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In this study, we re-examine the Darevskia parvula group comprehensively using morphology, osteology and mitochondrial phylogeny, and describe a new endemic species from Turkey: Darevskia tuniyevi sp. nov. A total of 257 adult specimens were evaluated for external morphology (scalation and biometry) with univariate (descriptive statistics and ANOVA with post-hoc tests) and multivariate (Discriminant Analysis and ANOSIM) analyses. In parallel, osteological data and molecular analyses using three DNA markers (mitochondrial 16S rRNA and Cyt-b, nuclear Rag-1) were used to complete the description of the new taxon. The molecular phylogenetic analyses indicated that the D. parvula group is composed of three taxa as D. parvula, D. adjarica and D. tuniyevi sp. nov., and showed that D. adjarica and D. tuniyevi sp. nov. are reciprocal sister taxa. On the other hand, D. adjarica is morphologically very different from other two forms, while D. parvula is hardly distinguishable externally from D. tuniyevi sp. nov. Therefore, we can consider that D. parvula and D. tuniyevi sp. nov. are cryptic species. These two cryptic species retain their primitive morphology within the group, while D. adjarica has changed, perhaps due to different bioclimatic conditions in its Pleistocene refuge and current area.

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The intraspecific variability of Darevskia parvula (which has two classical subspecies easily identifiable by external characteristics, D. p. parvula and D. p. adjarica), was studied using various approaches including morphology (scalation and biometry), multivariate analyses (PCA, CDA, ANOSIM, UPGMA and MST), osteology, and molecular techniques. High mitochondrial distance, differences at the nuclear level and morphological distinctiveness warrant the specific status of both taxa, Darevskia parvula (Lantz & Cyrén, 1913) and Darevskia adjarica (Darevsky & Eiselt, 1980) stat. nov. A lectotype for D. parvula, originally described with syntypes of both species -D. parvula and D. adjarica- is designated. The uncorrected genetic distance between D. parvula and D. adjarica in the cytochrome b mitochondrial gene is 14.4% ± 1.9%. Intraspecific variability within D. parvula is very small (1.5% ± 0.5%), and was not detected in our samples of D. adjarica. The two species further differ by two mutations in the nuclear melano-cortin 1 receptor (mc1r) gene. Interestingly, past introgression of D. parvula mitochondrial haplotypes (5% ± 1% different to those currently known) into some D. adjarica has been detected in one locality; all the studied specimens of D. adjarica with mtDNA of D. parvula are unmistakably D. adjarica at the morphological and nuclear levels. Morphologically, there is almost no overlap between D. parvula and D. adjarica. These results are corroborated by CDA, MST and UPGMA trees. Specimens of the inland high mountain population of Ardahan (clearly D. adjarica in CDA, MST and UPGMA trees) occupy a somewhat intermediate position between both taxa in the PCA (when specimens and not populations as a whole are considered), but this morphological closeness may be attributed to the influence of climatic factors (continental conditions) on scalation of the specimens. Males appear to be more differentiated than females. Overlap among samples within each species is very marked; none can be separated clearly from its conspecifics. This is even more marked in D. parvula, which has a fairly small area compared to D. adjarica. Darevskia parvula and D. adjarica samples appear to be homogeneously clustered within species and well separated between the two species in the UPGMA trees. In males and females all the D. parvula samples are very similar and moderately differentiated. In males of D. adjarica, the most differentiated seems to be adjBorçka, the others all being clustered together, with adjÇaykara showing slightly more differentiation from the rest (adjOrtacalar, adjArdahan, adjIkizdere and adjÇermik). Darevskia adjarica females are also similarly distributed into two subgroups, one including Borçka, Çermik and Ardahan and the other including Ortacalar, Ikizdere and Çaykara. In both sexes, the inland Ardahan sample clearly belongs to D. adjarica. From the most connected MST samples, speculations can be made about areas of origin and expansion of the different taxa. Ortacalar (D. adjarica) and Hatila (D. parvula) are the most connected (morphologically more “central” in both taxa); in fact, both populations are relatively close, living on the northern (Black Sea) and southern (Anatolian) facing slopes, respectively of the Doğu Karadeniz Mountains (Kaçkar Mountains). This highlights these mountains, which rise from sea level up to nearly 4000 m asl. and have wide buffering possibilities against climate changes, as a zone of refuge and posterior dispersion of this species, and even of the original splitting into two taxa adapted to these different conditions, D. adjarica on the coast and D. parvula on the continental slope. Osteologically D. parvula and D. adjarica are very similar, although Georgian specimens from an isolated population (Atskuri) have closed clavicles not found in Turkish D. adjarica. Also, inland Ardahan D. adjarica have an extra vertebra in both males and females, compared to the other studied specimens from both species. ARRIBAS ET AL. 72 · Zootaxa 4472 (1) © 2018 Magnolia Press The present study indicates that the situation in Turkey is that D. parvula is well differentiated and lives around the Çoruh River Valley, contoured by D. adjarica populations on the coastal-facing slopes of the Doğu Karadeniz Mountains on one side, and the Yalnızçam Mountains on the other side, where D. adjarica enters from Georgia as the opposite extreme of a geographic distribution. The attribution of more inland ranges to D. parvula or D. adjarica, as well as the detailed genetic structure of both taxa may be confirmed with more specific studies.

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

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This study was designed to determine the degree of morphological differentiation between the subspecies of Darevskia parvula from northwestern Anatolia. The meristic pholidolial characters, metric dimensions and color-pattern features of specimens of both subspecies obtained from 10 different localities in the northeastern Anatolia, Turkey are given in detail and compared with regard to literature. In addition, thirty-six morphological characters were compared statistically and 66.7% of them showed differences between two subspecies.

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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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The groups of red-bellied lizards had a small distribution area in the Pontic zone. The several studies performed on these lizard groups are based on taxonomy and systematics. Although there were several taxonomic or systematic researches on some species of this group, the phylogeographical pattern and species disturbing boundaries of this group is still not clear. In the present study, we aimed to resolve the taxonomic and phylogenetic relationships of the red-bellied lizards in Turkey, based on two combined mitochondrial gene fragments and one protein-coding nuclear gene (rag1). Also, we evaluated ecological niches differentiations among red-bellied lizard groups. The mitochondrial DNA genes were found to be highly polymorphic in this group. One hundred and one variable nucleotide sites were detected on the combined gene sequences. According to phylogenetic trees based on the maximum likelihood (ML) and Bayesian inference (BI), the red-bellied lizards group have three species groups; Darevskia parvula, D. adjarica and unnamed Darevskia sp. (candidate species for Darevskia genus). This situation was supported by high bootstrap and posterior probability values in the trees of mitochondrial DNA gene fragments. However, no genetic variation was detected according to nuclear DNA (rag1) sequence. Because the species groups have no overlaps in terms of their ecological niches, ecological niche modelling (ENM) results revealed differences among the groups of D. parvula, D. adjarica, and unnamed Darevskia sp. Besides, we detected no geographical overlaps among three species groups, since there were geographical isolation zones among the species groups of red-bellied lizard.

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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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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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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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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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A detailed description of cranial and postcranial osteology of Darevskia parvula and D. adjarica is provided osteological data for further comparative studies of other lizards. The descriptions and comparisons are based on 12 specimens, of which 10 adult and 2 juveniles for each species. Darevskia parvula and D. adjarica are highly conservative species and they have similar osteological features. The differences in the current study are the number of teeth for the cranial skeleton and the number of vertebrae for the postcranial. Darevskia parvula has a lower number of teeth than D. adjarica.