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TERRA TYPICA. Settlement Krasnaya Polyana in the south of the Krasnodar Territory, Russia. DISTRIBUTION. Endemic of the montane Caucasus. It is widely distributed in the western part of the Great Caucasus Range, on northern, western and south-western slopes of the Black Sea chain of the Great Range in the Krasnodar Territory, Abkhazia and north-western and south-western Georgia, to the canyons of the rivers Kodory and Ingury in lower and upper Svanetia in the east. The northern border of its distribution passes in the area of the middle current of numerous left tributaries of the Kuban River. The most northern habitats of the species are known in the vicinity of Goryachyi Klyuch on Psekupsa River and the vicinity of the town Maikop. The southern border of the distribution range stretches on the southern slopes of the Great Range and, gradually lowing down, on the western spurs of the Gagra range. It comes out to the sea in the south of the Krasnodar Territory and western Abkhazia, where it is sporadically recorded on the coast from the ALPINE LIZARD DAREVSKIA ALPINA (DAREVSKY, 1967) ARMENIAN LIZARD DAREVSKIA ARMENIACA (MEHELY, 1909) BRAUNER’S ROCK LIZARD DAREVSKIA BRAUNERI (MEHELY, 1909) 81 L A C E R T I D A E town Anapa in the north up to the canyons of the rivers Khoby and Riony in the south-west of Georgia. The nominative subspecies D. b. brauneri occurs in the south of the Krasnodar Territory, in Abkhazia and adjacent regions of Georgia. D. b. darevskii (Szczerbak, 1962) inhabits foothills of the western part of the Great Caucasus range, mainly within the Krasnodar Territory. A comparatively small range of the subspecies D. b. szczerbaki (Lukina, 1963) occupies a narrow strip of coastal rocky steeps, sporadically occurring along the coast of the Black Sea from the town Anapa in the north and for more than 100 km further to the south-east towards Novorossiisk. CONSERVATION STATUS. On the most part of the distribution range the number of the species is rather high and stable.

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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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In Georgia, 16 species of rock lizards of the Genus Darevskia are occuring (D. alpina, D. „armeniaca`, D. brauneri, D. caucasica, D. clarkorum, D. daghestanica, D. „dahli`, D. derjugini, D. mixta, D. nairensis, D. parvula, D. portschinskii, D. praticola, D. rudis, D. „unisexualis` and D. Valentini). Besides short presentations of the single species and hints on their distribution and habitats, also some systematic remarks are given.

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Georgia, its history and the nature are briefly described. This is supplied by a short overview of the country`s herpetological fauna, with particular attention to the especially diverse family of true lizards. At the end, brief description of our common trip to Georgia (May - Juny 2002) is given.

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Many regions of the former USSR still re- ceive little attention from Western visitors, herpetologists not excluded. For the Cau- casus Mountains this is all the more sur- prising since the area forms a zone where northern and Middle Eastern flora and fauna meet, and which is thus of considera- bly zoogeographical interest. Presently we find three larger states in the Caucasus: Georgia, Azerbaijan and Arme- nia. Turkey borders to the southwest, Iran to the southeast, and in the north it is the Russian federation. Various smaller parts claim independence. The Black Sea forms the natural western boundary, the Caspian Sea the eastern one. Herpetologically the Caucasus is historically well-known because of the work of Darev- sky and collaborators on the rock lizards (at the time still in the large genus Lacerta, now in Darevskia) and the discovery of the first case of parthenogenesis in vertebrates (DAREVSKY, 1967; DAREVSKII, 1978; DAREVSKY & KULIKOVA, 1961). Below we will concentrate on the results of our field trip to Georgia during the last two weeks of June 2003. Predictably, because of our common interest in the group, the lacertids will receive the most attention.

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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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The description of a new subspecies of rock lizards Darevskia saxicola complex, inhabited the territory of Pitsunda-Myussera Reserve and in the Gagra (Republic of Abkhazia) is given. Myussera lizard, Darevskia brauneri myusserica ssp. nov., differs from other taxa of the complex by the following combination of morphological characters: (1) large or very large masseteris scute; (2) interrupted series of ciliated grains between supraciliar and supraocular scutes; (3) presence of additional scutes lying on either side of occipital and interparietal scutes or splitting of the latter; (4) net color pattern on the back (in females fuzzy); (5) dominance of gray and light gray color in the coloration of the back of females; (6) white throat and belly. In addition, these subspecies differs by some peculiarities of biology: biotopical preference to coastal pebble conglomerates and a relatively low population size. Probably, the taxon was formed during the Pleistocene. Formation of the coastal plains of Pitsunda Peninsula by alluvial and marine accumulation in the Late Neopleistocene – Holocene divided the area of Myussera lizard into the Gagra and Myussera plots. This area is situated within the Black Sea refuge of Eastern Mediterranean species of herpetofauna.

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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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The greatest number of reptiles having melanism (the number of specimens and species) have been recorded in the Caucasus in populations of five Darevskia species: Darevskia alpina, D. brauneri, D. lindholmi, D. pontica and D. rudis. The occurrence of melanistic specimens in reptiles can be explained by the influence of geochemical anomalies, thermoregulatory adaptation, isolation, and occurence in the periphery of their range. Melanism of D. lindholmi appears to depend on geochemical factors. The melanistic specimens of D. rudis, D. pontica and D. brauneri were found in a city on the northeastern boundary of their range. D. alpina was recorded in the mountains on the eastern periphery of the range.

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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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New records of Darevskia alpina from the vicinities of Zemo Bachiani Village in South Ossetia are presented and discussed: D. brauneri from the place of the confluence of the Luhunistskale River into the Rion River in Georgia and vicinities of Ertso Lake in South Ossetia, D. pontica from the vicinities of Slavyansk-na-Kubani, Timashevsk, Novotitarovskaya Village, Poltavskaya Village and Burakovsky Village in Krasnodar Region, D. portschinskii nigrita from the vicinities of Kizilkilisa Village in Georgia, D. praticola from the vicinities of Tskhinval, Acrisheu Village, Malaya Gufta Village, Hetagurova Village and the southern slope of the Zonkar Ridge in South Ossetia. Information on an attempt to introduce D. pontica into the territory of Rostov-on-Don is provided.

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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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New data on the distribution of four species of lizards (Darevskia brauneri, D. daghestanica, Lacerta agilis ioriensis, L. strigata) in the Caucasus are presented.

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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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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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This publication on the herpetofauna oft he Caucasus is a compilation oft wo articles published in 1967 and 1970 (Zool. Abh. SStaatl. Mus. Tierk. Dresden 29, pp. 85-93 and Aquarien terrarien 17, pp. 120-123.

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A remarkable hindleg regeneration was observed in a Sharp-snouted Rock Lizard (La- certa oxycephala). The hindleg was replaced by a scaled elongate and conical appendage. The moving lizard kept the regenerate motionless. The specimen apparently was able to catch enough food in nature. Comparable regenerations were found in Caucasian Rock Lizards, in two L. saxicola brauneri and in one L. raddei raddei. Always a hindleg was rege- nerated.

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The life and work of the Alsatian Louis Amédée Lantz (1886-1953), a pioneer of European herpetology. A trained and professional research-chemist, the Alsatian Louis Amédée Lantz was an active and particularly dynamic herpetologist. His residence of nearly twelve years in Russia enabled him to make several field trips to regions such as the Caucasus and Iran. He always carried with him a home made telescopic net that he used to catch lizards. That net became symbolic of Lantz for most people around him. He realized significant herpetological collections mainly deposited in the Museum of the Zoology Institute of the Russian Academy of Sciences in St. Petersburg. Soon after the Russian Revolution in 1917, he had to leave the country and stayed for a few years in France before joining England. During this period, Lantz visited the French Pyrenees and the Mediterranean border (Hyeres Islands, Bandol), enabling him to discover for the first time oviparous populations of common lizard and also to describe a new species of lizard from the French Pyrenees. Lantz is also at the origin of the discovery of unisexual reproduction by parthenogenesis in reptiles. His collaboration with the Swedish amateur herpetologist Otto Cyrén, whom he met through the German herpetologist Willy Wolterstorff, was particularly successful throughout his career. Lantz died at the age of 66 years, in Switzerland, as he was about to join the team of herpetologists at the Natural History Museum in Paris.

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Several recent papers have reviewed the life and work of French herpetologist Louis Amédée Lantz. They have detailed the composition of his collections deposited in several museums. However, since then, several other important specimens from his collections deposited at the Natural History Museum (NHM, UK) have come to light and we here present all of them in detail. We discovered paralectotypes of Lacerta saxicola obscura Lantz & Cyrén (BMNH 1918.11.21.5–7), syntypes of Lacerta boemica Sukhov (BMNH 1960.1.4.26–30, BMNH 1965.337–342) and Lacerta viridis media Lantz & Cyrén (BMNH 1960.1.4.25, 1966.512). We also identified numerous specimens from the French Mediterranean islands in Lantz’s collection deposited at the NHM, some of which represent the first reported specimens of their species from certain islands. We here provide data on all these specimens. We also place the Mediterranean island specimens from the Lantz collection in their historical context and emphasize the historical and taxonomic value of these collections.

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Data on the common tick parasitism, Ixodes ricinus in populations of sintopic species of lizards (Brauner’s rock lizard, Darevskia brauneri; Black Sea lizard, D. pontica; sand lizard, Lacerta agilis) on Northwestern Caucasus during the spring period are submitted.

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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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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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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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Tis paper contains information on the records of bottom macroinvertebrates, Fish, amphibians and reptiles collected during a Field expedition to the valley of the river Tskhenistskali (in the vicinities of town of Lentekhi and village of Sasashi) in April 2018. According to the results of the study, three Fish species (Actinopterigii) of three families (Salmonidae, Cyprinidae and Nemacheilidae) inhabits three studied rivers (Kheledula, Devashi and Tskhenistskali Rivers). Seven species of reptiles (Squamata: Anguidae — 1, Colubridae — 1, Viperidae — 1, Lacertidae — 4), and ve species of amphibians (Anura: Hylidae — 1, Bufonidae — 1, Ranidae – 2; Caudata: Salamandridae — 1) were registered within the investigated area. Around Lentekhi town three species of the genus Darevskia Arribas, 1999, namely Darevskia rudis svanetica (Darevsky & Eiselt, 1980), Darevskia brauneri brauneri (Mehely, 1909) and Darevskia derjugini abchasica (Bischoff, 1982) were found to live almost sympatrically, with only river serving as a natural barrier between the species. e paper also contains descriptions of habitats characteristic of the region for the species found and the results of route surveys with D. d. abchasica and P. ridibundus being the most numerous species of herpetofauna (sensu lato) — up to 35 and 20 individuals per 100 m of route. In order to illustrate the feeding base of the registered vertebrates, the study of species composition (n = 114) of bottom macroinvertebrates, that form the main component of food source for them. The recorded invertebrates belong to 16 taxonomic groups of the higher rank (up to 98.1 % of all benthic invertebrates appeared to be insects, including Ephemeroptera — 51 %, Plecoptera — 17.4 % Сhironomidae — 14.2 %, Trichoptera — 11.5 %, Diptera — 3 %). Other groups, namely Turbellaria, Nematoda, Oligochaeta, Araneida, Acarina, Collembola, Heteroptera, Coleoptera, in total form a little less than 3 % of all recorded species. Registrations of fauna from mountain regions will help to update actual maps of natural ranges of the species within the country and tend to be a valuable addition to the data that can be potentially used in GIS-modelling of species’ distribution according to predicted climate changes.

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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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Rock lizards of the genus Darevskia are interesting research models due to their asexual reproduction. Ectoparasitic mites and ticks of these lizards are poorly known, despite some of these chelicerates being vector pathogens of humans and wildlife. Here we document and curate previously known data on ectoparasitic Acari of rock lizards and, based on our extensive survey, provide an annotated list of these ectoparasitic arthropods (six tick species, one macronyssid species, and seven chigger species). We also provide new host records (Ixodes ricinus on Darevskia caucasica, D. dryada, D. mixta, and D. szczerbaki; Haemaphysalis sulcata on D. rudis; Odontacarus saxicolis on D. brauneri); and new geographical records (O. saxicolis in Russia and Georgia).

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Rock lizards of the genus Darevskia are interesting research models due to their asexual reproduction. Ectoparasitic mites and ticks of these lizards are poorly known, despite some of these chelicerates being vector pathogens of humans and wildlife. Here we document and curate previously known data on ectoparasitic Acari of rock lizards and, based on our extensive survey, provide an annotated list of these ectoparasitic arthropods (six tick species, one macronyssid species, and seven chigger species). We also provide new host records (Ixodes ricinus on Darevskia caucasica, D. dryada, D. mixta, and D. szczerbaki; Haemaphysalis sulcata on D. rudis; Odontacarus saxicolis on D. brauneri); and new geographical records (O. saxicolis in Russia and Georgia).

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The mite Ophionyssus natricis (Gervais, 1844) (Macronyssidae) is a widespread ectopara- site of reptiles and a vector of some important veterinary diseases. This paper reviews the geographic distribution and host range of O. natricis and the available information about its clinical significance. Some incorrect identifications and host records are also identified. The snake mite has been collected from 144 species of reptiles, mostly snakes, as well as one amphibian and three species of mammals. It has been recorded in 36 countries in Europe, Asia, Africa, North and South America, and Australia. Its geographic range has undoubtedly been extended by human activities. Our records of O. natricis on the lizards Darevskia brauneri (Méhely, 1909) and Ophisops elegans Ménétries 1832, and on the snake Telescopus fallax Fleischmann, 1831, are new host records.

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The mite Ophionyssus natricis (Gervais, 1844) (Macronyssidae) is a widespread ectopara- site of reptiles and a vector of some important veterinary diseases. This paper reviews the geographic distribution and host range of O. natricis and the available information about its clinical significance. Some incorrect identifications and host records are also identified. The snake mite has been collected from 144 species of reptiles, mostly snakes, as well as one amphibian and three species of mammals. It has been recorded in 36 countries in Europe, Asia, Africa, North and South America, and Australia. Its geographic range has undoubtedly been extended by human activities. Our records of O. natricis on the lizards Darevskia brauneri (Méhely, 1909) and Ophisops elegans Ménétries 1832, and on the snake Telescopus fallax Fleischmann, 1831, are new host records.

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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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Phenotypic evolution can cause either divergent or convergent phenotypic patterns. Even adaptation to the same environment may result in divergence of some elements of phenotype, whereas for other morphological traits it could cause phenotypic convergence. We hypothesize that at least some phenotypic characters diverge monotonically, hence they evolve irreversibly even in very closely related species, and this happens in spite of multiple convergent adaptive patterns. We studied the evolution of phenotype in 13 closely related Caucasian rock lizards (Darevskia), whose phylogenetic relationships are well known. We used head shape and the outlines of three important scales, using geometric morphometrics. We studied the association of the overall head shape, individual principal components of head shape and scale outlines with four predictors: phylogeny, habitat, sex and size. The overall head shape was not correlated with any of these predictors, whereas some principal components were correlated with habitat or phylogeny. Habitat type explained the highest fraction of variation in head shape and anal scale area. The relatedness inferred from the components of phenotype not correlated with habitat was congruent with the phylogenetic tree inferred from molecular data. Although adaptation to local environments may obscure the phylogenetic signal present in phenotype, there are components of phenotype whose evolution is irreversible

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When a female goes over to the settled way of life, the territory of a male with a high frequency of affilia- tive behavior (A-behavior) with respect to this female always becomes the formation centre of the female’s permanent home range. As the frequency of the male’s A-behavior becomes stably high (or low) in the first days of existence of a dyad and does not depend on the female’s behavior, it can be concluded that the choice of the male, rather than that of the female, underlies any long-term male – female connection, and it is just social choice rather than sexual one. The male’s sexual and affiliative activities in re- sponse to his female do not correlate with each other. The affiliativity and sexual activity of the male to- wards several females are unequal, and the attitudes of several males to the same female are usually un- equal as well. Therefore, the probability of a female becoming a permanent sexual partner of the male is determined by two circumstances simultaneously: this female must be both socially attractive (which pro- vides her stable residence) and sexually attractive. In other words, any female must pass a «double filter» to become a permanent sexual partner of a male.

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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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Rock lizards were caught and marked in a beech-hombeam forest in the Navagir Mountains between the cities of Anapa and Novorossiisk in 1997-2006. The age of lizards was determined by the skeletochronological method using the 2nd and 3rd phalanges of amputated fingers. The age may be correctly determined only in 5-year-old lizards, since after 5 years, the growth of phalanges sharply slows down. The observations were conducted up to the end of the summer of 2006. The mean longevity in mature settled lizards was not less than 9 ± 2.7 years (n = 20). 50% of the settled individuals reached the age of 10 years. The maximal age recorded in the lizards was 13 years. All three individuals at this age were alive at the end of the observations. The death on hibernation is the main mortality factor.

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There exist long-term intersexual friendly relationships between males and females of Brauner’s lizard that are characterized by high levels of spatial association and affiliative behavior of partners (see report 1). In this article, alternations in the frequency of affiliative behavior in males (Am) and females (Af) after the first appearance of a female on the male’s territory are considered, as well as conditions for the transition of the lizard to a stable residency (SR). Am becomes stable during a few-day initial period and does not change later. This suggests that criteria of the male’s choice are some peculiarities of the female’s exterior. In the initial period, Af is always low, but if Am is high, it gradually rises up to the level of Am. The main condition of the female’s SR is high Am of the owner of the territory in which the female’s comfort areas come to be. Thus, social partner choice in the female is based on the male’s behavior, and a high Am is a trigger which starts up the formation of friendly interrelations. A significant correlation between social and sexual interrelations is absent. As we can judge by the female’s reactions to sexual behavior of the male, its sexual activity is rather a factor of social disintegration. Social strategies are well interpreted in terms of the optimal foraging theory.

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Both adult males and females react to juveniles like their reaction to females. The pre- vailing reaction of adult females are threat or attack, indifference was rarely observed, and orientation approaching was recorded extremely rare. Circumduction, as a common display stopping the aggression of territorial males (t-males), does not stop female`s ag- gression. The prevailing response of non-territorial males (p-males) has distinct features of sexual aggression. The prevailing response of t-males to juveniles` routine behavior is indifference or orientation approaching, it was a threat in rare cases. The main re- sponse to circumduction is orientation approaching, no aggression has been observed. In t-males, unlike females and p-males, such a type of aggression as attacking has been never observed as well. Juveniles, however, avoid all the adults equally. In habitats densely populated with adults, the abundance of juveniles is lower and the level of their nomadism is higher than in sparsely populated habitats. In densely populated habitats, the juveniles are being displaced from the plots favorable for adults. The main role in this displacement is played by adult females. It is doubtful, however, that this displace- ment has any influence on the survival of juveniles, since they are able to successfully use a much broader spectrum of ecological conditions than adults, especially insolation and refuges which are the limiting conditions in the area under survey. Key words: Lacertidae, behavioral ecology, social interrelations, social behavior.

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A natural settlement of rock lizard was observed on the Navagir Ridge (Krasnodar Territory) in 1997–2006. In different years, the settlement included from 18 to 28 adult individuals. The frequency of affiliative behavior in the course of intersexual contacts and the frequency of attendance of comfort female areas by male allowed us to reveal three types of intersexual dyad relationships. They are as follows: (1) the friendly, or A-interrelations (maximal values of both indices; maximal duration of interrelation; and the only reason for the termination of interrelations – death of one of the partners); (2) the R-interrelations (minimal values of all the indices and the main reason for the termination of interrelations – female resettlement); and (3) the Ninterrelations (intermediate values of indices and the main reason for the termination of interrelations – transition to another type of interrelation). Both males and females could simultaneously have several R- and Ninterrelations. In males, the A-attitude towards only single female was always observed. In females, the A-attitude may be formed simultaneously towards two males. Approximately one third of the friendly connected groups were polyandrous triads (female and two males), the rest were monogamous (male and female). Apparently, in this species there is a general tendency towards social polyandry, which is restrained by certain social and/or environmental factors.

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Long-termed intersexual friendly relationships between males and females of the Brauner’s lizard are characterized by high levels of spatial association and high frequency of affiliative behavior between partners (see report 1). In present article we consider changes in the frequency of affiliative behavior in males (Am) and females (Af) after the first appearance of female on the male’s territory and conditions for the transition of lizard to stable residency (SR). Generally, Am becomes stable during the first days of initial period and does not change later. This fact suggests that in most instances male’s choice criteria are some peculiarities of the female’s exterior. In the initial period, Af is always low, however, gradually rises to the level of Am if Am is high. The main condition of female’s SR is high Am of the owner of the territory where female establishes her comfort areas. Hence, social partner choice in female is based on male’s behavior, and high Am triggers the formation of friendly interrelations. Significant correlation between social and sexual interrelations is absent. As can be judged by female’s reactions to male’s sexual behavior, his sexual activity is rather a factor of social disintegration. Structure of costs for social partner search is different in males and females, and these differences well harmonize with differences in gender strategies of partner choice. The social strategies are well interpreted in terms of the optimal foraging theory.

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Friendly intersexual relationships in Brauneri lizard are characterized by high levels of spatial association (frequency of male attendance female’s comfort areas, ZA) and by high frequency of affiliative behaviour both male (Am) and female (Af). Territory of each male is used by several females simultaneously. If a male has Apartner, Am and ZA do not increase on relation to any other female up to death of A-partner. Among main causes of male monogyny are monopolisation by females their comfort areas and A-female’s moderation of affiliative behaviour of male and “females-claimants”, i.e., monogyny is a result of competition between females to a considerable extent. Initiation of A-interrelation is possible in only conditions of high population density, small home ranges and long-continued stay of females in open sites. Thus, social relationships are also determined with ecological conditions.

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In previous papers we have ascertained the existence of many-year social monogamy in Brauner`s lizard (A-relations). The data presented in this communication show that the territorial male’s aggression against the female, including sexual aggression, manifests itself during, or immediately after, a territorial skirmish between the male and the intruder. During an hour after this assault, an increased frequency of rejections of social contacts with the male remained in the assaulted female. In dyads with stable long-term Arelations, a male’s aggressiveness against his female and the frequency of the female’s rejections from contacts were significantly higher in the periods with the high tension of territorial relations of the male with his neighbors. The formation of A-relations demands frequent male-female amicable contacts throughout more than a year as a rule. Thus, the high tension of territorial relations in a settlement may hamper the formation of social monogamy and the corresponding stable sexual relation.

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The present study is the first review of the recent herpetofauna of the Greater Caucasian caves. Also, it is worth noting some phossylous material on amphibians and reptiles (mainly Pleistocene remains) from the caves in this area. The goal of the study is to fill knowledge gaps and to present long-term data on the cave herpetofauna of the Greater Caucasus. In 1992–2019, amphibians and reptiles were found in 61 Greater Caucasian caves located in an altitude range from 55 m a.s.l. to 1300 m a.s.l. in Russia, Abkhazia, and Georgia. A total of 272 specimens representing ten species have been found. The most frequently observed species were Rana macrocnemis (39.0% of all recorded specimens), Pelodytes caucasicus (17.3%), Bufo verrucosissimus (11.0%), Darevskia brauneri (8.8%), Hyla orientalis (8.1%). The total proportion of the remaining five species (Ommatotriton ophryticus, Hyla savignyi, Pelophylax ridibundus, Anguis colchica, and Darevskia derjugini) was 15.8%. As a rule, representatives of the herpetofauna in the Greater Caucasian caves are quite rare and only single or few individuals are usually found. Exceptions were Fyodorovskaya Cave (Black Sea Region near the city of Sochi), Akshasha Cave, Abshdza Cave and Marshania Verkhnyaya Cave (last three in Abkhazia) where 141 specimens of amphibians and reptiles were observed, i.e. more than half (51.8%) of the totally recorded specimens. They belong to eight species (80.0% of all species found). The caves of the Greater Caucasus foothills, where the vast majority of amphibians and reptiles were observed, are usually characterised by a rich faunal diversity of invertebrate animals – their potential food. Amphibians were recorded from both twilight and deep parts of the caves, whereas lizards were found mainly close to the entrances, where the light penetrates and a relatively high air temperature is observed in the warm period of the year. Probably, some of the Greater Caucasian amphibians and reptiles enter the caves purposefully for wintering, and/or for feeding. The other animals occur accidentally in the caves (especially on the bottom of vertical pits), becoming «prisoners of circumstances». Judging by the length of time spent in the caves and, as a consequence, the change in body pigmentation (lighting of the main colouration tone) and good external condition of many individuals, these amphibians can be considered subtroglophiles. Rana macrocnemis, capable of laying eggs in caves, shows a tendency for eutroglophily.

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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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The distribution ranges of Darevskia brauneri and D. szczerbaki are discussed. It is noted the independent speciation of stenotop hemixerophylous maritime-rocks species — Darevskia szczerbaki in the conditions of Black Sea refugia of the East-Mediterranean Province and mezophylous arboreal-rocks and ground dwellers forms of D. b. brauneri and D. b. darevskii in broadleaf mezophyl forests of Western Caucasus.