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The microornamentation of the scales (oberhautchen) has been studied by means of enamel casts (in negative), or with detached skin pieces (positive cast in the upper face or negative in their lower face of the scales) and its observation by transparency with optical microscopy at 40x, 400x and 1000x. In this study we have found a remarkable uniformity in all genera of Lacertini studied (already known, although significantly expanding the number of species previously studied). In light of the results obtained, it seems that the use of the microornamentation of the scales in taxonomy is limited. There is a remarkable degree of variation and probably of homoplasy, as it happens in so many characters of Lacertini. Nor is it clearly linked to certain conditions of the environment, although it may have a certain importance in mechanisms such as evapotranspiration and perhaps in the crypsis through its influence on the reflectance of the scales. These scales can present protuberances (like small tubercles) on their surface (derived character) in Algyroides (the four species), and in the case of Algyroides marchi also seems to present small pits on its scale surface. In most lacertids the cells of the “oberhautchen” are usually narrow (or short anteroposteriorly, if preferred), between 1 and 4 μm, having the appearance of ribbons or transverse straps. Smaller sizes (derivated) appear more or less marked in Algyroides spp, Darevskia praticola complex, Parvilacerta spp and in Dinarolacerta mosorensis (although not in D. montenegrina) and is difficult to interpret. Except in Dinarolacerta, in the other species it coincides that they are ground or ground-rock interface species, and the coarser stripes could have to do with reflectance (imitating the vegetation or the medium between which they move, according to the hypothesis of Allam et al.). In the case of Dinarolacerta, very saxicolous, it is probably the sister group of Algyroides and the common morphology could have a phylogenetic origin linked to its relationship with Algyroides. There is also no relationship with moisture since some come from dry places and others are from very wet areas. The posterior flange of the cells may be slightly mounted in the next cell (primitive character) but appears quite raised (derived character) in Algyroides nigropunctatus (much less in A. moreoticus and A. marchi, and apparently little or nothing in A. fitzingeri). The rear flange of the cells can be smooth, corrugated or notched. In our samples, irregular (and even dentate) posterior ridges appear in numerous Darevskia such as D. derjugini, D. parvula, D. raddei complex [D. raddei raddei, D. r. vanensis and D. nairensis], in part of the group of D. rudis [D. rudis sspp. and D. bithynica sspp.], several Iberolacerta (I. bonnali, I. horvathi and I. monticola astur), as well as in Teira dugesii. The character appears, although less marked in Algyroides (more in A. moreoticus, but less marked or barely visible in the others), in Hellenolacerta graeca, and other Iberolacerta (I. aurelioi, I. martinezricai, and scarcely marked in I. cyreni). Very little marked in Darevskia mixta, somewhat more in D. dryada and none in D. clarkorum, and also appears in the group of D. saxicola (D. saxicola, D. brauneri brauneri, D. b. darevskii, D. lindholdmi, D. szczerbaki), Dalmatolacerta oxycephala, Dinarolacerta mosorensis (only in the Biokovo sample, and not in Durmitor or D. montenegrina), Apathya cappadocica, in some places of the scales of Parvilacerta fraasii (but not in P. parva) and very little in D. (p) hungarica; also in the remaining group of D. rudis (D. valentini valentini, D. v. lantzicyreni, D. rudis bolkardaghica, D. portschinskii), and some parthenogenetic species (D. armeniaca, D. dahli, D. rostombekowi) but not in D. unisexualis, D. uzzelli, D. sapphirina, D. bendimahiensis or in the triploid hybrid between D. unisexualis and D. nairensis. In these parthenogenetic species it seems a general rule that the character of the crenulation or irregularity of the posterior flange of the cells resembles that of the paternal rather than the maternal species. The exception is the triploid hybrid [arisen from the hybridization of D. unisexualis (parthenogenetic female, without the character) and a male of D. nairensis (with fairly developed character)] that lacks character, and in this case is more similar to the maternal species of which has two genetic sets compared to only one of the paternal one.

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The basin of Lake Skadar with its drainage area represents one of the Balkan Peninsula’s hotspots regarding amphibian and reptile fauna. The value of the batracho- and herpetofauna of the Lake Skadar region is reflected in a rich and diverse composition of species. This is undoubtedly the result of the basin’s specific geographic position and complex geological history. Recent studies revealed that Lake Skadar’s watershed is inhabited by 15 amphibian and 36 reptile species. Among these, one amphibian (Pelophylax shqipericus – EN) and two reptile species (Dinarolacerta mosorensis and Vipera ursinii, both VU) are considered to be globally threatened according to IUCN criteria. An additional three reptile species (Emys orbicularis, Testudo hermanni, and Elaphe quatuorlineata) could become threatened in the future. As an attractive tourist area, a significant part of the Lake Skadar region could be impacted by fast and intense anthropogenic changes. Therefore, preserving the network of suitable habitats, maintaining continuous monitoring, and investing in additional research are essential for maintaining this rich local amphibian and reptile diversity.

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This paper presents an updated checklist of Amphibians and Reptiles of Montenegro. The checklist contains 52 species (including two species complexes), compiled from published literature and personal field surveys. Species complexes are the Anguis fragilis complex and the Lacerta viridis complex. Trachemys scripta is introduced, while Podarcis siculus is potentially introduced species. Migratory species include three sea turtles (Caretta caretta, Chelonia mydas and Dermochelys coriacea). Additionally, there are four potentially present species (Proteus anguinus, Mediodactylus kotschyi, Tarentola mauritanica and Podarcis tauricus), which presence is doubtful or still not confirmed with certainty in Montenegro. National and international protection as well as conservation status of Amphibians and Reptiles is given.

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The evolutionary diversification of many terrestrial vertebrate groups is strongly linked to climatic events in the Cenozoic, the period from 65 Million years ago to today when modern animals first appeared. I investigated the effects of Cenozoic climate change on the taxonomic and morphological diversification of the Old World lizard family Lacertidae, with particular emphasis on the African radiation. African lacertids exhibit an unusual pattern of diversification, in which their highest species richness occurs in deserts north and south of the equator, despite being spread throughout the continent. This disparity is particularly surprising given that desert lacertids are thought to be evolutionarily younger than their mesic-dwelling relatives, suggesting increased diversification rates in arid habitats. To identify the evolutionary factors underlying this pattern, I use a combination of phylogenetic, morphological and ecological techniques. In Chapter 1, I apply Bayesian methods and fossil-based calibrations to molecular sequence data to construct a time-calibrated phylogeny for Lacertidae. I estimate that the family arose in the early Cenozoic, with the majority of their African radiation occurring in the Eocene and Oligocene. In Chapter 2, I describe changes in lacertid body shape across biomes and substrates, and find widespread morphological convergence in similar habitat types. I suggest that in addition to foraging demands, fluctuating and extreme climatic conditions, largely driven by precipitation and temperature, contribute to morphological convergence across independent arid-dwelling clades. Finally, I test if ancestral transitions in ecology, morphology, and rates of diversification temporally coincide with paleoclimatic events in the Cenozoic. I use High Resolution X-ray Computed Tomography to characterize changes in the skull related to life in arid habitats, and apply maximum likelihood methods to test if the origins of those traits temporally coincide with significant shifts in habitat, diversification rates and climatic changes. My results show that African lacertids experienced three major peaks in diversification, accompanied by the evolution of suites of arid-adapted morphological traits. These changes coincide with climatic shifts in Africa, including the transition from closed forests to open grasslands and savanna in the late Oligocene, prior to the peak temperatures of the mid-Miocene Climatic Optimum, and following the formation of the Benguela current leading to hyper-aridity in southern Africa. I conclude that deserts are important centers for reptile evolution, but that expected changes in climate due to global warming may outpace the ability of arid-dwelling species to adapt and persist in the future.

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Sexual differences in size and shape of the Mosor rock lizard, Dinarolacerta mosorensis (Kolombatović, 1886), from Lovćen Mountain (Montenegro) were examined on the basis of the intersex variation pattern of nine morphomet- ric, eight pholidotic, and four qualitative traits. Sexual dimorphism was apparent for all morphometric characters except snout-vent length, while scalation and dorsal pattern exhibited small differences between sexes. The value of the sexual size difference (SSD) index based on snout-vent length was 1.028. The sex-specific allometric slopes for head dimensions and interlimb distance significantly diverged. Head dimensions, especially head height, showed strong positive allometry in males, while interlimb distance was the only character which showed positive allometry in females. Generally, males had significantly greater body size than females. This was true of all body measurements except interlimb distance. The influence of sexual and natural selection on the examined traits is discussed.

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A new species of lacertid lizard of the genus Dinarolacerta is described from the Prokletije Mountain Massif, Montenegro. This new species, Dinarolacerta montenegrina sp. nov., is characterized by its relatively small size, by usually having only one postnasal scale on one or on both sides of the head, a relatively lower number of temporal and postocular scales and a relatively high number of ventral scales. Osteologically, it is mainly characterized by the complete absence of the anteromedial process in the postocular bone, and more reduced supraocular osteoderms. The phylogenetic analysis using partial sequences of the mitochondrial 12S rRNA gene supports the specific status of D. montenegrina sp. nov. and shows that it represents an old independent lineage that separated from its sister species, D. mosorensis, in the late Miocene. The Morača river canyon may have acted as a geomorphological and climatic barrier causing the speciation between the two species of Dinarolacerta. The discovery of this new species endemic to the Balkan Peninsula highlights the importance of the Dinarides as one of the main European hotspots of biodiversity. This high level of endemicity in the Dinaric region is probably the result of both its geographic situation and its complex geological history and morphology.

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Dinarolacerta mosorensis and D. montenegrina are allopatric and closely related rock lizards endemic to the Dinaric Mountains of the Balkan Peninsula. We analysed their habitat characteristics and relative abundances in the mountains of Montenegro. We found significant differences in structural features of the microhabitat used both between populations of D. mosorensis and the two species. Dinarolacerta mosorensis was associated with relatively more shaded and damper spots on rocks closer to vegetation and leaf litter, while D. montenegrina was found in more open, rocky situations. There were significant differences in relative abundance between the sampling sites. Generally, the studied lizards were more abundant at sites with greater percentage of leaf litter and lower percentages of bare ground and small rocks, despite the lower frequency of available refuges in these places. Our results provide basic information that could assist in the adoption of adequate management practices for protection or restoration of habitat attributes relevant to these vulnerable (D. mosorensis) and endemic species.

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The southern part of Montenegro has been identified as an area with high diversity of herpetofauna. However, comprehensive studies of distribution and diversity patterns of reptiles on the country level are still missing. Such studies are essential in designating areas of special conservation importance and nature protection planning in a milieu of increased habitat loss and degradation due to rapid urbanization and tourism development in this small Mediterranean country. To make progress on this problem, we analyzed distribution and diversity patterns of the lizards in Montenegro using a large database consisting of literature data and our unpublished records. We found that fifteen lizard species inhabit Montenegro, and two additional species may be present. The lizards were most diverse in the Maritime biogeographic region of Montenegro, while low diversity was found predominantly along the state borders in the Mountain-valley region. The identified pattern of lizard diversity is at least partly influenced by sampling bias. The eastern mountainous subregion had a distinct species composition compared to all other parts of the country. The East-Mediterranean chorotype was the most dominant, represented by seven species. The great diversity of the lizard fauna of Montenegro can be attributed to its specific topographic position with great influence of Mediterranean climate, heterogeneity of biomes, complex geological history and diverse physiogeographic features. High lizard species richness in the Maritime region and a unique species composition in the eastern subregion of Montenegro indicate that these areas are of high conservation interest.

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Although Albania has a rich reptile fauna, efforts to reveal its diversity have so far been limited. To fill this gap, we collected available published and unpublished (museum collections, online sources) records of reptile occurrences and conducted several expeditions to search for reptiles in areas with few or no previous records. Our georeferenced database contains 3731 records of 40 species from between 1918 and 2015. Based on this comprehensive dataset, we prepared distribution maps for each reptile species of the country. Applying spatial statistics, we revealed that overall sampling effort was clustered, with hotspots associated with easily accessible areas and natural heritage sites. The maximum number of species per cell was 26 with an average of seven. Cells harbouring large reptile diversity were located along the Adriatic and Ionian coasts, on the western slopes of south Albanian mountains, i.e. in areas generally considered as Balkans biodiversity hotspots or potential historical refugia. We found that species presence and diversity is strongly influenced by landscape features. Diversity of land cover, altitudinal variation, temperature and precipitation variation explained the observed pattern in our models. Our study presents the largest database of reptile occurrences to date and is the first to analyse reptile diversity patterns in Albania. The database and the diversity patterns can provide a basis for future macroecological studies and conservation planning.

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Lacertinae is one of the three lacertid lizard subfamilies with a geographical distribution confined to the Palaearctic. Several past attempts to reconstruct its phylogeny resulted in unresolved bush-like topologies. We address the question of whether the lack of resolution is due to insufficient data or whether this lack reflects a rapid succession of speciation events. We analyzed four partial and one complete gene sequences from mitochondrial and nuclear genomes, totalling roughly 3600 bp. We included 29 species representing all 19 genera suggested in recent revision of Lacertinae [Arnold, E.N., Arribas, O., Carranza, S., 2007. Systematics of the palaearctic and oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430, 1–86]. The resulting phylogeny, first, corroborates monophyly at the genus level for the suggested genera, as well as the finding that Atlantolacerta andreanskyi, until recently part of Lacertinae, belongs to the subfamily Eremiadinae. Second, we find that increasing the sequence length and combining multiple nuclear and mitochondrial sequences did not resolve the polytomy, suggesting that the inferred topology indicates a multiple cladogenesis within a short geological period, rather than a methodical artefact. Divergence time estimates, based on previous estimates of several node ages, range from 13.9 to 14.9 million years for the radiation event, however with very broad confidence interval. To associate the radiation with a narrower geological time we consider palaeogeographic and palaeoclimatic data, assuming that the Lacertinae probably evolved in Central Europe andWAsia after the collision of Africa and Eurasia. We suggest that this radiation may date to the late Langhian (ca. 14–13.5 million years) when geological events caused abrupt changes in regional water–land distribution and climate, offering a window of distinct conditions.

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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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Research on the taxonomy of European amphibians and reptiles has increased noticeably over the last few decades, indicating the need for recognition of new species and the cancellation of others. This paper provides a critical review of recent changes and draws up a tentative species list.

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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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The area of the south-western Balkans hosts a remarkably high species richness and has been considered a biodiversity hotspot. The significance of the Balkan Peninsula as a biodiversity refugium during the Quaternary climatic fluctuations has been repeatedly pointed out in literature, yet the area remains quite understudied in terms of phylogeography. Contributing to the biogeography and phylogeography of the Balkan area, we herein present the phylogeographic relationships within the lizards of the genus Algyroides, focusing on the two species that occur in the Balkans (namely, A. nigropunctatus and A. moreoticus), including representatives of Dinarolacerta and Lacerta lizards as outgroups. We combined phylogenetic, phylogeographic and species distribution modelling analyses, using both mitochondrial and nuclear DNA data, in order to uncover the phylogeographic history of the genus and evaluate the validity of the extant taxonomy. Our results reveal three major clades within Algyroides in southern Balkans; one corresponds to A. moreoticus, having sister group relationships with the genus Dinarolacerta, and the other two to A. nigropunctatus, each corresponding to one of the morphological subspecies, which seem to be monophyletic, allopatric and reproductively isolated. These results expose an unprecedented phylogeographic pattern in the area of western Greece where the two major clades of A. nigropunctatus meet, and can contribute to a taxonomic re-evaluation of the species. Moreover, questions are raised on the validity of Dinarolacerta as a genus due to it clustering within the genus Algyroides.