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TERRA TYPICA. Town Kislovodsk, Stavropol Territory. DISTRIBUTION. The main distribution range of the species covers the northern slopes of the Rocky range within Karachaevo- Cherkesskaya Autonomous Region, south of the Stavropol Territory and Kabardino-Balkaria, from the valley of the upper current of Kuban’ River and its tributaries in the west up to the ravine of the middle current of the river Chegem in the east. The most northern isolated population (which was considered as extinct) lives in the southern foothills of the Stavropol Upland, in the limits of the Alexandrovsky region of the Stavropol Territory. The southern border of the distribution range passes on the north-eastern spurs of the mountain Elbrus in the ravine of the river Kuban, and, crossing the Rocky Range (known from the Mariinsky pass), continues further on its watershed in the upper reaches of the rivers Kuma, Podkumok, Alikanovka, Beryozovaya, Zolka and others. On the ravine of the river Baksan it comes up to the town Tyrny-Auza and goes from the Chegem waterfalls up along the Chegem River gorge. CONSERVATION STATUS. The number is high and stable.

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The extent of resistance of animal cells and tissues to injurious heat effect is a cytophysiological criterion of species. In poikilotherms it can be determined both by phylogenetic relations and ecological conditions of a species` formation and existence. To analyze the relative role of these factors in formation of the tissue heat resistance level as characteristic of a species, a study was made on 9 species of lizards of the `Lacerta saxicola` complex. A characteristic feature of the complex is that some of the species belonging it are of hybrid origin, which implies that genetically they are closely related to other, parental forms. Heat resistance level of muscle tissue in a hybrid species may differ from the parental forms if the conditions of its existence are different. On other hand, genetically unrelated species living under similar conditions may have the same level of heat resistance of muscle tissue. Interspecific differences in heat resistance of muscles cannot be explained by phylogenetic relations only. Of major importance in formation of the cellular heat resistance level in the process of speciation are ecological, and primarily, thermal conditions of the environment.

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Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta lepida, Lacerta princeps, Lacerta schreiberi, Lacerta strigata, Lacerta trilineata, Lacerta viridis, Lacerta andreanszkyi, Lacerta armeniaca, Lacerta bedriagae, Lacerta brandtii, Lacerta cappadocica, Lacerta caucasica, Lacerta chlorogaster, Lacerta cyanura, Lacerta danfordi, Lacerta derjugini, Lacerta dugesii, Lacerta fraasii, Lacerta graeca, Lacerta horvathi, Lacerta jayakari, Lacerta laevis, Lacerta monticola, Lacerta mosorensis, Lacerta oxycephala, Lacerta parva, Lacerta perspicillata, Lacerta praticola, Lacerta rudis, Lacerta saxicola, Lacerta vivipara, Gallotia atlantica, Gallotia galloti, Gallotia simonyi, Podarcis erhardii, Podarcis filfolensis, Podarcis hispanica, Podarcis lilfordi, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis pityusensis, Podarcis sicula, Podarcis taurica, Podarcis tiliguerta, Podarcis wagleriana, Psammodromus algirus, Psammodromus blanci, Psammodromus hispanicus, Psammodromus microdactylus.

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The final part of a series of papers on the lizards of the Caucasus, that have been published in previous years. The paper reports on the life habits, capture and keeping in terraria of several lizards of Soviet Transcaucasia. In these respects the agamid Phrynocephalus helioscopus persi- cus and three species of the lacertid genus Eremias occuring in that area (i. e. strauchi, velox, pleskei) are discussed. Finally a !ist of all lizard species and subspecies actually known to occur in this region is given.

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A phylogenetic-comparative approach was used to assess and refine existing secondary structure models for a frequently studied region of the mitochondrial encoded large subunit (16S) rRNA in two large lizard lineages within the Scincomorpha, namely the Scincidae and the Lacertidae. Potential pairings and mutual information were analyzed to identify site interactions present within each lineage and provide consensus secondary structures. Many of the interactions proposed by previous models were supported, but several refinements were possible. The consensus structures allowed a detailed analysis of rRNA sequence evolution. Phylogenetic trees were inferred from Bayesian analyses of all sites, and the topologies used for maximum likelihood estimation of sequence evolution parameters. Assigning gamma-distributed relative rate categories to all interacting sites that were homologous between lineages revealed substantial differences between helices. In both lineages, sites within helix G2 were mostly conserved, while those within helix E18 evolved rapidly. Clear evidence of substantial site-specific rate variation (covarion-like evolution) was also detected, although this was not strongly associated with specific helices. This study, in conjunction with comparable findings on different, higher-level taxa, supports the ubiquitous nature of site-specific rate variation in this gene and justifies the incorporation of covarion models in phylogenetic inference.

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We investigated age structure, body size and longevity in two breeding populations of Darevskia clarkorum inhabiting altitudes ranging from 450 m a.s.l. (Kamilet) to 2250 m a.s.l. (Basyayla) in Turkey by skeletochronology performed on the phalanges. The mean age was found to be 6 years in the Kamilet population and 7 years in the Basyayla population. The maximum life span was 10 years in the lowland population while it was 12 years in the highland population. Age at sexual maturity of both males and females was 1-2 years in the lowland population while it was 2-3 for both sexes in the highland population. Both age and SVL of specimens from the Kamilet population were significantly different between the sexes while age and SVL did not differ significantly between the sexes in Basyayla population. As a conclusion, we observed that the mean age, longevity and age at maturity were increased by altitude while there was a decrease based on the mean SVL in the highland population of D. clarkorum. Our data on body size, longevity and age at sexual maturity may contribute to conservation efforts for this endangered species.

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This paper reports the isolation and characterization of twoHindIII repetitive DNA families from the genome of two lacertid lizards,Podarcis sicula andLacerta saxicola. These satellites did not appear to be related to each other. The consensus sequences of their monomeric units did not show any similarity, though both DNAs were A-T rich. Moreover, each of them was found only in closely related species. The monomeric unit of theHindIII DNA family isolated fromP. sicula (pLHS) showed a close resemblance to pLCS, a centromeric satellite DNA previously isolated from the same species; it was, however, mainly localized at pericentromeric, interstitial and telomeric levels. The results also provide interesting information on the systematics of the lacertids studied.

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The structure and evolution of a satellite DNA family was examined in lizards from the genus Darevskia(family Lacertidae). Comparison of tandem units of repeated DNA (satDNA), CLsat, in all species from the genus Darevskiahas shown that their variability is largely explained by single-nucleotide substitutions, which form about 50 diagnostic positions underlying classification of the family into three subfamilies. Maximum differences between the subfamilies reached 25%. At this level of tandem unit divergence in the subfamilies, no cross-hybridization between them was observed (at 65°C). The individual variability within one subfamily within the species was on average 5% while the variability between species consensuses within a subfamily was 10%. The presence of highly conserved regions in all monomers and some features of their organization show that satellites of all Darevskia species belong to one satDNA family. The organization of unit sequences of satellites CLsat and Agi160 also detected by us in another lizard genus, Lacerta s. str. was compared. Similarity that was found between these satellites suggests their relatedness and common origin. A possible pathway of evolution of these two satDNA families is proposed. The distribution and content of CLsat repeat subfamilies in all species of the genus was examined by Southern hybridization. Seven species had mainly CLsatI (83 to 96%); three species, approximately equal amounts of CLsatI and CLsatIII (the admixture of CLsatII was 2–5%); and five species, a combination of all three subfamilies in highly varying proportions. Based on these results as well as on zoogeographic views on the taxonomy and phylogeny of the Darevskia species, hypotheses on the evolution of molecular-genetic relationships within this genus are advanced.

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The genetic relatedness of several bisexual and of four unisexual Lacerta saxicola complexlizards was studied, using monomer sequences of the complex-specific CLsat tandem repeats and anonymous RAPD markers. Genomes of parthenospecies were shown to include different satellite monomers. The structure of each such monomer is specific for a certain pair of bisexual species. This fact might be interpreted in favor of co-dominant inheritance of these markers in bisexual species hybridogenesis. This idea is supported by the results obtained with RAPD markers; i.e., unisexual species genomes include only the loci characteristic of certain bisexual species. At the same time, in neither case parthenospecies possess specific, autoapomorphic loci that were not present in this or that bisexual species.

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The polymorphic species Lacerta saxicola Eversmann forms 19 distinct forms in the Caucasus, four of which (armeniaca, dahli, rostombekovi, and unisexualis) are devoid of males and normally reproduce parthenogenetically. These parthenogenetic forms inhabit the mountains on the periphery of the range of the normal bisexual form, L. s. defilippii, thus suggesting a case of geographic parthenogenesis. Since the populations of parthenogenetic lizards are virtually devoid of males, the application of geographic and physiological species criteria to them is a matter of some difficulty. A discussion of this situation with regard to the problem of so-called `agamic species` enables us to regard the parthenogenetic relatives of L. saxicola as distinct species rather than subspecies, and these forms are raised to that rank. The parthenogenetic form of L. s. defilippii is described as a separate species (L. unisexualis). Cytological studies have shown that parthenogenetic species of Rock Lizards are diploid (2n = 38) and that oogenesis proceeds similarly in parthenogenetic and bisexual forms up to the second meiotic division. In regions of sympatry the parthenogenetic and bisexual forms occasionally produce natural hybrids, resulting in sterile triploid (3n = 57) females, the sterility of which is caused by anomalies in the development and structure of the gonads. In the eggs laid by parthenogenetic females, peculiar abnormal embryos, which commonly die in an early stage of incubation, are often developed. Most of them prove to be males, the development of which stops approximately in the middle of incubation. There is reason to suppose that the transition of Rock Lizards to parthenogenesis is associated with natural interspecific hybridization, which in turn favors unisexual reproduction. This tendency could be realized under the influence of selection and apparently took place in forest refuges during Quaternary glaciation of the Caucasus. The transition of Rock Lizards to unisexual reproduction is shown to be accompanied by the appearance of morphologically and biologically distinct species. Natural parthenogenesis may thus be regarded as one of the factors of speciation in the L. saxicola group.

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Four parthenogenetic subspecies of Lacerta . saxicola Eversmann (Transcaucasia) have closely overlapping sympatric ranges. The lizards show no intermediate forms in zones of their mutual habitat. On the border - line of their ranges parthenogenetic lizards can mate with males of neighbouring bisexual forms of Lacerta saxicola, producing hybrids that always are sterile triploid (3n=57) females. Therefore, application of geographical and physiological species criteria to parthenogenetic forms allows to regard them as independent species. However, geographical .and physiofogiical criteria are based on the effect of reproductive isolation mechanisms whose realization is possible upon the occurrence of both sexes in the population. Thus, both criteria should not be unconditionally applied to parthenogenetic and agamic species consisting of females alone. However, the absence of males also plays the role of a peculiar isolation mechanism which maintains the integrity of parthenogenetic species. Due to this, parthenogenetic forms of Lacerta saxicola Eversmann which can be easily distinguished in the morphological respect should be considered as independent agamic species: Lacertra armeniaca Meh., L. dahli Dar., L. rostombekovi Dar. and L. unisexualis Dar.

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The paper provide information on the current status of the type specimens of Darevskia (saxicola) complex as they were in December 2011. I order to stabilize nomenclature of this complex, the neotype of Lacerta saxicola Eversmann, 1834, lectotypes of Lacerta saxicola brauneri Méhely, 1909 and Lacerta saxicola lindholmi Szczerbak, 1962 are designated. History of description of these taxa is discussed.

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To study the habitats and potential spread of the rock lizards from the Darevskia (saxicola) complex, the Maxent 3.3.3e program was used. Our analysis has found that currently in the Ciscaucasia (on the Stavropol Upland) there are no favorable climatic conditions for habitation of D. saxicola but it can be found on the territory of North Ossetia-Alania (in the Digorskoe Gorge), there are no clear boundaries between the habitats of D. b. brauneri and D. b. darevskii. This is further evidence in favor of their conspecificity; the habitat of D. [brauneri] szczerbaki has been stable in recent decades and no significant movement of its borders has taken place, in Abkhazia there is very limited territory suitable for habitation of D. b. myusserica. Bioclimatic factors affecting the habitat of the lizards from the complex were analyzed.

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1. Sexual dimorphism and geographical variability in signs of external morphology are revealed in the species of the studied complexes of rock lizards. The most important for their diagnosis are the features of folidosis of the head. Interspecific and intraspecific differences in morphometric features, color and pattern of the body were detected. 2. In the phylogenetic analysis, the Darevskia (caucasica) samples were divided into the group of the northern slope of the Central Caucasus (D. caucasica), the East Caucasus (D. daghestanica), the Western Caucasus (D. alpina), the northern slope of the Central and Western Caucasus (D. alpina ). Reliable identification and significant genetic differences of D. caucasica and D. daghestanica samples favor their attribution to different species. Within D. alpina two groups with high genetic differences (distances) were identified. Inside Darevskia (saxicola) two lines are described, one of which is connected with the Crimea and the Western Caucasus (D. lindholmi-D. Brauneri), and the second - with the North and West Caucasus (D. saxicola-D. Szczerbaki). 3. As a result of the nomenclatural revision, the lectotype Lacerta praticola pontica, L. saxicola brauneri, L. saxicola daghestanica, L. saxicola lindholmi, and the neotype L. saxicola; Specific locations, years and authors of the description of a number of taxa have been refined. 4. It has been established that the studied supernumerary complexes include the following valid taxa: Darevskia (praticola) - D. pontica pontica, D. pontica hungarica, D. praticola praticola, D. praticola hyrcanica, D. praticola loriensis; Darevskia (caucasica) - D. alpina, D. caucasica caucasica, D. caucasica vedenica, D. daghestanica; Darevskia (saxicola) - D. brauneri brauneri, D. brauneri myusserica, D. saxicola, D. szczerbaki, D. lindholmi. Lacerta saxicola darevskii is regarded as a minor synonym for Lacerta saxicola brauneri. 5. For the first time accurate inventories of findings (1,443 locations) and GIS maps of species arealways were compiled, reliably characterizing the features of their distribution; The structure of areals, the current trends of their changes and the factors influencing the distribution of rock lizards are revealed. It is recommended to carry out conservation measures of international and national level in order to preserve the populations of D. alpina, D. brauneri myusserica and D. szczerbaki, especially in typical territories. 6. The first and only reliable paleontological find of Darevskia sp. - Darevskia sf. brauneri from the Pleistocene site, the caves of Mezmaiskaya. A hypothesis is proposed for the formation of species of complexes, according to which the isolation of their common ancestor from a hypothetical form it is logical to associate with the Miocene and Pliocene boundaries, or with the Cimmerian century of the Pliocene, 153 and the radiation of modern species - with the Akchagylian age, corresponding to the Piacenza and Gelazian tiers. The parallel formation of several Darevskia (praticola) and Darevskia (caucasica) lines around the Greater Caucasus is assumed in the early Pliocene. At the end of the Pontian age, the primary area of ​​the ancestor of the modern forms of Darevskia (saxicola) and the subsequent formation in the Mediterranean landscapes of the Black Sea coast, along the surviving cliff, D. szczerbaki, and the cuestas of the North Caucasus - D. saxicola, could have occurred at the periphery of the Caucasian peninsula.

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A reptile catalogue of the Zoological Museum, Stavropol State University, is presented. 1,019 specimens of 88 species collected within the territories of Russian Federation, Ukraine, Georgia, Kazakhstan, Uzbekistan, Turkmenistan, and some other areas are listed. The history of the Museum and the formation of its collection are given.

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The results of statistical analysis of morphological characters and the study of the cytochrome b gene variability (mtDNA) of rock lizards of the Darevskia (saxicola) complex distributed in the Caucasus and the Crimea, have shown significant differentiation within this complex, assuming specific independence of D. szczerbaki (Lukina, 1963) and subspecific status of recently described D. brauneri myusserica Doronin, 2011. According to our data, within D. brauneri (Méhely, 1909) only two forms of subspecific status can be distinguished, namely D. b. brauneri and D. b. myusserica. In our opinion, Lacerta saxicola darevskii Szczerbak, 1962 (= D. brauneri darevskii) should be considered to be a junior synonym of D. b. brauneri. The possible scenarios of range formation in the Darevskia (saxicola) complex are discussed.

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From the publisher: Before you a unique atlas-determinant. For the first time in one edition descriptions of all species of amphibians and reptiles living on the territory of Russia are presented, illustrated with beautiful color photographs. The book is written by E.A. Dunaev and V.F. Orlova, the famous Russian herpetologists, researchers of the Zoological Museum of Moscow State University named after MV Lomonosov. Lomonosov. A simple and accessible form of presentation, detailed descriptions of the external appearance, lifestyle and behavior, photographs reflecting all variants of the variability of these animals, will help the curious reader to become more familiar with turtles, snakes, lizards, frogs and other representatives of the herpetofauna. And special keys will allow you to determine who you met on a country walk, in a city park, at a dacha or while traveling to different corners of our country. The book is of great interest not only for nature lovers, but also for specialists in herpetology and students of biological faculties of universities. It lists the amphibians and reptiles of the native fauna, reflects the latest changes in the taxonomy of these groups, lists species included in the Red Book of Russia and regional Red Books, lists reference and scientific publications on reptiles and amphibians of Russian fauna.

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A specially optimized restriction analysis of highly repetitive DNA elements, called DNA taxonprint, was applied for phylogenetic study of primates and lizards. It was shown that electrophoretic bands of DNA repeats revealed by the taxonprint technique have valuable properties for molecular systematics. Approximately half of taxonprint bands (TB) are invariable and do not disappear from the genomes during evolution or change spontaneously. Presumably these invariable bands are restriction fragments of dispersed DNA repeats. Another group represents variable taxonprint bands that differ even between closely related species. These variable bands are probably represented by tandem DNA repeats and could be used as species-specific markers. It was shown that taxonprint bands are independent characters since the appearance of a new taxonprint band does not change the previous band pattern. Phylogenetic reconstruction carried out on taxonprint data demonstrated that this approach could be of general utility for molecular systematics and species identification.

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Hybridization is a common evolutionary process with multiple possible outcomes. In vertebrates, interspecific hybridization has repeatedly generated parthenogenetic hybrid species. However, it is unknown whether the generation of parthenogenetic hybrids is a rare outcome of frequent hybridization between sexual species within a genus or the typical outcome of rare hybridization events. Darevskia is a genus of rock lizards with both hybrid parthenogenetic and sexual species. Using capture sequencing, we estimate phylogenetic relationships and gene flow among the sexual species, to determine how introgressive hybridization relates to the origins of parthenogenetic hybrids. We find evidence for widespread hybridization with gene flow, both between recently diverged species and deep branches. Surprisingly, we find no signal of gene flow between parental species of the parthenogenetic hybrids, suggesting that the parental pairs were either reproductively or geographically isolated early in their divergence. The generation of parthenogenetic hybrids in Darevskia is, then, a rare outcome of the total occurrence of hybridization within the genus, but the typical outcome when specific species pairs hybridize. Our results question the conventional view that parthenogenetic lineages are generated by hybridization in a window of divergence. Instead,

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Considering all costs associated to sex, sexual reproduction could be expected to be rare. However, we find sexual reproduction to be pervasive in nature, and only a reduced number of taxa are completely asexual. In fact, the variety of sex reproductive systems, mating techniques and traits, reveals that the organisms’ reproduction has been an expressive target of selection and adaptation throughout evolution. Sex is selected because it facilitates adaptation in complex and changing environments (Weismann, 1889), and this has been already shown empirically in several models. However, studies and theories developed concerning the paradox of sex will not be able to clarify the current distribution and abundance of asexual lineages in nature. The understanding of rates of origin and extinction of asexual lineages, together with the putative differential evolution rates in organisms with sexual or asexual reproduction, should be considered in the main “origin of sex” question. The model used here, the genus Darevskia, is the first true parthenogenetic vertebrate described. It is a highly speciose genus, currently including 27 species, of which seven are parthenogenetic and of hybrid origin. Only a few of the sexual species contributed for the hybridization events which originated the parthenogens clonal lineages, and these were directional: the maternal species were always Darevskia raddei or D. mixta, and the paternal always D. valentini or D. portshinskii. Using a set of microsatellite and mtDNA markers, we first start to study the role of hybridization in the origin of vertebrate parthenogens, the extent of gene flow with sexual relatives and the relation between hybridization, asexuality and polyploidy, in the context of the main theories of asexual evolution proposed to date. We focus on the D. unisexualis, D. uzzelli and D. armeniaca parthenogens and polyploid backcrosses found in sympatric locations between the parthenogens and its sexual parentals. We find that only specific parental pairs are responsible for the origin of vertebrate parthenogenesis, regardless of their phylogenetic distance. Despite the recurrent hybridization presently reported in Darevskia, asexuality originated multiple times but only in a single temporal event the past. Parthenogenetic females are capable of backcrossing with sexual males, but can only produce polyploid individuals with reduced fertility that do not contribute for gene flow between parthenogens and sexuals. The sexual reproduction machinery can be lost with time in the parthenogenetic hybrids, which are most likely the result of a post-zygotic reproductive barrier to gene flow in the speciation continuum of Darevskia sexual species. In order to reconstruct the phylogenetic inference of this group, a set of capture sequence probes were designed from a transcriptome de novo assembly. Phylogenetic inference reconstruction has been extensively used to ask several evolutionary biology questions. However, despite the exponential use of high-throughput sequencing technologies and its promising applications in phylogenetics, next generation sequencing (NGS) techniques are still far from being as widely used in this field as in other areas of evolutionary genetics. The workflow developed in this thesis has proved to be not only cost-effective, but also to produce a very high number of cross genome phylogenetically informative markers in this non-model species. Finally, taking advantage of the hundreds of cross genome markers developed, a species tree was constructed and within genus relationships evaluated. Introgression tests showed a widespread pattern of gene flow across the genus, not only recent between recently diverged clades, but also both recent and ancient gene flow between early diverging Darevskia clades. Several evidences of hybridization with gene flow between diverging clades have been found in other systems. However, such widespread patterns where introgression has happened between all major clades, and in a very high number of species pairs, are not frequently found. It is also found that gene flow is completely absent between parthenogens sexual parental pairs. In groups with sexual-parthenogenetic reproducing species, pre- or post-zygotic reproductive barriers could be more difficult to attain and hybrid asexuality can, thus, effectively create reproductive barriers between diverging sexual taxa, that were likely faced with recurrent secondary contact. The work presented in this thesis shows that only a few sexual species pairs were responsible for the origin of the parthenogens, that some species acted always as the maternal species and others always as the paternal, that the phylogenetic distances between each pair are included in a wide range interval, and that introgression is found widespread across the genus but absent between the parental pairs. Regardless of the reproduction type, this study also shows the relevance and depth of hybridization during divergence. As has been shown intensively, gene flow is common during speciation and clades divergence. However, it would be interesting to analyse other groups together and test if deep branch gene flow is found only in some exclusive groups (such as sexual-asexual genera) or if it is something more widespread. This work presented here moves Darevskia towards providing a model that can be used to understand the origin of asexuality in vertebrates. Here the basis for future studies about sex and asexuality is provided, so that a focus on the differential evolution rates genomes of both reproduction mode can be analysed.

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A maximum parsimony phylogeny of 14 Caucasian species of rock lizards, genus Ijicerta, subgenus Archaeolacerta, was constructed from mitochondrial cytochrome b and ATPase 6 partial gene sequences. Congruence analyses were carried out between the two genes. A synthesis of the data sets reveals three well supported monophyletic groups: (1) the caucasica group including (Lacerta derjugini ((Lalpina, L. clarkorum) (L. caucasica, L. daghestanica))); (2) the rudis group including (L. parvula (L. portschinskii (L. vakntini, L. rudis))); and (3) the saxicola group including (L. mixta (L. nairensis (L. saxicola, L. raddeijj). Despite the diagnosis of three groups, the placement of L. praticola as a basal taxon is uncertain, as are the relationships among the three groups. The mitochondrial DNA sequence data suggested prior hybridization between L. mixta and L. alpina and possibly between L. saxicola and L. nairensis. Lacerta raddei was resolved as a paraphyletic species on the mtDNA tree; this may result from either hybridization or random gene sorting.

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Karyotype data are presented for 12 species of lacertid lizards. Of these, 4 Acanthodactylus, 2 Eremias, 1 Ophisops, and 4 Lacerta have n = 19, with all chromosomes acrocentric. This is the typical lacertid karyotype, as reported in previous literature. One species, Lacerta parva, is quite different from all the others. The haploid number is 12, consisting of 7 pairs of metacentric macrochromosomes and 5 pairs of microchromosomes. The fundamental number (number of arms in the karyotype) is the same in L. parva as in all the other species studied. The possibility of female sex chromosomal heteromorphism is raised, but data are not sufficient to confirm this.

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Satellite DNA repeats were studied in Caucasian populations of 18 rock lizard species of the genus Darevskia. Four subfamilies (Caucasian Lacerta satellites (CLsat)I–IV) were identified, which shared 70%–75% sequence similarity. The distribution of CLsat subfamilies among the species was studied. All the species could be divided into at least 3 clades, depending on the content of CLsat subfamilies in each genome: “saxicola”, “rudis”, and “mixta” lizards. CLsatI was found in all studied species, but in very different quantities; the “saxicola” group contained this subfamily predominantly. The “rudis” group also contained CLsatIII, and the “mixta” group carried considerable amounts of CLsatII. The highest concentrations of CLsatI and CLsatII were detected in 2 ground lizards — D. derjugini and D. praticola, respectively. D. parvula predominantly carried CLsatIII. CLsatIV was found only in the Crimean species D. lindholmi. The distribution patterns of satellite subfamilies show possible postglacial speciation within the genus Darevskia. A hybrid origin of species that possess 2 or 3 CLsat subfamilies and important clarifications to the systematics of the genus are proposed.

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The Caucasian parthenogenetic rock lizards of the L. saxicola complex probably originated by hybridisation of some bisexual species. We verified this hypothesis using a new approach based on comparison of repetitive DNA characters, which produce species-specific patterns (named `taxonprint`). The method relies on restriction endonuclease hydrolysis of genomic DNA, with the following 32P-end labeling and polyacrylamide gel electrophoretic separation of the fragments. The parthenogenetic species L. armeniaca and L. dahli possess specific features both of putative maternal L. mixta (in some taxonprints), and of paternal L. valentini (in other ones), whereas L. portschinskii and L. rudis also could be the paternal species. Parthenoclones L. unisexualis and L. uzzelli have the specific DNA features in several taxonprints of L. raddei or L. nairensis (which cannot be discriminated by taxonprints) which are supposed to be maternal species for L. unisexualis and paternal ones for L. uzzelli. The specific features of L. valentini (or L. portschinskii) are observed in L. unisexualis; L. uzzelli has not been investigated in this respect. Parthenogenetic L. rostombekovi also possesses properties of L. raddei or L. nairensis, but we have not be able to find any features of supposed paternal L. portschinskii in this parthenogenetic species.

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Multiple band patterns of DNA repeats in the 20–500-nucleotide range can be detected by digesting genomic DNA with short—cutting restriction endonucleases, followed by end labeling of the restriction fragments and fractionation in nondenaturing polyacrylamide gels. We call such band patterns obtained from genomic DNA ‘‘taxonprints’’ (Fedorov et al. 1992). Here we show that taxonprints for the taxonomic groups studied (mammals, reptiles, fish, insects—altogether more than 50 species) have the following properties: (1) All individuals from the same species have identical taxonprints. (2) Taxonprint bands can be subdivided into those specific for a single species and those specific for groups of closely related species, genera, and even families. (3) Each restriction endonuclease produces unique band patterns; thus, five to ten restriction enzymes (about 100 bands) may be sufficient for a statistical treatment of phylogenetic relationships based on polymorphisms of restriction endinuclease sites. We demonstrate that taxonprint analysis allows one to distinguish closely related species and to establish the degree of similarity among species and among genera. These characteristics make to use this phenomenon for phylogenetic aims (Cooke 1975; Christie and Skinner 1979; Brown and Dover 1979, 1980; Elizur et al. 1982; Hembelen et al. 1982; Buckland 1983; Dandieu et al. 1984; Lima de Faria et al. 1984; Shubina and Mednikov 1986; Turner et al. 1991), but this approach has not been extensively applied. In this paper we have reexamined the problem on the basis of our own preliminary results, partly published in Russian journals, confirming the ideas about possible correlations between evolution of DNA repeats and speciation (Fedorov et al. 1992; Grechko et al. 1993; Potapov and Ryskov 1993). We have demonstrated that the modified method designated ‘‘DNA taxonprint’’ (Fedorov et al. 1992) can be effectively used for revealing species—specific genomic markers in different taxonomic groups. These taxonprint markers are found to be helpful for evaluation of the degree of species similarity between species, genera, and families. More than 50 species from different taxa were studied.

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DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.

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Amphibia and Reptilia collections at the Zoological Museum, University of Istanbul (ZMUI) were studied and revised. The samples were collected from 1921 until today. The localities of most species are found in the different regions of Turkey. The collection have 75 species of which 21 species are amphibians and 54 species are reptiles. With this study a zoological collection belong to ZMUI was given for the first time.

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Data on the reproduction of the rock lizard (Darevskia saxicola (Eversmann 1834)) in the gorges of Bol’shaya Laba (Karachay-Cherkessia) and Malaya Laba (Krasnodar Territory) rivers are presented. Gravid females of this species were captured in nature and transported to the laboratory. The animals were kept in captivity until oviposition. All eggs were obtained between July 20 and August 20. Breeding females had a body length of 42.5–60.1 mm. Hatches comprised 1–4 eggs 11.2–18.0 × 5.8–8.5 mm in size and weighing 0.185–0.535 g. Incubation in the laboratory lasted 38–47 days. Young lizards after hatching had a body length of 21.6–30.8 mm, tail 39.9–60.4 mm long, and weight of 0.375–0.635 g. A comparison of the reproductive characteristics of D. saxicola and syntopic species (D. derjugini (Nikolsky 1898) and D. pontica (Lantz et Cyren 1919)) was carried out. The late oviposition and relatively low fertility in the rock lizards are compen- sated for by the large size of eggs and newborn lizards.

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Data on the reproduction of the rock lizard (Darevskia saxicola (Eversmann 1834)) in the gorges of Bolshaya Laba (Karachay-Cherkessia) and Malaya Laba (Krasnodar Territory) rivers are presented. Gravid females of this species were captured in nature and transported to the laboratory. The animals were kept in captivity until oviposition. All eggs were obtained between July 20th and August 20th. Breeding females had a body length of 42.5–60.1 mm. Hatches comprised 1–4 eggs sized 11.2–18.0 × 5.8–8.5 mm and weighting 0.185–0.535 g. Incubation in the laboratory lasted 38–47 days. Young lizards after hatching had a body length of 21.6– 30.8 mm, tail 39.9–60.4 mm long, and weight 0.375–0.635 g. A comparison of the reproductive characteris- tics of D. saxicola and the syntopic species (D. derjugini (Nikolsky 1898) and D. pontica (Lantz et Cyren 1919)) was carried out. The late oviposition and relatively low fertility in the rock lizards are compensated for by the large size of eggs and newborn lizards.

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To study the molecular genetic relationships and correlate them with the taxonomy within the complex of lacertid lizards of the genus Darevskia, the locus analysis of the copies of the SINE-type repeat (Squam1) specific for the order Squamata was used. It was demonstrated that one of the loci (No. 34) contained the Squam1 copy insert in all species and subspecies of the examined genus. SINE allelic copies in some of the loci contained large indels and specific sets of mutations. The allelic variant M (medium, about 340 bp) was found most frequently; it was detected in all subspecies of D. saxicola (saxicola, darevskii, szczerbaki, lindholmi) and in most of the other species of the genus. Two species, D. derjugini and D. praticola, differed from the other species in the presence of long (L) and short (S) alleles. The longest allele was characteristic of the D. derjugini population from the Northern Caucasus (L, 379 bp, ssp. silvatica), while the shortest allele (97 bp) united the derjugini and barani subspecies. The second allele S (279 bp) characterizes the subspecies D. praticola praticola, some individuals of which also carry allele M. The second subspecies, D. p. pontica, contains allele L2, which differs from all other medium alleles in the presence of strictly specific short indel. In addition to apomorphic indels, the specificity and mutation distribution patterns among the Squam1 alleles were also examined. An analysis of the NJ tree indicated the concordance between morphological and molecular genetic characters of the species derjugini, praticola, and saxicola. Furthermore, four subspecies of D. saxicola were much closer to each other than the subspecies within the first two species; D. d. silvatica and the group of D. d. derjugini + barani were clearly separated. It cannot be excluded that populations from Azerbaijan and Serbia can be treated as the independent subspecies of D. praticola.

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An overview of the cases of introduction (both accidental and intentional) of amphibians and reptiles in the Caucasus and Crimea is provided. The introduction cases are systemized in accordance with the goals and ways of introduction and with regards to the expected scale of the impact and its nature. No negative consequences of the introduction of amphibian and reptile species in the Crimea and Caucasus are currently observed on any of the known examples. No cases of species naturalization, which the “invasion” term can be applied to, have been revealed. The importance of allochthonous populations as experimental sites to study microevolutionary processes and ecological adaptations of the species in new environmental conditions is discussed.

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In this study, 17 reptile and amphibian species were recorded from 19 different localities in the Ordu-Giresun region. Of these, 3 belong to the urodelans species group, 6 belong to anurans, 4 belong to lizards and 4 belong to snakes.

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A survey of the caucasian rock-dwelling lizards is given with an account of the areal inhabitated and a short descrip- tion of most kinds. The author got from friends in the DDR some specimens of Lacerta saxicola brau- neri, Lacerta rudis obscura, Lacerta ar- meniaca and a Lacerta raddei. All these lizards were kept in outside terrariums of different sizes. They thrived well, some species even reproduc- ting in fairly large numbers. The good results are attributed to the probable resemblance of the dutch climate with that from their coutry of origin, as well as to the fact that the sun had free access to the terrariums. Reproduction also took place when the animals were hibernated without a real winter-sleep. Some remarks concern the systematic relationships of the species mentioned.

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Anatomical-morphological features of the postcranial skeleton of nine taxa of the rock lizards of the genus Darevskia Arribas, 1999 belonging to the Darevskia (caucasica) and Darevskia (saxicola) complexes were studied for the first time on representative samples. Specimens with the rib formula 3+1 were found in both complexes (previously, only two variants were known for the genus Darevskia 3+2 and 3+3). Differences between supraspe- cific groups and their members were described with the use of type specimens or topotypes of each taxon. The studied complexes are differentiated by the range of variability of presacral vertebrae (in the D. (caucasica) their minimum number is 26, in the D. (saxicola) – 27), the number of the posterior presacral vertebrae with ribs (the average value of this trait in the D. (caucasica) is less than 6, in D. (saxicola) – more than 6). B-type of the tail vertebrae in 5 times more common in the D. (saxicola) than in the D. (caucasica). In turn, the latter group is cha- racterized by a larger share of individuals with ossified ribs at the third cervical vertebra and a greater variability of the sternal-xiphisternal costal (rib) formula. Skeletal morphology of D. alpina indicates its greater similarity with representatives of the D. (caucasica) complex. A certain originality of D. c. vedenica was recorded. The study of D. saxicola specimens from the zone of sympatry with D. c. caucasica (Chegem Gorge) did not reveal their simi- larity with the latter, except for the presence of two variants of the sternal-xiphisternal formula. Cryptic forms within D. lindholmi differ at least in two traits: the minimum number of posterior presacral vertebrae with short ribs and the presence of individuals with caudal vertebrae of B-type.

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The study of genome size variation in cells of vertebrates using the DNA flow cytometry makes it possible to precisely determine polyploid individuals, which is extremely important when studying the processes of reticulate speciation. In addition, in many groups of vertebrates, closely related species often differ in the nuclear DNA content. Therefore, the purpose of our study was to explore the variability of genome size and ploidy in populations of 29 species and subspecies of mountain lizards of the genus Darevskia, as well as their hybrids. As a result of the study, the range of variability in individuals of different ploidy (91% of diploid and 9% of triploid individuals) was established not to overlap. Among diploid species, no correlation was found between the nuclear DNA content and phylogenetic relationships, geographic coordinates, altitude, average annual temperatures and precipitation. Representatives of all studied species complexes (with the exception of D. adjarica) had approximately the same limits of variability. Two species (D. derjugini and D. saxicola) show significant intraspecific variability. Comparisons of the genome size of parthenogenetic and bisexual species generally revealed no noticeable differences between them. Studies of triploid hybrids have shown that their genome size as a whole roughly corresponds to the sum of the average size of the diploid genome of the maternal parthenogenetic species and the haploid genome of the paternal species. The variability of genome sizes within samples of triploid hybrids was on average slightly higher than in most parthenogenetic species, but some lower than in bisexual species. The paper discusses peculiarities of reticulate speciation in this group of animals.

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Diversity at 35 allozyme loci was examined in nine populations of the taxa currently included in Darevskia saxicola. The data support the recognition of the subspecies Darevskia s. saxicola and D. s. lindholmi as species. The remaining subspecies D. s. brauneri, D. s. darevskii and D. s. szczerbaki are conspecific although they can no longer be referred to Darevskia saxicola, but must be referred to Darevskia brauneri, whose name has priority. Considerable population substructuring was observed among the subspecies of Darevskia brauneri.

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Investigated the albumins from 31 species and subspecies of the genus Lacerta s.1. by means of the immunoloical techniue Micro-Complement-fixation. Dendrograms were computed by means of the ktch-MargJiash algorithm. Moreover the taxa of the Lacerta saxicola and the Lacerta danfordi comlexes were analyzed by electrophoresis concerning 14 genetic loci. The results were discussel together with karyological and genitalmorphological data so far known yielding the following relationships: Lacerta graeca shows a closer relationshi to the subgenus Podarcis as does the quite closely related species air L. dugesii and L. perspicikata. Lacerta Iaevis probabl is connected closely to Podarcis, too. Lacerta parva shows a closer relationship to the `small izards` than to the sub-genus Lacerta s. str. A clear decision concerning the position of the Lacerta danfordi complex according to chemosystematical data was not possible. It is isolated from the other species probably about as long as L. puma. Biochemical differences between its `species` are too small to confirm their taxonomical revalorisation by Eiselt and Schmidtler Within the Lacerta saxicola complex the investigated taxa show a close relationship. Lacerta praticola and L. derjugini were definitly assigned to this group.

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Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk. Location World-wide. Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions. Results I obtained body size data for 99% of the world`s lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non-aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed. Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.

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The phylogeny of 12 species of Caucasian rock lizards and ground lizards, genus Lacerta, subgenus Archacolacerta, was determined from allozyme data. Three additional species of Lacerta Sorlned the outgroup. Of the 35 loci resolved, 20 were phylogenetically informative. Permutation tail probabilities calculated using Hennig86 and RandomCladistics and g, statistics obtained in PAUP revealed significant pattern in the data. Our analysis resolved 12 most parsimonious trees; among which were three well-supported monophyletic groups: I) (Lacerta alpina, L. caucasica, L. daglzestanica) [the L. caucasica group]; 2) (L. mixta. L. saxicola. L. raddei-nairensis) [the L. saxicola group]; and 3) (L. parvlila, L. portschinskii (L. valentini, L. rudis)) [the L. ruciis group]. The second and third groups formed a clade. The ground lizards L. praticola and L. derj~cginwi ere resolved as ingroup members. but their placement in the globally most parsimonious trees was ambiguous. A preferred tree was chosen by alternately pruning L. praticola and L. devugini from the tree. In doing so, L. praticola was resolved as the sister taxon of the L. calccasica group; and L, derjligini was resolved as the sister taxon of the clade containing the L. snxicola and L. rrldis groups. Our results are somewhat congruent with the morphology-based phylogeny of Darevsky (1967).

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A phylogeny of Caucasian rock lizards (genus Darevskia, formerly Lacerta) was reconstructed using mitochondrial DNA sequence and allozyme data. All 15 bisexual species grouped into three major clades: the caucasica, saxicola and rudis clades. Unisexual Darevskia originate from inter-clade hybridization, never from within clades. Only two clades, the caucasica clade and the rudis clade, were involved in forming unisexuals; the saxicola clade was never involved. Furthermore, the hybridization is directional in that the caucasica clade contributed only maternal parents and the rudis clade only paternal parents. The formation of unisexual species is best explained by sexually directional phylogenetic constraints. We hypothesize that the causative agents are likely to be genes linked with the sex chromosomes within the parental sexual species.

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In the autumn of 2015 current distribution of the lizards of Darevskia (saxicola) complex, introduced in 1960s in Zhytomyr administrative region of Ukraine was studied. During the period from introduction until now Darevskia (saxicola) complex has spread along the left bank of the Teteriv River on the territory about 3.7 km lengths. Beside this was found an isolated population of this species, which located about at 4 km direct distance or 8.5 km shore line distance from the main area of occurring this species. It is supposed that this secondary introduction took place occasionally or intentionally due to holidaymakers (e. g. climbers, fishermen). The publication provides description of the lizards on 15 morphological characteristics and information on the results of spatial modeling potential distribution the rock lizards.

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Many animals display complex colour patterns that comprise several adjacent, often contrasting colour patches. Combining patches of complementary colours increases the overall conspicuousness of the complex pattern, enhancing signal detection. Therefore, selection for conspicuousness may act not only on the design of single colour patches, but also on their combination. Contrasting long- and short-wavelength colour patches are located on the ventral and lateral surfaces of many lacertid lizards. As the combination of long- and short-wavelength-based colours generates local chromatic contrast, we hypothesized that selection may favour the co-occurrence of lateral and ventral contrasting patches, resulting in complex colour patterns that maximize the overall conspicuousness of the signal. To test this hypothesis we performed a comparative phylogenetic study using a categorical colour classification based on spectral data and descriptive information on lacertid coloration collected from the literature. Our results demonstrate that conspicuous ventral (long wavelength-based) and lateral (short wavelength-based) colour patches co-occur throughout the lacertid phylogeny more often than expected by chance, especially in the subfamily Lacertini. These results suggest that selection promotes the evolution of the complex pattern rather than the acquisition of a single conspicuous colour patch, possibly due to the increased conspicuousness caused by the combination of colours with contrasting spectral properties.

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As was shown earlier with reference to several animal taxa, characterization of highly repetitive DNA sequences by their susceptibility to restriction endonucleases (DNA taxoprinting) does not exhibit any differences between species populations. This method was used in our study on the populations and subspecies of some lacertid lizards: Lacerta raddei, L. saxicola, L. rudis, and L. agilis. These features were shown to be practically indistinguishable in two Armenian populations (Gosh and Khosrov) of L. raddei; the third population (Egegnadzor) demonstrates some differences. Two subspecies of Caucasian L. rudis and five subspecies of L. agilis inhabiting distant parts of the extensive range have minimal dissimilarities in the taxoprints studied. Comparison of L. saxicola subspecies, three of which inhabit the Caucasus and one the Crimea (L. s. lindholmi) presented no significant differences in any of the taxoprints of Caucasian subspecies studied up to now, while Crimean subspecies is significantly dissimilar, not less than other species are dissimilar from each other. It was suggested that L. s. lindholmi may be considered as an individual species. The molecular characteristics of genomic DNA used seem to be useful in animal taxonomy.

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Genomic DNA polymorphism in lizards was analyzed by means of PCR-based random amplification of polymorphic DNA (RAPD-PCR). Several species, subspecies, and populations of the species complex Lacerta saxicola from the Caucasus were tested. Intrapopulation RAPD polymorphism was not detected in parthenogenetic species and was insignificant in bisexual species. RAPD patterns of Caucasian allopatric L. saxicola populations were almost identical and corresponded to the level of intrapopulation polymorphism. However, they differed from the RAPD patterns of the Crimean population of the same species, which separated earlier. A possibility of using RAPD markers to distinguish between taxonomic concepts of population and subspecies is discussed.

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The last species list of the European herpetofauna was published by Speybroeck, Beukema and Crochet (2010). In the meantime, ongoing research led to numerous taxonomic changes, including the discovery of new species-level lineages as well as reclassifications at genus level, requiring significant changes to this list. As of 2019, a new Taxonomic Committee was established as an official entity within the European Herpetological Society, Societas Europaea Herpetologica (SEH). Twelve members from nine European countries reviewed, discussed and voted on recent taxonomic research on a case-by-case basis. Accepted changes led to critical compilation of a new species list, which is hereby presented and discussed. According to our list, 301 species (95 amphibians, 15 chelonians, including six species of sea turtles, and 191 squamates) occur within our expanded geographical definition of Europe. The list includes 14 non-native species (three amphibians, one chelonian, and ten squamates).

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Elements of the herpetofauna are in the author`s opinion of great importance for the zoogeographical division of the Palaearctic region. It is suggested that this region be divided into the following four subregions: 1/ arctic, 2/ boreal Euro-Siberian, 3/ bo- real Himalayan-Manchurian and 4/ Mediterranean-Central-Asiatic.

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The rock lizards of the Darevskia saxicola complex are found exclusively in the west of the Greater Caucasus and in southern Crimea. The earliest split within this group occurred between D. saxicola from the northern and D. brauneri from the southern slopes of the Greater Caucasus, followed by the split between D. brauneri and the Crimean D. lindholmi, and the expansion of D. saxicola to the westernmost slopes of the Greater Caucasus. We collected nominal D. brauneri from the two easternmost populations of the species range: the valleys of the Tekhuri and Enguri rivers in Georgia. Analysis of mitochondrial DNA showed that the lizards from both valleys are deeply differentiated from each other and from previously characterised D. brauneri . Lizards from Tekhuri keep a basal position within the clade D. saxicola (excluding D. praticola), and lizards from Enguri are closer matrilineally to the northern Caucasian D. saxicola than to either of the populations of nominal D. brauneri. Tekhuri lizards have broader heads and more small scales between the inner and outer rows of supraoculars than the other populations of the group. We suggest that the taxonomy of the group requires revision, considering the multiple deeply divergent mitochondrial lineages and introgressive gene flow between the continental populations of nominal D. saxicola and D. brauneri. The D. saxicola complex in the Caucasus resembles a `ring species` arrangement as described for other taxa and mountain regions.

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The lizard Lacerta saxicola inhabits all oak forests on valley slopes in the investigated area. It occurs permanently in beam-beech forests on valley bottoms only in gaps forming as a result of falling trees. The lizard population in the gaps amounted to about 8% of the total one in the forests. The population density in gaps is several times as high as on slopes. The gap populations are stable closed settlements of territorial adults. The juveniles are forced out of the gaps to valley slopes. The abundance of food and shelters on valley bottoms is much higher than on its slopes. In forests, the conditions for lizards are unfavorable due to lower temperature. They can bask only in gaps. The lizards from slopes visit periodically valley bottoms, but in gaps they do not leave their settlements. In hot periods lizards migrate from slopes to valley bottoms. This situation is repeated every year. In the period of mass miggration in gaps lizard populations remain closed and permanent by their composition. Settled males from gaps mate with local settled females as well as with those from slopes coming to gaps. Males from slopes have then opportunity to mate only with females from the same biotope.

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Changes of the space use pattern by males and females of the rock lizard (Lacerta saxi- cola) and the process of establishing socio-sexual interrelations are described. The or- dered pattern of space use enables the development of a complicated system of interre- lations between males and females but it does not completely determine the structure of this system. Our long-term observations make it clear that, contrary to the generally ac- cepted conceptions, the system of intersexual relations does not directly refer to repro- ductive success. Whereas females, keeping in touch with males, obtain the possibility to use the habitat resources in the best way and the possibility to choose their sexual part- ners, males do not obtain this. Aggressive males, defending their territories, obtain the possibility of communicating with females, which is much more attractive than copula- tion itself.

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The territory of male was formed around centers of the activity of settled females. A young male obtained an access to these centers only in the case when the territorial male died and new centers had been formed within his home range. A direct aggression towards other males appeared when the male obtained an access to regular contacts with females. This access might be unlimited when all or the majority of the centers of female activity were within the male territory (α-males). The access might be partly limited when the territory contained only one or few such centers (β-males). In α-males, the obligatory aggression of high intensity was provoked by the behavior of an intruder, which as improper for its status, or by the attempts to rape any female. The intruder`s courting to female provoked aggression only in 80% of instances. The most intense aggression of β-male was caused by the intruders` attempts to court the female that was preferable for him or by his invasion into the her activity center. The territories of males with different status proved to differ in both the motivation of their behavior and their position in a system of social relationships and use of resources.

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A group of marked lizards was observed for three years in natural conditions. In every marked individual, the last phalanx was amputated, and its age was determined by the bone layers. All males come to maturity in the 3d year of life independently of their coloration. Different males become green in the 3d, 4t h, 5t h or 6t h year of life. All green males were settled and displayed a territorial behavior. The territorial behavior in brown males was never recorded. The mails, which gained green coloration in the 3d year, had small and regular increment. The males gained green color in the 4`1 year or later had a greater increment in the 1s t eyar of life and small growth in the future. The absence of green coloration does not prevent brown males from aggression of green individuals. The size and quality of home range and the degree of pressure of conspecifics are not related to the brightness of coloration of possessor of home ranges. However, females in encounters with bright-green males take initiative in contacts much more frequently, than in those with light-green ones. In contacts with brown, ales, the initiative of females was never ovserved. In home ranges of bright-green males, the density of females` population was 4 times as high as in home ranges of light-green males.

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It is discovered that green colour of the back of adult male is determined with the time of exposing to sun. In beech-hornbeam forest (shady habitat), not all the males have green coloration and coloration of a male is an unambiguous sign of his territorial status. Coloration is one of the main communicative signals, which regulate lizards behaviour during both male-female and male-male interactions. The result of this regulation is the stable long-term system of interrelations, which provides: a) different access to habitat resources and mate in the males of different statuses; b) free mate choice in females; c) reduction of frequency and intensity of aggressive confl icts between the members of society. In oak forest (sunlight habitat), all adult males have green coloration which has only one meaning — it serves for identifi cation of conspecifi c sex. An access of males of different status to resources and mate is weakly differentiated and in great extent depends on a chance. Frequency and intensity of aggressive confl icts are great. These interactions are very important for formation of social structure there.

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A new species of rock lizards of the genus Darevskia Arribas, 1999 — Darevskia arribasi sp. nov. is described from vicinity of Ertso Lake in South Ossetia. In comparison of Darevskia arribasi sp. nov. with other species of the “saxicola-brauneri” complex [D. lindholmi (Szczerbak, 1962), D. saxicola (Eversmann, 1834), D. brauneri (Mehely, 1909) (including D. b. brauneri, D. b. myusserica Doronin, 2011, D. b. darevskyi (Szczerbak, 1962), and D. szczerbaki (Lukina, 1963)], the new species is characterized by the smallest body size among all representa- tives of this complex Lxx — 55 – 62 mm; }} — 58 – 63 mm. The number of scales around the middle of the body (Sq.) in D. arribasi sp. nov. (52 – 58, more often 54 – 56) also significantly less than for D. saxicola sensu lato in generally. Along the throat from ear to ear fold there are 36 – 42 scales, less than in D. saxicola sensu lato. Two large preanal shields present in front of the anal shield in half of the individuals examined; in a quarter of in- dividuals on both sides of the small central preanal there are by one enlarged lateral preanal shield, in another quarter all preanal shields are of the same size. Central temporal (Massetericum) always developed; always six pair of mandibulars, three in contact; collar straight, extremely rarely slightly serrated; ribs on the caudal scales are expressed to some extent; rostral usually separated from frontonasal by nasal shields. The coloration of adult males is olive-lettuce, or olive-brown from above; adult females are grey-brown; juvenile coloration coffee-brown with the blue and black spots on the edge of the outer ventral shields at the middle of trunk. Taxonomical status of new species is well supported with result of analysis of the mitochondrial cytochrome b data.

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Five proteins (mannosephosphate isomerase, glucosephosphate isomerase, lactate dehydrogenase, creatine kinase, hemoglobin) were examined for six bisexual Transcaucasian taxa related to Lacerta saxicola (Lacerta valentini, L. portschinskii, L. raddei raddei, L. r. nairensis, L. mixta, L. parvula) and four unisexual taxa (Lacerta armeniaca, L. dahli, L. rostombekovi, L. unisexualis). Heterozygosity in the bisexual taxa was very limited for this sample of proteins. All individuals examined of each unisexual were heterozygous at loci specifying at least two of these proteins (L. armeniaca: mannosephosphate isomerase and creatine kinase; L. dahli, mannosephosphate isomerase and hemoglobin; L. rostombekovi, glucosephosphate isomerase, mannosephosphate isomerase and hemoglobin; L. unisexualis, glucosephosphate isomerase, mannosephosphate isomerase and creatine kinase). High levels of heterozygosity in unisexuals appears to result from the hybrid origin of the unisexuals. The exact combinations of alleles present in unisexuals would readily result from certain crosses among the bisexuals, and from no others. The first point supports the hypothesis that hybridization accounts for the observed heterozygosity, the second identifies, biochemically at least, the probable parents. On these biochemical grounds, L. armeniaca arose by hybridization of L. valentini and L. mixta; L. unisexualis from L. valentini and L. r. nairensis; L. rostombekovi from L. r. raddei and L. portschinskii; and L. dahli from L. portschinskii and L. mixta. With regard to altitudinal distribution, vegetational associations and geographic distribution, each unisexual species appears to be intermediate between its putative parental species. L. armeniaca, L. unisexualis and L. rostombekovi all live in drier situations than either parental species. L. dahli is an exception since it occupies slightly more moist habitats than L. portschinskii. Biogeographical considerations appear to place the age of L. unisexualis and L. rostombekovi at greater than 5000 years. It is possible that their occupancy of more extreme habitats than their parental species represents a relic ecology, reflecting the adaptations of the parental bisexual species when the unisexual species arose, rather than a weed habitat into which they moved to escape from competition with their parental species. The fixed heterozygosity of the unisexual species of Lacerta indicates that the restitution of somatic diploidy results either from a premeiotic endoduplication without cytokinesis, or from fusion of the female pronucleus with one of the second division meiotic products of the first polar nucleus. The first mechanism is inconsistent with the number of bivalents reported, the second is otherwise unknown in organisms in which the first polar nucleus becomes a polar body. The bisexual taxa are treated as several distinct species, four pairs of which are partly sympatric with little or no hybridization. Morphological and ecological differences between the other taxa are so great that there seems little question about specific distinctness. The degree of biochemical and morphological difference between these non-sympatric taxa is as great as that between those that do occur sympatrically without fusing. The formation of parthenogenetic species as a result of past hybridization between some pairs indicates a great selective disadvantage to these pairs of hybridizing, and is thus compelling evidence that the two members of each pair are not conspecific.