L. chemovi, L. ery- 1843. throgastra, L. himalayana, L. lehmanni, L. stolicz- The analysis of phylogenetic relations of Lau­ kana in 1975 — 1992 during field work in Armenia, dakia species was accomplished on the basis of both Georgia, Dagestan, Turkmenistan, Tadzhikistan, morphological and biochemical data (Joger and Arano Uzbekistan and Mongolia. We also used all the avail­ 1987; Ananjeva et al. 1990; Anan’eva and Sokolova able published information on the histoiy of landscape 1990; Joger 1991). These results are partly contradic­ formation typical for Laudakia areal and biotopical tory and can be considered only as preliminary ideas. distribution of these lizards in Iran Afghanistan, Paki­ They can not definitely determine the relations within stan, Turkey, Greece, Israel, and China. this groups of lizards. The construction of more clear Asian rock agamids endemic for Palearctic have the distribution looking like a integral unit (Fig. 1, Ta­ Zoological Institute, Russian Academy of Sciences, St. Peters­ ble 1). Within their diversity we can distinguish at burg, Russia. I least two complexes of species with a number of mor­ ' Caucasian State Biosphere Reserve, Sochi, Russia phological and ecological differences: “caucasia” О 1993 Folium Publishing Company Some Aspects of Historical Biogeography of Asian Rock Agamids 43 complex and “himalayana” complex. The first one in­ cludes L. caucasia, L. microlepis, L. nuristanica, Black L. tuberculata, and probably L. sacra, L. lehmanni, Caspian Sea and L. erythrogastra. “Himalayana” complex con­ sists of related species L. himalayana, L. chernovi, L. badakhshana, L. stoliczkana (Peters 1971; Anan’e­ va etal. 1981; Anan’eva and Peters 1990). It is possi­ ble, that L. pakistanica that is most closely related to Arabian Sea L. agrorensis and L. melanura (Baig 1989) can be at­ tributed to this complex. We realize that it is a specu­ lative scheme, and probably future genetic examina­ F1S. I. General distribution of Asiatic rock agamids of Laudakia tion will permit to distinguish more groups and com­ genus in Palearctic. plexes of species within Laudakia. It was ilreadv noted (Anan’eva and Peters 1990), that long limbs, presence of a small gular sac and Despite the absence of clear phylogenetic hy­ nuchal crest, the poly annular structure of the caudal pothesis for Laudakia all the herpetologists who stud­ segments, juvenile color patterns should be consid­ ied their taxonomy and ecology are unanimous in the opinion, that speciation of these lizards has been con­ ered as plesiomorphic characters. But the interpreta­ nected in the past and at present with Hindukush, tion of separate morphological characters (including Pamir and Himalayas mountains (Peters 1971; Anan- scalation) as indication of species relationships is un­ jeva et al. 1981; Anan’eva and Peters 1990; Baig acceptable, and we use here such notions on plesio- 1992). Strong heterogeneity of dissected mountain re­ morphic characters of scalation only in context of are- lief probably promoted the isolation of populations, an alographic patterns of Laudakia. As it was noted ecological differentiation of these lizards is now nor­ above, the parallel trends occur in different develop­ mally expressed in different altitudinal distribution. In mental lines within Laudakia genus, expressed for ex­ this context it seems to be useful to discuss our hy­ ample in parallel changing of the number of annuli in pothesis for geographic distribution of the two com­ caudal segments in L. stellio, L. stoliczkana, and plexes of Asian mountain agamas and formation of L. caucasia (Peters 1971; Anan’eva and Ataev 1984). their contemporary ranges. TABLE 1. The distribution of the species of Laudakia genus. Species Distribution Laudakia agrorensis (Stoliczka 1872) Afghanistan, Pakistan, India Laudakia badakhshana (Anderson and Leviton 1969) Afghanistan Laudakia caucasia (Eichwald 1831) Armenia, Georgia, Azerbajan, Tajikistan, Turkmenistan, Turkey, Iraq, Iran, Afghanistan, Pakistan Laudakia chernovi (Ananjeva et al. 1981) Tajikistan, Turkmenistan, Uzbekistan Laudakia erythrogastra (Nikolskyi 1896) Iran, Turkmenistan, Afghanistan Laudakia himalayana (Steindachner 1869) Tajikistan, Uzbekistan, Kirgizstan, Afghanistan, Pakistan, India Laudahu lehmanni (Nikolskyi 1896) Tajikistan, Uzbekistan, Kirgizstan, Turkmenistan, Afghanistan Laudakia melanura (Blyth 18S4) Iran, Pakistan Laudakia microlepis (Blanford 1874) Iran, Pakistan, Afghanistan Laudakia nupta (De Filippi 1843) Iraq, Iran, Afghanistan, Pakistan Laudakia nuristanica (Anderson and Leviton 1969) Afghanistan, Pakistan Laudakia sacra (Smith 1935) Tibet (China) Laudakia stellio (Linnaeus 1758) Greece, Southwest Asia, Northern Egypt Laudakia stoliczkana (Blanford 1875) Mongolia, China Laudakia tuberculata (Hardwicke and Gray 1827) India, Nepal, Afghanistan, Pakistan Laudakia pakistanica (Baig 1989) Pakistan 44 N. В. Anan’eva and В. S. liiniev We have veiy - ant;' paleontological data that there arose acceptable arid areas as distinct from luxu­ cannot elucidate the problems of origin and ancient riant tropical vegetation of the Himalayas proper in diversity of agamid lizards of Laudakia genus appropriate for colonization by these arid lizards. (Moody 1980; Estes 1983) especially because we Ouickly rising and reaching already cold desert condi­ have no reliable arguments for allocation of fossil re­ tions Tibet could not serve as a conductor for the ra­ cords to Agama, Trapelus, Laudakia or Ananthocer- diation of Asian mountain agamas which retain even cus. now the main recently the chorological patterns in The analvsis of present diversity of Asian moun­ subtropical area of Asia, Europe, and northern Africa tain ring-tailed agamas permits to assume the area of (L. stellio). recent Hindukush and Himalayas mountain systems We shall consider two complexes of species of to be the center of Laudakia origin. Their speciation Laudakia genus in the framework of this hypothesis. was influenced by aridization of the southern part of Our observations are based mainly on the field Asian continent and Alpic orogenesis. The allozyme work results in the areas of distribution of L. caucasia, based phylogenetic hypothesis for agamid genus L. himalayana, L. chernovi, L. lehmanni, L. erythro- Phrynocephalus (Macey et al. 1991, 1992) argues for gastra, L. stoliczkana. These observations and eco­ ancient divergence of