ISSN 1022-7954, Russian Journal of Genetics, 2011, Vol. 47, No. 7, pp. 842–856. © Pleiades Publishing, Inc., 2011.
Original Russian Text © E.N. Solovyeva, N.A. Poyarkov, E.A. Dunaev, T.N. Duysebayeva, A.A. Bannikova, 2011, published in Genetika, 2011, Vol. 47, No. 7, pp. 952–967.
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INTRODUCTION
Sunwatcher toad-headed agama Phrynocephalus
superspecies helioscopus is a wide-range species complex
(hereafter designated as the helioscopus–persicus com-
plex), which is found in CentralAsia, northwestern China,
northern Iran, and Transcaucasia [1]. From the time of its
first description (P. S. Pallas, 1771) till nowadays, no stable
opinions on the species phylogeny have been formed. To
solve this issue, mostly morphological characters, the
indices of which strongly overlap, were used. Sunwatcher
toad-headed agamas are characterized by polymorphism
in the chromosome number and the ratio between macro-
and microchromosomes [2–4]. According to V. K. Erem-
chenko and A.M. Panfilov [5], karyotype of 48 chromo-
somes (M/m = 24/24) is ancestral to Ph. helioscopus.
Karyotype of Persian toad-headed agama (Ph. persicus
horvathi) was found to be most different from ancestral
variant. Populations of true sunwatcher toad-headed
agama (Ph. helioscopus) are indistinguishable in the mus-
cle tissue thermostability. However, Ph. p. horvathi from
the south of Azerbaijan possess the same cytophysiological
characteristics as Ph. helioscopus from Central Asia [6].
Development of molecular genetic approaches made
it possible to put in order the concepts on the species com-
plex Ph. helioscopus–Ph. persicus. However, genetic vari-
ation of this group was the subject of investigation in only
few studies. In these studies, from three [5, 7] to 5–12 [8–
14] phyletic clades are discussed. Thus, according to cur-
rent ideas, within the species of Phrynocephalus heliosco-
pus three subspecies, Ph. helioscopus helioscopus,
Ph. h. saidalievi, and Ph. h. varius are distinguished. In
turn, within Phrynocephalus persicus two subspecies,
Ph. p. persicus and Ph. p. horvathi, are identified. In gen-
eral, taxonomic relationships within the species complex
of toad-headed agamas remain unresolved.
Since the range of sunwatcher and Persian toad-
headed agamas covers nearly all Central Asiaand adja-
cent territories, it seems likely that analysis of phylo-
genetic relationships within this wide-range species
complex will shed light on the events, which influ-
enced the dispersal and speciation of many other
organisms, inhabiting this region.
MATERIALS AND METHODS
Sample Description
Molecular genetic analysis was performed using tissue
samples of 86 Phrynocephalus helioscopus lizards repre-
senting 53 populations covering the whole range (Fig. 1,
Table 1). Tissue samples of Ph. axillaris (ZMMU
R-12302), Ph. mystaceus (ZMMU R-12149), and
Ph. interscapularis (ZMMU R-12260) were taken as
outgroups. Most of the samples examined were held in
herpetological collections of the Zoological Museum
of Lomonosov Moscow State University (ZMMU,
70 samples) and the Institute of Zoology and Parasi-
tology, Uzbek Academy of Sciences, Republic of
Molecular Differentiation and Taxonomy of the Sunwatcher
Toad-Headed Agama Species Complex Phrynocephalus
Superspecies helioscopus (Pallas 1771) (Reptilia: Agamidae)
E. N. Solovyeva
a
, N. A. Poyarkov
a
, E. A. Dunaev
b
, T. N. Duysebayeva
c
, and A. A. Bannikova
a
a
Department of Vertebrate Zoology, Lomonosov Moscow State University, Moscow, 119991 Russia
e-mail: anolis@yandex.ru
b
Zoological Museum, Lomonosov Moscow State University, Moscow, 125009 Russia
c
Institute of Zoology, Almaty, 480060 Kazakhstan
Received September 20, 2010
Abstract—Lizards of the sunwatcher toad-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-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-
groups, many of which were characterized by a narrow range and genetic isolation. Monophyly of sunwatcher
(Ph. helioscopus) and Persian (Ph. persicus) toad-headed agamas was confirmed. However, both of these spe-
cies probably represent the species complexes. Zoogeography of Central Asiais discussed.
DOI: 10.1134/S1022795411070155
ANIMAL GENETICS