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Interactions between the environment and internal regulation drive the biophysiological dynamics of lizards. Although diurnal lizards are usually heliothermic, they can sometimes be active in the absence of sunlight. Here, we report, for the first time, a case of nocturnal behavior (aided by the artificial light) in the spiny-tailed lizard - Darevskia rudis (Bedriaga, 1886) – a species that normally exhibits diurnal activity.

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TERRA TYPICA. Town Trapezund (Trabzon), north-eastern Turkey. DISTRIBUTION. The species is widely distributed, mainly in the western Transcaucasia, and in the north – in the montane Ingooshetia, Chechnya, neighbouring Dagestan and in northern Ossetiya. Outside of the Caucasus it is widely distributed in the northern half of Turkey. Of eight described subspecies the five occur in the Caucasus. D. r. bischoffi (Boehme et Budak, 1977) occurs in Ajaria. D. r. obscura (Lantz et Cyren, 1936) inhabits the valley and adjoining slopes of mountains of the upper current of Kura river within Georgia. D. r. macromaculata (Darevsky, 1967) is recorded in the ravine of the right tributary of Kura River, river Akhalkalakichai in southern Georgia. D. r. svanetica (Darevsky et Eiselt, 1980) occurs in the ravine of the upper current of the river Inguri in Svanetia (western Georgia). D. r. chechenica (Eiselt et Darevsky, 1991) is widely distributed on the southern slopes of the Great Caucasus from lower Svanetia in the west up to northwestern Azerbaijan in the east, and also isolatedly on the northern slopes within the limits of the montane Chechnya and Dagestan. CONSERVATION STATUS. The number of the species is high and stable everywhere.

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Revealing biodiversity allows the accurate determination of the underlying causes of many biological processes such as speciation and hybridization. These processes contain many complex patterns, especially in areas with high species diversity. As two of the prominent zoogeographic areas, Anatolia and Caucasus are also home to the genus Darevskia, which has a complex morphological structure and parthenogenetic speciation. Darevskia valentini and D. rudis are two largely distributed taxa of this genus, both of which have a controversial taxonomic delimitation. Here we performed both a highly detailed morphological comparison and a molecular evaluation for the populations in both species groups. The most comprehensive taxonomic revision of this complex was carried out to determine the cases where the data obtained were compatible or not with each approach. As a result of the obtained outputs, it seems that D. spitzenbergerae stat. nov., D. mirabilis stat. nov. and D. obscura stat. nov. should be accepted as the species level, this later with subspecies D. o. bischoffi comb. nov. and D. o. macromaculata comb. nov.. Also, we propose two new taxa: D. josefschmidtleri sp. nov. and D. spitzenbergerae wernermayeri ssp. nov.. It has also been shown that “lantzicyreni” subspecies belong to D. rudis instead of D. valentini. The extensive revision has contributed to subsequent studies to more accurately understand the past histories of species in the genus Darevskia.

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A broad sample of Darevskia rudis from the main part of its range was reviewed with regard to external morphology (dis-criminant, UPGMA, MST and ANOVA analyses) and osteology. Darevskia bithynica is raised to species rank, with two subspecies: D. b. bithynica and D. b. tristis. The other subspecies are fairly similar (D. r. rudis being the most different). Two singular populations are described as subspecies: D. r. mirabilis ssp. nov. from Kaçkar Mountains, geographically adjoins the otherwise different D. r. bischoffi and D. r. bolkardaghica ssp. nov., which is geographically isolated but that seems to be very closely related to D. r. obscura.

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Ilya S. Darevsky co-described 70 taxa (three genera, 46 species, 21 subspecies) in 44 publications belonging to five orders, eight families of amphibians and reptiles during his career in herpetology. Of this number, three taxa are fossil and 57 taxa are currently considered as valid. By the regions where new taxa were discovered Southeast Asia and Western Asia (includes Caucasus and Asia Minor) dominates. The largest number of descriptions was published in the Russian Journal of Herpetology.

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Fourty two lizard species are distrubuted in Turkey. In this study, ıt is aimed that seven host species of Lacertidae family are determined helminth fauna -D. armeniaca (Hemşin lizard), D. derjugini (Artvin lizard), D. parvula (Gürcü lizard), D. raddei (Radde lizard), D. rudis (Trabzon lizard), D. unisexualis (Ağrı lizard), D. valentini (Valentin lizard). It was applied firstly classical taxonomy to morphological characters and then DNA sequence analysis in determining fauna. Ten species identified are member of phylum: Nematoda. These species are: two of genus; Skrjabinodon medinae, S. alcaraziensis, Spauligodon sp., S. carbonelli, S. aloisei, Skrjabinelazia hoffmanni, Strongyloides darevsky, Oswaldocruzia filiformis, Thubunae sp., Acuaridae larva; 2 Sestod Oochoristica tuberculata and Mesocestoides spp. and 1 Akantosefal Sphaerirostris scanensis. Totally 13 helminth species are reported. It has been confirmed that present of parasite species identified according to morphological, anatomical and ecological characteristics with determining target genom of helminthes. The results of morphologic and molecular support each other. Four species- one of them is larva-of identified 13 helminth species are new record of Turkish helminth fauna.

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The present study investigated the composition of helminth parasites of Darevskia rudis, Spiny Tailed Lizard from Turkey. One hundred and two samples (49♀♀, 53♂♂) from the Tokat, Trabzon, Rize, Gümüşhane and Artvin Provinces were collected and examined for helminth parasites. New host and locality records were recorded. As a result of the present study, seven species of Nematoda, Skrjabinodon medinae, Spauligodon sp., Spauligodon carbonelli, Spauligodon aloisei, Skrjabinelazia hoffmanni, Strongyloides darevsky, Oswaldocruzia filiformis; one species of Cestoda, Mesocestoides spp. (tetrathyridium) and one species of Acanthocephala Sphaerirostris scanensis were reported from the lizard samples. Sp. carbonelli and Sphaerirostris scanensis are here recorded for the first time in Turkey. D. rudis is the new host recorded for Sk. medinae, Spauligodon sp., Sp. carbonelli, Sp. aloisei from Nematoda, Mesocestoides spp. from Cestoda and Sphaerirostris scanensis from Acanthocephala. This host has been studied for the first time for the helminth parasites from the Tokat and Gümüşhane Provinces. There is, to our knowledge, only one report of helminthes for D. rudis in Turkey.

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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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In the herpetological collection of ZFMK 528 scientific species group names are represented by type materi- al. Of these, 304 names are documented by primary type specimens (onomatophores) while for 224 further names sec- ondary type specimens (typoids) are available, ranging chronologically from 1801 to 2010. The list is a shortened pred- ecessor of a comprehensive type catalogue in progress. It lists name bearing types with their catalogue numbers includ- ing information on further type series members also in other institutions, while secondary types are listed only by pres- ence, both in ZFMK and other collections including holotype repositories. Geographic origin and currently valid names are also provided.

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1. Reinvestigation of the syntypes of Lacerta rudis BEDRIAGA 1886 (=nomen novum pro Lacerta depressa CAMERANO 1878, cf. BuDAK & BöHME 1977) and the comparison with a topotypical series show, that only one form is involved. As lectotype for the species, thus for the nominate form, too, the cf` specimen MIZS (Torino) 2737:1 is designated. 2. The populations of NE-Anatolia and Transcaucasia, so far misinterpreted as being the nominate form, represent a distinct form and are described as Lacerta rudis bischoffi n. subsp.

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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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New material of Turkish rock-lizards of the rudis-group (Lacerta semcoZa-complex) can be assigned to the forms r. rudis, r. tristis and r. obscura; thus, the last named form is recorded for the first time for Turkey. Geographically intermediate samples could be interpreted as clinal, connecting r. rudis and r. tristis, but it turns out to be more likely, that there are two sympatric different taxa overlapping in the intermediate zone. This possibility which could not be proved until now leads to several extraordinaryly difficult nomenclatorial conclusions. They will be discussed further in a following contribution.

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It is a fact that the amphibians and reptiles respond to unfavourable weather conditions by searching an underground refuge to survive during winter. The current literature has shown that some species of reptilians and amphibians may become active before their known end of hibernation periods, especially in lowland areas. In the present study, it has been detected that the changing weather conditions in a highland area in Gümüshane, Turkey can cause similar effects on Lacerta media, Bufo bufo and Rana macrocnemis that share the same habitats. One of the reasons of the early activities of some reptilian and amphibian species before their known end of hibernation periods may be the high sensitivity of these species to the changing temperature conditions. Effects of global warming on reptilians and amphibians obvious, and some members of these animals can even be active during winter. This conclusion brings into mind that L. media, B. bufo and R. macrocnemis can continue to be active at highland areas as long as air temperature values allow them.

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Background. From the genus Darevskia in the territory of Azerbaijan Republic noted 11 species (D. armeniaca, D. caucasica, D. chlorogaster, D. daghestanica, D. derjugini, D. portschinskii, D. praticola, D. raddei, D. rostombekovi, D. rudis, D. valentini). Information on the condition of rock lizards in the republic has not been updated for a long time. The aim of the work was to clarify the data on the distribution and abundance of these species in Azerbaijan.
Materials and methods. The research was carried in the Talysh Mountains (Lenkoran, Lerik and Yardimli districts of Azerbaijan in 2008–2011), in the Great Caucasus (Guba, Gusar, Gabala and Zagatala districts in 2013–2016), in the Minor Caucasus (Dashkesen, Gadabay and Tovuz districts in 2017–2018).
Results. On the basis of own data and analysis of literature sources the data on the distribution and number of rock lizards from the genus Darevskia in Azerbaijan are given. The authors note that the areas of forest lizards of the Talysh (D. chlorogaster and D. praticola hyrcanica) are being reduced and these species need legisla- tive protection. The extinct species for the territory of the Republic should be considered D. derjugini.

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In this investigation, we aimed to determine the amphibian and reptile species distributed in Bartın province (Northwest Blacksea Region of Turkey). As a result of previously and current studies, we recorded two Urodela amphibian species (Ommatotriton ophryticus and Triturus ivanbureschi), six Anura amphibian species (Bufo bufo, Bufotes variabilis, Hyla orientalis, Pelophylax ridibundus, Rana macrocnemis and R. dalmatina), three turtle species (Emys orbicularis, Mauremys rivulata and Testudo graeca), four lizard species (Anguis fragilis, Darevskia rudis, Lacerta viridis and Podarcis muralis) and eight snake species (Coronella austriaca, Dolichophis caspius, Eirenis modestus, Natrix natrix, N. tessellata, N. megalocephala, Vipera transcaucasiana and Zamenis longissimus) in Bartın Province, which totally 23 species. There are no endemic species in our country among them. Natrix megalocephala is also a new record for Bartın province.

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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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Präsentation von Lacerta rudis svanetica ssp. nov. vom Südabfall der zentralen Kaukasus-Hauptkette in Nordwest-Grusinien einschließlich einer isolierten Population in Nordwest-Abchasien und von Lacerta parvula adjarica ssp. nov. von der türkischen Schwarzmeerküste samt ihrem Hinterland zwischen Trabzon und Batumi sowie im (Çoruh- (Tschoroch-)tal ab Borçka, in Adjarien und in den angrenzenden Gebieten Grusiniens zwischen Meria (SE Poti) und Borzomi; bei Bakuriani existiert ein isoliertes Vorkommen.

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The paper provides data on the current location of the type specimens of Darevskia (rudis) complex as of July 2017: Podarcis depressa Camerano, 1878 (syntypes – MZUT R2737.1-3, NHM 1946.9.1.31-32), Lacerta depressa var. rudis Bedriaga, 1886 (lectotype – MZUT 2737.1, paralectotypes – MZUT 2737.2-3, NHM 1946.9.1.32), Lacerta depressa var. modesta Bedriaga, 1886 (syntype – NHM 1946.9.1.31), Lacerta saxicola Subsp. bithynica Méhely, 1909 (lectotype [here designated] – ZISP 17128, paralectotype – NMW 11685), Lacerta saxicola tristis Lantz et Cyrén, 1936 (lectotype – GNM 2481, paralectotypes – GNM 2482-2485, 2487 and GNM one unnumbered specimen), Lacerta saxicola obscura Lantz et Cyrén, 1936 (lectotype [here designated] – ZISP ZISP 17171.1, paralectotypes – NHM 1966.769.773, ZISP 14435, 14436, 14437, 14438, 14439, 14440, 14442, 15402, 16290, 17057.1- 3, 17058, 17059.1-3, 17171.2-26, 17172.1-15), Lacerta rudis macromaculata Darevsky, 1967 (holotype – ZISP 17940.6, paratypes – ZISP 17440.1-18, the location of three paratypes is unknown), Lacerta rudis bischoffi Böhme et Budak, 1977 (holotype – ZDEU 183/977, paratypes – ZDEU 222/976:1, 3-6, 8-9, ZDEU 183/977.2-10, ZFMK 22062.63), Lacerta rudis svanetica Darevsky et Eiselt, 1980 (holotype – ZISP 17875.1, paratypes – ZISP 17875.2-8, 17814.1-18, NMW 24138.1-6), Lacerta rudis chechenica Eiselt et Darevsky, 1991 (holotype – NMW 33504.1, paratypes – ZISP 17878.1-13, 17882.2-4, ZISP 18374.1-5, 18781.1-6, 19545.1-13, ZFMK 16448.52, 29262, 38037.45, 40431-33, NMW 32619.1,2, 33504.2,4,5), Darevskia rudis mirabilis Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et Üzüm, 2013 (holotype – ZDEU 145/2002.9, paratypes – ZDEU 145/2002.1-8, 10-18, O. Arribas private collection (1-6)), Darevskia rudis bolkardaghica Arribas, Ilgaz, Kumlutaş, Durmuş, Avci et Üzüm, 2013 (holotype – ZDEU 144/2009.6, paratypes – ZDEU 144/2009.1-5, 7-8, O. Arribas private collection (1-4)). Strictly following the ICZN the name Lacerta depressa var. rudis Bedriaga, 1886 should not be considered as the replacement name (nomen novum) for Podarcis depressa Camerano, 1878, but the only available name. The history of description of all known forms of the complex is given. Localities, collectors and dates of collection of the type specimens are clarified.

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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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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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Conventional and multivariate taxonomic methods allow the description of a new subspecies of Lacerta rudis BEDRIAGA from the Central Caucasus: L. rudis chechenica ssp. n. - In this connection the intraspecific relations between the northeastern groupings of Lacerta rudis are demonstrated. - The data-analyses used are described.

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Series of newly collected specimens of Lacerta valentini BOETTGER in Eastern Turkey were closely investigated in order to demarcate the areas of the subspecies L. v. valentini and L. v. lantzicyreni. Intermediary populations were found to exist between them, one was described as a new subspecies, Lacerta valentini spitzenbergerae, living in the Cilo-Dag Mountains, Vilayet Hakkâri, Turk

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

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

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

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Obligate parthenogenesis is found in only 0.1% of the vertebrate species, is thought to be relatively short lived and is typically of hybrid origin. However, neither the evolutionary persistence of asexuality in vertebrates, nor the conditions that allow the generation of new parthenogenetic lineages are currently well understood. It has been proposed that vertebrate parthenogenetic lineages arise from hybridisation between two divergent taxa within a specific range of phylogenetic distances (the ‘Balance Hypothesis’). Moreover, parthenogenetic species often maintain a certain level of hybridisation with their closest sexual relatives, potentially generating new polyploid hybrid lineages. Here we address the role of hybridisation in the origin and evolutionary lifespan of vertebrate parthenogens. We use a set of microsatellite markers to characterise the origins of parthenogens in the lizard genus Darevskia, to study the distinctiveness of sexual and asexual taxa currently in sympatry, and to analyse the evolutionary consequences of interspecific hybridisation between asexual females and sexual males. We find that parthenogens result from multiple past hybridisation events between species from specific lineages over a range of phylogenetic distances. This suggests that the Balance Hypothesis needs to allow for lineage-specific effects, as envisaged in the Phylogenetic Constraint Hypothesis. Our results show recurrent backcrossing between sexual and parthenogenic Darevskia but neither gene flow nor formation of new asexual lineages. We suggest that, along with their demographic advantage, parthenogens gain additional leverage to outcompete sexuals in nature when the retention of sexual reproductive machinery allows backcrossing with their sexual ancestors.

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

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The genus Darevskia comprises over 20 species of small-bodied lizards, mainly occurring in the Caucasus Mountain Region. They show differences in body size, scalation and coloration, however, fully diagnostic characters that could separate the species with a high confidence level are still lacking. The early phylogenetic hypothesis of this group was based on ‘traditional’ analysis of morphology, based on multiple body and head measurements and scalation traits. Later, a molecular phylogeny of the genus rejected some of the proposed topology based on morphological traits. In this paper, we used quantitative morphological data (outline-based shape data) to test phylogenetic similarities, as proposed by these earlier hypotheses. We analyzed the pileus shape and the anal area of more than 200 individuals, representing six species of Darevskia, using outline based elliptic Fourier analysis. The analysis did confirm a clustering of the individuals and species (using UPGMA) from the same mitochondrial DNA clade. Hence, the phylogenetic affinity of the major clades in Darevskia could be identified using both molecular methods and outline analysis, whereas a traditional quantitative morphological analysis could not.

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The Caucasian rock lizard Darevskia mixta was sampled and studied from throughout its range, using mitochondrial cytochrome b sequences and scalation. The populations of the Greater and the Lesser Caucasus are reciprocally monophyletic matrilineally, and the respective lineages have been separated since the mid-Pleistocene. The lizards from the Greater Caucasus commonly have an unpaired preanal scale, whereas the lizards from the Lesser Caucasus have an additional scale behind the central temporal and subdivided interparietal scale more commonly than those from the Greater Caucasus. The Lesser Caucasus populations are further subdivided into two geographically distinct matrilineages, and are more diverse genetically and morphologically than the Greater Caucasus populations. The central part of the Lesser Caucasus is suggested to be the ancestral area for the entire D. mixta lineage. Successive Pleistocene periods of glaciation appear to be responsible both for the isolation of D. mixta from its common relatives, and for subdivision within D. mixta. Presence of D. mixta in NE Turkey is challenged.

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Geometric morphometrics is a better tool to evaluate the variation of shape than ‘traditional’ morphometrics. In reptiles, it outperforms morphometrics based on linear measurements and scalation. In an earlier study, two-dimensional outline-based geometric morphometrics in six species of rock lizards (Darevskia) showed that their shapes reflected the species divergence pattern. This allowed us to separate species from different clades, but we could not distinguish closely related species within a clade. We hypothesized that three-dimensional (3D) head shape data would be sufficiently discriminative to identify closely related species. To test this hypothesis, we compared head shapes of three closely related species of the ‘rudis’ clade, Darevskia portschinskii, Darevskia valentini and Darevskia rudis, with the last species treated as two distinct groups (D. r. obscura vs. D. r. rudis), using 3D landmark data. The 3D analysis isolated the species of the ‘rudis’ clade from each other and even separated individuals from populations that showed a genetic introgression pattern. The analysis showed that D. r. obscura is morphologically as distinct from D. r. rudis as from the other nominal species. For this reason, we suggest elevating the status of D. r. obscura to species level, i.e. Darevskia obscura Lantz & Cyrén, 1936.

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To characterize the molecular genetic diversity of the genus Darevskia, several populations were examined by the inter-SINE-PCR method, reporting the number and sizes of the spacers between individual copies of SINE-like interspersed repeats. Examination of 17 D. raddei geographical populations and several reference species revealed unequal genetic differences, measured as Nei and Li’s genetic distances (DNL), for different groups of samples. The highest homogeneity was observed for the apparently panmictic D. raddei nairensis population from the basin of the Hrazdan River: genetic differences within each of the five samples and between them were similarly low (less than 0.1). The difference between ten samples of D. raddei raddei from Armenia and Karabakh (0.2–0.3) was somewhat higher than the interindividual difference within each sample (0.1–0.2), indicating that the samples belonged to different populations. The assumption was supported by the phylogenetic tree topology and multidimensional scaling. The differences between samples from the morphological subspecies D. raddei raddei and D. raddei nairensis ranged 0.3–0.4. The difference of two D. raddei raddei samples of Talysh (Azerbaijan) from other samples of the same subspecies corresponded to the subspecific level. The genetic distances between the good species D. raddei and D. rudis was 0.6–0.7. In terms of DNL, a questionable population from northwestern Turkey (“D. tristis”) was closer to D. rudis (DNL = 0.45), probably representing its subspecies. The phylogeography of D. raddei is discussed.

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

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Darevskia rudis (BEDRIAGA, 1886) zeigt im Gebiet der türkischen Schwarzmeerregion eine weite vertikale Vebreitung. Insgesamt wurden 62 Individuen dieser Eidechse von drei Populationen/Fundorten aus unterschiedlichen Höhenlagen (Provinz Artvin: Ardanuç, 2137 m ü.M., Borçka, 1277 m ü.M., Çermik, 700 m ü.M.) skeletochronologisch altersbestimmt und hinsichtlich ihrer Kopf-Rumpf-Länge vermessen. Im Mittel waren Kopf-Rumpf-Länge und Alter in beiden Geschlechtern bei der höchst gelegenen Population geringer als bei den beiden tiefer gelegenene Populationen. Ein signifikant positiver Zusammenhang fand sich zwischen Alter und Kopf-Rumpf-Länge bei den Weibchen der beiden hoch gelegenen Populationen, jedoch nicht für die aus Çermik und die Männchen aller drei Populationen. Der Geschlechtsdimorphismus in der Körpergröße war in der tiefstgelegenenen (Çermik) Population durch größere Männchen und in den höher gelegenen Popultionen (Borçka und Ardanuç) durch größere Weibchen gekennzeichnet. Die skeletochronologischen Untersuchungen zeigten deutliche Unterschiede in der Körpergröße und Alterstruktur zwischen tiefer und höher gelegenen Populationen von D. rudis auf.

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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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The current literature and our field observations have shown that spiny-tailed lizard may also be active in the winter season. Activity of Darevskia rudis in the February was recorded for the first time in the province of Giresun, Eastern Black Sea Region, Turkey. In conclusion, we assessed that the spiny-tailed lizard could be active throughout the year due to global climate changes.

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The current literature and our field observations have shown that spiny-tailed lizard may also be active in the winter season. Activity of Darevskia rudis in the February was recorded for the first time in the province of Giresun, Eastern Black Sea Region, Turkey. In conclusion, we assessed that the spiny-tailed lizard could be active throughout the year due to global climate changes.

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The spiny-tailed lizard, which has a series of taxonomic revisions, is one of the most common lizard species in Turkey. In this study, sequence data derived from three microsatellite loci (Du215, Du281, and Du323), two mito chondrial (16S rRNA and Cyt-b) genes and combined data were used to evaluate the taxonomic status of Darevskia rudis and Darevskia bithynica with new samples from all subspecies populations in Turkey. Our results indicated that the genetic variations of microsatellite loci were not specific to populations within species, and only minor differences separated D. rudis and D. bithynica populations. Furthermore, the markers we used for phylo genetic analyses (NJ, ML, MP, and BI) produced topologically similar trees based on 16S rRNA and Cyt-b while the combined data produced conflicting trees with the separate gene analyses. Finally, the basal relationships among the populations in Turkish populations D. rudis and D. bithynica were not resolved with this dataset, and we found a hard polytomy at the basis of the phylogeny.

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The spiny-tailed lizard of the genus Darevskia have a series of taxonomic revisions, but still their phylogenetic relationships remain uncertain. In the present study, we have assessed taxonomic relationships among Darevskia bithynica and Darevskia rudis populations through estimation of phylogenetic relationships among 96 specimens using microsatellite DNA (Du215, Du281, Du323 and Du418 loci) and 53 specimens using mtDNA (16S rRNA and cytb) from main populations in Turkey. Although D. b. bithynica and D. r. mirabilis were separated based on the PCoA analysis at low level from other D. rudis and D. bithynica populations, the distance values of Nei’s genetic distance, Nei’s unbiased genetic distance, Fst and Linear Fst were not high among taxa in microsatellite DNA. On the other hand, our phylogenetic analyses (NJ, ML, MP and BI) did not separate D. rudis and D. bithynica populations. Finally, most of the topologically identical trees of phylogenetic analyses and microsatellite results showed that the extant populations of D. rudis and D. bithynica were found to be polytomy. Based on our molecular phylogenetic study, D. rudis complex is still ongoing revisions.

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Three rock lizard species (genus Darevskia) occur in the same mountain gorge of the Greater Caucasus mountains located in Northern Georgia. Whereas Darevskia caucasica and Darevskia derjugini belong to the same phylogenetic clade known as “caucasica”, Darevskia rudis is associated to the “rudis” clade. The mountain lizards differed interspecifically in their scalation and ventral colouration patterns, but the body proportions of D. caucasica and D. rudis were more similar to each other than to D. derjugini as both had flatter heads and longer limbs than D. derjugini. Females of all three species had longer trunks and shorter limbs than males. We hypothesized that hybrids occur more likely between D. derjugini and D. caucasica rather than between species belonging to different clades. However our analysis of microsatellite genotyping clearly revealed no traces of hybridisation and/or gene flow among any of the coexisting lizard species. The absence of individuals with intermediate morphological and genetic features reflects the presence of strong reproductive barriers at the prezygotic level.

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

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Turkey is biogeographically diverse and consequently has a rich herpetofauna. As a result of active herpetological research, the number of species has steadily increased in recent years. I present here a new checklist of amphibian and reptile species distributed in Turkey, revising the nomenclature to reflect the latest taxonomic knowledge. In addition, information about the systematics of many species is also given. In total 35 (19.4%) amphibian and 145 (80.6%) reptile species comprise the Turkish herpetofauna. Among amphibians, 16 (45.7%) anurans and 19 urodelans (54.3%) are present. Among reptiles, 11 (7.6%) testudines, 71 (49%) saurians, 3 (2.1%) amphisbaenians and 60 (41.3%) ophidians are considered part of the herpetofauna. The endemism rate in Turkey is considered relatively high with a total of 34 species (12 amphibian species – 34.3% and 22 reptile species – 15.2%) endemic to Turkey, yielding a total herpetofaunal endemism of 18.9%. While 38 species have not been threat-assessed by the IUCN, 92 of the 180 Turkish herpetofaunal species are of Least Concern (LC), 13 are Near Threatened (NT), 10 are Vulnerable (VU), 14 are Endangered (EN), and 7 are Critically Endangered (CR). In addition, 6 species are in the DD (Data Deficient) category.

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Darevskia rudis (Bedriaga, 1886) and D. bithynica (Méhely, 1909) are morphologically different species, but they are phyloge-netically same species in the D. rudis complex. These species are distributed in northern of Turkey. In the present study, we have evaluated taxonomic status of D. rudis and D. bithynica using ecological niche differentiation. All occurrence data of these species were used to predict and evaluate the suitable areas where they may be expected to be found in Turkey. In order tocompare their ecological niches and explain ecological differentiation, niche similarity tests (niche identity and background tests)and point-based analyses were performed. According to results of ecological niche modeling, we found niche differentiation between D. rudis and D bithynica. Ecological niches for the two species differentiated along environmental variables, asprecipitation of driest quarter for the two species were most important in determining habitat suitability. This factor is importantin niche differentiation between the two species. Finally, our results confirmed the niche differentiation between D. rudis and D. bithynica and added new insights into the taxonomic distinction between D. rudis and D. bithynica that they are two species.

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

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Genetic diversity at 37 allozyme loci was surveyed from Lacerta valentini (4 populations), L. portschinskii and L. rudis (1 population each). The number of polymorphic loci ranged from 1 (L. valentini) to 11 (L. rudis). Mean heterozygosity (direct count) ranged from 0.003 (L. valentini) to 0.071 (L. rudis). Nei`s (1978) genetic distance ranged from 0-0.03 among populations of L. valentini, 0.127-0.163 between L. valentini and L. rudis and 0.366-0.487 between L. portschinskii and the two other taxa. Indices of genetic variability for species having disjunct distributions were lower than in species with contiguous distributions, similar to the case of insular populations, which have lower values than do mainland populations.

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Eggs of Lacerta rudis bischoffi developed norrnally at fluctuiating, temperatures from 13,4°C to almost 40°C in an outdoor terrarium.

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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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Parts of the mitochondrial genes coding for 12SrRNA and 16SrRNA (together about 960 bp) were sequenced for all Mediterranean species of `Mountain lizards` of the genera Archaeolacerta (sensu lato) and Iberolacerta. All subspecies of the Iberian species Iberolacerta cyreni and I. monticola were included in this study. In addition, samples of Apathya cappadocica and Darevskia rudis were analysed to elucidate the relationships of the European `Mountain lizards` to their possible relatives in the Near East. Maximum parsimony and neighbour joining analyses lead to the following major conclusions: (i) the monophyly of the genus Iberolacerta is very well supported; (ii) Archaeolacerta bedriagae (the type species of the genus) is most basal with respect to the ingroup taxa. If we accept Iberolacerta as a genus, Archaeolacerta becames paraphyletic. Therefore, we propose to restrict Archaeolacerta to the type species and to treat A. mosorensis and A. oxycephala provisionally as members of the collective genus Lacerta; (iii) within the genus Iberolacerta three groups were found: a Pyrenean group, an Iberian group and I. horvathi. The relationships among these groups remain unresolved; and (iv) the Peña de Francia lizards, described originally as a subspecies of I. cyreni, are in fact more closely related to I. monticola.

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

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The paper describes the history of biogeographic views on the herpetofauna of both Dagestan and the Caucasus as a whole. The species are grouped into eight zoogeographic groups on the basis of our de- tailed chorological analysis. Mesophilous and xerophilous refuges of the Dagestan herpetofauna are de- scribed. Our herpeto-geographical subdivision of the territory is given. The territory of Dagestan is lo- cated in the Palaearctic subkingdom of Holarctic kingdom and divided between two areas, namely, the Turanian and Mediterranean ones. The boundary between the regions is the junction of the northern foot- hills and plains of the Eastern Caucasus. A possible reconstruction of the regional herpetofauna genesis is considered.

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

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

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

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Einst individuenstarke Populationen von Darevskia rudis chechenica wurden in Georgien (Mtskheta-Mtianeti) durch intensive Straßenbaumaßnahmen vernichtet.

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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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Short description of Lacerta cf. rudis and its habitat at Bolkar Dağları in the Middle Taurus Mts. (Turkey). Report on management and breeding of L. cf. rudis from the neighbouring Ala Dağları. Striking is the extremely long duration of the copula - max. 2 hours and 4 minutes. The clutch consists of up to three eggs. From one egg 1 got though deformed, but viable twins.

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After a brief description of Georgia’s landscape the author reports on the lizards of the family Lacertidae that were found during a herpetological trip in May 2012. In addition, a list of the recorded herpetological taxa is presented.

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

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Anatolia is characterized by a complex topological structure and a wide diversity of climatic regions. This geological context has been crucial during the evolution of the rock lizard genus Darevskia, promoting the origin of several species within this group of lizards. Our study focuses on the evolutionary history of the rudis species complex, comprised by Darevskia rudis, D. valentini and D. portschinskii, distributed across Anatolia and the Caucasus, with overlapping ranges. Previous studies have suggested the existence of ancient gene flow between these taxa, which, allied with poor sampling, led to weakly resolved phylogenies and mismatching topologies. Here, we employ two mitochondrial (cytb and ND4) and two nuclear (MC1R and C‐mos) DNA fragments, coupled with ecological niche modelling and a significant representation of the species’ wide distribution range to further assess relationships. The specimens identified either as Darevskia rudis or as D. valentini appear as paraphyletic and substructured into four clades, which started to differentiate during the lower Pleistocene (around 1.3 Mya). We also differentiate genetically the recently described Darevskia bithynica and identify the Kizil River valley as a strong vicariant candidate to explain the origin of this species by allopatric speciation. Moreover, the eastern Black Sea Mountains seem to have played an important role as glacial refugium for several populations of D. rudis. Overall, the present study confirms the circum‐Black Sea as a potential Pleistocene refuge and supports the refugia‐within‐refugia hypothesis with the finding of several geographically coherent evolutionary units in the region.

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Populations of the lizard Darevskia rudis (Bedriaga, 1886) from northern Anatolia were examined for intestinal parasites in adult specimens. One cestode, Nematotaenia tarentolae López-Neyra, 1944 and four nematode species, Spauligodon saxicolae Sharpilo, 1962, Skrjabinelazia hoffmanni Li, 1934, Oswaldocruzia filiformis (Goeze, 1782) and Strongyloides darevskyi Sharpilo, 1976, were found. Three of these nematodes, S. saxicolae, S. hoffmanni and S. darevskyi are suggested to be part of a module in the network of Darevskia spp. and their parasites. Only one, S. darevskyi, was identified as a Darevskia spp. specialist. The very low infection and diversity parameters are indicative of the depauperate helminth communities found in this lacertid lizard, falling among the lowest within the Palaearctic saurians. Nevertheless these values are higher than those found in parthenogenetic Darevskia spp. Interpopulation variation in the intensity of S. saxicolae and N. tarentolae is attributable to local changes in ecological conditions. On the other hand, parasite abundance and richness increased in the warmer localities, while the effect of lizard sex and size on infection was negligible. The structure of these helminth communities in D. rudis are compared with those observed in other European lacertid lizards.

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

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The taxonomic categories such as population and subspecies were studied on the example of three Caucasian lizard species of genus Darevskia - D. praticola, D. derjugini, and D. rudis by comparing the morphological data and results inferred from nuclear DNA markers. RAPD and new inter-MIR-PCR (IM-PCR) methods were used. The IM-PCR was used to characterize the lacertid DNA fragments located between dispersed SINE type repeats which occurred to be ortologous to mammalian repeats of the same type. It was shown that separation of the Northern population of D. derjugini (subspecies silvatica) is supported by the comparison with two Southern populations (derjugini and barani). The latter ones, in their turn, are very similar and hardly can be considered as good subspecies by the genetic distance. The subspecific division of D. praticola (praticola and pontica) also requires more specification. For example, several populations from North Caucasus (ssp. praticola) occurred to be heterogeneous. The level of differences between ssp. praticola and pontica is of the same order as in some of the praticola populations. Low level of molecular differences between two subspecies of D. rudis (obscura and bischoffi) does not confirm their validity as a full subspecies.

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The present study investigated the reptilian fauna in Amasya Province, Turkey. Reptile species were identified from collections made during field studies or recorded in literature, with some new locality records obtained. Field studies were undertaken over two consecutive years (2016 and 2017). Two lacertid species, one skink species, two colubrid species and one viper species were officially recorded for the first time or their information was updated. In addition to species locality records, chorotypical and habitat selection were also assessed and the International Union for Conservation of Nature Red List of Threatened Species criteria included. Data on the distribution and locality information for each taxon is also provided. Our findings demonstrate that Amasya might be an ecotone zone between the Mediterranean, Caucasian, and European ecosystems. Although there are some concerns for the sustainable dynamics of reptilian fauna, relatively rich and different European nature information system habitat types provide basic survival conditions for reptilian fauna in the province.

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Detailed investigations carried out on Darevskia rudis (Bedriaga, 1886) populations from Northeastern Anatolia, Turkey to identify sexual dimorphism. 11 morphometric and 30 meristic features of 317 specimens were analyzed. Seven meristic (Ventral width, Dorsals attached ventrals at mid-trunk, Femoral pores, Subdigital lamellae left, Tibial scales, Dorsalia). and four metric (Pileus length, Pileus width, Head length, Head width) characters were significant in sexual dimorphism pattern (p<0.05). Males have relatively longer head sizes, and related to this snout-vent length than females. Furthermore, it was found that the number of femoral pores are also higher in males than females because signaling compounds might be released in breeding seasons.

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In this study, which was carried out in July 1998, erythrocyte sizes of Lacerta rudis were examined. The L. rudis specimens were collected from Zigana Pass (Trabzon) at an altitude of 1820 m. Blood smears were prepared at Zigana Pass. Mature erythrocytes (entire cell) and their nuclei lengths and widths were measured by means of an ocular micrometer at a total magnification of 1600x

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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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Given sufficient time and limited gene flow, evolutionary lineages tend to transform into separate species. Mechanisms preventing assimilation during repeated gene-flow events include divergent adaptations and the development of pre- or postzygotic isolation. We analysed the morphological and genetic boundaries of three species of the rock lizard clade Darevskia ‘rudis’ (Darevskia rudis, Darevskia valentini, and Darevskia portschinskii) in relation to the environment, and tried to reconstruct evolutionary pathways underlying the observed separation among the species. We studied the geographic distribution of the scalation traits, microsatellite genotypes, and mitochondrial haplotypes. Our analyses showed consistent morphological and genetic patterns at the centre of the ranges for each species, but asymmetric distribution of alleles and scalation characters within the current contact zones among the species. The genetic and morphological diversification of the clade has been shaped during glacial isolation in an area of Southern Caucasus, away from the Black Sea Coast. The ancestral lineage of D. portschinskii separated from the common D. rudis–D. valentini lineage in the middle Pleistocene, and the two latter lineages separated in relatively recent geological time. Neither of the lineages attained complete lineage sorting; moreover, isolation and migration modelling have helped to detect recombinant gene flow from D. rudis to D. portschinskii (but not to D. valentini). This is most likely linked with climatically more similar suitable habitats between D. rudis and D. portschinskii than between D. valentini and the other two species. In itself, the isolation period was insufficient for the development of intrinsic isolation mechanisms in the system studied. Thus, differential landscape-dependent selection within the contact zones is likely to have triggered the rapid development of isolation mechanisms.

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The authors submit a supplement to the 1994 catalogues of type specimens of the herpetological collection at the Natural History Museum, Vienna (Austria). Additions comprise amphibian and reptile type specimens designated after January 1st, 1994 as well as specimens which were not recognized as types earlier.

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The results of five expeditions (1995, 1996, 2007, 2011, and 2012) to the north-eastern and eastern Turkey are presented. These data were collected from the Pontic (Lazistan) Ridge, Choroch River basin, and the upper flow basins of the Aras River, Kura River, Euphrates River, Tiger River, Dogankent (Kharshit) River, and the Van Lake basin. In total, 2 Caudata, 6 Anura, 2 Testudines (3 subspecies), 25 Sauria (30 subspecies) and 21 Serpentes species were found.

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We genotyped multiple populations of all seven parthenogenetic species of rock lizards, genus Darevskia, as well as their putative sexual parental populations, using double digest RAD sequencing genomic markers. Taking advantage of the conserved homology of the ZW/ZZ sex chromosomes among lacertid lizards, we aligned our short sequence reads to a reference Z chromosome assembly of the common wall lizard Podarcis muralis. This provided unique insight into the origin of all-female hybrid populations, which inherited their single Z chromosome exclusively from a paternal ancestor. The mapped Z-linked loci were used to construct a robust time-calibrated phylogeny. In each parent–offspring species pair, the geographically nearest population of the paternal species was identified as the most likely ancestor of the respective parthenogen, mirroring the trend observed previously on the maternal side in mitochondrial DNA. The estimated splits between the sampled paternal sexual ancestors and their daughter species occurred much earlier than suggested previously and during two narrow time periods: (1) the parthenogenetic Darevskia armeniaca, D. dahli, D. uzzelli and D. rostombekowi dated back to ~0.5 or ~0.9 Mya, depending on the calibration point used, while (2) D. bendimahiensis, D. sapphirina and D. unisexualis appear to have diverged ~1 or ~2 Mya.

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Knowledge on the spatial distribution of taxa is crucial for the decision-making processes in conservation and management of biodiversity that rely on precise distribution data. We present an annotated list for a total of 37 amphibian (20 caudatans and 17 anurans) and 141 reptile species (11 chelonians, 70 lizards, 3 amphisbaenians and 57 snakes) in Turkey, using both available scientific literature up to December 2020 and our own fieldwork data from 1987 to 2020. We provide a comprehensive listing of taxonomy, names, distribution and conservation status of Turkish amphibians and reptiles. The herpetofauna list will be particularly useful for establishing national conservation priorities as well as for placing Turkish fauna into phylogenetic and biogeographic contexts. We compiled information published in books, journals and various web sources and added our personal data. We projected the data in the WGS84 coordinate system and created an overlay grid with cells of 50x50 km2. The database comprises more than 500 grid cells and 11,913 records. As a result, the distribution of Turkish amphibians and reptiles has been extensively mapped with geographical information systems and a database has been created. The obtained data will be useful in planning future studies on taxonomy, ecology and conservation of Turkish amphibians and reptiles.

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Totally, 106 lacertid lizards of two species, Anatololacerta anatolica (n=63) and Darevskia rudis (n=43), from Bursa Province, Turkey, were examined for helminth parasites. In Anatololacerta anatolica, one species of Cestoda, Mesocestoides sp. (tetrathyridium), four species of Nematoda, Skrjabinodon medinae, Spauligodon saxicolae, Skrjabinelazia hoffmanni and Ascarops strongylina (larva in cysts), and one species of Acanthocephala, Plagiorhynchus sp. (cystacanth), were found. In Darevskia rudis, two nematode species, Skrjabinodon medinae and Spauligodon saxicolae, were recorded. The helminth species reported in this study are generalists infecting more than one host species.