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Die vorliegende Studie behandelt die Nahrungszusammensetzung dreier nichtsympatrischer Eidechsenarten von den Ägäischen Inseln, Griechenland. Der Mageninhalt von 157 Exemplaren von Podarcis milensis (62), Podarcis gaigeae (50) und Podarcis erhardii (45) wurde auf Beuterückstände untersucht. Alle Tiere kamen aus Museumssammlungen und waren im Sommer gefunden worden. In dieser Jahreszeit nehmen Podarcis milensis und Podarcis gaigeae ähnliche Nahrung auf, hauptsächlich Ameisen, Coleoptera und Insektenlarven, während Podarcis erhardii Coleoptera und Orthoptera erbeutet. Die Ergebnisse werden im Zusammenhang mit der verfügbaren Beute diskutiert.

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Although the insular distribution of the rich herpetofauna of Greece has been studied more extensively, the mainland one is still underexplored. The region of Attica in central Greece represents one of these “black spots” in the mainland. Thus, in 2019 we surveyed for the first time the herpetofauna of Attica in a systematic way. We collected 794 field records of 31 species (4 amphibians, 27 reptiles); Testudo graeca and Trachemys elegans were documented for the first time for Attica.

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Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta lepida, Lacerta princeps, Lacerta schreiberi, Lacerta strigata, Lacerta trilineata, Lacerta viridis, Lacerta andreanszkyi, Lacerta armeniaca, Lacerta bedriagae, Lacerta brandtii, Lacerta cappadocica, Lacerta caucasica, Lacerta chlorogaster, Lacerta cyanura, Lacerta danfordi, Lacerta derjugini, Lacerta dugesii, Lacerta fraasii, Lacerta graeca, Lacerta horvathi, Lacerta jayakari, Lacerta laevis, Lacerta monticola, Lacerta mosorensis, Lacerta oxycephala, Lacerta parva, Lacerta perspicillata, Lacerta praticola, Lacerta rudis, Lacerta saxicola, Lacerta vivipara, Gallotia atlantica, Gallotia galloti, Gallotia simonyi, Podarcis erhardii, Podarcis filfolensis, Podarcis hispanica, Podarcis lilfordi, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis pityusensis, Podarcis sicula, Podarcis taurica, Podarcis tiliguerta, Podarcis wagleriana, Psammodromus algirus, Psammodromus blanci, Psammodromus hispanicus, Psammodromus microdactylus.

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The basic anatomy of the lacertid hemipenis (intromittent organ) and methods for its investigation are described. In many members of the Lacertidae, the hemipenis has a structure quite unlike that of other squamate reptiles: the distal lobes of the retracted organ are complexly folded and there is a well-defined supporting structure of dense connective tissue, the armature. This incorporates blood sinuses and has an intramuscular portion embedded in the m. retractor penis magnus and two club-shaped bodies, the clavulae, that support the lobes in the erect organ. Unarmatured hemipenes occur in some lacertids and, like those of other squamates, possess sac-like lobes in the retracted state, but they are singular in having the lobes invested by the m. retractor penis magnus. It is argued that many of these apparently primitive hemipenes are in fact secondary derivatives of the armatured type. There is considerable inter-specific variation in hemipenial structure which is described systematically. In some cases this involves differences in size, asymmetry and simplification, which may arise as physical isolating mechanisms and is useful in distinguishing otherwise very similar species, particularly in the genus Mesalina (p. 1253). Other shared derived hemipenial features provide useful information about relationships between species and higher taxa and a summary of the hypotheses that they support is given (p. 1254).

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Twenty-four species of lacertid lizards were examined at 31 sites in western Yogoslavia, Greece and Iberia. Comparative observations were made on over 4500 individual lizards, noting such features as times of activitiy, hunting methods, diet, micro- and macrohabitat, refuges used and body temperature. These data are used as a basis for assessing resource partition in related sympatric species, for find out whether different systematic groups have characteristic types of niche, and for comparing cumminity structures in the peninsulas of southern Europa. Acanthodactylus erythrurus, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta graeca, Lacerta horvathi, Lacerta lepida, Lacerta monticola, Lacerta oxycephala, Lacerta mosorensis, Lacerta schreiberi, Lacerta trilineata, Lacerta viridis, Lacerta vivipara, Podarcis bocagei, Podarcis erhardii, Podarcis hispanica, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis sicula, Podarcis taurica, Psammodromus algirus, Psammodromus hispanicus.

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The longevity of several lacertid lizard species in captivity is reported. The conditions of keeping, which have been successful throughout many years, are briefly presented.

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First week of October 2004 a group of nature lovers visited the southern continental part of Hellas (Greece): the Peloponnesos. Nesos means island. According to mythology Pelops, son of Tantalos, was awarded with an island, in his case a pseudo-island. The Island of Pelops or Pelopo-nesos is almost an island: it is THE original peninsula connected to the Balkans via the original Isthmos! From a biogeographical point of view this peninsula is interesting, with plants and animals found elsewhere in Europe, but also with island characteristics and with some endemical forms. Cyclamens, Peloponnesos Colchicum and several other plants were encountered. The author gives a short travelogue. He describes batrachological and herpetological results of the excur- sions made. The situation of chameleons and slowworms on the peninsula receives some extra concern. Also a list of herpspecies found and photographed is added.

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related to individual quality; or a combination of both. Understanding the true nature of this colour variation may thus help to identify the possible selective mechanism producing it. The common wall lizard (Podarcis muralis) exhibits polymorphic colouration, both sexes showing three pure morphs differing in throat and belly colouration (white, yellow and red) and three intermediate morphs (white-yellow, white-red, and yellow-red). In this study we used digital photography and image analysis (RGB system) to investigate colour variation among morphs, sexes and populations. We found that colouration of the six phenotypes results from the combination of three discrete colour tinges (white, red and yellow): pure morphs express a single colour, and no continuous variation exists among them. Similarly, ventral parts of the intermediate white-red and yellow-red morphs present a mixture of two discrete colours resulting from the concurrent expression in different scales of the same colours showed by the corresponding pure morphs. Finally, the white-yellow intermediates show the same colour tinge than pure yellow morph, but with a lower intensity. The finding that white-yellow lizards are smaller than all other ones suggests that they might be subadults not yet expressing the full yellow colouration. Morph colouration significantly varied among sexes, suggesting a possible role for hormone plasma levels in controlling colour expression. It is noteworthy that RGB values varied among sites, indicating that colour expression in this species holds some level of environmental plasticity. Therefore, our results confirmed the presence of discrete colour morphs, which may be genetically based in both sexes of common wall lizards. However, our findings also showed intra-individual variation in colour expression within a morph that is associated with habitat occupancy, which suggests phenotypic plasticity in morphs.

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Global change, including habitat isolation and climate change, has both short- and long-term impacts on wildlife populations. For example, genetic drift and inbreeding result in genetic impoverishment in small, isolated populations, while species undergo range shifts or adaptive phenotypic change in response to shifts in environmental temperatures. In this study, we utilize a model system in which Holocene landscape changes have occurred to examine long-term effects of population isolation. To examine how isolation may constrain responses to climate change, we characterized ecophysiology across land-bridge island populations of Erhard`s wall lizard Podarcis erhardii. We hypothesized that 1) small, isolated populations that are likely genetically depauperate would exhibit lower phenotypic variability; and 2) populations would be adapted to local microhabitat conditions. We compared a population at a low elevation site on the large island of Naxos with two small populations on nearby islets to determine the effects of population fragmentation. We further compared the low elevation Naxos population with two high elevation sites characterized by disparate microclimates to examine the effects of microclimate. To assess the thermal biology and ecophysiological limits of the study species we measured operative body temperatures (Te), field body temperatures (Tb), preferred temperatures (Tp), thermal tolerances (CTmax and CTmin), and evaporative water loss (EWL). Our results indicate that small, isolated populations did not exhibit thermal biology or evaporative water loss, while EWL and thermoregulatory effort varied according to microhabitat characteristics. This study integrates fine-scale measurements with environmental data to provide a holistic view of the relationships between ecophysiology, fragmentation, and microclimate. Our methods can be applied to other ectotherms to gain a better understanding of potential impacts of global change on natural populations.

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Während eines dreitägigen Aufenthaltes wurde die Herpetofauna von Skiathos (Nördliche Sporaden, Griechenland) untersucht. Insgesamt konnten drei Schlangen- und drei bzw. vier Echsenarten (Coluber najadum, Elaphe situla, Malpolon monspessulanus, Ophisaurus apodus, Podarcis erhardii, Lacerta triliniata und vermutlich Lacerta viridis) beobachtet werden. Eine Schlangenart (Elaphe situla) und zwei Echsenarten (Ophisaurus apodus und Podaras erhardii) sind Erstnachweise für Skiathos.

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Cyrtodactylus kotschyi tinensig n. subsp. is olescribed as ä hew subspecies from Tinos. This subspecies is distinguished from C. k. saronicus by the larger number of tubercles on the thigh. lt` is distributed on Andros too, while Ce k. kotschyi occurs on the Syros-Archipelago, and intermediate popülations between C. k, tinensis n. subsp. and C. k. saronicus ranging over Seriphos, Jyaros, Myconos, Naxe, Ikaria and the adjacent Islands. From-Ikaria, Agama stellio daani n. subsp. is described. It differs from A. s. stellio in its blackish grey instead of yellow or red head, the dark spotted throat and in serological characters. All ^^ A» s`^1^0 whi0^ h-as tieen added to the nominate sub- sp ecTe^~Toj^iirly belong to this subspecies, beside those of the Mycpnos-Archipelago which have to remain to A.s, stellio. The Podarcis erhardi from Syros and Stapod`ia do not belong to P. e. `mykoriensis; they intermediate between this subspecies and` P.e. naxensis. Also Matrix natriy from the Northern Cyclades`does not belong to` N. n^persa but intermediate between this rac?e and H. n> schw`eizerT7 The green`lizards of Tinos are Lacgrta triliheata; this race, now named L. t. citrovittata occurs also on Andros and probably on Mycöno s toQ,`while the L. trilineata of Syros belong to the same race as those from Naxe.` The records of ERHARD, V. HELPREICH and ERBER (partly) were omitted out of the qhecklist as very doubtful. Tab. 2 Shows which kinds of amphibians and reptiles occur on the Northern Cyclades`. The following species were first recorded fropi several Islands in th^s paper: Cyrtodactylus kotschyi (Tragonisi), Hemidae-frylus turcicus (Aridros, Tinos, Hyconos, Jyaros), Agamä^atglljo (Riniaj, PocLarcIs erhardi (Tragonisi, Stapodia, Cherrönisi, Strongylo~7 Schinonisi, Aspronisi, Hegalo and Hikro Gaidaronisi), Ablepharus kitaibelii (Jyaros, Rinia), Elaphe situla (Tinos), Elaphe quatuor—^ `Ij.nea.tj9. (Tipps), Natrlx`.hätriy (Tinos), Confirination is givep` `to many recörds, especially from the last Century (s, £ig. 1, tab. 2). The herpetofaunas of Andros, Tinos and the Myconos- Archipelago are y^ry similar, while that of Syros resembles more those of the Centra,! Cyclades. In summing up the differences between the Northern Cyclades and the •Western or Central Cyclades are rather small. The herpetofauna of Ikaria is rather similar to that of the Northern Cyclades too. But there are large differences between the herpeto- fauna of the Cyclades and that of the mainland and Euboee. In the case of the amphibians and reptiles, the wetland and the terrassed fields, many of them are abandoned now and give rise to phryganas, ma,cchias and secondary `woodlands with all stages of succession, are the most important habitats. But phryganas and intensively cultivated areas were settled by few species only, and in the towns only Hemidactylus turcicus occurs. More extended rocky areas ^ithbüf äny` Vegetation do not bear any amphibians or reptiles. Those species which are restricted merely to wetlands like Bufo bufo, Bufo viridis, Rana ridibunda, M caspica, L`acer.ta trilineata ahd Nafrlx natrix are rather `endangered `on the` Northern Cyclades by habitat destruction, and some populations of these species may have disappeared in the meantime.

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Intraspecific colour morphs usually differ in more traits than just colour. These traits can manifest as differences in morph physiology, behaviour, and ecology. Ecological differences among colour morphs, such as the degree of parasitism, can influence the evolution, maintenance, and loss of morphs from populations. High ectoparasite load can directly and deleteriously impact host fitness, and thus could influence colour morph persistence in populations if certain morphs are more frequently exposed to parasites or are more susceptible to parasitism. The Aegean wall lizard, Podarcis erhardii, is a colour polymorphic island lizard that is parasitized externally by ticks and mites. These ectoparasites can affect aspects of host lizard behaviour and physiology – including thermoregulation and body mass – and therefore are an important factor influencing the ecology and fitness of P. erhardii. We find that among sympatric colour morphs, ectoparasite loads differ; namely, monochromatic orange morphs have the highest numbers of ectoparasites, and in general, morphs with orange alleles (orange, orange-white, and orange-yellow) are more heavily parasitized by ticks and mites than the other morphs. Our results indicate that morphs with orange alleles tend to occupy microhabitats with significantly more vegetation cover and thus may increase their exposure to ticks and mites. Ecological differences between morphs could be an important factor contributing to demonstrated patterns of orange morph rarity and loss in P. erhardii populations.

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The study of island taxa can help reveal the mechanisms of natural selection when it acts on small, isolated populations. To elucidate how small populations evolve in high-competition, low-predation environments, we examined differences in morphological characteristics, tail autotomy rates, and home range sizes in several populations of Aegean Wall Lizards (Podarcis erhardii; Lacertidae) living on one large-island site and three small islet sites. While there was relatively little consistent morphological divergence between the study populations, we did observe significantly higher rates of shed and regenerated tails on predator-free, small-island sites, a counter intuitive pattern for a known antipredator defense. Rather than a sign of failed predation events, this pattern is best explained by the fact that tail-shedding can be precipitated by intraspecific aggression and tail cannibalism, which are known to be particularly frequent on small islands. We also found that small-island female lizards, but not males, maintained significantly smaller home ranges than large-island females. Our results indicate that differences in prevailing ecological conditions (most likely reduced food availability and relaxed predation pressure) drive distinct changes in tail shedding rates and home range size in these island lizards.

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The current book provides a synthesis of the information concern- ing field identification of all the species and subspecies of Amphibia and Reptilia hitherto registered in Bulgaria. It comprises 61 species and 23 subspecies, among which 5 species (Pelophylax lessonae, P. kurtmuelleri, Hemidactylus turcicus, Eremias arguta, Montivipera xanthina) that have not been found yet on the territory of the country, but which are known from sites in proximity to Bulgar- ian frontiers. The frogs and toads are represented with 14 spe- cies, newts and salamanders with 6 species, turtles and tortoises with 7 species (Trachemys scripta introduced; Caretta caretta and Chelonia mydas considered extinct), and snakes with 19 species (Vipera aspis and V. ursinii considered extinct). The structure of each species’ description includes the following headings: Name, Diagnosis, General Distribution, Distribution in Bulgaria, Habitats, and Biology. Besides in Bulgarian and Latin, all species are sup- plemented with their English, French, German and Russian names. All synonyms in the main herpetological literature are listed after the currently accepted Latin name. Original keys for identification of the families, genera and species, based entirely on morphologi- cal characters of adult specimens, as well as on the structure of amphibians’ eggs and larvae, are elaborated to facilitate the users. A photo gallery, comprising original colour photos shot in nature, and numerous line drawings of different key characters are made for better distinction between the species. For purely conservation- al reasons, the distribution maps not only indicate the scientifically confirmed species distribution in the country, but also show the areas with suitable habitats where a given species is likely to oc- cur. The degree of suitability is indicated with a different degree of shading: optimal (black); suboptimal (80% gray); suitable (60% gray); less suitable (20% gray); white means unsuitable. The guide is also introducing the readers to some general aspects of the tax- onomy, systematics, morphology, biology and ecology of herpeto- fauna. Special chapters are devoted to the methods of observation and capture of amphibians and reptiles, first aid after a snake’s bite, and the relationship between herpetofauna – humans. The status of each species according to main biodiversity conservation acts and conventions is given in Table 1. Being the first publication of its kind, the book is intended for professional herpetologists, students or nature lovers.

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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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Ambientali, Università degli Studi di Milano, Milano, Italy 3Laboratoire d’Ecologie Alpine, University Grenoble Alpes, University Savoie Mont Blanc, CNRS, LECA, Grenoble, France 4Kosmos – Museo di Storia Naturale dell’Università di Pavia, Pavia, Italy 5Department of Earth and Environmental Sciences, University of Milano-Bicocca, Milan, Italy 6I.P.L.A., Istituto per le Piante da Legno e l`Ambiente, Torino, Italy Correspondence Gentile Francesco Ficetola, Dipartimento di Scienze e Politiche Ambientali, Università degli Studi di Milano, Via Celoria 10, 20126 Milano, Italy. Email: francesco.ficetola@gmail.com Funding information Italian Ministry for Research, Grant/Award Number: 2017KLZ3MA Editor: Ana Margarida Coelho dos Santos Abstract Aim: Analyses of biogeographical patterns and macroecology of islands require large datasets reporting the occurrence of species. The Mediterranean region is a biodiver- sity hotspot, which hosts a large number of reptile species and has been the focus of many studies. Nevertheless, comprehensive inventories describing the features and biodiversity of these environments are lacking. We gathered a dataset summarizing data on reptile distribution on islands of the Mediterranean basin and Macaronesia, also including detailed information on the geographical features. Location: Islands from the Mediterranean Basin, the Atlantic Ocean within the Mediterranean biogeographical region, and Macaronesia (Canary and Savage Islands, Azores, Madeira and Cape Verde). Time period: Present. Taxon: Reptiles (squamates and turtles). Methods: Initially, we developed a geographical database describing islands of the study region, then gathered information on reptile occurrences from 757 bibliograph- ical sources, including atlases, published papers and the grey literature. Through a critical review of these sources, we also obtained information on the status of popula- tions (native, island endemic or non-native) and on the reliability of occurrence data. Results: We obtained basic geographical information from 1875 islands covering the whole study region and with a very broad range of geographical features. We gath- ered >4150 records of reptile occurrence on islands, referring to 198 taxonomic units (species or species complexes); information on population status was available for 84.9% of records. Data are provided as comma-delimited text files. Main conclusions: The database provides a key resource for biogeographical analyses and can also serve as a backbone for conservation studies. The availability of a large database on island features can also be useful for biogeographers working on other taxonomic groups. Nevertheless, more data are required for some geographical areas, in order to ascertain the status (e.g., native vs. non-native) of many populations and to understand the interplay between natural and human-driven processes.

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The interspecific relationships among female size, clutch size, egg size, and hatchling size were examined for 64 European lacertids. The eggs of all species increased linearly in both linear dimensions and mass during incubation. Across species initial egg mass was positively correlated with juvenile mass, with an allometric relationship exponent of 0.87. Initial egg mass across species increased proportionally with female mass to the power of 0.57. Moreover, an increase in maternal mass was also accompanied by an increase in clutch size. The number of eggs per clutch across species scaled with female mass to the power of 0.39. Removing the effect of female mass resulted in a negative correlation between egg mass and clutch size. Species for which the average egg size was lower than expected on the basis of female mass, tended to have larger relative clutch size. The total egg mass per clutch was about one third of female mass (exponent 0.94).

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Eleven species of amphibians and 22 species of reptiles are recorded from the mountainous Florina prefecture on the Greek borders of Albania and the Republic Macedonia. Some ecological notes and the distribution of these species are also reported.

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‘costly’ antipredator behaviors are to be selected against in the event a prey becomes isolated from predators. Predator naïveté, the ignorance of prey to threats imposed by predator species, is a phenomenon often observed on islands where limited space and resources lead to the extinction of predator species. Here, we took advantage of a Pleistocene land-bridge island system with varying degrees of predator diversity and period of isolation to explore the maintenance of antipredator behaviors in a widely distributed prey species. We report on the evolution of antipredator defenses of a model island prey species (Podarcis erhardii; Squamata, Lacertidae,), and identify the factors responsible for the expression or loss of antipredator defenses in island populations. We focus on two antipredator behaviors (flight initiation distance and caudal autotomy), which were quantified for 37 different islands and one mainland location that vary in predator diversity, isolation period, and area. The results suggest that as predator diversity is lost, both flight initiation distance and laboratory caudal autotomy defenses decrease steadily with loss of predators. Contrary to previous studies of herpetofaunal autotomy in this system, we found that field autotomy rates were significantly higher on predator-free islands, and laboratory-induced autotomy was not explained solely by the presence of vipers as reported in an earlier study. While we found that behaviors could erode relatively quickly after isolation from some predators (4 years), in general, behaviors eroded progressively with a duration of isolation with the longest isolated populations having the least expressed antipredator behaviors.

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Color polymorphism defies evolutionary expectations as striking phenotypic variation is maintained within a single species. Color and other traits mediate social interactions, and stable polymorphism within a population is hypothesized to be related to correlational selection of other phenotypic traits among color morphs. Here, we report on a previously unknown throat color polymorphism in the Aegean Wall Lizard (Podarcis erhardii) and examine morph-correlated differences in traits important to social behavior and communication: maximum bite force capacity and chemical signal profile. We find that both sexes of P. erhardii have three color morphs: orange, yellow, and white. Moreover, orange males are significantly larger and tend to bite harder than yellow and white males. Although the established color polymorphism only partially matches the observed intraspecific variation in chemical signal signatures, the chemical profile of the secretions of orange males is significantly divergent from that of white males. Our findings suggest that morph colors are related to differences in traits that are crucial for social interactions and competitive ability, illustrating the need to look beyond color when studying polymorphism evolution.

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Organisms generally have many defenses against predation yet may lack effective defenses if from populations without predators. Evolutionary theory predicts that ‘costly’ antipredator behaviors will be selected against when predation risk diminishes. We examined antipredator behaviors in Aegean wall lizards, Podarcis erhardii, across an archipelago of land-bridge islands that vary in predator diversity and period of isolation. We examined two defenses, flight initiation distance and tail autotomy. Flight initiation distance generally decreased with declining predator diversity. All predator types had distinctive effects on flight initiation distance with mammals and birds having the largest estimated effects. Rates of autotomy observed in the field were highest on predator-free islands yet laboratory-induced autotomy increased linearly with overall predator diversity. Against expectation from previous work, tail autotomy was not explained solely by the presence of vipers. Analyses of populations directly isolated from rich predator communities revealed that flight initiation distance decreased with increased duration of isolation in addition to the effects of current predator diversity, whereas tail autotomy could be explained simply by current predator diversity. Although selection against costly defenses should depend on time with reduced threats, different defenses may diminish along different trajectories even within the same predator-prey system.

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Space is a limited resource in which many animals need to perform basic functions such as feeding and reproducing. Competition over access to space can induce a variety of behaviours that may result in differential access to crucial resources related to survival and fitness. The Aegean wall lizard, Podarcis erhardii, is a colour-polymorphic lizard that inhabits dry stone walls where they access food, safely thermoregulate, shelter from predators and interact with other lizards. Many colour-polymorphic species have morphs with distinct behavioural strategies, which may play a role in morph evolution and maintenance. Here, we conducted the first behavioural experiments on P. erhardii colour morphs. Our goal was to compare morph competitive ability and characterize morph differences in social behaviours using laboratory contest experiments over limited heated space on a stone wall in a neutral arena. Contest experiments revealed that colour morph, not size, predicted intermorph contest outcomes. White and yellow morphs were associated with winning and the orange morph was associated with losing contests. Male colour morphs exhibited different levels of aggressive, boldness, chemical signalling and visual signalling behaviours depending on which morph they were in contest with. White morphs always performed aggressive and scent-marking behaviours more frequently during contests with other morphs. Yellow morphs performed aggressive, bold, chemical signalling and visual signalling behaviours at intermediate frequencies relative to other morphs. Orange morphs performed aggressive behaviours equally often when in contest with yellow morphs but performed all other behaviours less frequently against yellow and white morphs. Considering these results, behavioural variation among P. erhardii colour morphs may promote morph maintenance.

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Variation in color morph behavior is an important factor in the maintenance of color polymorphism. Alternative anti-predator behaviors are often associated with morphological traits such as coloration, possibly because predator-mediated viability selection favors certain combinations of anti-predator behavior and color. The Aegean wall lizard, Podarcis erhardii, is color polymorphic and populations can have up to three monochromatic morphs: orange, yellow, and white. We investigated whether escape behaviors differ among coexisting color morphs, and if morph behaviors are repeatable across different populations with the same predator species. Specifically, we assessed color morph flight initiation distance (FID), distance to the nearest refuge (DNR), and distance to chosen refuge (DR) in two populations of Aegean wall lizards from Naxos island. We also analyzed the type of refugia color morphs selected and their re-emergence behavior following a standardized approach. We found that orange morphs have different escape behaviors from white and yellow morphs, and these differences are consistent in both populations we sampled. Orange morphs have shorter FIDs, DNRs, and DRs; select different refuge types; and re-emerge less often after being approached compared to white and yellow morphs. Observed differences in color morph escape behaviors support the idea that morphs have evolved alternative behavioral strategies that may play a role in population-level morph maintenance and loss.

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Color polymorphism—two or more heritable color phenotypes maintained within a single breeding population— is an extreme type of intraspecific diversity widespread across the tree of life. Color polymorphism is hypothesized to be an engine for speciation, where morph loss or divergence between distinct color morphs within a species results in the rapid evolution of new lineages, and thus, color polymorphic lineages are expected to display elevated diversification rates. Multiple species in the lizard family Lacertidae are color polymorphic, making them an ideal group to investigate the evolutionary history of this trait and its influence on macroevolution. Here, we produce a comprehensive species-level phylogeny of the lizard family Lacertidae to reconstruct the evolutionary history of color polymorphism and test if color polymorphism has been a driver of diversification. Accounting for phylogenetic uncertainty with multiple phylogenies and simulation studies, we estimate an ancient origin of color polymorphism (111 Ma) within the Lacertini tribe (subfamily Lacertinae). Color polymorphism most likely evolved few times in the Lacertidae and has been lost at a much faster rate than gained. Evolutionary transitions to color polymorphism are associated with shifts in increased net diversification rate in this family of lizards. Taken together, our empirical results support long-standing theoretical expectations that color polymorphism is a driver of diversification.

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The natural history of the island of Alonissos in the Northern Sporades is briefly described. The existing herpetological literature is presented and the new finds listed, namely Pelophylax ridibundus, Testudo marginata, Telescopus fallax and Zamenis situla. A melanistic snake, presumably Hierophis caspius, was also found. A total of 13 species of amphibians and reptiles has been reported for Alonissos.

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Nach gegenwärtiger Kenntnis umfaßt die Herpetofauna von Serifos zwölf Arten. Die insel ist wasserreich, die Verbreitung der vier dort vorkommenden semi-aquatischen Arten wurde untersucht. Die Populationen von Bufo viridis, Pelophylax kurtmuelleri und Mauremys rivulata sind als vital anzusprechen, hingegen konnte Natrix tessellata nicht mehr nachgewiesen werden. Es werden einige Aspekte des Naturschutzes im Hinblick auf diese Arten behandelt. Auf Serifos verschlechtern zwei infrastrukturprojekte die Situation der semiaquatischen Herpetofauna zusammen mit Schadeinflüssen durch Tourismus und Landnutzung. Weiters wird auf Austrocknungstendenzen in der Ägäis verwiesen, die für die insel-Herpetofaunen inskünftig Probleme verursachen werden.

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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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Relationships among representatives of five genera of lacertid lizards from Iberia, Morocco, and South Africa were studied using quantitative micro-complement fixation analysis of serum albumin evolution. Using the albumin molecular clock to establish divergence times we suggest (1) South African Ichnotropis and North African Psammodromus diverged from the lineage representing Lacerta lepida-L. monticola during the Oligocene, (2) South African Pedioplanis and Heliobolus diverged from this lineage during the late Miocene, and (3) ancestral representatives of L. andreanszkyi, L. perspicillata and Podarcis hispanica diverged from lineages leading to L. monticola and L. lepida during the mid-Miocene. Radiation within the Palearctic Lacertidae has clearly been extensive, yet fewer than twenty percent of the species in this radiation have been examined biochemically. Until additional data can be gathered, the current classification of the Palearctic Lacertidae cannot be much improved and we recommend adherence to the taxonomy proposed by Arnold (1973).

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Presence of Podarcis erhardii naxensis in Antiparos (central Cyclades) and field notes on the herpetocenosis of the island (Reptilia). -- Presence of Podarcis erhardii naxensis (Werner, 1899) is reported by the author in Antiparos, a small island south-west of Paros (central Cyclades); the lacertid however appears very localized. To explain the reason of this fact the following supposition is suggested: feeding on large invertebrata, Lacerta trilineata limits to small invertebrata the trophic niche of Agama stellio, which owing to alimentary competition, confines P. erhardii to the built up area and high places. The apparently early time (May) of births in Eryx jaculus turcicus local population is also reported.

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A list of reptiles and amphibians occurring on the Dodecanese Archipelago is provided with the latest taxonomic updates. Notes relating to animals’ ecology and to critical habitats are also given. Observations are also made regarding the subspecies Dolichophis jugularis zinneri Cattaneo, 2012 endemic to Rhodes and its Archipelago, and a survey on the Amphibians and Reptiles species known from the southwestern Turkey, whose presence in the Dodecanese Islands has not yet been detected, is discussed.

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The Greek lizard fauna consists of 26 species from which 5 are monotypic and the other ones are represented by a total of 86 subspecies. Five species and 61 subspecies are endemic of Greece. A checklist including the geographical distribution of each taxon in the Greek region is presented.

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A list of the herpetofauna of Samothraki is given in Table 2. In this list I have tried to include all amphibia and teptilia to which I can find certain references. Only two amphibians are known, R. ridibunda and B. viridis. I looked persistently for Brown Frogs but to no avail although conditions were suitable. The streams that run across to the north coast rise at fairly low altitude but the water is cool enough to support the life style of, say, R. graeca. Higher mountain streams might well contain Brown Frogs but these were not visited. The co-existence of two wall lizards that do not normally come together on islands is noteworthy: P. erhardii and P. muralis. Both seem uncommon or even rare and probably represent populations in decline. The snake-eyed Lizard, Ophisops elegans, is absent from Samothraki. This fact, and the occurrence of P. erhardii, demonstrates that this island did not receive faunal elements from Asia Minor. Regarding the snakes all species are what one might anticipate for this coastal island. An exception is E. longissima (Buttle 1989) which otherwise is not known from islands in the Aegean region. The presence of E. quatuorlineata sauromates (Wettstein 1953 p.800) is based on a sight identification but can be taken to be sufficient documentation to allow its inclusion, as with Buttle`s siting of E. longissima. Viperine snakes have not been found.

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1. The herpetofauna of the Argo-Saronic islands includes two amphibians, one tortoise, six lizards, and six snakes. There are no indigenous species, 2. Owing to the lack of previous reports for the area most species are recorded for the first time. However, of particular interest are Chalcides o. ocellatus, Coluber najadum dahlii and Malpolon monspessulanus insignitus, as these are basically mainland forms and absent from the Cycladcs, although the first two are found on Kea, close to the Attic peninsula . The record for Eryx jaculus turcicus is valuable as knowledge of its range in Greece is very incomplete.

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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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Global climate change is altering the ecology of organisms across all major biomes and is likely to contribute to a rapidly increasing number of species extinctions. The effects of climate change are further exacerbated in fragmented landscapes, where isolated populations are known to be losing genetic diversity. This loss of genetic diversity is thought to impact the physiological flexibility (termed `plasticity`) that a species needs to survive the warmer, more fluctuating temperatures that are associated with global climate change. In this study we examined the thermal plasticity of adult male Aegean wall lizards (Podarcis erhardii, Lacertidae) occurring on Cycladic land bridge islands (Aegean Sea, Greece). Populations were sampled from three different size islands ranging from 0.01 km2 to 448 km2. Previous studies have shown that P. erhardii exhibits a predictable gradient in genetic diversity correlating with island area and time since isolation. After collection, lizards were acclimated under identical thermal lab conditions for three weeks after which lizards were divided into control and treatment groups. Treatment groups were subjected to an elevated temperature regime for three weeks corresponding to local conditions under a warming climate while control groups were left under the initial cooler lab conditions. Thermal preference (Tp) and critical thermal maximum (CTmax) were quantified after the initial three week lab acclimation period and then again after a three week experimental manipulation period. Changes in these parameters were then used as measures of thermal plasticity. Overall conclusions from this study indicate that (i.) P. erhardii has surprisingly rigid thermal preferences, and (ii.) level of genetic impoverishment is not related to the extent of thermal plasticity in the species. Understanding how global warming might impact reptile populations isolated in fragmented landscapes will be critically important for evaluating a population’s extinction risk and aid in guiding appropriate management decisions.

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Lizards and Eleonora’s falcon occur on many Mediterranean islets. Data given in literature and new observations concerning their asymmetrical interactions, which have been reviewed and illustrated, allow to regard those as a commensal relationship typical on these micro-insular ecosystems. Some considerations on the ecological, ecomorphological and phenological traits involved on this commensalism are also briefly discussed.

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Comparative analyses were carried out about the variability of the pileal shields of different species belonging to the family Lacertidae. The results of the comparisons have been evaluated in respect to the phylogenetical relationships existing between the different genera and species of the family.

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Microgeographic variation in fitness-relevant traits may be more common than previously appreciated. The fitness of many vertebrates is directly related to their locomotor capacity, a whole-organism trait integrating behavior, morphology, and physiology. Because locomotion is inextricably related to context, I hypothesized that it might vary with habitat structure in a wide-ranging lizard, Podarcis erhardii, found in the Greek Cyclade Islands. I compared lizard populations living on human-built rock walls, a novel habitat with complex vertical structure, with nearby lizard populations that are naive to human-built infrastructure and live in flat, loose-substrate habitat. I tested for differences in morphology, behavior, and performance. Lizards from built sites were larger and had significantly (and relatively) longer forelimbs and hindlimbs. The differences in hindlimb morphology were especially pronounced for distal components—the foot and longest toe. These morphologies facilitated a significant behavioral shift in jumping propensity across a rocky experimental substrate. I found no difference in maximum velocity between these populations; however, females originating from wall sites potentially accelerated faster over the rocky experimental substrate. The variation between these closely neighboring populations suggests that the lizards inhabiting walls have experienced a suite of trait changes enabling them to take advantage of the novel habitat structure created by humans.

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Foraging mode is a functional trait with cascading impacts on ecological communities. The foraging syndrome hypothesis posits a suite of concurrent traits that vary with foraging mode; however, comparative studies testing this hypothesis are typically interspecific. While foraging modes are often considered typological for a species when predicting foraging-related traits or mode-specific cascading impacts, intraspecific mode switching has been documented in some lizards. Mode-switching lizards provide an opportunity to test foraging syndromes and explore how intraspecific variability in foraging mode might affect local ecological communities. Because lizard natural history is intimately tied to habitat use and structure, I tested for mode switching between populations of the Aegean wall lizard, Podarcis erhardii, inhabiting undisturbed habitat and human-built rock walls on the Greek island of Naxos. I observed foraging behavior among 10 populations and tested lizard morphological and performance predictions at each site. Furthermore, I investigated the diet of lizards at each site relative to the available invertebrate community. I found that lizards living on rock walls were significantly more sedentary—sit and wait—than lizards at nonwall sites. I also found that head width increased in females and the ratio of hindlimbs to forelimbs in both sexes increased as predicted. Diet also changed, with nonwall lizards consuming a higher proportion of sedentary prey. Lizard bite force also varied significantly between sites; however, the pattern observed was opposite to that predicted, suggesting that bite force in these lizards may more closely relate to intraspecific competition than to diet. This study demonstrates microgeographic variability in lizard foraging mode as a result of human land use. In addition, these results demonstrate that foraging mode syndromes can shift intraspecifically with potential cascading effects on local ecological communities.

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Species are capable of adapting to new and rapidly changing environmental contexts. An emerging research frontier in ecology and evolutionary biology is predicting the cascading effects of these adaptations on ecological dynamics and ecosystem services. Making these predictions first requires an understanding of how, when, and why traits change. A `functional trait` approach may provide a framework for understanding context-driven changes in traits and their cascading impacts. My dissertation has focused the argument for rapid trait changes in human contexts, and has empirically tested drivers of functional trait changes in the Aegean wall lizard, Podarcis erhardii. Functional traits are those that mediate a biotic interaction or ecosystem function of interest. Maximum bite capacity is a functional trait in many lizards as it dictates the lizard`s ability to capture and consume prey and protect valuable resources from competitors. In chapter 1, I made use of a natural inter-island gradient in P. erhardii bite force across the Cyclade Islands to determine that variation in this functional trait was more related to intra-specific competition than shifts in diet. In chapters 2 and 3 I tested the impact of human-built stone walls and terraces on another lizard functional trait, foraging mode. I found that lizards on walls adopted a relatively sit-and-wait foraging mode, in contrast to lizards in undisturbed habitats, which actively foraged for their prey. I went on to test a predicted syndrome of morphological, behavioral, and performance traits associated with foraging mode. I found that, according to predictions, the sit-and-wait predators had wider heads and longer legs, facilitating jumping, and capturing a higher proportion of active prey. My final chapter lays out a conceptual argument for the utility of human-dominated settings, specifically cities, for furthering our understanding of these ecoevolutionary dynamics. Because cities are functioning ecosystems that present myriad adaptive drivers, they may serve as replicate macrocosms for understanding how species adapt to new ecological contexts and what effect those adaptations may have on ecological dynamics. My dissertation has documented substantial variability in several lizard functional traits across biogeographic and anthropogenic contexts. Collectively, this dissertation demonstrates the importance of understanding intraspecific variability on ecological dynamics, makes the case for humans as instigators of functional trait changes, and lays the groundwork for future advances in the study of evolutionary-ecological feedbacks in human-dominated landscapes.

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1.Body size often varies among insular populations relative to continental conspecifics – the “island rule” – and functional, context-dependent morphological differences tend to track this body size variation on islands.2.Two hypotheses are often proposed as potential drivers of insular population differences in morphology: one relating to diet, and the other involving intra-specific competition and aggression. We directly tested whether differences in morphology and maximum bite capacity were explained by inter-island changes in hardness of both available as well as consumed prey, and levels of lizard-to-lizard aggression among small-island populations.3.Our study included 11 islands in the Greek Cyclades and made use of a gradient in island area spanning five orders of magnitude. We focused on the widespread lizard Podarcis erhardii.4.We found that on smaller islands, P. erhardii body size was larger, head height was larger relative to body size, and maximum bite capacity became proportionally stronger.5.This pattern in morphology and performance was not related to differences in diet, but was highly correlated with proxies of intra-specific aggression – bite scars and missing toes.6.Our findings suggest that critical functional traits such as body size and bite force in P. erhardii follow the predictions of the island rule and are changing in response to changes in the competitive landscape across islands of different sizes.

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Over-water dispersal to small islets is an important eco-evolutionary process. Most often, new arrivals on islets find the environment harsh or mate-less, making their footholds on these islets fleeting. Occasionally, introduced animals are able to survive the strong selection following their arrival, leading to subsequent propagation and, in several famous cases, adaptive radiation. What traits predict that initial survival? We established a replicated island introduction experiment to investigate this process in lizards. In 2014, we introduced 20 Podarcis erhardii lizards to each of five small islets in the Greek Cyclades Islands. We found that the lizards that survived were those with better initial body condition, longer distal portions of their limbs and a greater propensity for jumping. Contrary to our expectations, neither body size nor the strength of the lizards’ bite – two traits positively related to competitive ability, which becomes important later in the colonization process in lizards – predicted survival. This is the first selection study of its kind investigating an experimental introduction of Podarcis, and whether the traits that determined initial survival are important in driving the future evolutionary trajectories of these populations remains to be determined.

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1. Studies of animal communication have documented myriad rapid, contextdependent changes in visual and acoustic signal design. In contrast, relatively little is known about the capacity of vertebrate chemical signals to rapidly respond, either plastically or deterministically, to changes in context. 2. Four years following an experimental introduction of lizards to replicate experimental islets, we aimed to determine if chemical signal design of the experimental populations differed from that of the source population. 3. In 2014, we translocated Podarcis erhardii lizards from a large, predator-rich island to each of five replicate predator-free islets. Mean population densities increased fivefold over the following 4 years and bite scars suggest significantly more intraspecific fighting among these experimental populations. In 2018, we analysed the chemical signal design of males in each of the experimental populations and compared it to the chemical signals of the source population. 4. We found that males consistently presented a significantly more complex chemical signal compared to the source population. Moreover, their chemical signals were marked by high proportions of octadecanoic acid, oleic acid and α-tocopherol, the three compounds that are known to be associated with lizard territoriality and mate choice. 5. Our island introduction experiment thus suggests that the chemical signal design of animals can shift rapidly and predictably in novel ecological contexts

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Resource-limited environments may drive the rapid evolution of phenotypic traits and ecological preferences optimizing the exploitation of resources. Very small islands are often charac- terized by reduced food availability, seasonal fluctuations in resources and strong unpredictability. These features may drive the evolution of phenotypic traits such as high bite forces, allowing animals to exploit a wider variety of the available resources. They may also lead to more generalist dietary patterns in response to food scarcity. However, the lack of predators and competitors on such small islands often also leads to high densities and the evolution of strong sexual dimorphism, which may also drive the evolution of bite force. Here, we take advantage of a unique replicated introduction experiment to test whether lizards introduced into very small islands alter their feeding ecology and use different resources, resulting in the evolution of a large body size, large head size and large bite forces. Our results show that three years after their introduction, the island lizards were larger and had greater bite forces and more pronounced sexual dimorphism. However, the diets were only marginally different between animals from the source population on a very large nearby island and those on the islets. Moreover, distinct differences in diet between animals on the different islets were observed, suggesting that the local environment is a strong driver of resource use. Overall, lizards with absolutely and relatively (adjusted for body size) large bite forces did eat larger and harder prey. Taken together, our data suggest that intraspecific competition is an important driver of the rapid evolution of bite force, which may allow these lizards to exploit the scarce and fluctuating resources on the islets. Whether or not lizards will evolve to include other types of food such as plants in their diet, facilitated by their large bite forces, remains to be explored in future studies.

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Resource-limited environments may drive the rapid evolution of phenotypic traits and ecological preferences optimizing the exploitation of resources. Very small islands are often charac- terized by reduced food availability, seasonal fluctuations in resources and strong unpredictability. These features may drive the evolution of phenotypic traits such as high bite forces, allowing animals to exploit a wider variety of the available resources. They may also lead to more generalist dietary patterns in response to food scarcity. However, the lack of predators and competitors on such small islands often also leads to high densities and the evolution of strong sexual dimorphism, which may also drive the evolution of bite force. Here, we take advantage of a unique replicated introduction experiment to test whether lizards introduced into very small islands alter their feeding ecology and use different resources, resulting in the evolution of a large body size, large head size and large bite forces. Our results show that three years after their introduction, the island lizards were larger and had greater bite forces and more pronounced sexual dimorphism. However, the diets were only marginally different between animals from the source population on a very large nearby island and those on the islets. Moreover, distinct differences in diet between animals on the different islets were observed, suggesting that the local environment is a strong driver of resource use. Overall, lizards with absolutely and relatively (adjusted for body size) large bite forces did eat larger and harder prey. Taken together, our data suggest that intraspecific competition is an important driver of the rapid evolution of bite force, which may allow these lizards to exploit the scarce and fluctuating resources on the islets. Whether or not lizards will evolve to include other types of food such as plants in their diet, facilitated by their large bite forces, remains to be explored in future studies.

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Mediterranean islands have complex reptile assemblages, but little is known about the factors that determine their organization. In this study, the structure of as-semblages of Squamata was evaluated based on their species richness and two measures of phylogenetic diversity (variability and clustering). I evaluated the com-position of the assemblages comparing distinct biogeographic subregions within the Mediterranean: Adriatic, Aegean, Balearic, Corsica–Sardinia, Crete, Gulf of Gabés, Ionian Sea, Ligurian Sea, Malta, Sicily, and Tyrrhenian Sea. The effect of island en-vironments and geographical isolation on the diversity metrics was assessed using generalized linear models. The analyses indicated that species richness was mostly influenced by island area and geographical isolation. Assemblages on smaller islands were poorer in species and phylogenetically dispersed, possibly as an effect of inter-specific competition. The species composition of the assemblages was determined by similar environmental drivers within the biogeographic subregions, including is-land area, island elevation, geographical isolation, and aridity. In several subregions, significant patterns of phylogenetic attraction were found in species co-occurrences, caused by the limits imposed by the island size on large predatory species

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Background Lizards of the genus Podarcis are widespread in the Mediterranean region, including islands and island archipelagos. These small-bodied lizards have a predominantly protective green-brown colouration. However, some populations display unusual patterns, in which the colouration is predominantly blue or uniformly black. This study explores the factors that influence this chromatic variation, whether environmental (climate and island conditions) or evolutionary (phylogenetic trait conservatism). The colouration of 1400 individuals (27 species) was analysed in the CIELAB colour space. Results Pagel’s λ indicated that colouration is weakly conserved within phylogenetic lineages. Although the island surface plays a key role in the chromatic variability of these lacertids, geographic isolation and climate hold less influence. The colouration of some small island populations tends to be uniform and dark, possibly due to intense intraspecific competition and lower predatory pressure. Conclusions This study highlights the importance of island populations in understanding the processes that favour the emergence of extreme phenotypes in small ectothermic vertebrates.

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Nonylphenol (NP) is an endocrine disruptor widely distributed in the environment. It accumulates in the lipids of living organisms and enters the human food chain. The main source of human exposure is expected to be food, drinking water and foodstuff contaminated through leaching from packaging or pesticide formulation applications. NP acts as an estrogenic compound and it is able to mimic the action of estradiol 17β (E2) by binding to the estrogen receptor (ER). The aim of the present study was to investigate the NP effects on the hypothalamic–pituitary–adrenal gland (HPA) axis of the bioindicator Podarcis sicula lizard. A time-dependent stimulation of the HPA axis and variations of both catecholamine plasma levels were showed. Moreover, NP effects on adrenal gland morphology were evaluated by light and transmission electron microscopy. Clear morphological signs of adrenal gland stimulation such as an increase of steroidogenic cord diameter and vascularization, a strong escalation of adrenaline cell number and a decrease of noradrenaline cells were observed. The notably elevated levels of adrenal hormones suggested a permanent turning on of hypothalamic corticotropin releasing factor (CRF) secretion together with a lack of the negative feedback of HPA axis, perturbing systemic responses of the organism. Our data may help to predict the biological alterations induced by NP and to extend its impact upon adrenal function.

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I assessed the effect of island characteristics and population isolation on the endemic insular lizard Podarcis erhardii and its native hemogregarine parasite Hepatozoon spp. I analyzed the relationships of prevalence, infection likelihood, and parasitemia to several factors at the island (time of isolation, area, distance to nearest larger land mass), population (host density), and organismal (load of hematophagous ectoparasites) levels. My results suggest that smaller islands, as well as islands that have been isolated for longer periods of time, show higher infection rates and higher parasitemia in hosts than others. I also found that distance between a focal island to the nearest larger land mass, as well as the load of hematophagous ectoparasites on an individual, were poor predictors of infection variables in P. erhardii. These results indicate that island area, host population density, and island age are likely to be significant drivers of changes in host-parasite interactions in fragmented populations.

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I assessed the effect of island characteristics and population isolation on the endemic insular lizard Podarcis erhardii and its native hemogregarine parasite Hepatozoon spp. I analyzed the relationships of prevalence, infection likelihood, and parasitemia to several factors at the island (time of isolation, area, distance to nearest larger land mass), population (host density), and organismal (load of hematophagous ectoparasites) levels. My results suggest that smaller islands, as well as islands that have been isolated for longer periods of time, show higher infection rates and higher parasitemia in hosts than others. I also found that distance between a focal island to the nearest larger land mass, as well as the load of hematophagous ectoparasites on an individual, were poor predictors of infection variables in P. erhardii. These results indicate that island area, host population density, and island age are likely to be significant drivers of changes in host-parasite interactions in fragmented populations.

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The biogeography of host-parasite dynamics is an area that has received little attention in studies of island ecology. While a few studies have shed insight on patterns of parasitism in insular host populations, more empirical evidence is needed to ascertain how isolation impacts parasites. Biogeography generally theorizes that the physical size of islands and the duration of each island’s isolation can be driving geographic factors controlling species interactions and populations dynamics. To test this, we assessed the effect of island structure and population isolation on the endemic insular lizard Podarcis erhardii and its native hemogregarine parasite (Apicomplexa: Adeleorina) in the Cyclades (Aegean Sea). We analyzed the relationships of prevalence and parasitemia of hemogregarine infection with several factors concerning the island (size, time of isolation, spatial isolation, population density) and host (body size) levels using regression and structural equation models, respectively. Regressions indicate that islands with greater host density and islands which have been isolated for shorter timespans tend to have higher hemogregarine prevalences; structural equation models suggest a similar pattern for parasitemia. We hypothesize this may be driven by insular density compensation. Hosts on islands that are more temporally and spatially isolated also tend to have higher prevalence and parasitemia of hemogregarines. Our results indicate that island area, island isolation, and host population density are likely to be significant drivers of changes in host-parasite interactions in fragmented populations.

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On islands worldwide, lizards are frequently the most important vertebrates, both in terms of aggregate biomass and total population size. While some research has been done on the effects of predation on lizard population size, little is known on the effects of marine nutrient subsidies or herbivory on lizard population size. In this study we investigate the effects of seabird-delivered marine subsidies, herbivory by introduced goats, and predation by native snakes on the populations of the Aegean Wall lizard (Podarcis erhardii, Lacertidae, Reptilia) living on islands in the Aegean Sea (Greece). Our results indicate that Podarcis population densities are determined by a hierarchical interplay between predation on the one hand, and nutrient availability - mediated through goat browsing and seabird subsidies- on the other. Predation by snakes significantly depresses lizard densities on those islands that are big enough to support resident snake populations. On snake-free islands only, lizard numbers are positively associated with breeding seabird populations, presumably because of seabird subsidies. Also only on snake-free islands, lizard numbers are negatively associated with density of stocked goats. Experimental goat removal and addition experiments lead to the same results. Goats appear to impact lizard populations through the destruction of the vegetation and through the introduction of invasive generalist ticks that switch hosts from goats to parasitize lizards.

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The life histories of island species have been the focus of evolutionary studies ever since Darwin visited the Galapagos. While much progress has been made in understanding the drivers of these patterns, comparatively little is known on the corresponding patterns in lacertid lizards. Here we summarize and review the results of more than 30 years of investigations on a model island species, the Aegean Wall lizard Podarcis erhardii. We discuss how the interactions between predation, parasitism and availability of natural resources shape the behavior, reproduction and morphology of insular populations based on data from dozens of islands from the Aegean Sea Region (Greece).

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Parasites are ubiquitous in wildlife populations and are regularly present in free-ranging vertebrates. Little research has been done on the impact of insularity on the patterns of parasitism. Islands, known for low genetic diversity and poor parasitic infracommunities, offer a tractable model to better understand the factors that shape parasitism. This study analyses helminth parasite loads from eight populations of the Erhard’s Wall lizard (Podarcis erhardii, Lacertidae) occurring on islands of varying size, age and isolation in the Aegean Sea (Greece). By comparing burdens of infection among the different populations, we aimed to elucidate how island features shape levels of parasitism. Parasite communities on the islands were very species-poor, with only four genera of nematodes (Spauligodon sp., Skrjabinodon sp., Parapharyngodon sp. and Skrjabinelazia sp.) detected among the lizard populations. We found that aggregate worm infection level was positively correlated with island area, whereas total parasite prevalence was negatively correlated with island age, though marginally. This suggests that numbers of parasites are determined by the loss of genetic diversity as consequence of long-term population bottlenecks. Given that the island populations constitute an important model for the effects of habitat fragmentation on biodiversity, our findings raise concerns about the long-term prospects for many specialized parasite populations in the face of widespread anthropogenic habitat fragmentation.

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The Sporades are one of the most biologically important archipelagos in the Aegean Sea (Greece) and have received priority conservation over the last 50 years. However, despite numerous early efforts, its herpetofauna is only partially described, resulting in many distributional gaps that have prevented adequate understanding and management of the resident species communities. We review one century of bibliography from the Northern Sporades and combine this information with a review of museum specimens and insights from numerous extensive field surveys over the last near-decade to provide for the first time a comprehensive picture of the reptiles and amphibians of the archipelago. We report here on 26 new island records and find that the herptile communities of the region are largely derived from the herpetofauna of the nearby Thessaly mainland, with only a few introduced taxa. There is also a small but significant set of endemic taxa in the archipelago. Island species richness declines with decreasing island size and increasing duration of island isolation. Herptile communities on smaller islands are progressively nested subsets of the communities on larger islands. The presence of reptile species depends sensitively on the condition and management of native ecosystems. While non-aquatic species maintain largely healthy populations, most populations are under pressure from the combined effects of rampant tourist development, the destruction and degradation of rare wetland habitats, and the abandonment of traditional agricultural landscapes. We provide recommendations regarding sustainable management of the local reptile and amphibian populations.

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Reproductive investment, including the number of offspring produced, is one of the fundamental characteristics of a species. It is particularly important for island vertebrates, which face a disproportionate number of threats to their survival, because it predicts, among other things, a species’ resilience to environmental disruption. Taxa producing more offspring recover more quickly from environmental perturbations and survive environmental change better. However, ecologists do not understand which primary drivers shape a species’ reproductive investment well. Here, we compare the reproductive efforts of 14 island populations of the Aegean Wall Lizard (Podarcis erhardii), which lives across widely diverging environmental conditions. We test three hypotheses, namely that reproductive investment (measured as clutch size, clutch volume) is (1) positively associated with predation risk [‘Predation Risk Hypothesis’]; (2) positively associated with the presence of reliable vegetation cover that provides shelter [‘Gravid Female Protection Hypothesis’]; and (3) limited by (and hence positively correlated with) food availability [‘Food Limitation Hypothesis’]. Although field data are somewhat consistent with all three hypotheses, statistical analyses provide strong support for the Predation Risk Hypothesis. The results not only shed light on which fundamental forces shape reproductive investment in island vertebrates, but can also help shape conservation priorities.

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Recent climate change has caused the distributions of many species to shift poleward, yet few empirical studies have addressed which species will be vulnerable to longer-term climate changes. To investigate past consequences of climate change, we calculated the population extinction rates of 35 reptile species from 87 Greek land-bridge islands in the Mediterranean that occurred over the past 16,000 years. Population extinction rates were higher for those species that today have more northern distributions. We further found that northern species requiring cool, mesic habitats had less available suitable habitat among islands, implicating loss of suitable habitat in their elevated extinction rates. These extinctions occurred in the context of increasing habitat fragmentation, with islands shrinking and separating as sea levels rose. Thus, the circumstances faced by reptiles on the islands are similar to challenges for numerous species today that must cope with a changing climate while living in an increasingly human-fragmented landscape. Our island-biogeographical approach to investigating historical population extinctions gives insight into the long-term patterns of species responses to climate change.

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During naturalistic research performed in August of 2022 and 2023 in several Aegean islands, three new herpetological records occurred: Hemorrhois nummifer on Telendos Island, Laudakia stellio on Pserimos Island and Dolichophis caspius on Kitriani Islet.

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Between April 22 and May 30, 1980 herpetological field studies and collecting trips were undertaken on the Greek islands Kea, Spanopoula, Kithnos, Sifnos, Kitriani (Cyclades) Alonissos and Piperi (Northern Sporades). Collected specimens as well as data taken from the literature lead to an actual list of occuring species, completed by some biological observations. Collecting sites are shown in maps. Ten species of amphibians and reptiles are recorded from these islands for the first time.

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Due to its species richness, Kresna Gorge is relatively well investigated in herpetological aspect. However, almost all of the researchers concentrate their efforts in the well-known biodiversity hot spots situated in the middle and the southern parts of the area, thus leaving the northern parts almost unexplored. Most of the locations of the researched species presented in this work are newly identi ed, since the areas of investigation remained out of the scope of the majority of scientists. In addition the research presents some of the conservation measures in the area, taken by the non-governmental organization Fund for Wild Flora and Fauna (FWFF), which directly affected the prosperity of the researched species populations. The identi ed nature conservation bene ts and possible fails of these measures were evaluated.

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In diesem Naturführer werden auf über 100 Farbfotos alle mitteleuroipäischen Arten und darüber hinaus auch süd- und osteuropäische vorgestellt: Frösche, Kröten, Molche und Salamander, Schildkröten, Echsen und Schlangen. Fachkundige Texte informirern über Kennzeichen, Vorkommrn, Verhalten, Fortpflanzung, Ernährung und Besonderheiten. Der bewährte Kosmos-Farbcode teilt die Amphibien und Reptilien nach ihrer Verwandtschaft in 5 Gruppen ein und erleichtert so die Bestimmung. Die praktischen Klappen bieten interessante Zusatzinformationen über Froschlurchentwicklung, Laichformen und anatomischen Besonderheiten bei Reptilien.

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Im Verlauf einer von der Deutschen Forschungsgemeinschaft finanzierten Reise nach Griechenland bereisten die Verfasser vom 23. bis 29. April 1972 den Paros-Archipel in der Zentral-Ägäis. Der Deutschen Forschungsgemeinschaft sei an dieser Stelle für ihre großzügige Unterstützung gedankt. Danken möchten wir auch Herrn Dr. W. BÖHME, der Vergleichsmaterial aus den umfangreichen Beständen des Zoologischen Forschungsinstituts und Museums A. Koenig in Bonn zur Verfügung stellte, die F. BUCHHOLZ auf mehreren Griechenlandreisen gesammelt hatte. Neben Paros und Antiparos, den beiden größten Inseln, konnten wir mit einem gemieteten Fischerboot den folgenden unbewohnten Inselchen des Archipels einen Besuch abstatten: Despotiko, Strongylo, Tourlos, Preza, Glaropunta und Panteronisi in der südlichen Inselgruppe sowie Ovriokastro und Gaidaronisi vor der Nordküste von Paros. Insgesamt konnten wir hier 13 Arten von Amphibien und Reptilien nachweisen: Rana ridibunda, Mauremys caspica, Cyrtodactylus kotschyi, H emidactylus turcicus, Agama stellio, Podarcis erhardii, Lacerta trilineata, Ablepharus kitaibelii, Eryx jaculus, Elaphe quatuorlineata, Natrix natrix, Telescopus fallax und Vipera ammodytes. Diese von uns im Jahre 1972 eingebrachten Informationen werden ergänzt durch Angaben anderer Autoren (BUCHHOLZ 1955; CLARK 1965, 1967, 1968a, 1968b, 1970, 1971, 1972; WERNER 1938; WETTSTEIN 1953, 1957). Dadurch wird zwar die Anzahl der untersuchten Inseln vergrößert (Spiridonisi, Tria Nisia-Inseln; WETTSTEIN 1957), das Artenspektrum aber nur um Testudo marginata (CLARK 1970) und Elaphe situla (BUCHHOLZ 1955) erweitert.

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Attention is given to the reptiles and amphibians of the island Lesbos in Greece. The observations were made during a holiday in July 1987.

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Landbridge islands offer unique opportunities for understanding the effects of fragmentation history on genetic variation in island taxa. The formation of islands by rising sea levels can be likened to a population bottleneck whose magnitude and duration is determined by island area and time since isolation, respectively. The Holocene landbridge islands of the Aegean Sea (Greece) were formed since the last glacial maximum and constitute an ideal system for disentangling the effects of island area, age and geographic isolation on genetic variability. Of the many reptile species inhabiting this island system, the Aegean wall lizard Podarcis erhardii is an excellent indicator of fragmentation history due to its widespread distribution and poor over-water dispersal abilities. In this study, we utilize a detailed record of Holocene fragmentation to investigate the effects of island history on wall lizard mitochondrial and nuclear microsatellite diversity. Findings show that the spatial distribution of mitochondrial haplotypes reflects historical patterns of fragmentation rather than geographic proximity per se. In keeping with neutral bottleneck theory, larger and younger islands retain more nuclear genetic variation than smaller, older islands. Conversely, there is no evidence of an effect of isolation by distance or effect of distance to the nearest larger landmass on genetic variability, indicating little gene flow between islands. Lastly, population-specific measures of genetic differentiation are inversely correlated with island area, suggesting that smaller islands exhibit greater divergence due to their greater susceptibility to drift. Taken together, these results suggest that both island area and time since isolation are important predictors of genetic variation and that these patterns likely arose through the progressive fragmentation of ancestral diversity and the ensuing cumulative effects of drift.

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Aim Isolation is a key factor in island biology. It is usually defined as the distance to the geographically nearest mainland, but many other definitions exist. We explored how testing different isolation indices affects the inference of impacts of isolation on faunal characteristics. We focused on land bridge islands and compared the relationships of many spatial and temporal (i.e., through time) isolation indices with community‐, population‐ and individual‐level characteristics (species richness, population density and body size, respectively). Location Aegean Sea islands, Greece. Time period Current. Taxon Many animal taxa. Methods We estimated 21 isolation indices for 205 islands and recorded species richness data for 15 taxa (invertebrates and vertebrates). We obtained body size data for seven lizard species and population density data for three. We explored how well indices predict each characteristic, in each taxon, by conducting a series of ordinary least squares regressions (controlling for island area when needed) and a meta‐analysis. Results Isolation was significantly (and negatively) associated with species richness in 10 of 15 taxa. It was significantly (and positively) associated with body size in only one of seven species and was not associated with population density. The effect of isolation on species richness was much weaker than that of island area, regardless of the index tested. Spatial indices generally out‐performed temporal indices, and indices directly related to the mainland out‐performed those related mainly to neighbouring islands. No index was universally superior to others, including the distance to the geographically nearest mainland. Main conclusions The choice of index can alter our perception of the impacts of isolation on biological patterns. The nearly automatic, ubiquitous use of distance to the geographically nearest mainland misrepresents the complexity of the effects of isolation. We recommend the simultaneous testing of several indices that represent different aspects of isolation, in order to produce more constructive and thorough investigations and avoid imprecise inference.

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Body size evolution on islands is widely studied and hotly debated. Gigantism and dwarfism are thought to evolve under strong natural selection, especially on small remote islands. We report a curious co-occurrence of both dwarf and giant lizards on the same small, remote island (Plakida): the largest Podarcis erhardii (Lacertidae) and smallest Mediodactylus kotschyi sensu lato; Gekkonidae--the two commonest insular reptiles in the Aegean Sea. The geckos of Plakida have a peculiar tail-waving behavior, documented here for the first time in this genus. We suspect that P. erhardii evolved large size to consume geckos and the geckos evolved a unique tail-waving behavior as a defensive mechanism.

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Aim: Animal body sizes are often remarkably variable across islands, but despite much research we still have a poor understanding of both the patterns and the drivers of body size evolution. Theory predicts that interspecific competition and predation pressures are relaxed on small, remote islands, and that these conditions promote body size evolution. We studied body size variation across multiple insular populations of 16 reptile species co-occurring in the same archipelago and tested which island characteristics primarily drive body size evolution, the nature of the common patterns, and whether co-occurring species respond in a similar manner to insular conditions. Location: Aegean Sea islands. Time period: 1984–2016. Major taxa studied: Reptiles. Methods: We combined fieldwork, museum measurements and a comprehensive literature survey to collect data on nearly 10,000 individuals, representing eight lizard and eight snake species across 273 islands. We also quantified a large array of predictors to assess directly the effects of island area, isolation (both spatial and temporal), predation and interspecific competition on body size evolution. We used linear models and meta-analyses to determine which predictors are informative for all reptiles, for lizards and snakes separately, and for each species. Results: Body size varies with different predictors across the species we studied, and patterns differ within families and between lizards and snakes. Each predictor influenced body size in at least one species, but no general trend was recovered. As a group, lizards are hardly affected by any of the predictors we tested, whereas snake size generally increases with area and with competitor and predator richness, and decreases with isolation. Main conclusions: No factor emerges as a predominant driver of Aegean reptile sizes. This contradicts theories of general body size evolutionary trajectories on islands. We conclude that overarching generalizations oversimplify patterns and processes of reptile body size evolution on islands. Instead, species’ autecology and island particularities interact to drive the course of size evolution.

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In June 2007, 12 species of Amphibia and 19 species of Reptilia were found during investigations at 18 localities situated from sea level up to 1507 m a. s. l. in Albania. The highest number of amphibians (5) was recorded at Shelegurë, 1032 m a. s. l. and of reptiles (9) at Diviakë, 1–5 m a. s. l. The highest total number of species (5 amphibians and 6 reptiles) was recorded at Shelegurë. Testudo hermanni was the most abundant species of reptile recorded. Of the amphibians, Bombina variegata and Pelophylax ridibundus were also abundant and Podarcis muralis occurred at the highest locality (Llogarë, Cikës Mts., 1507 m a. s. l.). Herein, are presented new faunistic records for 11 species of Reptilia. From the zoogeographical point of view, 50.0% of the amphibians recorded are Mediterranean, 33.3% European and 16.7% Palaearctic species. Of the reptiles, 52.6% are Mediterranean, 15.8% European and 31.6% Palaearctic species. Mediterranean faunal elements are predominant in the batrachofauna and herpetofauna of Albania. Four of the amphibians recorded (Triturus macedonicus, Pelophylax epeiroticus, P. shqipericus, Rana graeca) and three reptiles (Algyroides nigropunctatus, Podarcis erhardii, Anguis graeca) are endemic to the Balkan Peninsula. An annotated list of all the species recorded in Albania in 2007 is presented.

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This note presents photographic material and a detailed description of a concolor morph in Podarcis erhardii (Bedriaga, 1882) and the first published record of this morph on Alonissos. Data on the observation and a short review of the literature are also given.

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Exotic predators such as feral cats (Felis catus), have been the driving force behind the extinction of many endemic species of island mammals, birds and reptiles. Island endemics appear to be exceptionally susceptible to invasive predators because of small population size and frequent lack of anti-predator defenses. The goal of this study was to determine the impacts of feral cats on the island populations of Aegean Wall lizards (Podarcis erhardii, Lacertidae) in relationship to the expression of anti-predator behaviors. I estimated lizard population densities in areas with low cat density sites (LCD) versus high cat density (HCD) sites by conducting 100-m transect along dry-stone walls, on the island of Naxos, as well as on surrounding islets (Cyclades, Greece). Degree of expression of antipredator behaviors was determined by measuring flight initiation distance (FID) and rates of tail autotomy both in the field and in the lab for six populations in HCD, LCD sites and four satellite islets without cat presence. I also staged controlled encounters with mounted cats decoys and quantified escaping responses from lizards from these populations. I found that feral cats had a strong negative effect on lizard population densities. Lizards adapted their antipredator behaviors in response to cat predation by extending their FIDs, increasing their capacity for tail autotomy, and by staying closer to refugia. In laboratory predation simulations, lizards from cat-free islets had significantly shorter FIDs than LCD site lizards and in particular than HCD site lizards. Furthermore, some unique islet behaviors, presumably evolved in response to lack of predators and to ameliorate chronic conditions of food shortage, appear to render islet lizards strongly susceptible to cat predation. These behaviors include rarely utilizing available refugia, and moving towards anything new, including cat decoys. Nonetheless, I found that repeated exposures over three trials led to significant increases in FIDs for all populations, indicating at least some behavioral plasticity. My results suggest that although lizards may adapt their antipredator behaviors to cope with introduced predators, this offers at best only partial protection, so that there remains strong concern about their survival in the face of expanding feral cat populations.

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Exotic predators have driven the extinction of many island species. We examined impacts of feral cats on the abundance and anti-predator behaviours of Aegean wall lizards in the Cyclades (Greece), where cats were introduced thousands of years ago.We compared populations with high and low cat density on Naxos Island and populations on surrounding islets with no cats. Cats reduced wall lizard populations by half. Lizards facing greater risk from cats stayed closer to refuges, were more likely to shed their tails in a standardized assay, and fled at greater distances when approached by either a person in the field or amounted cat decoy in the laboratory. All populations showed phenotypic plasticity in flight initiation distance, suggesting that this feature is ancient and could have helped wall lizards survive the initial introduction of cats to the region. Lizards from islets sought shelter less frequently and often initially approached the cat decoy. These differences reflect changes since islet isolation and could render islet lizards strongly susceptible to cat predation.

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In June 1973 the Greek islands Amorgos, Thira and Milos were visited. On Amorgos numerous Lacerta erhardii amorgensis and Gymnodactylus kotschi kotschi were seen. Two Eryx jaculus turcicus were met, they are slow animals, one was picked up and stayed very quite. On Thira a lizard, probably Lacerta erhardii naxensis, was photographed. On Milos Vipera lebetina schweizeri and telescopus fallax fallax were seen, caught by a snakedealer.

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Though biological invasions constitute one of the biggest threats for global biodiversity, our understanding of the mechanisms that enable invasive species to outperform native species is still limited, especially, in terms of behavior. Most available studies have examined behavioral traits which favor invasive species on the later stages of invasion, however, our knowledge on earlier stages, namely, when alien species face novel environments and must exploit new resources, remains obscure. Here, we focus on one crucial behavioral trait, finding food. The Italian wall lizard (Podarcis siculus) has been widely introduced and established viable populations in S. Europe and N. America. We examined whether P. siculus has enhanced exploratory behavior and abilities to find food compared to two native congeneric species with which it may come in contact in the near future, an insular endemic (P. milensis) and a widely distributed lizard (P. erhardii). We performed a Y-maze experiment, in which we varied arm markings in a standard way to prevent learning. Podarcis siculus was more efficient than its congenerics in finding and consuming food. This exploitative superiority was persistent, more frequent and repetitive. Interesting behavioral differences were also detected within the native species. Some P. milensis individuals showed no interest in exploring the maze, while few P. erhardii individuals remained rather indifferent to food even after detecting it. Our results suggest that the invasive P. siculus displays behavioral traits that could provide better opportunities for survival in the new environment and thus facilitate establishment even in the presence of congenerics. This provides further support to the idea that behavior plays a crucial role in animal invasions.

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Given agriculture’s outsized proportion of global land use, food production systems must be examined for their role in conservation during this era of anthropogenic global change. Central to this objective is investigating opportunities where goals of food security and biodiversity preservation are complimentary rather than incongruent. Biological control of herbivorous pests by natural enemies is one such area of focus, where accruing evidence suggests landscape complexity fosters functional diversity of predators who offer free pest control services. Despite being widespread generalist predators, reptiles have infrequently been considered for their role in suppressing invertebrate populations. In this study, we investigated an insular population of the common Mediterranean lacertid, Podarcis erhardii (Aegean wall lizard), for its impact on invertebrate communities and herbivory damage through field enclosure experiments. We further studied P. erhardii’s functional response to known agricultural pests in a laboratory setting. Our field results revealed a significant negative interaction effect between lizard inclusion and time on the abundance of ground-dwelling arthropods sized 2-5 mm (! = -0.017, SE = 0.006, p = 0.006), and a marginally significant negative effect of lizard inclusion on arthropods > 5 mm (! = -0.303, SE = 0.153, p = 0.052), irrespective of time. We detected no effect of lizards on the abundance of airborne arthropods or prey < 2 mm. However, our functional response trials revealed lizards’ capacity to consume copious amounts of small prey when naturally aggregated, as well as large amounts of prey biomass relative to predator body size. We found no influence of lizard inclusion on herbivory damage, potentially a result of intraguild predation by lizards on large predatory arthropods and small sample size. Our study illustrates the capacity of lizards to suppress large or naturally clumped ground-dwelling prey, especially later in the growing season when resources become scarce. We argue that biocontrol services are best realized when P. erhardii is among a diverse assemblage of predators, in order to maximize control of herbivorous insects over appropriate spatiotemporal scales. Accordingly, we recommend future studies involve long-term investigation of food web level interactions, including traitmediated effects.

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Chemosystematic investigations on the position of Lacerta jayakari and the genera Gallotia and Psammodromus (Sauria; Lacertidae) 13 species of the genera Lacerta, Psammodromus and Gallotia (L. erhardii, L. filfolensis, L. bedriagae, L. cappadocica, L. jayakari, L. vivipara, L. trilineata, L. agilis, P. hispanicus, P. algirus, G. atlantica, G. galloti, G. stehlin:) have been examined by means of micro complement fixation tests (albumin) as well as by proteinelectrophoresis concerning 14 genetic loci. The results and their con-sequences are the follows: 1. L. cappadocica, L. bedriagae, and L. erhardii/L. filfolensis are representatives of phylogeneticly equivalent groups (subgenera). Their isolation in the Middle Miocene corresponds with the first contacts of the landmasses of Eurasia and Africa. 2. Lacerta jayakari may be considered as a representative of a further group equivalent in this aspect; though the immunological results would not exclude a position near the green lizards (subgenus Lacerta s. str.). 3. Because of this isolated position of L. jayakari for this species (and Lacerta cyanura) the subgenus Omanosaura subgen. nov. has been established. 4. Psammodromus hispanicus and P. algirus have probably been isolated of each other since the Middle Miocene, too. 5. The radiation of the genus Gallotia corresponds closely with genesis of the Canary Isles.

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Albumins from several species assigned to Lacerta s. 1. and Gallotia were investigated by means of the Micro-Complement fixation technique. The phylogenetic trees evaluated according to Fitch & Margoliash (1967) and Beverley & Wilson (1982) led us to infer a division, which is in accordance with protein electrophoretic results (Mayer & Tiedemann, 1982) as well as with the subgeneric division used before the studies of Böhme (1971) and Arnold (1973).

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Recently, several works have focused on the lacertid lizards of the genus Podarcis, revealing cases of hidden diversity and paraphyly, and offering evidence that suggests the revision of the extant taxonomical arrangements within the genus. Hidden diversity and paraphyly have been shown to exist in the relationships betweentheBalkan species P. peloponnesiacaandP. erhardii as well. Herewecouple a molecular (mtDNA) dataset with a corresponding morphological one, consisting of morphometric and pholidotic characters, to check for concordance between the two. Phylogenetic analyses reinforced previous suggestions for paraphyly of P. erhardii with respect to P. peloponnesiaca.We found the variation of certain pholidotic characters concordant with the relationships inferred from partial mtDNA sequences, whereas morphometric characters were not. The latter is possibly due to greater influence of morphometric characters by environmental factors. To avoid the observed paraphyly we proceed with the description of the populations from Crete and the islet of Pori, until now designated as P. erhardii, as separate taxa at the species level.

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Variation in the reproductive traits of Greek populations of Podarcis erhardii, P. peloponnesiaca, and P. taurica living in similar habitats along the same latitude was examined. Female body size, clutch size, and egg volume were determined. In all three species, clutch size was positively correlated with maternal body size. P. erhardii appears to modulate its reproductive effort by means of variation in both egg size and number. On the other hand, egg size in P. peloponnesiaca and P. taurica has probably been optimized, and an increase in reproductive effort would result in the production of a larger number of eggs.

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Within-species colour variation is widespread among animals. Understanding how this arises can elucidate evolutionary mechanisms, such as those underlying reproductive isolation and speciation. Here, we investigated whether five island populations of Aegean wall lizards (Podarcis erhardii) have more effective camouflage against their own (local) island substrates than against other (non-local) island substrates to avian predators, and whether this was linked to island differences in substrate appearance. We also investigated whether degree of local substrate matching varied among island populations and between sexes. In most populations, both sexes were better matched against local backgrounds than against non-local backgrounds, particularly in terms of luminance (perceived lightness), which usually occurred when local and non-local backgrounds were different in appearance. This was found even between island populations that historically had a land connection and in populations that have been isolated relatively recently, suggesting that isolation in these distinct island environments has been sufficient to cause enhanced local background matching, sometimes on a rapid evolutionary time-scale. However, as expected, there was less evidence for heightened local matching in populations with a lower risk from avian predators and a prolonged history of volcanic activity. Overall, these results show that lizard coloration is tuned to provide camouflage in local environments, either due to genetic adaptation or changes during development. However, the occurrence and extent of selection for local matching may depend on specific conditions associated with local ecology and biogeographic history. These results emphasize how anti-predator adaptations to different environments can drive divergence within a species, which may contribute to reproductive isolation among populations and lead to ecological speciation.

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conspicuous sexual signals, which often have conflicting effects on survival. Here, we tested whether color variation between two island populations of Aegean wall lizards (Podarcis erhardii) is due to sexual dimorphism and differential survival of individuals varying in appearance. On both islands, we measured attack rates by wild avian predators on clay models matching the coloration of real male and female P. erhardii from each island population, modeled to avian predator vision. Avian predator attack rates differed among model treatments, although only on one island. Male-colored models, which were more conspicuous against their experimental backgrounds to avian predators, were accordingly detected and attacked more frequently by birds than less conspicuous female-colored models. This suggests that female coloration has evolved primarily under selection for camouflage, whereas sexually competing males exhibit costly conspicuous coloration. Unexpectedly, there was no difference in avian attack frequency between local and non-local model types. This may have arisen if the models did not resemble lizard coloration with sufficient precision, or if real lizards behaviorally choose backgrounds that improve camouflage. Overall, these results show that sexually dimorphic coloration can affect the risk of predator attacks, indicating that color variation within a species can be caused by interactions between natural and sexual selection. However, more work is needed to determine how these findings depend on the island environment that each population inhabits.

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Visual signals are often under conflicting selection to be hidden from predators while being conspicuous to mates and rivals. Here, we investigated whether 3 different island populations of Aegean wall lizards (Podarcis erhardii) with variable coloration among diverse island habitats exhibit simultaneous camouflage and sexual signals. We examined whether signals appear better tuned to conspecific vision as opposed to that of avian predators, and whether background-matching camouflage and sexual signals are partitioned to specific body regions. This could facilitate both covert sexual signaling and camouflage according to the viewing perspectives of predators and conspecifics. We found that lizards typically appeared twice as conspicuous to conspecifics than to avian predators against the same visual background, largely due to lizards’ enhanced sensitivity to ultraviolet, suggesting that P. erhardii signals are tuned to conspecific vision to reduce detection by predators. Males were more conspicuous than females to both predators and conspecifics. In 2 populations, male backs were relatively more camouflaged to predators compared to signaling flanks, whereas in females, exposed and concealed surfaces were camouflaged to predators and generally did not differ in background matching. These findings indicate that lizard coloration evolves under the competing demands of natural and sexual selection to promote signals that are visible to conspecifics while being less perceptible to avian predators. They also elucidate how interactions between natural and sexual selection influence signal detectability and partitioning to different body regions, highlighting the importance of considering receiver vision, viewing perspectives, and signaling environments in studies of signal evolution.

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The albumins from 41 species of the family Lacertidae representing a substantial part of genera as well as infrageneric groups of the collective genus Lacerta s.l. were investigated by means of the MCF technique. The data allow a chronological estimation of the most important radiation events and their correlation with paleogeographical facts. Our results are in remarkable contradiction to the phylogeny presented by ARNOLD (1989a) based mainly on morphological characters. The most important phylogenetic relationships are: 1. The genera Gallona and Psammodromus are already isolated from the other taxa since the Oligocene. We assess these main branches as subfamilies Gallotiinae and Lacertinae. 2. Lacerta s.str. and Lacerta subgenus Zootoca form the sister group of the remaining Lacertinae. 3. A colonization of Africa in the Lower Miocene has led to a separation in two lines, with mainly Eurasian and African members respectively. 4. ARNOLD`S (I.e.) `Ethiopian and advanced Saharo-Eurasian clade` proved to be a paraphyletic unit, one group of genera being more closely related to European taxa. All techniques applied to the systematics of Lacertidae hitherto are critically discussed. An area-time- hypothesis concerning the phylogeny of Lacertidae is presented. Key words: Lacertidae, phylogeny of; Gallotiinae subfam. nov.; Lacertinae subfam. nov.; micro complement fixation; albumin evolution.

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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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Small-lizards from the Greek islands Milos and Skiros are compared with populations from the continent of Podarcis taurica ionica, Podarcis muralis albanica and Podarcis erhardii by employing protein-electrophoretic data for 18 genloci. The gen frequencies of the polymorph loci are given and the genetic distances are calulated according to Nei. Extension of genetic distances is discussed. Podarcis milensis can be confirmed as a species. Populations of Milos and Skiros have shown to be closest related, both have to be considered as closely related to Podarcis taurica and not to Podarcis muralis.

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The collection of Amphibians and Reptiles of the Museum of Natural History of Ferrara has characteristics of historical representation, taxonomic and zoogeography that may be considered as examples of medium-sized types of collections more preva- lent in Italian museums. It consists of a historic part of the study and a donation from Guido Campadelli. The collection consists of 1739 specimens of amphibians (1734 of which held in 437 containers with ethanol and 5 taxidermized specimens) for a total of 78 amphibian species. There are 811 specimens of Reptiles (769 of which are stored in 526 containers with ethanol and 42 taxidermized specimens) representing a total of 108 species. The findings that form the collection of Amphibians come from 153 loca- lities of 15 states and 12 regions and 29 provinces of Italy. In the batrachological col- lection 62.1% of the total European species, and 94.3% of the total species in Italy, are represented. Zoogeographic regions most represented are the Palaearctic with 67% of the species of Amphibians in the collection and the Neotropical with 29%, fol- lowed by the Ethiopian (6%) and Nearctic (5%). Reptiles specimens come from 247 lo- calities of 15 states and 15 regions and 36 provinces of Italy. In the collection 46,2% of European species and 91,5% of the Italian ones are represented. The most represen- ted zoogeographic region are the Palaearctic with 89% of species in the collection, followed by the Ethiopian (14%), the Eastern (13%), Neotropical (11%) and the Nearctic (7%); australian and malagasy species do not exceed 3%. The herpetology collection was the subject of taxonomic and biogeographic analysis.

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Recently, biologists have become increasingly interested in cognitive variation among individuals and how it relates to differences in fitness. However, very few studies so far have studied the long-term repeatability and heritability of cognitive performance in wild animals. This is nevertheless crucial information to fully understand the potential ecological and evolutionary impact of individual variation in cognitive performance. In 2019, we assessed exploration, problem solving and spatial and reversal learning in 66 Aegean wall lizards, Podarcis erhardii, then released them in seminatural enclosures consisting of either simple or complex habitat. One year later, we recaptured and retested the surviving lizards and their offspring to estimate the long-term repeatability and heritability of these behavioural and cognitive characteristics. We found that exploration and spatial learning were moderately repeatable, but reversal learning only marginally and learning flexibility and problem solving not at all. Reversal learning ability declined over time in lizards kept in simple habitat, but not in those kept in complex habitats – suggesting habitat-dependent cognitive plasticity. To our knowledge, this is the first study demonstrating (long-term) consistency in cognitive traits within a nonavian reptile. The combination of modest repeatability and low heritability does suggest that, within our study species, personality and cognitive variation among individuals and populations is mostly moulded by environmental effects.

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Harsh and variable environments have been hypothesized to both drive and constrain the evolution towards higher cognitive abilities and behavioural flexibility. In this study, we compared the cognitive abilities of island and mainland Aegean wall lizards (Podarcis erhardii), which were expected to live in respectively a more variable and a more stable habitat. We used four proxies of behavioural flexibility: a neophobia assay, a problem-solving test and a spatial + reversal learning task. Surprisingly, the two populations did not differ in neophobia or problem-solving. Insular lizards, however, outperformed mainland conspecifics in an initial spatial learning task, but were less successful during the subsequent reversal learning. Our results thus seem to indicate that the effect of environmental variability on cognition is complex, as it may favour some, but not all aspects of behavioural flexibility.

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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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Aim Temperature influences most components of animal ecology and life history – but what kind of temperature? Physiologists usually examine the influence of body temperatures, while biogeographers and macroecologists tend to focus on environmental temperatures. We aim to examine the relationship between these two measures, to determine the factors that affect lizard body temperatures and to test the effect of both temperature measures on lizard life history. Location World-wide. Methods We used a large (861 species) global dataset of lizard body temperatures, and the mean annual temperatures across their geographic ranges to examine the relationships between body and mean annual temperatures.We then examined factors influencing body temperatures, and tested for the influence of both on ecological and life-history traits while accounting for the influence of shared ancestry. Results Body temperatures and mean annual temperatures are uncorrelated. However, accounting for activity time (nocturnal species have low body temperatures), use of space (fossorial and semi-aquatic species are ‘colder’), insularity (mainland species are ‘hotter’) and phylogeny, the two temperatures are positively correlated. High body temperatures are only associated with larger hatchlings and increased rates of biomass production. Annual temperatures are positively correlated with clutch frequency and annual longevity, and negatively correlated with clutch size, age at first reproduction and longevity. Main conclusions Lizards with low body temperatures do not seem to have ‘slower’ life-history attributes than species with high body temperatures. The longer seasons prevalent in warm regions, and physiological processes that operate while lizards are inactive (but warm enough), make environmental temperatures better predictors of lizard life-history variation than body temperatures. This surprisingly greater effect of environmental temperatures on lizard life histories hints that global warming may have a profound influence on lizard ecology and evolution.

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Islands are usually species-poor and insular animal communities likewise usually harbour fewer species than equivalent mainland communities. Intraspecific competition on islands is therefore usually weaker than it is on the mainland. Predation pressure often also decreases on islands, and some in many instances some insular organisms face no predators at all. Faced with such relaxed predation and competition pressures insular animals can evolve to exploit the wider resource spectrum than they can access in more species-rich environments. An extension of this scenario, called the “niche variation hypothesis”, results in populations of depauperate communities evolving greater morphological variability. This variability is often thought to manifest itself in enhanced sexual size dimorphism, with the larger sex growing larger still, whereas individuals of the smaller sex grow smaller, so as to reduce intra-specific competition. A clear prediction of this theory is therefore that communities with more competitors or predators will be less dimorphic. We tested this by measuring the sexual size dimorphism (SSD) of two Podarcis species in two Aegean archipelagos. We measured SVLs of female and male lizards across many islands and examined it against the number of other lacertid lizards, the number all lizards, and the numbers of snake species on each island. For each species (Podarcis gaigeae, 16 islands; P. erhardii, 27 islands) we also examined whether the presence, rather than the number of lacertids, lizards, and snakes, affects the degree of SSD. Podarcis gaigeae was, on average, more dimorphic than P. erhardii (males 14% longer vs. 6%). In neither species, however, was the degree of SSD affected by either the number or the presence of snakes, lacertid lizards or lizards in general. We conclude that on small islands lacking other reptiles the resource base is likely more limited as well, and thus does not allow the sexes to diverge to specialize on different-sized prey.

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Im Anschluß an die in der ersten Folge (1950) veröffentlichten Fälle von Reptilienbastarden werden weitere Bemerkungen über 33 Hybriden gemacht; davon betreffen 5 die Schildkröten und je 1 4 die Eidechsen und Schlangen. Nicht weniger als 14 sind Artbastarde, Die meisten von diesen zeigen einen intermediären Charakter. Es ist bemerkenswert, daß das „erytbronotus`-Merkmal bei Lacerta agilis sich offenbar dominant, das `concolor`-Merkmal bei Lacerta melisellensis fiumana und Lacerta s. sicula dagegen rezessiv vererbt. Eine besondere Beachtung verdient der hypothetische Bastard zwischen Lacerta a. agilis und v. viridis aus Maria Taferl in Niederösterreich sowie die gezogenen Bastarde zwischen Lacerta v. viridis und schreiberi. F1-Bastarde von Natrix n. natrix und n. persa sprechen für die Dominanz des Merkmals „gestreift`.

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Purpose During the last three decades, the number of devices that emit non-ionizing electromagnetic radiation (EMR) at the wireless communication spectrum has rapidly increased and possible effects on living organisms have become a major concern. The purpose of this study was to investigate the effects of radiofrequency EMR emitted by a widely used wireless communication device, namely the Digital Enhanced Communication Telephony (DECT) base, on the immune responses of the Aegean wall lizard (Podarcis erhardii). Materials and methods Adult male lizards were exposed 24 h/day for 8 weeks to 1880–1900 MHz DECT base radiation at average electric field intensity of 3.2 V/m. Immune reactivity was assessed using the phytohemagglutinin (PHA) skin swelling and mixed lymphocyte reaction (MLR) tests. Results Our results revealed a noticeable suppression (approximately 45%) of inflammatory responses in EMR-exposed lizards compared to sham-exposed animals. T cell-mediated responses were marginally affected. Conclusion Daily radiofrequency EMR exposure seems to affect, at least partially, the immunocompetence of the Aegean wall lizard.

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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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The present study is a review of the herpetofauna of the sourtheastern part of the Black Sea Coast and Strandzha Nature Park. A total of thirty nine species were recorded for both regions (12 amphinian species, 27 - reptiles), of which two newt species (Lissotriton vulgaris, Triturus ivanbureshi), ten species of anurans (Bombina bombina, B. variegata, Bufo bufo, Bufotes viridis, Pelobates syriacus, Hyla orientalis, Rana dalmatina, Pelophylax ridibundus, P. cf. esculentus, P. bedriage), eleven species of lizards (Mediodactylus daniliewskii, Angius colchicus, Pseudopus apodus, Ablepharus kitaibelii, Lacerta agilis, L. trilineata, L. viridis, Podarcis muralis, P. tauricus, P. erhardii, Darevskia praticola), four species of native turtles and tortoises (Testudo graeca, T. hermanni, Emys orbicularis, Mauremys rivulata) and twelve species of snakes (Xerotyphlops vermicularis, Platyceps najadum, P. collaris, Natrix natrix, N. tessellata, Dolichophis caspius, Elaphe sauromates, Zamenis longissimus, Zamenis situla, Coronella austriaca, Malpolon insignitus, Vipera ammodytes). A comparison between the species compositon of the herpetofauna between South Black Sea Coast and Strandzha NP is made and analyzed.

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The consequences of fragmentation may have severe effects on the survival of isolated populations. The importance of this phenomenon is of a major importance in conservation biology and has initiated new disciplines such as conservation physiology and immunoecology. Insularity provides an excellent system for the study of fragmentation. Island`s features, such as area and age, have been reported to exert a direct impact on organisms: genetic diversity is positively correlated with island area but negatively correlated with island age and exactly the same pattern stands for the efficiency of immune system, since the last is depended directly on genetic variation. We worked in a system three islands of Central Aegean that differ in age and area: Naxos, a large (430 km2) and old island (16.350 y.a.), Kopries, a small, young islet (0.3 km2, 4.200 y.a.) and Daskalio, a tiny (0.01 km2) and very young (1.500 y.a.) islet. Genetic variation in these populations is described by the following relenence: Naxos > Daskalio > Kopries. Using this study model, we tried to assess the impact of fragentation on the immune system of Aegean wall lizard (Podarcis erhardii). We expected that reduced genetic variation in small and old islands would impair immune response increasing thus the susceptibility to infections by parasites. We used exclusively adult males in order to eliminate the impact of female particularities (oviposition, gravidity). Infection was made artificially using the hemoparasite Hepatozoon sp., which is a natural parasite of P. erhardii. We have to mention that this parasite comes from another taxon (P. cretensis), that considers as sister species to P. erhardii. We quantified parasite burden through light microscopy in blood smears every 4 days for two weeks. Parasite load was considerably lower in the case of Naxos, suggesting that lizards from this population are able to resist parasite infection. The reverse results were obtained from Kopries where infection was realized easily and quickly whereas the population from Daskalio shows an intermediate response. We believe that our findings could be attributed to the restricted genetic variation of Kopries, due to the small size and the age of the island. Hence island`s features seem to shape immune efficiency as result of decreased genetic diversity.

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Aim We discuss biogeographical hypotheses for the Mediterranean lizard species Podarcis and Teira within a phylogenetic framework based on partial mitochondrial DNA sequences. Methods We derived the most likely phylogenetic hypothesis from our data set (597 aligned positions from the 12S rDNA and phenyl tRNA) under parsimony, distance and maximum likelihood assumptions. Results The species usually included in Teira do not form a strongly monophyletic clade. In contrast, the monophyly of the genus Podarcis is rather well supported. Seven lineages are identified in the genus; in order of appearance within the tree, these are: the Balearic pityusensis and lilfordi pair, the sicula complex, a Tyrrhenian tiliguerta and raffonei pair, muralis, the Siculo-Maltese filfolensis and wagleriana pair, the Balkan group (erhardi, peloponnesiaca, milensis, melisellensis and taurica), and the Ibero-Maghrebian group (bocagei, atrata, hispanica and vaucheri). Conclusions The origin of the three European genera of lacertid assayed (Lacerta, Teira and Podarcis) is hypothesized to have occurred in the Oligocene. For Podarcis, a possible scenario of a Miocene diversification is derived from the sequence data, and the zoogeography of the lineages are discussed in relation to the palaeogeography of the Mediterranean. It is hypothesized that in the early history of the genus the main lineages separated by rapid, numerous and close events that produced a starting point very similar to a polytomy, hard to resolve by parsimony analysis of the data set.

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Temperature sensitivity of digestive processes has important ramifications for digestive performance in ectothermic vertebrates. We conducted a comparative analysis of temperature effects on digestive processes [gut passage times (GPTs) and apparent digestive efficiencies (ADEs)] in five lacertid lizards occurring in insular (Podarcis erhardii, P. gaigeae), and mainland (P. muralis, P. peloponnesiaca, Lacerta graeca) Mediterranean environments. GPTs were negatively correlated to temperature with mainland taxa having 10–20% longer GPTs than island taxa. In contrast to previous studies that estimate ADEs using bomb calorimetry, we compare ADEs by analyzing discrete efficiencies for lipids, sugars and proteins at three temperature regimes (20, 25, and 30°C); each of these categories produces different results. ADEs for lipids and sugars showed a monotonic increase with temperature whereas ADEs for proteins decreased with temperature. Island taxa had consistently higher ADEs than their mainland counterparts for lipids and for proteins but not for sugars. They are characterized by superior energy acquisition abilities despite significantly shorter GPTs. Their increased digestive performance relative to the mainland species appears to allow them to maximize energy acquisition in unproductive island environments where food availability is spatially and seasonally clustered.

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The ability of an animal to shed its tail is a widespread antipredator strategy among lizards. The degree of expression of this defense is expected to be shaped by prevailing environmental conditions including local predation pressure. We test these hypotheses by comparing several aspects of caudal autotomy in 15 Mediterranean lizard taxa existing across a swath of mainland and island localities that differ in the number and identity of predator species present. Autotomic ease varied substantially among the study populations, in a pattern that is best explained by the presence of vipers. Neither insularity nor the presence of other types of predators explain the observed autotomy rates. Final concentration of accumulated tail muscle lactate and duration of movement of a shed tail, two traits that were previously thought to relate to predation pressure, are in general not shaped by either predator diversity or insularity. Under conditions of relaxed predation selection, an uncoupling of different aspects of caudal autotomy exists, with some elements (ease of autotomy) declining faster than others (duration of movement, lactate concentration). We compared rates of shed tails in the field against rates of laboratory autotomies conducted under standardized conditions and found very high correlation values (r > 0.96). This suggests that field autotomy rates, rather than being a metric of predatory attacks, merely reflect the innate predisposition of a taxon to shed its tail.

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Both environmental temperatures and spatial heterogeneity can profoundly affect the biology of ectotherms. In lizards, thermoregulation may show high plasticity and may respond to environmental shifts. In the context of global climate change, lizards showing plastic thermoregulatory responses may be favored. In this study, we designed an experiment to evaluate the extent to which lizard thermoregulation responds to introduction to a new environment in a snapshot of time. In 2014, we captured individuals of the Aegean Wall lizard (Podarcis erhardii) from Naxos Island (429.8 km2) and released them onto two small, lizard-free islets, Galiatsos (0.0073 km2) and Kampana (0.004 km2) (Aegean Sea, Greece). In 2017, we returned to the islets and estimated the effectiveness (E), accuracy and precision of thermoregulation measuring operative, preferred (Tpref) and body temperatures. We hypothesized that the three habitats would differ in thermal quality and investigated the extent to which lizards from Naxos demonstrate plasticity when introduced to the novel, islet habitats. Thermal parameters did not differ between Galiatsos and Naxos and this was reflected in the similar E and Tpref. However, lizards from Kampana deviated in all focal traits from Naxos, resulting in higher E and a preference for higher Tpref. In sum, Naxos lizards shifted their thermoregulatory profile due to the idiosyncratic features of their new islet habitat. Our results advocate a high plasticity in lizard thermoregulation and suggest that there is room for effective responses to environmental changes, at least for Podarcis lizards in insular habitats.

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Caudal autotomy is one of the most effective and widespread defensive mechanisms among lizards. When predators grasp the tail, lizards have the ability to shed this limb from the point of the attack and further. Numerous factors have been reported to affect tail shedding performance such as temperature, age, predation pressure, intraspecific competition etc. Interestingly, the impact of sex on tail loss remains greatly understudied. This is interesting as there are theoretical reasons for why autotomic performance should vary with sex. Here we analyzed tail autotomy performance in the lab for 12 species of lacertid lizards belonging to six genera (Algyroides, Anatololacerta, Hellenolacerta, Lacerta, Ophisops, Podarcis) and the field frequencies of broken or regenerated tails of a single species (Podarcis erhardii) from a diversity of various insular populations. Our aim was to investigate whether sex affects caudal autotomy, as well as the duration of post-autotomic tail movement. Tail loss performance in the lab was assessed through a standardized predation simulation experiment. When trials resulted in autotomy, we recorded the movement of the thrashing tail with a digital timekeeper until cessation of movement. In the case of P. erhardii frequencies, the condition of the tail (intact, broken or regenerated) was recorded for each individual captured in the field. We found no effect of sex on tail loss in any of the species, either in the lab or in the field. We also failed to detect any influence of sex on the duration of tail movement. Our findings indicate that tail autotomy is a rather conservative strategy that is used in the same way and extent by both sexes. However, we have to stress out that our lab experiments took place during the non-reproductive period. It would be interesting to repeat this experiment in gravid females, as pregnancy alters most aspects of reptilian biology.

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Caudal autotomy is one of the most effective and widespread defensive mechanisms among lizards. When predators grasp the tail, lizards are able to shed it from the point of the attack and further. Numerous factors have been reported to affect tail-shedding performance such as temperature, age, predation pressure, intraspecific competition etc. Interestingly, the impact of sex on tail loss remains greatly understudied. Here, we analyzed tail autotomy performance, simulated in the lab, in 12 species of lacertid lizards belonging to five genera (Algyroides, Anatololacerta, Hellenolacerta, Ophisops, Podarcis). Our aim was to investigate whether sex affects caudal autotomy and/or the duration of post-autotomic tail movement. We failed to detect any effect of sex on tail loss in the species examined. Also, we did not find any sexual impact on the duration of tail movement after autotomy, with a single exception. Our findings suggest that autotomy serves as a defensive tactic equally in both sexes and is used in the same extent.

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Tail autotomy, the self‐induced tail separation from the body, is a common and effective antipredator mechanism in lizards. In this study, we examine the muscle energetics of tail shedding in six lacertid lizard species (Podarcis erhardii, Podarcis peloponnesiaca, Podarcis muralis, Podarcis gaigeae, Podarcis milensis, and Lacerta graeca) from the northeast Mediterranean region. Very long periods of postautotomy tail movement were demonstrated for all species ( min), and differences among species were not statistically significant. Postautotomy tail movement, powered by anaerobic muscle activity, resulted in a strong increase in lactate concentrations and a concomitant depletion of muscle glycogen of exhausted tails relative to resting tails. No significant differences were found in either lactate or glycogen concentrations among the species examined. Duration of movement was negatively correlated with final lactate concentrations. The lack of differentiation in postautotomy energetic physiology in this group of species that have evolved under very different predation environments indicates that postautotomy muscle metabolism involves an overall conservative suite of characters.

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Der Artikel beschreibt die griechische Insel Alonnisos als liebenswertes idyllischen Urlaubsziel für Naturfreunde abseits der großen Touristenströme. Eine Bildserie zeigt die ganze Variationsbreite der Färbung und Zeichnung von Podarcis erhardii ruthveni, und als „Bonbon“ wird noch ein Dolichophis caspius präsentiert. 80 % der beobachteten Eidechsen wiesen deutlich sichtbaren Milbenbefall auf. GUNTRAM DEICHSEL nimmt im Anhang Stellung.

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This article describes Alonnisos as a lovely and idyllic holiday destination for people interested in nature away from the crowd. An extensive series of pictures shows the whole range of variations in the colouring and marking of Podarcis erhardii ruthveni, and as a highlight a Dolichophis caspius is presented. 80% of the lizards observed displayed clearly visible mite infestation. GUNTRAM DEICHSEL comments on this observation in the appendix.

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This article describes Alonnisos as a lovely and idyllic holiday destination for people interested in nature away from the crowd. An extensive series of pictures shows the whole range of variations in the colouring and marking of Podarcis erhardii ruthveni, and as a highlight a Dolichophis caspius is presented. 80% of the lizards observed displayed clearly visible mite infestation. Guntram Deichsel comments on this observation in the appendix.

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Podarcis wall lizards mainly feed on coleopterans, orthopterans, arachnids, and other small invertebrates. However, Aegean wall lizards, Podarcis erhardii, are widely distributed across Aegean islands and are increasingly observed eating non-traditional food items ranging from plant material to conspecific eggs and body parts. Here, we report the first documented case of P. erhardii consuming a large centipede, Scolopendra cingulata. The predator-prey relationship between these species has appeared to go both ways and may intensify on islands.

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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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Different kinds of interactions between lizards and plants in the Mediterranean basin have been described. Lizards have shown to `use` plants as a refuge against predators, as a foraging site, as a thermal microhabitat or as a food resource. In the last case, they can either act as herbivores (+/- interaction, consuming vegetative and/or reproductive parts, and damaging the plant to a higher or lower degree) or as mutualists (+/+ interaction, by behaving as legitimate pollinators and/or seed dispersers). The importance of plants as a food resource for Mediterranean lizards has been found to be high in insular habitats, espescially in small islands, where the paucity of arthropod prey availability redirects the foraging activity of lizards towards other nutrient resources. In such small islands, plants may play a major role in the trophic ecology of lacertid lizards, as suggested by recent studies. Likewise, lizards may be important both as efficient pollen transporters and seed dispersersfor a variety of plants. Some of these lizard-plant interactions are unique. The effect lizards have on the reproductive and dispersal succes of some species may be very relevant for plant preserveation, and in turn, the preserveation of prticular behavioral and ecological traits of the particular lizard populations depends, at least partly, on the plants they interact with.

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Although the herpetofauna of Greece is well known (see for example, Werner, 1938; Wettstein, 1953, 1957; Ondrias, 1968; Chondropoulos, 1986, 1989), several distributional questions still remain unclear, mainly for the mainland but also for islands. ` In this paper we present some chorological information from a field trip of one month (April 1990) to mainland and insular Greece.

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The geographical position, mild climate, mountainous landscape, and habitat heterogeneity determine Bulgaria’s rich diversity of herpetofauna. Currently, 17 species of amphibians (20 subspecies) and 36 species of reptiles (45 subspecies) are known from Bulgaria. Due to its transitional position, Bulgaria lies at the southern or northern distribution limits of many species of amphibians and reptiles. Vertical distribution is presented for all species found in Bulgaria. The richest in terms of diversity and abundance are the areas below 400 m, where nearly 95 % of taxa occur. The geographical ranges of almost all amphibians and reptiles in Bulgaria are well-known, although on a local scale the information is often limited. Zoogeographical analysis demonstrates that amphibian species belong to nine chorotypes, of which the Eastern Mediterranean and the European chorotypes are represented with three species each. Out of the ten chorotypes recognized for the reptiles, 51.4 % of the species belong to the Eastern Mediterranean (25.7 %) and Turanian–Mediterranean (25.7 %) chorotypes. Only two subspecies are Bulgarian endemics (Salamandra salamandra beschkovi and Mediodactylus kotschyi rumelicus). Due to the high diversity and mosaic distribution of habitats in Bulgaria, the herpetofauna has a rather continuous occurrence and high relative abundance. Although 80 % of the Bulgarian herpetofauna is legally protected by the Biodiversity Protection Act (2002) and other conventions, some species are threatened through collection for trade or habitat changes due to construction. Several sites in the southern part of Bulgaria are very rich in herpetofauna (nine to ten species of amphibians and 19 to 21 species of reptiles).

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The Western Rhodopes (8732 km2) encompass vast forested areas and transborder mountain ridges up to 2191 m a.s.l between Bulgaria and Greece. The paper outlines all published herpetofaunal data for Bulgarian and Greek Western Rhodopes and supplements many original records. The herpetological diversity is remarkably high: 12 species of amphibians (4 tailed and 8 anurans) and 27 species of reptiles (2 tortoises, 2 terrapins, 12 lizards and 11 snakes). Altogether 34 species were found in the Bulgarian part, 32 were proved for the Greek part of the mountain. Two species of snakes, Typhlops vermicularis and Zamenis situla are considered extinct for the studied region. The most common and abundant amphibians are Salamandra salamandra, Rana temporaria, Rana graeca and Bufo bufo. The most widespread reptiles are Podarcis muralis, Lacerta viridis, Anguis fragilis, Natrix natrix and Coronella austriaca. Brief zoogeographical analysis and distribution maps for the majority of the species are provided. Herpetological conservation issues and threats are also discussed.

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The complex geological history of the Balkan Peninsula and especially of the Hellenic area (multiple geological events of land connections) during the late Tertiary have likely contributed to the diversification and distribution of many terrestrial animals. Located at the margin of the Eurasian and African plates, this area has experienced tremendous geological alterations since the late Tertiary. Connections offered the opportunities for biological dispersal, while submergence of land bridges brought about distributive isolation among related taxa. If this was a major element influencing the formation of the rich fauna of the Hellenic area, then the phylogenetic relationships of terrestrial taxa should reflect these paleogeographic events. Compared with what is known for the geological history of this region, the impact of those geological events on the historical biogeography of terrestrial animals is less well understood. The reconstruction of phylogenies is of primary importance in the understating of the dynamic patterns of evolution, i.e. the biogeography of a group and the bases of its biological diversity at any level. Although the phylogeny of the genus Podarcis has been the subject of much discussion the relationships among the species are still unclear. Because morphology is so uniform, it provides few characters for phylogenetic analysis, and these tend to be conflicting. Various karyological, immunological and protein electrophoretic studies have been made, but these usually involve only a minority of species, and results from different species combinations are equivocal. Wall lizards of the genus Podarcis (Sauria, Lacertidae) comprises 17-18 currently recognized species in southern Europe, where they are the predominant reptile group. The taxonomy of Podarcis is complex and unstable. Based on DNA sequence data the species of Podarcis falls into four main groups that have substantial geographic coherence (Western Island group, South western group, Italian group and Balkan group). The Balkan species are divided in two subgroups: the subgroup of P. taurica (P. taurica, P. milensis, P. gaigeae and perhaps P. melisellensis), and the subgroup of P. erhardii (P. erhardii and P. peloponnesiaca). We addressed the question of phylogenetic relations among the species of Podarcis encountered in Balkan Peninsula, as they can be inferred from partial mtDNA (cyt b and 16S) and nuclear (c-mos) sequences. Our data support the monophyly of Podarcis and suggest that there are three phylogenetic clades: the clade of P. taurica (P. taurica, P. gaigeae, P. milensis, and P. melisellensis); the clade of P. erhardii (P. erhardii and P. peloponnesiaca) and the clade of P. muralis (P. muralis and P. sicula). The phylogenetic affiliations produced from the molecular data do not agree with the species and especially subspecies groupings predicted from the morphological classification of the populations sampled. By examining intraspecific variability we have found that extant populations of P. erhardii are paraphyletic. The high level of genetic differentiation observed in the mtDNA data between the P. erhardii populations from the Crete and Pori islands and the rest of P. erhardii samples from Cyclades and continental Greece and its phylogenetic position raises questions regarding the inclusion of those populations (Crete and Pori) within P. erhardii. Since, based on morphology, P. peloponnesiaca is a well defined species found in sympatry with P. e. livadhiaca and accepted today as such, then Podarcis from Crete and Pori would also need to be raised to specific rank to avoid the paraphyly of P. erhardii. However, although the differentiation of populations from Crete and Pori islands is clear, we suggest the use of PhyloCode in order to avoid taxonomic confusion at least until further research mainly on morphology becomes available. Thus, for the time being, we refer to wall lizards of Crete and Pori P. erhardii*. With regard to morphological subspecies of P. taurica, P. gaigeae and P. erhardii, it is clear that they do not represent monophyletic units and should be re-evaluated in the light of new evidence. The topology of the phylogenetic trees produced and the genetic distances among the clades of the Podarcis may describe a biogeographic hypothesis for these species in the Balkan area. The pattern of the distribution of the proposed taxa is congruent with existing paleogeographic evidence, as inferred by the application of a molecular clock, which supports the view that DNA sequences may be a useful tool for the study of paleogeography. The data analyzed, stress the need for a reconsideration of the evolutionary history of Balkan Podarcis species and help overcome difficulties that classical taxonomy has encountered at both the species and subspecies level. As a whole, the examination of mtDNA lineages in the lizards of genus Podarcis may contribute substantially to the refining of its taxonomic status. Phylogenetic information can now be added to the knowledge of their morphology and distribution, producing a more accurate taxonomy for this group. Our results also confirm that the molecular information in conjunction with geological data can be used to resolve questions about the paleogeography of a region or on the phylogeography of a species.

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Five polymorphic microsatellite loci (containing di- and penta-nucleotide repeats) were developed for the Erhard`s wall lizard, Podarcis erhardii, an endemic species of southern Balkan Peninsula. The number of alleles ranged from five to 17. Observed and expected heterozygosities ranged from 0.28 to 0.92 and 0.57 to 0.91, respectively. These markers will provide a valuable tool for population genetics analyses and can contribute to the evaluation of conservation programs.

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Erhard’s wall lizard, Podarcis erhardii (Sauria: Lacertidae), is highly diversified in Greece and especially in the southern Aegean region. Out of the 28 recognized subspecies, 27 are found in Greece from the North Sporades island-complex in the North Aegean (grossly south of the 39th parallel) to the island of Crete in the South. The species exhibits great morphological and ecological plasticity and inhabits many different habitats from rocky islets and sandy shores to mountaintops as high as 2000 m. By examining intraspecific variability at a segment of the mitochondrial gene cytochrome b we have found that that extant populations of P. erhardii are paraphyletic. Furthermore, we have found that subspecies previously defined on the basis of morphological characteristics do not correspond to different molecular phylogenetic clades, so that their status should be reconsidered. The DNA based biogeographical and phylogenetic history of Podarcis in Southern Greece is congruent with available paleogeographic data of the region, which supports the view that DNA sequences may be a useful tool for the study of palaeogeography.

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Wall lizards of the genus Podarcis (Sauria, Lacertidae) comprise 17 currently recognized species in southern Europe, where they are the predominant nonavian reptile group. The taxonomy of Podarcis is complex and unstable. Based on DNA sequence data, the species of Podarcis falls into four main groups that have substantial geographic coherence (Western island group, southwestern group, Italian group, and Balkan Peninsula group). The Balkan Peninsula species are divided into two subgroups: the subgroup of P.taurica (P. taurica, P. milensis, P. gaigeae, and perhaps P. melisellensis), and the subgroup of P. erhardii (P. erhardii and P. pelo- ponnesiaca). In the present study, the question of phylogenetic relationships among the species of Podarcis encountered in the Balkan Peninsula was addressed using partial mtDNA sequences for cytochrome b (cyt b) and 16S rRNA (16S). The data support the mono- phyly of Podarcis and suggest that there are three phylogenetic clades: the clade A (P. taurica, P. gaigeae, P. milensis, and P. melisell- ensis); the clade B (P. erhardii and P. peloponnesiaca), and the clade C (P. muralis and P. sicula). By examining intraspeciWc relationships it was found that extant populations of P. erhardii are paraphyletic. Furthermore, subspecies previously deWned on the basis of morphological characteristics do not correspond to diVerent molecular phylogenetic clades, suggesting that their status should be reconsidered. The distinct geographic distribution of the major clades of the phylogenetic tree and its topology suggest a spatial and temporal sequence of phylogenetic separations that coincide with some major paleogeographic separations during the geological history of the Aegean Sea. The results stress the need for a reconsideration of the evolutionary history of Balkan Podarcis species and help overcome diYculties that classical taxonomy has encountered at both the species and subspecies level.

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Wall lizards of the genus Podarcis (Sauria, Lacertidae) comprise 17 currently recognized species in southern Europe, where they are the predominant reptile group. The taxonomy of Podarcis is complex and unstable. Based on DNA sequence data the species of Podarcis falls into four main groups that have substantial geographical conherence (western island group, southwestern group, Italian group and Balkan group). The Balkan species are divided in two subgroups: the subgroup of Podarcis taurica ( P. taurica, P. milensis, P. gaigeae and perhaps P. melisellensis ), and the subgroup of Podarcis erhardii ( P. erhardii and P. pelopon- nesiaca ). We addressed the question of phylogenetic relations among the species of the P. taurica subgroup encountered in Greece, as they can be inferred from partial mtDNA (cyt b and 16S) sequences. Our data support the monophyly of P. taurica subgroup and suggest that P. gaigeae, P. milensis and P. melisellensis form a clade, which thereinafter connects to P. taurica . Within the previous clade, P. gaigeae is more closely related to P. milensis than to P. melisellensis . However, the specimens of P. taurica were subdivided in two different groups. The first one includes the specimens from northeastern Greece, and the other group includes the specimens from the rest of continental Greece and Ionian islands. Because the molecular clock of the cyt b and 16 rRNA genes was not rejected in our model test, it is possible to estimate times of speciation events. Based on the splitting of the island of Crete from Peloponnisos [ c. 5 million years ago (Ma)], the evolutionary rate for the cyt b is 1.55% per million years (Myr) and for the 16S rRNA is 0.46% per Myr. These results suggest that the evolutionary history of P. taurica in Greece is more complex than a single evolutionary invasion. The data analysed, stress the need for a reconsideration of the evolutionary his- tory of Greek Podarcis species and help overcome difficulties that classical taxonomy has encountered at both the species level.

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Wall lizards of the genus Podarcis (Sauria, Lacertidae) are the predominant reptile group in southern Europe, including 24 recognized species. Mitochondrial DNA data have shown that, with the exception of P. muralis, the Podarcis species distributed in the Balkan peninsula form a species group that is further sub-divided into two subgroups: the one of “P. tauricus” consisting of P. tauricus, P. milensis, P. gaigeae, and P. melisellensis, and the other of “P. erhardii” comprising P. erhardii, P. levendis, P. cretensis, and P. peloponnesiacus. In an attempt to explore the Balkan Podarcis phylogenomic relationships, assess the levels of genetic structure and to re-evaluate the number of extant species, we employed phylogenomic and admixture approaches on ddRADseq (double digested Restriction site Associated DNA sequencing) genomic data. With this efficient Next Generation Sequencing approach, we were able to obtain a large number of genomic loci randomly distributed throughout the genome and use them to resolve the previously obscure phylogenetic relationships among the different Podarcis species distributed in the Balkans. The obtained phylogenomic relationships support the monophyly of both aforementioned subgroups and revealed several divergent lineages within each subgroup, stressing the need for taxonomic re-evaluation of Podarcis’ species in Balkans. The phylogenomic trees and the species delimitation analyses confirmed all recently recognized species (P. levendis, P. cretensis, and P. ionicus) and showed the presence of at least two more species, one in P. erhardii and the other in P. peloponnesiacus.

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The monophyletic species subgroup of Podarcis tauricus is distributed in the western and southern parts of the Balkans, and includes four species with unresolved and unstudied inter- and intra-specific phylogenetic relationships. Using sequence data from two mitochondrial and three nuclear genes and applying several phylogenetic methods and species delimitation approaches to an extensive dataset, we have reconstructed the phylogeny of the Podarcis wall lizards in the Balkans, and re-investigated the taxonomic status of the P. tauricus species subgroup. Multilocus analyses revealed that the aforementioned subgroup consists of five major clades, with P. melisellensis as its most basal taxon. Monophyly of P. tauricus sensu stricto is not supported, with one of the subspecies (P. t. ionicus) displaying great genetic diversity (hidden diversity or cryptic species). It comprises five, geographically distinct, subclades with genetic distances on the species level. Species delimitation approaches revealed nine species within the P. tauricus species subgroup (P. melisellensis, P. gaigeae, P. milensis, and six in the P. tauricus complex), underlining the necessity of taxonomic re-evaluation. We thus synonymize some previously recognized subspecies in this subgroup, elevate P. t. tauricus and P. g. gaigeae to the species level and suggest a distinct Albanian-Greek clade, provisionally named as the P. ionicus species complex. The latter clade comprises five unconfirmed candidate species that call for comprehensive studies in the future.

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The Aegean wall lizard Podarcis erhardii, is widely distributed across the islands of the Aegean Sea and has emerged as an important model for studying the evolution of life histories under a diversity of ecological and isolation conditions. While we have a relatively good understanding of the ecology and life history of the species, the parasite communities of the taxon remain almost completely unknown. Quantifying the composition of these communities in P. erhardii is not only important for autoecological reasons, but also because inter-island comparisons of the parasite communities of this host can shed light on the factors that structure parasite diversity in general. Here we investigate the gastrointestinal parasite communities of P.erhardii ruthveni populations occurring on 17 islands of the Sporades group in the NW Aegean Sea. We dissected the GI tracts of 118 lizards and all helminths encountered were identified and recorded. In all, 8 species of helminthes were found, 1 Trematode (Paradistomum mutabile), 1 Cestode (Oochoristica sp.) and 6 Nematodes (Parapharyngodon micipsae, Parapharyngodon bulbosus, Parapharyngodon echinatus, Spauligodon sp., Abbreviata sp., and Skrjabinelazia sp.). The prevalence, mean intensity, and mean abundance of infection were respectively 61%; 5.74 (± 11.43; range 1-90); and 3.5 (± 9.3; range 0-90). The Brillouin’s index of diversity was 0.048 (± 0.13; range 0-0.6). These values were lower than those obtained for P. erhardii from Cyclad Islands, and suggest that the investigated populations (i) harbour only impoverished helminth communities relative to other mainland and insular lacertid populations; (ii) persist on the non-herbivorous diet. These features can probably be best attributed to the insular conditions related to the trophic availability and the long periods of isolation of these island lizard populations.

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The Aegean wall lizard Podarcis erhardii, is widely distributed across the islands of the Aegean Sea (Greece). While there exists a relatively substantial body of knowledge on the ecology and life history of the species, the parasite communities of the taxon remain almost completely unknown. Quantifying the composition of these communities in P. erhardii is not only important for autoecological reasons, but also because inter-island comparisons of this lizard`s parasite communities can shed light on the factors that structure parasite diversity in general. Here we investigate the gastrointestinal parasite communities of P. erhardii populations occurring on 16 landbridge islands of the Sporades group in the NW Aegean Sea by examining the gastrointestinal tracts of 113 lizards. In all, 8 species of helminths were found: 1 Trematode (Paradistomum mutabile), 1 Cestode (Oochoristica sp.) and 6 Nematodes (Parapharyngodon micipsae, Parapharyngodon bulbosus, Parapharyngodon echinatus, Spauligodon sp., Abbreviata sp., and Skrjabinelazia sp.). The prevalence, mean intensity, and mean abundance of infection were respectively 63.71%; 6.01 (±11.71; range 1-90); and 3.57 (±9.5; range 0-90). Brillouin`s index of diversity for the Sporades was 0.048 (±0.13; range 0-0.142). These values were lower than for most other mainland and insular lacertid populations, and suggest that the investigated island populations harbor very depauperate helminth communities. The severe impoverishment of the parasite communities and the differential persistence of generalist parasite species with simple life cycles is most likely the result of a combination of insular environmental conditions (spatial and temporal isolation, arid climate, small host population sizes) and host life history characteristics (diet, simple gastrointestinal tract architecture). The paucity of parasites in these relictual island populations suggests that small reptile populations fragmented by anthropogenic activities may not be able to sustain their native parasite communities over the long term.

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Tissue regeneration is a fundamental evolutionary adaptation, which is well known in lizards that can regenerate their entire tail. However, numerous parameters of this process remain poorly understood. Lizard tail serves many functions. Thus, tail autotomy comes with many disadvantages and the need for quick regeneration is imperative. To provide the required energy and materials for caudal tissue building, lizards are expected to undergo a number of physiological and biochemical adjustments. Previous research showed that tail regeneration induces changes in the digestive process. Here, we investigated if and how tail regeneration affects the digestive performance in five wall lizard species deriving from mainland and island sites and questioned whether the association of tail regeneration and digestion is affected by species relationships or environmental features, including predation pressure. We expected that lizards from high predation environments would regenerate their tail faster and modify accordingly their digestive efficiency, prioritizing the digestion of proteins; the main building blocks for tissue repair. Second, we anticipated that the general food shortage on islands would inhibit the process. Our findings showed that all species shifted their digestive efficiency, as predicted. Elongation rate was higher in sites with stronger predation regime and this was also applied to the rate with which protein digestion raised. Gut passage time increases during regeneration so as to improve the nutrient absorbance, but among the islanders, the pace was more intense. The deviations between species should be attributed to the different ecological conditions prevailing on islands rather than to their phylogenetic relationships.

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In caudal autotomy, lizards shed their tail to escape from an attacking predator. Since the tail serves multiple functions, caudal regeneration is of pivotal importance. However, it is a demanding procedure that requires substantial energy and nutrients. Therefore, lizards have to increase energy income to fuel the extraordinary requirements of the regenerating tail. We presumed that autotomized lizards would adjust their digestion to acquire this additional energy. To clarify the effects of tail regeneration on digestion, we compared the digestive performance before autotomy, during regeneration, and after its completion. Tail regeneration indeed increased gut passage time but did not affect digestive performance in a uniform pattern: though protein income was maximized, lipid and sugar acquisition remained stable. This divergence in proteins may be attributed to their particular role in tail reconstruction, as they are the main building blocks for tissue formation.

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The major histocompatibility complex (MHC) plays a key role in disease resistance and is the most polymorphic gene region in vertebrates. Although habitat fragmentation is predicted to lead to a loss in MHC variation through drift, the impact of other evolutionary forces may counter this effect. Here we assess the impact of selection, drift, migration, and recombination on MHC class II and microsatellite variability in 14 island populations of the Aegean wall lizard Podarcis erhardii. Lizards were sampled from islands within the Cyclades (Greece) formed by rising sea levels as the last glacial maximum approximately 20,000 before present. Bathymetric data were used to determine the area and age of each island, allowing us to infer the corresponding magnitude and timing of genetic bottlenecks associated with island formation. Both MHC and microsatellite variation were positively associated with island area, supporting the hypothesis that drift governs neutral and adaptive variation in this system. However, MHC but not microsatellite variability declined significantly with island age. This discrepancy is likely due to the fact that microsatellites attain mutation-drift equilibrium more rapidly than MHC. Although we detected signals of balancing selection, recombination and migration, the effects of these evolutionary processes appeared negligible relative to drift. This study demonstrates how land bridge islands can provide novel insights into the impact of historical fragmentation on genetic diversity as well as help disentangle the effects of different evolutionary forces on neutral and adaptive diversity.

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During a two weeks stay in the west of Crete (25. 09. - 09. 10. 1996) the following species were observed: Lacerta trilineata polylepidota, Podarcis erhardii, Hemidactylus turcicus, Coluber gemonensis, Mauremys caspica rivulata, Trachemys scripta elegans and Bufo Viridis.

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Endemic wildlife species confined to islands are both ecologically unique and tend to be severely endangered. More than 60% of the species known to have gone extinct to date used to occur on islands (Spatz et al. 2017). While there are several reasons why island taxa tend to be threatened, susceptibility to exotic species, such as invasive predators, is an important contributor. Earlier research has suggested that loss of antipredator defenses in island species is at least partially responsible for this. Locomotor abilities, including sprint speed and stamina, are traits that are essential to escape predation. Here we tested the hypothesis that island species that have evolved in predator-poor environments have lost their ability to run fast – a key trait for escaping invasive predators. We examined maximal sprint speed and maximal stamina in the Aegean wall lizard (Podarcis erhardii). The species is widespread throughout the Aegean archipelago (Greece, Mediterranean Basin) and is found on islands with varying levels of predation pressure making it an ideal study organism. We show that on islands with predators, lizards run at higher speeds relative to lizards on predator-free islands. Not all predator categories however are of equal importance: presence of mammalian - but not other - predators was significantly associated with higher sprint speeds in island lizards. In contrast to sprint speed, stamina was not related to predation environment suggesting that this is not a trait that is under selection by predators. Stamina may therefore be less useful for predicting endangerment by introduced predators. Differences in locomotor performance were not attributable to differences in limb lengths. The results of this study suggest that conservation funding may be best allocated to protect the most susceptible wildlife populations that live explicitly on low-predation islands as well as on islands isolated for long periods of time. Key Words: Antipredator defenses, locomotor performance, sprint speed, predation, islands.

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In recent years, great attention has been paid to many Podarcis species for which the observed intra-specific variability often revealed species complexes still characterized by an unresolved relationship. When compared to other species, P. siculus underwent fewer revisions and the number of species hidden within this taxon may have been, therefore, underestimated. However, recent studies based on genetic and morphological data highlighted a marked differentiation of the populations inhabiting the western Pontine Archipelago. In the present work we used published genetic data (three mitochondrial and three nuclear gene fragments) from 25 Podarcis species to provide a multilocus phylogeny of the genus in order to understand the degree of differentiation of the western Pontine populations. In addition, we analyzed new morphometric traits (scale counts) of 151 specimens from the main islands of the Pontine Archipelago. The phylogenetic analysis revealed five principal Podarcis groups with biogeographic consistency. The genetic distinctiveness of the Podarcis populations of the western Pontine Islands is similar or even more ancient than those observed in numerous other pairs of Podarcis sister species. In the light of these evidences we raise the western Pontine lizards to specific rank; thus they should be referred to as Podarcis latastei.

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Caudal autotomy is a widespread antipredator mechanism among lizards. Thrashing the tail distracts the predator’s attention, facilitating the lizard’s escape. Regeneration occurs rapidly, but the regenerated tail differs from the original in many ways. In the present study, we examine probable physiological differences between, and the performance of, regenerated and original tails in Aegean Wall Lizard from Aegean islets. Autotomy was induced artificially. There was no difference in duration of tail movement. Moreover, we failed to detect any differences in lactate and glycogen concentrations. Differences in LDH activity and protein levels were not significant. However, lipid levels were higher in regenerated tails. Finally, regenerated tails show higher autotomy percentages when predation was simulated. We failed to detect large-scale physiological differences in regenerated tails (with the exception of lipids) in these particular populations.

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Caudal autotomy is a widespread antipredator mechanism among lizards. Thrashing the tail distracts the predator’s attention, facilitating the lizard’s escape. Regeneration occurs rapidly, but the regenerated tail differs from the original in many ways. In the present study, we examine probable physiological differences between, and the performance of, regenerated and original tails in Aegean Wall Lizard from Aegean islets. Autotomy was induced artificially. There was no difference in duration of tail movement. Moreover, we failed to detect any differences in lactate and glycogen concentrations. Differences in LDH activity and protein levels were not significant. However, lipid levels were higher in regenerated tails. Finally, regenerated tails show higher autotomy percentages when predation was simulated. We failed to detect large-scale physiological differences in regenerated tails (with the exception of lipids) in these particular populations.

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Palatal dentition in lizards is incompletely known, especially data on its variability are scarce. We studied variation in the number of pterygoid teeth in three species of Podarcis, a species-rich genus of lacertid lizards: terrestrial, P. siculus and saxicolous, P. erhardii and P. cretensis. In contrast to some previous studies, we found no sexual dimorphism in the number of palatal teeth in any of these species. The number of teeth was not correlated to lizard size. In our sample, P. cretensis on average had more teeth than did P. erhardii but fewer than did P. siculus. In addition, some specimens of P. cretensis and P. siculus showed asymmetry in the number of pterygoid teeth, which may be a result of anthropogenic pressure. The observed variability in the occurrence of palatal dentition illustrates the importance of scoring this character in phylogenetic analyses only on the basis of a sufficient sample.

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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 naming of species and of all systematic entities of living things are dynamic con- cepts. Through scientific research, changes in the systematics and names of amphibi- ans and reptiles are constantly being pro- posed, much to the chagrin of many profes- sional and amateur herpetologists. Yet most changes are necessary if taxonomy and systematics are to reflect evolutionary his- tory and phylogeny, rather than letting user–friendliness and conservatism prevail. The European herpetofauna and its taxo- nomy have received increasing attention through molecular (e.g. DNA) studies. Once most animal groups have been studied in this way, stability may be established. In the following pages, a concise overview of changes in the taxonomy of European amphibians and reptiles is presented. We have restricted ourselves to the geographi- cal boundaries of Europe, albeit excluding former Soviet states. Thus, politically Euro- pean areas in Asia (e.g. Cyprus and Greek islands in front of the Aegean Turkish coast) and Africa (e.g. Canary Islands and Ma- deira) have not been included. For a time frame we have used the species and names as presented in the well–known European field guide by ARNOLD (2002), although numerous changes were proposed and published prior to 2002. We refer to the cited literature for more comprehensive explanations. Only species or higher level changes concerning European taxa are listed. Subspecific changes and intraspeci- fic variability are noted only when contra- dicting long–established monotypy of a species, or when subspecies are being rejected. Vernacular names are mostly adopted from ARNOLD (2002). Some exo- genous species that are well–established and are reproducing on European soil, are included and are listed separately. Species believed to have been introduced to Europe over 100 years ago and persisting until today, have been included in the list of endogenous species. Many other non– native species have, however, been en- countered in the wild in Europe. Final content changes were made on De- cember 1, 2007. This type of update and the proposed species list will most likely be outdated at its date of publication. Further updating by means of a consecutive series of similar papers seems desirable.

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In den Jahren 2005 bis 2007 fanden im Becken von Feneos (Peloponnes, Griechenland) umfangreiche Untersuchungen zur Avifauna und Herpetofauna statt. Bei der erstmaligen Erfassung der Vogelwelt im Bergtal konnten 98 Vogelarten nachgewiesen werden, davon 81 Brutvögel, 14 Durchzügler und drei Nahrungsgäste. Für die Herpetofauna gelang der Nachweis von sieben Amphibien- und 26 Reptilienarten.

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Biologists have noted since the days of Darwin and Wallace how islandpopulations of vertebrates differ in body size from their mainland relatives. While much progress has been made in understanding global patterns in body size evolution across different species of wildlife (Benítez-López et al., 2021) scientists still don’t have a goodunderstanding of the ecological and environmental processes that generate these patterns. While many studies concerning vertebrate body size evolution have been performed, a hypothesis-driven approach that concerns life history differences in females and males is lacking, and it is not even known whether any putative factors influence female and male body size in the same way. We examined 36 populations of Aegean wall lizards (Podarcis erhardii, Lacertidae), a species widely distributed across the Balkan mainland and hundreds of Aegean islands. The species is unusual in that it not only occurs in a broad range of ecological conditions, but also that it varies substantially in body size (Males: 37%; Females: 33.9%) and that it occurs in numerous, clearly isolated and well-characterized island populations. It therefore constitutes an excellent model for determining the drivers of island body size across varying island environments. Here we evaluate multiple hypotheses ranging from predation pressure (predator species richness), resource availability (Normalized Difference Vegetation Index (NDVI), seabird density, island area), and competition for mates (lizard density) as possible factors affecting lizard body size. We found that the factors affecting both female and male lizard body size depend on the presence of nesting seabird colonies on an island. Female body size on seabird islands was most strongly associated with seabird colony density, while male body size on seabird islands was only associated with conspecific lizard population density, suggesting a role for seabird subsidies and increased food resources for the evolution of female body size and intraspecific competition in the evolution of male body size. On non-seabird islands, female body size was positively correlated with island size, while male body size was not significantly associated with any of the factors we measured. This suggests that different factors were associated with female and male body size evolution in island populations of P. erhardii, and we propose that a general pattern like the island rule is subject not only to selective forces that are sex-specific, but also that there will important differences dictated by island ecology.

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While global patterns in body size evolution in island vertebrates have been described extensively, the ecological processes that generate these patterns are not well understood. Here we used variation among lizard populations occupying an archipelago to test hypotheses about body size evolution. We examined 35 populations of Aegean wall lizards (Podarcis erhardii, Lacertidae), a species widely distributed across the Balkan mainland and hundreds of Aegean islands. We evaluated measures of resource availability (island area and seabird density), intraspecific competition (lizard abundance) and predation risk (presence of rats, carnivorous mammals, vipers, and birds of prey) as possible factors affecting lizard body size. Lizard body size increased with island size overall, as well as with seabird colony density, suggesting a role for increased food resources and especially seabird subsidies in the evolution of body size. Lizards were larger where lizard population density was higher, suggesting a possible role for intraspecific competition in the evolution of body size. In our sample, rats, carnivorous mammals, and vipers did not have obvious effects on lizard body size, and lizards were smaller on islands with resident birds of prey. Males were larger than females on average, yet sexual dimorphism did not vary consistently with measures of resource availability and competition. Overall, our results suggest that local resource levels predict population density and body size in these lizards, and that nesting seabirds in particular can substantially affect lizard body size.

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Podarcis is a genus of lacertid lizards. It contains over twenty species and represents one of the largest radiations of reptiles in Europe. Podarcis lizards show high molecular diversity but are relatively similar morphologically. Here, I describe cranial osteology of P. muralis, P. siculus, P. hispanicus*, P. melisellensis, P. erhardii, P. peloponnesiacus and P. cretensis. P. erhardii and P. cretensis – until recently regarded as conspecific and considered very similar morphologically – differ in several cranial characters. The studied population of P. cretensis, coming from Elafonisos island, shows some apparently heterochronic traits, such as tripartite occipital condyle, low ascending process of the supraoccipital, open parietal fossa and less well pronounced sexual dimorphism. Heterochrony could also play role in the evolution of P. hispanicus*. This is supported by the absence of median crest of the parietal. Morphometric analyses confirm sexual dimorphism in Podarcis lizards – males have relatively longer and higher skull which results in a higher bite force. This allows them to eat harder prey, enhances their chances in combats and helps maintain a secure grip during copulation.

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Although reptile fauna of FYR of Macedonia (FYROM) was the matter of interest since more than a century, detailed distributions of most of the species are still missing. According to published data, 32 reptile species inhabit FYROM. In this paper, detailed distribution data of reptile species in FYROM are presented. Research of herpetofauna of FYROM was conducted from 1999 until 2012 and included 393 localities. For 32 recorded reptile species, exact localities and altitudes, as well as published records are given. Analyses of the reptile diversity of the FYROM showed that the regions with the highest species diversity are: the Prespa and Ohrid Lake Region, Skopje region, Veles region and the Dojran Lake Region, with 18-21 reptile species per UTM 10 x 10 squares. These regions should be considered for the future designation of important herpetological areas at national level. Results of comparative analysis of reptile faunas from all Balkan countries showed that Macedonian fauna is the most similar with Albanian (92.537) and Bulgarian (90.909) reptile faunas. Zoogeographic analysis showed that reptile fauna of FYROM consists of eight chorotypes in total, the most dominant chorotype being the Eastern-Mediterranean with 13 species, followed by the Turano-Mediterranean with six and Southern-European with five species. Detailed distribution data of this study could be basis for future action plans, protection and conservation measures of reptiles in FYROM, and should be implemented in the national legislation and acts. In addition, this data are imperative for the Red Data Book of reptilians and designation of the Important Herpetological Areas in the country.

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Ammouliani is a small inhabited island of an area about 6.9 km2, located in the Gulf of Agion Oros, Chalkidiki, Greece. Hitherto, there have been no herpetological surveys on the island and thus its herpetofaunal composition is still unknown. In the current short note, records of amphibians and reptiles from Ammouliani Island are presented for the first time. A two-day expedition in Spring 2022 resulted in one amphibian and nine reptile species records, all found to occurin relatively low densities. The small number of species of the island and the low densities are discussed.

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During a field study in 1983-1984 in a 200 km 2 study area in the south of Evros province (Greece) the presence of 11 lizard species was established. For every species a distribution map within the area is supplied. In different parts of the area thc lizards formed different species combinations. Ophisops elegans appeared to vicariate in certain terrains with both Podarcis erhardii and P. taurica. By studying habitat selection and activity patterns of all species data on spatial and temporal niche segregation could be obtained. The activity patterns of all species were strongly influenced by exposition and altitude. In some species a strong altitude influence was also measured on growth and some other life history data. Some pairs of morphologically or ecologically related species are compared in more detail. Some species occurred in very high densities (Lacerta viridis up to 500, Ablepharus kitaibelii up to 1100 ind./ha). Tail loss frequencies of all species are given. For many birds of prey they formed an essential food source.

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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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As ecological conditions are transformed by a changing global climate, it is becoming increasingly essential to understand the vulnerability and response of organisms to these altered environmental circumstances. Whereas some strides have been made in understanding the effects of global climate change on endotherms, woefully little is known about ectotherms, which constitute the bulk of the diversity of species in natural ecosystems. Though ectothermic organisms can compensate for fluctuations in the thermal environment by modifying their behavior and physiology, these adjustments likely represent a tradeoff between maintaining an optimal body temperature and allocating energy to other important life history processes (i.e. foraging, anti-predator behaviors and mate-finding). While global climate change is likely to affect many aspects of a species’ life history and ecology (e.g. susceptibility to disease, food availability, etc.) some of the most direct impacts are likely to come from a possible mismatch between newly prevailing thermal conditions and the species’ thermal preferences. We investigated the thermal biology of four ecologically important reptile species in the Central Aegean Sea (Greece): Podarcis erhardii (Linnaeus 1756, Lacertidae), Laudakia stellio (Linnaeus 1758, Agamidae), Mediodactylus kotschyi (Steindachner 1870, Gekkonidae), and Hemidactylus turcicus (Linnaeus 1758, Gekkonidae). We determined the preferred body temperature for each species using a controlled thermal gradient in the lab. Combining these average thermal preferences with morphological measurements, field body temperatures, and microhabitat characteristics, our work sheds light on the thermal ecology of each species.

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Presence and impacts of pathogen infections in wildlife are known to be determined by a hierarchical set of factors including host, habitat, and general environment characteristics. While an increased number of studies have been documenting host-parasite interactions in natural systems, little is known about these impacts in island ecosystems. To elucidate the impacts of infection in island populations, we investigate the effects of Hepatazoon (and apicomplexan hemoparasite) wall lizard (Podarcis erhardii) populations, on 17 Cycladic islands (Aegean Sea, Greece). Specifically, we evaluated the effects of infection on multiple aspects of lizard life history from lizards from these islands. We found significant declines of condition in infected populations, though only in male lizards, suggesting a sex-specific response to infection. Mixed model analyses also suggest that some island level characteristics can affect the health and locomotion of lizards. These results indicate variability in the impacts of malarial infection across populations of P. erhardii.

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Temperature rules the lives of ectotherms. To perform basic biological functions, ectotherms must make behavioral adjustments to keep their body temperatures near a preferred temperature (Tpref). Many color polymorphic lizards are active thermoregulators and exhibit morph differences in traits related to thermoregulation, such as color, body size, and microhabitat use. The Aegean wall lizard, Podarcis erhardii, is a heliothermic lizard with orange, white, and yellow color morphs that differ in size, behavior, and microhabitat use. Here, we tested whether P. erhardii color morphs from the same population from Naxos island, Greece, differ in Tpref. We hypothesized that orange morphs would prefer lower temperatures than white and yellow morphs because orange morphs are often found on cooler substrates and in microhabitats with more vegetation cover. We obtained Tpref for 95 individuals using laboratory thermal gradient experiments of wild-caught lizards and found that orange morphs do, indeed, prefer cooler temperatures. Average orange morph Tpref was 2.85 C lower than average white and yellow morph Tpref. Our results add support to the idea that P. erhardii color morphs have multivariate alternative phenotypes and present the possibility that thermally heterogeneous environments play a role in the maintenance of color polymorphism in this species.

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In this paper we present confirmed and potential distribution ranges of all native Serbian reptile species. The information provided herein presents the combination of the newly collected faunistic data and previously published records. The centres of reptilian diversity in Serbia were evaluated, in order to focus future conservation efforts on the regions of particular importance for the protection of this understudied group of animals. We found four main centres of species richness: three in Metohija and one in Šumadija, with 17–21 species per 50 × 50 km square. Analysis of the similarity of species composition in different regions of Serbia showed that South-eastern Serbia is the most distinct from other regions of Serbia, due to high number of Mediterranean species, with two being found exclusively in that region (E. quatuorlineata and P. najadum). Metohija is also very distinct, due to the highest number of species and significant number of specific (Mediterranean) faunal elements. The remaining geographic regions are grouped into three clusters, with Kosovo-Southern Serbia cluster being especially distinct in the terms of species richness and the presence of Mediterranean species. We also compared Serbian herpetofauna with those in other Balkan countries, considering species’ numbers and zoogeographic herpetofaunistic elements. Serbian herpetofauna is closest to the Romanian. Zoogeographic analysis showed that reptilian fauna of Serbia consists of eight chorotypes, with the Eastern-Mediterranean (nine species) and Southern-European (five species) as the most dominant ones.

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We present the updated distribution of batracho- and herpetofauna in southern and south-eastern Serbia, based on literature and new field records for 16 amphibian and 22 reptile species. In these two biogeographic regions, already recognized as biodiversity hotspots, we discovered two new amphibian species with limited distribution: Hyla orientalis and Pelobates balcanicus. Also, one species (Triturus ivanbureshi) has restricted distribution, while one (Bombina bombina) is rare. Four reptile species have limited distributions and six are rare in these regions. The presence of hybrid zones and the influence of the Mediterranean climate make southern and south-eastern Serbia areas of great importance and the target for future ecological and conservation studies in Serbia.

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Benny Trapp, Fotograf, Griechenlandkenner und Leiter des Schutzprojektes `Afrikanisches Chamäleon` auf dem Peloponnes, stellt in diesem Buch alle 62 Reptilien- und Amphibienarten des griechischen Festlandes in faszinierenden Bildern vor. Die Informationen zu Beschreibung, Verbreitung, Lebensraum, Fortpflanzung und Verhalten jeder Art werden durch praktische Tipps zu Beobachtungsmöglichkeiten der oftmals scheuen und heimlich lebenden Tiere ergänzt. Neben Informationen über Land und Leute sowie Naturschutz in Griechenland erfährt man Nützliches über Giftschlangen und zur Naturfotografie. Mit seinen bestechenden, zahlreichen Fotos sicherlich eines der prächtigsten Bücher über europäische Amphibien und Reptilien.

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During a two week stay to the Cyclade island Tinos in May 2006 we were able to observe the Balkan Gieen Lizard in is natural habitat and take some photographs as well. Lacerta trilineata citrovittata in some cases showed a high flight distance of up to 5o m.

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During our vacation on the Greek islands Paros and Antiparos in May 2019 (10.5-20.5. on Paros and 21.5.-25.5. on Antiparos) we were able to photograph the Aegean Wall Lizard (Podarcis erhardii) at several places of which some of them are new records for this lizard. Our photographs illustrate the high variability of the subspecies Podarcis erhardii naxensis, which is widespread on these islands. Despite intense search we were unfortunately not able to find any Cyclades Giant Green Lizard (Lacerta citrovittata). Surprisingly, we found a population of the Yellow bellied Toad (Bombina variegata) near Lefkes. This is the first record for Paros.

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With this photo documentation we give a herpetological insight into three vacation trips to the Greek islands of Paros and Antiparos in the years 2019, 2020 and 2021. The focus of our interest was the search for further occurrences of the Aegean wall lizard (Podarcis erhardii naxensis) and the scarce and very shy Cycladic Green Lizard (Lacerta citrovittata). During our excursions, we always had an eye for the accompanying herpetofauna and nature in general, which allows us to introduce some more interesting species in this article.

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During our vacation on the Greek islands Paros and Antiparos in May 2019 (10.5-20.5. on Paros and 21.5.-25.5. on Antiparos) we were able to photograph the Aegean Wall Lizard (Podarcis erhardii) at several places of which some of them are new records for this lizard. Our photographs illustrate the high variability of the subspecies Podarcis erhardii naxensis, which is widespread on these islands. Despite intense search we were unfortunately not able to find any Balkan Green Lizards of the subspecies Lacerta trilineata citrovittata. Surprisingly, we found a population of the Yellow-bellied Toad (Bombina variegata) near Lefkes. This is the first record for Paros.

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During a stay on the Cyclades island of Andros, in addition to the ubiquitous Erhard’s Wall Lizard (Podarcis erhardii mykonensis), some specimens of the magnificent Cyclades Giant Green Lizard (Lacerta citrovittata) could also be photographed. The exceptionally high variability of coloring and markings in the Erhard’s Wall Lizards is remarkably. Photographs of the accompanying herpetofauna, habitats and landscapes offer further interesting insights into the impressive nature of this island.

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Caudal autotomy, the self-induced tail shedding from the body, is a common defensive strategy in lizards. Tail loss is followed rapidly by regeneration, revealing the importance of a fully functional tail. Predation pressure has been traditionally correlated with autotomy performance. However there is a lack of evidence regarding the impact of predation regime on tail regeneration. Another important factor that has been neglected is the population density. Though it is well established that crowding alters various life-history traits, the impact of density on caudal regeneration remains understudied. In this paper we compared three island populations of the Aegean Wall Lizard (Podarcis erhardii) that have evolved under different levels of predation pressure: whereas the Naxos population is exposed to numerous predators the Kopries and Daskalio islet populations experience low predation pressure. To evaluate the effect of density-induced stress on caudal regeneration, lizards were treated under two conditions of housing, in single and crowded (six individuals) terraria. Tail sheding occurred prior to this treatment following a standardised method. The length of regenerated tail was recorded weekly. Regeneration rate was higher in single terraria when compared to crowded, reflecting the negative impact of crowding. However we failed to detect any statistically significant difference between single and crowded terraria in the case of Naxos. We believe that the underlying reason must be the heavy predation pressure under which rapid tail regeneration, even under unfavourable conditions, is crucial for survival. It seems that the imperious need for regeneration counteracts density-induced stress.

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Tail autotomy is one of the most efficient antipredator mechanisms in lizards. Tail condition is crucial to the overall fitness of a lizard and has important impacts on many seminal life history functions (energy storage, locomotion, social status). Regrowth of an autotomised tail is a complex and energy-consuming mechanism that deprives resources from other substantial processes such as reproduction or immune response. It has been established that regeneration rate and effectiveness may be influenced by many different factors. In the present study we tried to assess the impact of haemoparasites (Hepatozoon sp.) and intraspecific competition on regeneration performance. We conducted this study on the Aegean wall lizard deriving from Naxos complex biotopes (main island and surrounding islets) where population densities vary significantly and parasite loads may be different (Haemophysalis sp. ticks introduced on some islands by goats). In order to simulate these diverse densities lizards were housed in individual and group (8 animals per each) terraria. The length of the tail was measured weekly, Hepatozoon parasitemia was quantified by examining thin blood smears under the microscope whereas corticosterone levels, the main stress hormone, were recorded in all cases. We failed to detect any difference regarding parasite load in the regeneration rate among studied populations. In contrast, regeneration rate was lower in the case of lizards that were housed in group terraria. This suggests that though infection status has a minor impact on tail regrowth, population density affects thoroughly the feature under study.

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Autotomy consists in the voluntary shedding of an expendable body part under conditions of defense. The broad phylogenetic expression of this trait represents the evolutionary success of this strategy. Caudal autotomy in lizards is followed rapidly by regeneration, revealing the importance of a fully functional tail. Predation pressure has been traditionally correlated with the tail autotomy performance. However there is a lack of evidence regarding the impact of predation regime on tail regeneration. Another important factor that has been neglected is the population density. Though it is well established that intraspecific competition in dense (usually insular) populations increase tail shedding performance, the impact of density on caudal regeneration remains understudied. In this paper we worked with three populations of the Aegean wall lizard (Podarcis erhardii) experiencing different predation pressure: while Naxos island (the biggest in Cyclades group) hosts numerous predators, including snakes (and one species of viper), Kopries and Daskalio islets are snake-free biotopes. In order to evaluate the effect of density-induced stress on caudal regeneration, lizards were treated under two conditions of grouping, in one- individual and in six-idnividual terraria, and always in respect to their origin forming hence 6 different groups. Tail sheding occurred prior to this treatment following a standardised method. The length of the regenerated tail was recorder weekly. We used exclusively adult males in order to eliminate the impact of female particularities (oviposition, gravidity). Regeneration rate was higher in the case of one-individual terraria for both islets. We believe that this finding could be attributed to the stress conditions exerted in the “crowded” terraria. Nonetheless no significant difference was found in the rates between one- and six-individual terraria in Naxos Island. We believe that the underlying reason must be the heavy predation regime under which fast tail regeneration is of a crucial importance, even under unfavourably overcrowded conditions.

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In this paper, the updated checklist of Serbian amphibians and reptiles is presented. The updated checklist of Serbian amphibians and reptiles was compiled from the literature on distribution, taxonomy and phylogeny. The checklist contains 48 autochthonous and two introduced species. Five species underwent taxonomic revisions (Triturus ivanbureschi, T. macedonicus, Pelobates balcanicus, Bufotes viridis and Mediodactylus kotschyi). Two species were added after recent taxonomic splits of species’ complexes (Hyla orientalis and Anguis colchica). Presence of new (Lacerta trilineata) and introduced species (Hemidactylus turcicus) was confirmed during recent fieldwork. There are at least eight more species that can potentially be present in Serbia. The national protection legislative should be updated according to the current checklist.

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We explored the ontogenetic dynamics of the morphological and allometric disparity in the cranium shapes of twelve lacertid lizard species. The analysed species (Darevskia praticola, Dinarolacerta mosorensis, Iberolacerta horvathi, Lacerta agilis, L. trilineata, L. viridis, Podarcis erhardii, P. melisellensis, P. muralis, P. sicula, P. taurica and Zootoca vivipara) can be classified into different ecomorphs: terrestrial lizards that inhabit vegetated habitats (habitats with lush or sparse vegetation), saxicolous and shrub-climbing lizards. We observed that there was an overall increase in the morphological disparity (MD) during the ontogeny of the lacertid lizards. The ventral cranium, which is involved in the mechanics of jaw movement and feeding, showed higher levels of MD, an ontogenetic shift in the morphospace planes and more variable allometric patterns than more conserved dorsal crania. With respect to ecology, the allometric trajectories of the shrub-climbing species tended to cluster together, whereas the allometric trajectories of the saxicolous species were highly dispersed. Our results indicate that the ontogenetic patterns of morphological and allometric disparity in the lacertid lizards are modified by ecology and functional constraints and that the identical mechanisms that lead to intraspecific morphological variation also produce morphological divergence at higher taxonomic levels.

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We used Procrustes-based geometric morphometrics to explore morphological variability in dorsal and ventral cranium shape of twelve lacertid lizard species from the Balkan Peninsula (), at the juvenile (neonatal) and at the adult stage (females and males separately). These species differed in phylogenetic relatedness, overall size and habitat preference (terrestrial in overgrown habitats, terrestrial in sparsely vegetated habitats, saxicolous and semiarboreal). General pattern of shape variation appeared to be preserved throughout ontogeny, especially for the dorsal cranium - adult female and male morphospaces corresponded to the neonate morphospace. The inspection of morphospaces showed that the general pattern of shape variability was along the gradient from the smaller to the species with larger body size. Along this gradient, species clustered according to habitat preference. The main difference between neonates and adults was the position of semiarboreal species, which completely separated from the rest of the species at the adult stage. The overall morphological disparity (MD) increased during the course of ontogeny (from neonates to adult females and males). Ventral cranium, with its structures involved in mechanics of jaw movement and feeding, showed greater increase in MD, as well as the shift in the morphospace hyperplanes. On the generic level, Lacerta spp. showed significantly higher MD than Podarcis spp. In contrast to other ecological groups (terrestrial and semiarboreal) saxicolous lizards showed a tendency of decreasing shape disparity during ontogeny. The species from saxicolous group were phylogenetically heterogeneous but morphologically convergent due to the specific habitat constraints, and their distinctive MD pattern could be achieved by different allometric paths. The patterns of shape variation and MD were modified by ecology, functional constraints and different ontogenetic trajectories. Further studies on the ecomorphology, allometric diversity and morphological integration, as well as the reassessment of the problematic lacertid phylogeny are needed in order to shed more light on the complex relationships among morphology, ecology and phylogeny in lacertid lizards.

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Different factors and processes that produce phenotypic variation at the individual, population, or interspecific level can influence or alter the covariance structure among morphological traits. Therefore, studies of the patterns of integration and modularity at multiple levels—static, ontogenetic, and evolutionary, can provide invaluable data on underlying factors and processes that structured morphological variation, directed, or constrained evolutionary changes. Our dataset, consisting of cranium shape data for 14 lizard species from the family Lacertidae, with substantial samples of hatchlings and adults along with their inferred evolutionary relationships, enabled us to assess modularity and morphological integration at all three levels. Five, not mutually exclusive modularity hypotheses of lizard cranium, were tested, and the effects of allometry on intensity and the pattern of integration and modularity were estimated. We used geometric morphometrics to extract symmetric and asymmetric, as well as allometric and nonallometric, components of shape variation. At the static level, firm confirmation of cranial modularity was found for hypotheses which separate anterior and posterior functional compartments of the skull. At the ontogenetic level, two alternative hypotheses (the “anteroposterior” and “neurodermatocranial” hypotheses) of ventral cranial modularity were confirmed. At the evolutionary level, the “neurodermatocranial” hypothesis was confirmed for the ventral cranium, which is in accordance with the pattern observed at the ontogenetic level. The observed pattern of static modularity could be driven by functional demands and can be regarded as adaptive. Ontogenetic modularity and evolutionary modularity show the same developmental origin, indicating conservatism of modularity patterns driven by developmental constraints.

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In this study, we present updated distributional data for all Serbian lacertids, having taken into account the available previously published information, and the so far unpublished authors’ field records and evidence from other sources. According to our current knowledge, eight lacertid species inhabit Serbia. Analysis of lacertid diversity within the country showed that biogeographic regions with highest species diversity are Metohija (eight species) and Kosovo (seven species). Regions in Serbia are mutually similar concerning lacertid faunas (average Bray-Curtis dissimilarity index is 0.82). Comparative analysis of lacertid faunas among all Balkan countries showed that Serbian lacertid fauna is the most similar to those of Albania and the FYR of Macedonia, and then to faunas of Romania and Bulgaria. Zoogeographic analysis showed that Serbian lacertids belong to three distinct chorotypes, with the East-Mediterranean being the richest (four species). Generally, diversity of lacertids in Serbia is most strongly determined by topographic diversity, habitat heterogeneity, and climatic peculiarities, with the inflow of the sub-Mediterranean climate along several river valleys in the south being especially important. This paper is intended to serve as a basis for further research, since data on distribution of lacertid species in Serbia are far from complete, with large distributional gaps of recorded species and with unconfirmed presence of two species.

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We used geometric morphometrics to explore the influence of phylogenetic and allometric constraints as well as ecology on variation in cranium shape in five species of monophyletic, morphologically similar Podarcis lizards (Podarcis erhardii, Podarcis melisellensis, Podarcis muralis, Podarcis sicula and Podarcis taurica). These species belong to different clades, they differ in their habitat preferences and can be classified into two distinct morphotypes: saxicolous and terrestrial. We found (i) no phylogenetic signal in cranium shape, (ii) diverging allometric slopes among species, and (iii) a significant effect of habitat on cranium shape. The saxicolous species (P. erhardii and P. muralis) had crania with elongated parietals, elongated cranium bases, shortened anterior parts of the dorsal cranium, reduced chambers of the jaw adductor muscles and larger subocular foramina. These cranial features are adaptations that compensate for a flattened cranium, dwelling on vertical surfaces and seeking refuge in crevices. The crania of the terrestrial species (P. melisellensis, P. sicula and P. taurica) tended to be more elongate and robust, with enlarged chambers of the jaw adductor muscle, reduced skull bases and shortened parietals. Terrestrial species exhibited more variation in cranium shape than saxicolous species. Our study suggests that shape variation in Podarcis sp. lizards is largely influenced by ecology, which likely affects species-specific patterns of static allometry.

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Most recent molecular studies revealed the phylogeny of Greek Podarcis species, which for years remained elusive, due to discordant data produced from various chromosomal, complement fixation and protein studies. In this report, we analyzed cellular immune responses of spleen- derived lymphocytes from six allopatric Podarcis species encountered in Greece, by assessing two-way mixed lymphocyte reaction (MLR)- induced proliferation. On the basis of stimulation indices (S.I.) as determined from cultures set up from xenogeneic splenocytes coincubated in pairs, we generated a phylogenetic tree, fully consistent with the phylogenetic relationships of Podarcis as determined by parallel analyses based on partial mitochondrial (mt) DNA sequences. Although the exact mechanisms triggering lymphocyte responses in lizard two-way xenogeneic MLR are not fully understood, our results show the potential use of cell-mediated immune responses as an additional approach to mtDNA analysis, for species delimitation within specific lizard taxa.

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The aim of this study was to determine if divergence in habitat use among lacertid lizards is paralleled by morphological differentiation. For 35 lacertid species, we measured body, head and limb dimensions. Habitat use was inferred from the literature: ground-dwelling on open terrain, ground-dwelling in vegetated areas, shrub-climbing, tree-climbing, saxicolous (i.e. rock-climbing). Traditional (i.e. non-phylogenetic) statistical analyses suggest morphological differences among species groups with different habitat use. Ground-dwelling species from open habitats tend to have longer femurs, tibiae and humeri (relative to body length) than other groups. Cursorial (i.e. level-running) species have relatively high heads and trunks compared to climbing species. These differences follow biomechanical predictions and it is tempting to consider tham as adaptations to habitat use. However, phylogenetic analyses of the data fail to establish a clear relationship between habitat use and morphology in the data set considered. There is a weak indication that the differences in head and trunk height have evolved as an adaptation to different habitat use, but the differences in relative limb dimensions among species groups with different habitat use vanish. Either adaptation of limb dimensions to habitat use has not occurred in lacertid lizards, or our methods are unable to demonstrate such an adaptation. We show that uncertainties in the topology of the phylogenetic tree used are unlikely to influence the outcome of our study. We also address the fact that habitat use is often similar in different branches of the phylogenetic tree, and the consequences this may have for the power of our statistical analyses.

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The Aegean Wall lizard (Podarcis erhardii) has a wide distribution across the islands of the Aegean Sea. The seemingly unexplained absence of the species from Paros (Central Cyclades), as well as from most (but importantly not all) of the surrounding archipelago, has been noted for over a century and has remained one of the most tantalizing mysteries in Aegean herpetology. Over the last 10 years we visited Paros and all of the surrounding islands mapping out the distribution of the species and quantifying the local ecological conditions. Based on field observations, laboratory measurements and analysis of historical records and utilizing a tree regression approach we offer for the first time an explanation for the likely processes that produced the unusual distributional pattern of P. erhardii.

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In this paper we present distributional data for reptiles from different localities in the Western Balkans. During an 11 year period (2002-2012) we collected new data in the frame of spring student research camps organized by the Biology Students’ Society (DŠB). Surveys were conducted by members of the herpetological groups that were mostly biology students. Surveys were located on five islands Dugi otok, Brač, Korčula, Mljet, and Pag, one peninsula Pelješac (Croatia), in the continental area around Kamenovo and Lake Skadar (Montenegro), Dojran Lake (Macedonia), at lower Neretva Delta (Croatia) and around Niš (Serbia). Altogether 34 different reptile species were recorded in the surveys. The species lists comprised of 7-22 different species per single locality. The highest reptile biodiversity was recorded around Dojran Lake in Macedonia.

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Die Eidechsen Bulgariens und deren Lebensräume werden in Text und Bild vorgestellt. Die Autoren konnten auf mehreren Reisen neun der insgesamt zehn vorkommenden Lacertiden-Arten finden.

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Reproductive investment, including the average number of offspring produced by an organism, is one of the fundamental characteristics of a species. Among other things, it predicts a species’ resilience to environmental disruption: taxa that produce more offspring are able to recover more quickly from environmental perturbations and survive long-term environmental change. Despite the clear importance of this trait, ecologists do not have a good understanding of the primary drivers shaping the reproductive investment of each species. To answer this question, I compare the reproductive efforts of numerous island populations of the Aegean Wall Lizard (Podarcis erhardii), which differ in multiple key environmental characteristics. I test three hypotheses, namely that reproductive investment (measured as clutch size, clutch volume and egg volume) is: 1) positively associated with predation risk [‘Predation Risk Hypothesis’]; 2) positively associated with the presence of reliable vegetation cover that provides shelter [‘Gravid Female Protection Hypothesis’], and 3) limited by (and hence positively correlated with) food availability [‘Food Limitation Hypothesis’]. Although field data are consistent with all three hypotheses, statistical analysis shows strong support for the Predation Risk Hypothesis. The result not only shed light on which fundamental forces shape reproductive investment in island vertebrates, but can also help set conservation priorities by identifying the most sensitive populations because reduced reproductive ability can be predicted based on easily quantifiable island characteristics (number of sympatric predator species).