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TERRA TYPICA. The Crimean Peninsula. DISTRIBUTION. The distribution range of the species is limited by the mountain Crimea from Sevastopol in the west up to the cape Kiik-Atlam near the town Feodosia in the east. The northern border of the distribution range runs on the northern slopes of the Crimean Mountains through the towns Bakhchisarai, Simferopol and the vicinity of the settlement Vishennoe in the Belgorod region. A small isolated population lives on the rocks Adalary near Gurzuf. CONSERVATION STATUS. The number is rather high and stable.

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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 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 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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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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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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A part of the COI mitochondrial barcoding gene was sequenced from seven species of diff erent taxonomical groups: Ambystoma mexicanum (Amphibia, Ambystomatidae), Darevskia lindholmi, Lacerta agilis exigua (Reptilia, Lacertidae), Erinaceus roumanicus (Mammalia, Erinaceidae), Macrobiotus sp. 1 and 2 (Eutardigrada, Macrobiotidae) and Cameraria ohridella (Insecta, Gracillariidae). Th e sequences were compared with available sequences from databases and positioned on phylogenetic trees when the taxa had not yet been sequenced. Th e presence of Mexican axolotls in herpetoculture in Ukraine was confi rmed. Th e partial COI genes of the Crimean rock lizard and an eastern sub-species of the sand lizard were sequenced. We demonstrated the presence of two tardigrade mitochondrial lineages of the Macrobiotus hufelandi group in the same sample from the Zeya Natural Reserve in the Far East: one was nearly identical to the Italian M. macrocalix, and the other one is similar to M. persimilis and M. vladimiri. We also confi rmed the presence of the invasive haplotype “A” of the horse chestnut leaf miner in Ukraine, in line with the hypothesized route of invasion from Central Europe.

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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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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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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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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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This article includes a brief outline of herpetological studies in Crimea over the past 50 years, checklists of taxa of amphibians and reptiles, and a general description of the herpetofauna of Crimea. Some formal issues for the protection of amphibians and reptiles are also considered.

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The Lindholm rock lizard, Darevskia lindholmi, is the only member of the genus Darevskia whose range is restricted solely to Europe, representing a local endemism found only in the Crimean Mountains. In our study, we investigated the cytochrome b gene (mtDNA) of 101 D. lindholmi sequences from 65 Crimean localities, representing its entire range. We found that D. lindholmi is highly genetically structured, and its range is divided into populations belonging to three mitochondrial lineages. The Lindholm rock lizard populations inhabiting the middle part of the Crimean Mountains (further referred to as the Central lineage) are sharply differ entiated from the other two lineages (the Common and the Southwestern lineages), which are present in most of the species range. The genetic distance between the Central lineage and the other two taken together is 4.6%, according to our results, suggesting that the divergence occurred during the Early Pleistocene. The narrowly distributed Southwestern lineage and the widespread Common lineage, on the other hand, are differentiated by 1%. Field observations on the representatives of the main evolutionary groups show that their ecology is also different: the Central lineage is a mesophilic and cold-resistant form, while the other two closely related lineages are more xerophilic and thermophilic. Results of the potential ranges modeling and ecological niche analysis confirm that the genetic lineages occupy different niches of the Crimea. Furthermore, the area of inhabitation of the Central lineage splits the western and eastern parts of the Common lineage range, while the Southwestern lineage is restricted along the coast of the southwestern coast of the peninsula. The long-term co-existence of deeply divergent sister mitochondrial lineages in a relatively small (circa 7,000 km2) isolated mountain system serves as a mesocosm for understanding the speciation process. Our data suggest that the Central lineage warrants further taxonomic investigation.

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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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This work summarizes information on the distribution and status of the populations of amphibians and reptiles of the city of Sevastopol. Data obtained over a quarter of a century were refined by a targeted herpetological examination of the entire territory of Sevastopol (over 1000 km2) in 2018 and early 2019. Most species of am- phibians and reptiles known in Crimea are recorded from the Sevastopol Region, with the exception of some taxa that inhabit only or mainly plains environments (Pelobates vespertinus, Eremias arguta, and Lacerta agilis exigua). Most taxa included in the Red Book of Sevastopol, to date, retain stable populations. Apparently, Vi- pera renardi has disappeared from the region. Analysis of the taxon chorotypes indicates a dominance of spe- cies of Mediterranean (sensu lato) origin. The mild climate of the southwestern part of the Crimean Peninsula determines the unique spatial distribution of the most thermophilic reptile species (Mediodactylus danilewskii, Pseudopus apodus, and Zamenis situla) and, in particular, their wide distribution on the northern macroslope of the Crimean Mountains and (or) the highest elevations in Crimea. The zoning of the territory of Sevastopol, according to herpetological data, made it possible to identify eight districts that differ clearly in species compo- sition and population density of background and rare species. On a national scale, the territory of Sevastopol is important for the conservation of the genetic diversity of species such as Triturus karelinii, M. danilewskii, Ps. apodus, Z. situla, Dolichophis caspius, and Elaphe sauromates. Currently, the state of the populations of T. karelinii, Emys orbicularis, and El. sauromates is most alarming. Scientifically important natural and some synanthropic (in the Khersonesos of Taurida) populations of M. danilewskii, as well as relic populations of the Crimean endemic Lacerta agilis tauridica, require close attention. The “Baydarskyi”, “Cape Aya” and “Laspi” state regional wildlife sanctuaries play the most significant role in preserving the herpetofauna of the region, covering the upper part of the Chernaya River basin (Main Range) and the extreme southwestern part of the Southern Coast of Crimea, as well as the Mekenzievskoe Forestry in the foothills.

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New data on the boundaries of the range of the Lindholm lizard (Darevskia lindholmi), an endemic of the Crimean peninsula, are presented. The species is petrophilic and inhabits a wide range of biotopes in various landscape layers of the Crimean Mountains. The upper boundary of the distribution of D. lindholmi in the southwest of the Main Ridge of the Crimean Mountains reaches 1520 m above sea level. m. (Ai-Petrinskaya yayla, Mount Kemal-Egerek), while on other highlands with maximum heights of over 1.5 km, but colder climate (Babugan, Chatyrdag), the species was traced only up to 1250-1320 m above sea level. m. The northern border of the D. lindholmi range in the western part of the Crimean Mountains runs along the Outer piedmont ridge (right bank of the Alma River), while in the eastern one, along the last rock massifs of the Inner piedmont ridge north of 45 ° N. Isolated peripheral populations found in the forest-steppe or frigan-steppe landscapes of the Crimean foothills and the arid South-East coast differ significantly in their remoteness from the main range, the number and density of the lizard population. A hypothetical history of the formation of the current range of D. lindholmi is discussed.

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New data on the boundaries of the range of the Lindholm lizard (Darevskia lindholmi), an endemic of the Crimean peninsula, are presented. The species is petrophilic and inhabits a wide range of biotopes in various landscape layers of the Crimean Mountains. The upper boundary of the distribution of D. lindholmi in the southwest of the Main Ridge of the Crimean Mountains reaches 1520 m above sea level. m. (Ai-Petrinskaya yayla, Mount Kemal-Egerek), while on other highlands with maximum heights of over 1.5 km, but colder climate (Babugan, Chatyrdag), the species was traced only up to 1250-1320 m above sea level. m. The northern border of the D. lindholmi range in the western part of the Crimean Mountains runs along the Outer piedmont ridge (right bank of the Alma River), while in the eastern one, along the last rock massifs of the Inner piedmont ridge north of 45 ° N. Isolated peripheral populations found in the forest-steppe or frigan-steppe landscapes of the Crimean foothills and the arid South-East coast differ significantly in their remoteness from the main range, the number and density of the lizard population. A hypothetical history of the formation of the current range of D. lindholmi is discussed.

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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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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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Background. Reliable identification of animal taxa characterized by weak mor-phological differences and, at the same time, by significant genetic differentiation is essential condition for the study of biological diversity. The authors set the task of developing test systems using the PCR-RFLP method for the identification of mitochondrial lineages of the Crimean rock lizard Darevskia lindholmi (Szczerbak, 1962), an endemic of Crimea, which is distributed exclusively in the mountainous part of the peninsula. Materials and methods. Based on the analysis of the nucleotide sequences of the cytochrome b gene of the mitochondrial DNA of D. lindholmi, a search for marker nucleotide substitutions specific for each lineage and recognition sites for the HaeIII and TasI restriction endonucleases was carried out. Results. The use of the HaeIII restriction enzyme makes it possible to identify only one, the most differentiated lineage, while the TasI restriction enzyme allows an iden-tification of all three known mitochondrial lineages. Conclusions. The test system proposed by the authors is a simple and reliable method for identifying mitochondrial lineages of the Crimean rock lizard and can be successfully applied in screening studies.

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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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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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Based on results obtained by investigation of morphological differentiation of individuals of different age and sex, a highly correlated variation of 33 morphometric traits has been revealed in Lindholm’s rock lizard, Darevskia lindholmi (Szczerbak, 1962). It has been proved that the increase of linear body dimensions reflects the basic (ontogenetic) trend of variation of traits during late ontogenesis of D. lindholmi. It has been shown that sexual differences of some traits in D. lindholmi are almost absent, while by the linear body dimensions (the value of the first canonical root) males are larger than females. The age-related morphological differentiation of this lizards significantly exceeds sexual differences among adult individuals.

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Principal trends and methods of investigating the postnatal growth in reptiles are discussed. Particular attention is paid to the development of the method of skeletochronology, which enables age determination of individuals, back-calculation of their body length in years before the capture, and reconstruction of growth curves based on such data.

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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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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 current paper summarizes old and recent herptiles records in the karst cavities of the Crimean Mountains, highlighting also the means of entry of poikilothermic vertebrates into the karst. It was found that the underground habitats are on purpose utilized by some amphibians and reptiles, characterized by mostly crepuscular and/or night activity as well as by the pronounced ability to move on the vertical surfaces and, in many cases, by the synanthropization tendency. One of the prerequisites for the animals’ entry to the karst might be their habitat’s conditions such as arid climate, absence of the forest, harsh winters at the upper distribution limit.

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The paper deals with the study of the comparative osteological collection of extant amphibians and reptiles deposited in the Department of Palaeontology of the National Museum of Natural History, NAS of Ukraine. The collection has been recently catalogued and here is described for the first time. It consists of 407 storage units represented by complete or fragmented dry skeletons, skulls, and isolated bones belonging to 104 species of 65 genera, 23 families, and five orders (Caudata, Anura, Crocodilia, Testudines, Squamata). The collection also includes bones of 12 species currently listed in the Red Data Book of Ukraine: Salamandra salanandra, Lissotriton montandonti, Bombina (bombina) variegata, Rana dalmatina, Pseudopus apodus, Mediodactylus kotschyi, Lacerta viridis, Coronella austriaca, Elaphe dione, Zamenis situla, Zamenis longissimus, and Vipera renardi. Collection materials were collected at different times (from the last third of the 19th century until the present day) in 16 countries in Europe, Asia, and Africa. In terms of further enrichment, the collection can be used in batrachological and herpetological studies, and will be of interest for palaeontologists, archaeozoologists, and morphologists who work with osteological materials.