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Many animals use their excrements to communicate with others. In order to increase signal efficacy, animals often behaviourally select for specific defecation sites that maximize the detectability of their faecal deposits, such as the tip of rocks by some lizard species. However, the field conditions in which these observations are made make it difficult to reject alternative explanations of defecation site preference; rock tips may also provide better opportunities for thermoregulation, foraging, or escaping predators, and not solely for increasing the detectability of excrements. In addition, we still know little on whether lizard defecation behaviour varies within-species. In this laboratory study, we take an experimental approach to test defecation site preference of Podarcis melisellensis lizards in a standardized setting, and assess whether preferences differ between sexes, and among populations. Our findings show that in an environment where all stones provide equal thermoregulatory advantage, prey availability, and predator pressure, lizards still select for the largest stone in their territory as preferred defecation site. Moreover, we demonstrate that lizards’ defecation preference is a strong conservative behaviour, showing no significant intraspecific variation. Together, these findings corroborate the idea that lizards may defecate on prominent rocky substrates in order to increase (visual) detectability of the deposited faecal pellets.

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Skrjabinelazia Sypliaxov, 1930 comprises 10 species distinguished by several characters typical of the genus including, among the most important, the presence/absence of spicules, cuticle ornamentation and vesicles, head-shape, the presence/absence of a leaflet crown in the buccal cavity, female tail-shape and male cone-shape. The three samples studied are new species: S. boomkeri n. sp., a parasite of Pachydactylus turneri, Gekkonidae, from South Africa (Klaserie Reserve); S. vozae n. sp., a parasite of Lacerta vivipara, Lacertidae, from France (Cévennes), which is close to two lacertid parasites, S. taurica Sypliaxov, 1930 and L. hoffmanni Li, 1934, respectively from the Crimea and North China (Peking); and S. mawsangelae n. sp. (male unknown), a parasite of Christinus marmoratus, Gekkonidae, from Australia (Pearson Island), which is, surprisingly, distinct from Skrjabinelazia sp. of Angel & Mawson (1968) from the same host in another region (North of Adelaide) of South Australia. Two main groups are distinguished in Skrjabinelazia: the species with spicules which are parasitic in the Lacertidae, and the species with a gubernaculum only which are parasitic in the Gekkonidae. The unique species described from the Iguanidae, S. intermedia (Freitas, 1940) from Brazil (Para), also without spicules, seems to be derived from gekkonid parasites, as it also has an evolved oesophagus with a glandular region, unlike the simple oesophagus seen in the larval stages of Skrjabinelazia.

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Animals modulate intraspecific signal shape and intensity, notably during reproductive periods. Signal variability typically follows a seasonal scheme, traceable through the expression of visual, acoustic, chemical and behavioral patterns. The chemical channel is particularly important in lizards, as demonstrated by well-developed epidermal glands in the cloacal region that secrete lipids and proteins recognized by conspecifics. In males, the seasonal pattern of gland activity is underpinned by variation of circulating androgens. Changes in the composition of lipid secretions convey information about the signaler’s quality (e.g., size, immunity). Presumably, individual identity is associated with a protein signature present in the femoral secretions, but this has been poorly investigated. For the first time, we assessed the seasonal variability of the protein signal in relation to plasma testosterone level (T), glandular activity and the concentration of provitamin D3 in the lipid fraction. We sampled 174 male common wall lizards (Podarcis muralis) over the entire activity season. An elevation of T was observed one to two months before the secretion peak of lipids during the mating season; such expected delay between hormonal fluctuation and maximal physiological response fits well with the assumption that provitamin D3 indicates individual quality. One-dimensional electrophoretic analysis of proteins showed that gel bands were preserved over the season with an invariant region; a result in agreement with the hypothesis that proteins are stable identity signals. However, the relative intensity of bands varied markedly, synchronously with that of lipid secretion pattern. These variations of protein secretion suggest additional roles of proteins, an issue that requires further studies.

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The study of animal communities is mainly concerned with how the groupings of species are distributed in nature and the ways in which the members of these communities assemble and interact with each other. Pianka suggested that animals partition environmental resources in three basic ways: temporally, spatially, and trophically. Such differences in activities separate the niches, reduce competition, and presumably allow the coexistence of a variety of species in the communities. The present work compares the spatial, trophic and temporal niches of four reptile species, two geckos (Tarentola mauritanica and Hemidactylus turcicus) and two lizards (Podarcis muralis and Podarcis siculus) living in a same roman-age archeological park situated area inside Rome metropolitan area (central Italy), and sharing the same two-dimensional habitat (the roman aqueduct walls). The results showed an overall overlap among species higher than expected by chance for all considered ecological dimensions except for the spatial resource related to the vertical position on the wall. Surprisingly, despite living in a two-dimension habitat would expect an increase of interspecific interaction rate, no partition was observed between species with the most convergent ecological requirements. Although our hypotheses about lizard communities structuring and interactions in the urban habitat require further investigation, we think that our lizard community is based more on the ecological needs of each species rather than on species’ interactions. The observed spatial segregation between diurnal geckos and lizards would not support alone the interaction hypothesis. We speculated that the coexistence of ecologically overlapping species at the study area is allowed by a non-limiting availability of resources (above all food) that would result in the reduction of heterospecific competitive interactions and an overall wide overlap of resource use.

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To investigate the reservoir role of the lizard Psammodromus algirus for the Lyme disease spirochete, 199 lizards were trapped from April to October 2003 in El Jouza, northwestern Tunisia. In this site, the infection rate of free-living Ixodes ricinus (L.) by Borrelia was evaluated by immunofluorescence as 34.6% for adult ticks and 12.5% for nymphs. Eighty percent of P. algirus (117/146) captured during this study were infested by I. ricinus, the predominant tick species collected from lizards. The intensity of tick infestation of this host by larvae and nymphs ranged from 0.14 to 7.07 and from 1.5 to 6.58, respectively. These immature stages of I. ricinus were found on lizards in spring and the beginning of summer, with a peak of intensity during June (10.16 immature ticks by lizard). Tissue cultures from lizards and xenodiagnosis with larval I. ricinus were used to assess the infection and the ability, respectively, of infected lizards to transmit Borrelia to naive ticks. Seventeen percent of xenodiagnostic ticks (40/229) acquired B. lusitaniae while feeding on P. algirus. Therefore, we demonstrated the ability of the lizards to sustain Borrelia infection and to infect attached ticks, and we proved that P. algirus is a reservoir host competent to transmit B. lusitaniae.

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Remarks on some reptiles from Apulia (SE Italy), especially on Coluber viridi­flavus, and description of two new microinsular subspecies: Coluber viridiflavus antoniimanueli from S. Andrea Island, near Gallipoli, and Podarcis sicula aemiliani from the Apani Rocks, near Brindisi.

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Preliminary description of a new microinsular Italian wall lizard from Dino Island (Thyrrhenian Sea; Calabria): Podarcis sicula amparoae n. subsp.

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Auctor scrutatur- 1273 lacertas e paeninsula et insulis Italiciis (praetermissis insulis Etruscis, de quarum lacertis iam antea disseruerat), in quibus undecim subspecies ante ignotas invenit; de nonnullarum distributione disputat; et singulas species et subspecies accurate describit, rationem quamdam dicotomicam etiam tradens, qua facilius recognosci possint.

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Sexual size dimorphism (SSD) among species from the Lacertidae family is expressed mainly in body and head size, with males being larger than females, having larger heads but shorter trunks in relation to body size. Most lacertid species display male-biased SSD, but in few species, such as Lacerta agilis, females display larger size. The sand lizard Lacerta agilis is one of the most widespread lizard species in Eurasia and inhabits an area from central Greece and south Armenia to south Sweden in the North, and from Great Britain to the lake Baikal in the East. At least nine subspecies are currently recognized and two of them occur in Bulgaria. In the present study we analysed SSD in the poorly known subspecies Lacerta agilis bosnica, which inhabits the mountain areas on the Balkan Peninsula. We examined 9 somatometric traits, which were transformed into 10 indices, describing body proportions. All of the measurements were taken in live animals captured in mountains in Western Bulgaria (the mountains of Vitosha, Osogovo, Stara Planina, and Plana). We established statistically significant differences between the sexes in 9 of the indices. The differences could be described as follows: males have longer tails, longer legs (fore and hind legs), and larger heads (both in length and width). The similarities and differences in regards to SSD in other taxa from the genus Lacerta (for which similar research was done in Bulgaria) are further discussed.

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Lacerta pamphylica and Lacerta trilineata are two currently recognized green lizard species with a historically problematic taxonomy. In cases of tangled phylogenies, next‐generation sequencing and double‐digest restriction‐site‐associated DNA protocols can provide a wealth of genomic data and resolve difficult taxonomic issues. Here, we generated genome‐wide SNPs and mitochondrial sequences, and applied molecular species delimitation approaches to provide a stable taxonomy for the Aegean green lizards. Mitochondrial gene trees, genetic cluster delimitation and population structure analyses converged into recognizing the populations of (a) L. pamphylica, (b) east Aegean islands, Anatolia and Thrace (diplochondrodes lineage), (c) central Aegean islands (citrovittata), and (d) remaining Balkan populations and islands (trilineata), as separate clusters. Phylogenomic analyses revealed a split into two major clades, east and west of the Aegean Barrier, unambiguously showing a sister–clade relationship between pamphylica and diplochondrodes, rendering L. trilineata paraphyletic. Species delimitation models were tested in a Bayesian framework using the genomic SNPs: lumping all populations into a single ‘species’ had the lowest likelihood but the current taxonomy was also outperformed by all other models. All lines of evidence support the Pamphylian green lizard as a valid species; thus, east Aegean L. trilineata should also be considered a distinct species under the name Lacerta diplochondrodes. Finally, evidence from the mitochondrial and nuclear genomes is overwhelmingly in favour of recognizing the morphologically distinct Cycladian green lizards as a distinct species. We propose their elevation to full species under the name Lacerta citrovittata. All remaining insular and continental populations of the Balkan Peninsula represent the species L. trilineata.

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Numerous studies of urban environment impact on wildlife imply urbanization can have both negative and positive effects. Phenotypic variation of pileus in the common wall lizard (Podarcis muralis) was analysed to determine whether urbanization levels can be associated with developmental instability induced by environmental stress. Pileus developmental pathways and instability in natural, suburban and urban populations were quantified by patterns of size and shape, fluctuating asymmetry (FA), modular organization and integration, allometric trajectories and frequency of phenodeviants. Our results show high asymmetry and modular structure of pileus with the high frequency of phenodeviants for natural, suburban and urban populations indicating elevated developmental instability in all three habitat types. However, some peculiarities were observed comparing habitats – the lowest level of FA and integration in urban populations and unexpectedly high level of FA and frequency of phenodeviants in the natural population. In addition, significant correlations between symmetric and asymmetric shape patterns, and presence of modular organization for all three habitat types suggest that genetic/environmental and developmental parcellation are somewhat aligned. Our results indicate that pileus morphology varies in a complex manner and future studies that link physiological, behavioural and morphological parameters to demographic parameters and fitness are necessary to fully understand how environmental stress affects developmental instability.

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We calculated growth rate for the spiny-footed lizard (Acanthodactylus erythrurus) inhabiting coastal eastern Spain from long-term mark-recapture data. Growth curves differ between sexes, with males growing faster than females and achieving larger size maximums. In this population each sex reaches maturity at about 300 days of age, approximately 34% faster than males, and 28% faster than females studied in a population further south and west in Iberia. Our logarithmic growth model has an accuracy of 96.8% and high statistical significance (r = 0.96 for males and r = 0.97 for females). Although both the exponential curve of “best fit” for growth estimated for males (r = 0.81), and the linear curve of “best fit” estimated for females (r = 0.77) in a population from Cádiz (Busack and Jaksic, 1982) are also significant. The overlap between growth curves and the general data cloud of the population showed that at the end of the year the proportion of individuals younger than one year was 80% and the proportion of individuals older than one year was 20%. Our data, in agreement with calculated maximum life spans for males (1.9 years) and females (2.1 years) in Cádiz, suggest a life span of approximately two years for both sexes. Females seem to reproduce only in a unique season in their lives, at this locality, as in Cádiz (Busack and Klosterman, 1987) they likely lay only one clutch, whereas females in Morocco (Bons, 1962) may produce two.

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Mammals and birds are capable of navigating to a goal using learned map-like representations of space (i.e. place learning), but research assessing this navigational strategy in reptiles has produced inconclusive results, in part due to the use of procedures that do not take account of the peculiarities of reptilian behavior and physiology. Here I present a procedure suitable for testing spatial cognition that exploits a naturally evolved, ethologically relevant ability common to many lizards (i.e. refuge seeking behavior). The procedure requires lizards to learn the location of an open refuge inside a rectangular arena containing artificial refuges in every corner, using distal extramaze visual cues and with no local cues marking the location of the open refuge. The procedure probes the lizards’ place learning ability and effectively rules out the use of egocentric and response-based strategies. The described procedure was successfully used to demonstrate place learning in a lacertid lizard (Podarcis liolepis). Over the course of two weeks of training both the latency to entering the open refuge and the number of corners visited in each trial decreased gradually, indicating that learning had taken place in over 60% of the lizards tested. These results confirm that, under certain circumstances, lizards are capable of navigating to a goal using a place learning strategy.

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Food habits of Psamodrommus algirus Linnaeus, 1758 (Reptilia: Lacertidae) in Djurdjura, northern Algeria. -- In this study, we present the first data on the diet of Psammodromus algirus of Tala Guilef sector (Djurdjura National Park). The analysis of 43 feces allowed us to identify 245 food items, spread over 15 Operational Taxonomic Units (OTUs). Insects are the most consumed and, among them, the five most ingested categories of preys are: Coleoptera, Hymenoptera, Heteroptera, Homoptera, and Orthoptera. During the study period, the food spectrum is slightly diversified (BA = 0,41). Statistics analysis applied to prey categories proportions shows a significant influence of seasonal variation on the diet: x² = 42,94 p < 0,0001; ddl = 14. It presents a specialized diet in spring (BA = 0,25) concentrated essentially on Coleoptera and Hymenoptera. In summer, it became more opportunistic (BA = 0,46), but Coleoptera remain the most consumed category of preys.

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This thesis deals with the nature conservation of reptiles and amphibians in the Netherlands, the present practices and what should be done to improve them. Most of the species of the herpetofauna, as these groups together are known, are in a state of continuing decline. Obviously something is wrong; apart from the role played in this decline by loss of habitat, measures in the field are not effective for the sustainable survival of these species. This thesis has been written to rectify this; based on current knowledge of the ecology of these protected animals and on own experiences with nature management, it is meant to provide a guide and practical tool when carrying out the appropriate measures in the field. The introduction describes work that has been done internationally, especially at European level, to protect the herpetofauna. Further, how this has influenced policy plans and legislation in the Netherlands. It also points out the shortcomings, both in the approach of the policymakers, as well as in the current training of nature managers, and how these are reflected by the measures taken in the field. This information provides a background for chapter 7, the last chapter of this thesis, where the practice of management is discussed. The remaining chapters include five articles on ecological research; they also show how much time is needed to improve our knowledge on the ecology of these animals. Also, not understanding it can unwittingly create marginal conditions for a species, as was the case for the Slow-worm and Sand Lizard with the forest management on the Utrechtse Heuvelrug in the 1980s; this is described in the first two articles. That long-term surveys of population dynamics are more than necessary, is shown by the great fluctuations in the presence and activity of the European tree frog over the six years it was studied in the southwest of the Netherlands. Methodology is also discussed in this third article, and a practical formula given for estimating population size. Pond characteristics form the subject of the next article, wherein statistical analysis illustrates how they can indicate the possible presence and therefore the suitability of a pond for amphibians. Practical indications are given, useful when constructing new ponds as part of the Pond Action Plans. The last article has implications for the reintroduction of species into new or former habitats. That there is phenotypic variation within species of amphibians occurring within 200 kilometres of each other, shows that, to avoid failure, this should be taken into account in the selection both of the source population and the location of the habitat into which the animals are to be introduced. All these aspects are brought together in the final chapter where the practice of the conservation of both groups is discussed in turn. The habitats are the point of departure, and it is shown how conservation measures with plants or landscapes in mind can bring about deterioration or loss of habitat for a particular species of reptile or amphibian. The integration of management goals should be the approach for as far as is possible, but legal obligations may force us to take species specific measures into account for herpetofauna. Heathlands are of utmost importance for reptiles and the most relevant features of the management for this type of habitat are highlighted. For their part, amphibians find their major aquatic habitat in ponds. Details are given of the management for various amphibian species, both in ponds, and as far as we know it, in their habitat on land. The extensive literature on these subjects, found at the end of Chapters 1 and 7, as well as that of the articles, provides opportunity for going further into depth. The recommendations from Chapter 7, dealing with the structure of the vegetation to a considerable extent, should improve the situation of the herpetofauna in the Netherlands.

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Maastricht herbergt ’s lands enige natuurlijke populatie muurhagedissen. Kernpopulaties van de soort zijn aanwezig in de Hoge en Lage Fronten (vestingwerken) en op een spoorlijn en emplacement. Tevens zijn de hazelworm en levendbarende hagedis in het Belvédèregebied (Maastricht Noord) aanwezig. In het gebied staan diverse stedelijke en infrastructurele ontwikkelingen op stapel, welke in veel gevallen zullen leiden tot vernietiging, versnippering en/of verstoring van reptielenleefgebied. De aanwezige reptielsoorten en hun leefgebieden zijn streng beschermd en aantasting van de populaties is zonder ontheffing op de Flora en faunawet niet toegestaan. In overleg met het ministerie van EL&I, de initiatiefnemers van de plannen en RAVON is besloten te zoeken naar een integrale oplossing voor de diverse projecten. Het doel dat hierbij is gesteld, is de instandhouding van duurzame en robuuste metapopulaties van de aanwezige reptielen. Door het creëren van een „robuust raamwerk” wordt de bescherming van de reptielen in Maastricht Noord geborgd, waarbij ook de genoemde ruimtelijke ontwikkelingen doorgang kunnen vinden. In het voorliggende rapport wordt de actuele verspreiding van reptielen in het gebied, de huidige kwaliteit van de leefgebieden en de mogelijke maatregelen voor realisering van duurzame metapopulaties reptielen beschreven. Uit het veldonderzoek naar reptielen blijkt dat de muurhagedis voorkomt in de Hoge Fronten, Lage Fronten, emplacement Boschpoort, spoorlijn Maastricht – Lanaken, Maasoever en het bedrijventerrein Bosscherveld. Aanvullende waarnemingen op de bekende verspreiding zijn gedaan op diverse bedrijventerreinen in het Bosscherveld. Door de beperkte populatieomvang (<500 adulte dieren) en de versnippering van leefgebieden wordt de staat van instandhouding van de populatie muurhagedissen niet duurzaam geacht. De hazelworm komt in grote delen van het Belvédèregebied in hoge dichtheid voor en de populatie wordt in de huidige staat duurzaam geacht. De levendbarende hagedis is uitsluitend aanwezig op een klein stuk van de spoorlijn aan de Belgische grens. De populatie is klein, kwetsbaar en niet in een duurzame staat. Voor de drie reptielsoorten wordt het raamwerk beschreven voor robuuste metapopulaties. Een duurzame metapopulatie bestaat uit minimaal 500 adulte dieren, verspreid over meerdere kernpopulaties in primaire leefgebieden die met elkaar in contact staan door (satellietpopulaties in) verbindingszones. Voor de realisering van een robuuste metapopulatie van de muurhagedis is de ontwikkeling van nieuw leefgebied noodzakelijk (Fort Willem, uitbreiding Frontenpark en Belvédèreberg). Diverse verbindingszones dienen nog gerealiseerd te worden door aanpassing van inrichting en beheer en de aanleg van passages. Per leefgebied en verbindingszone worden in het voorliggende rapport de noodzakelijke maatregelen beschreven en wordt een prioritering aangebracht hierin. Door realisering van de voorgesteld maatregelen kan de populatie muurhagedis groeien tot 500 – 900 adulte exemplaren. Nieuw leefgebied voor met name de muurhagedis kan gerealiseerd worden door het aanbrengen van stenige en houtige structuren (stapelmuren, steenkorven, steenhopen, steenbedden en houtstapels). Hergebruik van materialen uit het gebied (zoals sloopafval) voor de bouw van deze voorzieningen is goed mogelijk Reptielvriendelijk vegetatiebeheer is van cruciaal belang om leefgebieden geschikt te houden. De wijze waarop wegen passeerbaar gemaakt kunnen worden voor reptielen, middels robuuste voorzieningen, wordt beschreven. De wijze waarop de deelprojecten in het gebied uitgevoerd worden dient vastgelegd te worden in een uitvoeringsplan, met aandacht voor werkwijze, materiaalkeuze, tijdstip van uitvoering en een goed afstemming in tijd van deelprojecten. Ook is ecologische begeleiding van de werkzaamheden van groot belang. De bestaande plannen in het gebied, zoals de vestingvisie, bestemmingsplan Noorderbrug en de voorbereidingsbesluiten Bosscherveld Noord en RMP Noord, bieden naast bedreigingen zeker ook kansen voor de aanwezige reptielen. Met name de vestingvisie heeft een hoge ecologische ambitie. Binnen de stedelijke ontwikkelingsplannen zijn er voldoende mogelijkheden om verlies en verstoring van leefgebied te compenseren. De realisering van robuuste metapopulaties biedt voor met name de muurhagedis goed uitzicht op duurzaam voortbestaan. Een netwerk van met elkaar verbonden deelpopulaties functioneert echter alleen wanneer alle onderdelen gerealiseerd worden.

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The use of information from chemical cues about the presence of conspecifics and their social interactions may be advantageous because it allows individuals to assess the social environment in the absence of the signallers. We tested experimentally whether the selection of nocturnal shelters by juveniles of the common lizard was influenced by the scent marks from three isolated or three socially housed adult males or females, keeping constant the number of donors for all treatments. We gave each juvenile a choice between a shelter containing odours from adults and a shelter with no odour and we compared the response to odours from three adults housed singly with that to odours from three adults that had the opportunity to interact. The shelter site selection of juveniles was influenced by the odour of socially housed adult males, but not by that of isolated males, and partly depended on the mother’s site of origin and the juvenile’s body condition. This study shows that juveniles use social information from conspecific chemical cues and that various phenotypes may use this information in different ways.

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Apicomplexan haemoparasites of the genera Schellackia Reichenow, 1919, and Karyolysus Labbé, 1894, seem to be common in lizards and widespread across the world. For decades, their identification has been based on morphological descriptions and life cycle patterns, with molecular characterizations, applied only recently. We used molecular characterization to confirm the identification of haemoparasites detected by microscopy in blood smears of Lacerta schreiberi Bedriaga, 1878, a lizard of the Iberian Peninsula. Since blood samples other than blood smears were not available from the studied lizards, 264 engorged ticks Ixodes ricinus (Linneaus, 1758) collected from them were used as an alternative non-invasive source of haemoparasite DNA for molecular genetic analyses. Of the 48 blood smears microscopically examined, 31 were positive for blood parasites (64.6% prevalence). We identified trophozoites and gamonts similar to Karyolysus lacazei (Labbé, 1894) (24/48; 50%) and Schellackia-like sporozoites (20/48; 41.7%). Mixed infections with both species occurred in 13 blood smears (27.1%). Sequence data were obtained for both parasites from engorged ticks. Phylogenetic analyses placed our unique haemogregarine sequence within the Karyolysus clade, nevertheless, within substantial polytomy. Thus, according to its morphology and effect on the host cell, we refer to this haemogregarine as Karyolysus cf. lacazei. Besides the Schellackia sequences being identical to a previously identified haplotype, we also obtained sequences of three new closely related haplotypes.

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Lizards and small rodents were live captured in Tuscany, central Italy, from May through August 2005. Prevalence of infestation by larval Ixodes ricinus L. (Acari: Ixodidae) and mean numbers of larvae per host were not significantly different for common wall lizards, Podarcis muralis Laurenti, and Apodemus spp. mice, whereas infestation levels by nymphs were significantly greater on lizards. Borrelia lusitaniae, which was previously shown to be dominant in host-seeking I. ricinus in the same study area, was detected by polymerase chain reaction (PCR) in 19.8% (95% confidence interval: 14.4, 26.0) of larval ticks and in 52.9% (27.8, 77.0) of nymphs that were collected from lizards. Moreover, 18.8% (7.2, 36.4) and 25.0 (3.2, 65.1) of lizards` tail biopsies and blood samples, respectively, were positive for B. lusitaniae. Conversely, attached ticks and ear biopsies from Apodemus spp. mice were PCR negative. Passerine birds belonging to 10 species were live captured in March 2005, and Borrelia valaisiana was detected in 57.1% (18.4, 90.1) of I. ricinus nymphs feeding on Eurasian blackbirds, Turdus merula L. Results of this study suggest that lizards play an important role as reservoirs for B. lusitanae and may affect the dominance of this genospecies in the Mediterranean area.

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Studies of geographical distribution of parthenogenetic Darevskia armeniaca and its parental species D. mixta and D. valentini can extend our knowledge in terms of their biology and prediction of expansion of their ranges owing to increasing anthropogenic impact and global climate change. We studied the rock lizards of the genus Darevskia because their ecology, biology and potential ranges have not been well studied. Our objectives were: (1) to create maps of the potential geographical distribution of the lizard species; (2) to identify bioclimatic, topographic and landscape variables that determine their potential ranges; (3) to establish similarities and differences in ecological conditions of lizard localities between the parthenogenetic lizard and its parental species. We extended the available published data set with our own field survey records to assess effects of environmental factors on ecological niches of the lizards and their distribution in Transcaucasia and Asia Minor using methods of ordination and species distribution models (SDMs). Our results show that the main drivers regulating the distribution of Darevskia spp. are climatic, topographic and land cover/use. High SDM performance estimated as a Boyce index ~0.917 proved that we had incorporated the most important predictor variables for describing ecological niches of the lizard species. We used an ordination method to compare niches in terms of their overlap, similarity, and niche shifts of the parthenogenetic species relative to its parental species. Habitat preference of D. mixta was determined by altitude, vegetation type, high precipitation during the warmest season, low humidity variation, and low insolation. Habitats of D. valentini were predicted to be located at the highest altitude in the zone of subalpine meadows, low temperature during the dry months, low precipitation during the warm months, an average coefficient of seasonal variation in humidity, high insolation, and a short distance to roads. We suggest that the centroid of the niche of the “daughter” species, D. armeniaca, occupies an intermediate position along environmental gradients relative to that of its parental species with only one exclusion. The coefficient of humidity variation in D. armeniaca differed from that of its parental species. In general, we predicted that ecological niches of parthenogenetic and parental species overlapped by 36%, however, the ecological niche of D. armeniaca was similar to that of its “paternal” species D. valentini. We demonstrate that the parthenogenetic D. armeniaca prefers mountain forest, meadow, steppe and human-disturbed habitats, while the “maternal” species D. mixta inhabits mainly forest habitats and the “paternal” species D. valentini commonly occurs in mountain meadow and mountain steppe habitats in the subalpine zone. These results can be used by environment conservation organizations to preserve the most important habitats of these lizards.

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In different months during 2015-2017, the abundance and sex-age structure of the Sand lizard (Lacerta agilis) were studied on two transects. Parallel transects were established in the southeastern part of the cluster area «Burtinskaya steppe» of the Orenburg State Nature Reserve (Russia). The length of each transect was 3 km. The width of the mineralised plowed strip between the transects was 9-11 m. Transect 2 was located in the area burnt in August 2014, while transect 1 was situated in the area burnt in August 2016. In terms of size and external morphological traits, the captured Sand lizards were divided into three groups: juvenile individuals, adult males, and adult females. After measurements, the marked animals were put back to the places of capture. The number and sex-age structure were restored on transects in May June, after the first winter following the fire impact. We found that on transect 1, the species abundance at late April early May 2015 was almost three times higher (18.67 individuals / km) than on transect 2 (6.67 individuals / km). However, at late May 2015, the species abundance on both transects became almost the same (12.67 individuals / km in transect 1 against 11.33 individuals / km in transect 2). The juvenile individuals suffered from the fire influence more than adults. After the 2016 fire (i.e., in mid-May 2017), the proportion of one-year-old individuals was 21.2% of the total number of Sand lizards recorded in transect 1, while it counted 67.3% of the total number of Sand lizards recorded in transect 2, burnt in 2014. We believe that the rapid recovery of the abundance and sex-age structure in the Burtinskaya steppe was associated with the migration of Sand lizards through the mineralised strip separating the burnt and non-burnt transects.

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Prey often respond to predation risk by increasing refuge use, but this strategy may entail a loss of body condition. Factors responsible for this loss of body condition remain unclear. Also how prey deal with refuge use to cope with predation risk without incurring costs of body condition, and how initial body condition affects refuge use remain barely known. We analyzed in the field whether adult Iberolacerta cyreni lizards modify their escape strategies and refuge use in areas with different levels of habitat deterioration and ecotourism pressure, which represent different levels of predation risk, and the consequences of changes in antipredator behavior to body condition. Lizards inhabiting deteriorated areas, where risk is higher, remained closer to refuges, but decreased time spent hidden in refuges after attacks, probably to maintain similar body condition than lizards inhabiting natural areas. We performed two laboratory experiments to isolate potential costs of refuge use that might affect the body condition of male lizards: a) a decrease of the efficiency of digestion due to low temperatures inside refuges and/or b) a reduction in food intake. Results suggest that refuge use is costly in terms of body condition due to reduced food intake, but the loss of efficiency of digestion seems unimportant. Lizards modified refuge use in relation to their body condition, with lizards with worse condition decreasing time hidden after predatory attacks. We concluded that lizards compensated for increased predation risk with flexible antipredatory strategies, coping with risk without incurring costs for body condition.

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Deforestation may increase predation risk for prey because it may make prey more conspicuous and limit the number of refuges suitable to avoid predators. Therefore, prey may need to increase the magnitude of escape responses. However, excessive antipredatory effort might lead to a loss of body mass and a decrease in defense against parasites, with important consequences for short- and long-term fitness. We analyzed whether Psammodromus algirus (L., 1758) lizards that inhabit patches with different levels of deterioration of the vegetation within the same oak forest differed in relative abundance numbers, microhabitat use, antipredatory strategies, and health state. Results showed lizards selected similar microhabitats regardless of the level of deterioration of the vegetation and relative abundance of lizards was similar in both areas. However, habitat deterioration seemed to increase predation risk, at least for females, because they were detected at longer distances in deteriorated areas. Females seemed to adjust their antipredatory behavior accordingly to high risk of predation by increasing approach distances allowed to predators. The costs associated with frequent antipredatory displays might explain why females in deteriorated habitats had lower body condition and greater blood parasite loads than females in natural areas. This loss of body condition and increased parasitemia might have deleterious consequences for female fitness and therefore affect the maintenance of lizard populations in the long-term.

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The low-altitude populations of the threatenedlacertid lizard Iberolacerta monticola in A Coruña province have suffered population declines in the last decades. The present study has been done on the main population of this species in the Mandeo river basin, located in the river reserve of Chelo. Population size and density have been estimated, as well as other population parameters. We used the MARK program, which allows to use different variants of the Jolly-Seber method. In addition, we did an analysis of the sex-ratio and the structure of the population using size classes.The results indicated a high density, but limited to a reduced area with favourable habitat, surrounded by inadequate zones (forests with dense tree coverage),as well asa high mortality of immature individualsand an unequal proportion betweenimmaturesand adults. It seems to point out that this population shows insular characteristics. Other relevant data forthe survival of this population were obtained, as the apparent lack of predators, based on the low percentage of individuals with tail loss, or the reduced presence of individuals affected by ectoparasites of the genus Ixodes. The low number of observed movements points out that the individuals of this population are essentially sedentary. Some consequences for the survival of this population are set and some measures are proposed for its conservation.

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Light is essential for embryonic development in many oviparous animals including fish, amphibians, and birds. However, light may be harmful for reptile embryos developing underground where they are in complete darkness and beneath thin eggshells. Nonetheless, how embryonic light conditions affect reptile development and offspring remains largely unknown. Here we incubated eggs in dark and light conditions to determine the effects of light exposure on embryonic development and offspring visual ability, spatial cognitive ability and growth in a lacertid lizard, Eremias argus. Our experiments demonstrated that light stimulation shortened incubation duration of eggs, but did not affect hatching success, offspring size, visual ability or survival. More interestingly, light exposure during incubation decreased spatial cognitive ability and post-hatching growth of offspring. On the basis of negative effects on offspring growth rates, our study indicates that in squamate reptiles with thin eggshells, light exposure in early development has negative effects on offspring cognitive ability.

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

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Sexual organisms should be better suited than asexual ones in a context of continuous evolution in response to opposite organisms in changing environments (“Red Queen” hypothesis of sex). However, sex also carries costs associated with the maintenance of males and mating (sex cost hypothesis). Here, both non-mutually excluding hypotheses are tested by analysing the infestation by haemogregarines of mixed communities of Darevskia rock lizards composed of parthenogens generated by hybridisation and their bisexual relatives. Prevalence and intensity were recorded from 339 adult lizards belonging to six species from five syntopic localities and analysed using Generalized Mixed-Models (GLMM). Both infestation parameters depended on host-size (like due to longer exposure with age), sex and, for intensity, species. Once accounting for locality and species, males were more parasitized than conspecific females with bisexual species, but no signal of reproductive mode itself on parasitization was recovered. Essentially, male-male interactions increased haemogregarine intensity while females either sexual or asexual had similar reproductive costs when in the same conditions. These findings deviate from the predictions from “Red Queen” dynamics while asymmetric gender costs are here confirmed. Thus, increased parasitization pressure on males adds to other costs, such as higher social interactions and lower fecundity, to explain why parthenogenetic lizards apparently prevail in the short-term evolutionary scale. How this is translated in the long-term requires further phylogenetic analysis.

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Während zweier Reisen in den Jahren 2005 und 2011 konnten insgesamt vier Arten der Echten Eidechsen (Lacertidae) für den Nationalpark Plitvicer Seen (Kroatien) fotografisch nachgewiesen werden. Die Waldeidechse (Zootoca vivipara vivipara), die Smaragdeidechse (Lacerta viridis viridis), die Mauereidechse (Podarcis muralis muralis) und die Kroatische Gebirgseidechse (Iberolacerta horvathi) kommen in unterschiedlichen Dichten und Lebensräumen im Nationalpark vor.

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Es wird über die Erfahrungen bei der Haltung und Nachzucht der Grünen Baumeidechse oder Afrikanischen Baumeidechse (Gastropholis prasina) berichtet.

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Neben den autochthonen Vorkommen der Westlichen Smaragdeidechse existieren in Baden-Württemberg seit einigen Jahren mindestens zwei auf Aussetzungen beruhende Populationen der Art. Diese beiden reproduktionsfähigen Bestände sind in Stuttgart und bei Tübingen angesiedelt.

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Während unseres Urlaubs auf den griechischen Inseln Paros und Antiparos im Mai 2019 (10.5. – 20.5. Paros und 21.5. – 25.5. Antiparos) konnten wir die Ägäische Mauereidechse (Podarcis erhardii) an mehreren und zum Teil bis dato unbekannten Stellen nachweisen. Mit Fotografien wird die hohe Variabilität der auf diesen Inseln verbreiteten Unterart Podarcis erhardii naxensis dargestellt und kommentiert. Die Riesensmaragdeidechse, hier in der Unterart Lacerta trilineata citrovittata verbreitet, haben wir trotz intensiver Suche leider nicht finden können. Bei Lefkes fanden wir aber überraschenderweise eine Population Gelbbauchunken (Bombina variegata). Hierbei handelt es sich um den Erstnachweis für Paros.

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

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Contribution to the knowledge of the distribution of Horvath’s rock lizard Iberolacerta horvathi and Common wall lizard Podarcis muralis in the Friuli Venezia Giulia and in the Veneto regions (North-eastern Italy) – Data collected between 2002 and 2008, during an investigation aimed to fi nd unknown populations of Iberolacerta horvathi and at extending the knowledge about the distribution of Podarcis muralis in Northern Friuli Venezia Giulia and Veneto, are given. Iberolacerta horvathi has been found in 22 new localities, increasing of about 45% of those previously known in Italy, between 520 and 1825 metres a.s.l.. Podarcis muralis has been found in 103 localities, most of them in UTM squares (10X10 Km) not covered by the national atlas. In Friuli Venezia Giulia the species has been found regularly over 900 metres a.s.l. where suitable habitats are present, and locally it reaches 1800 metres a.s.l.. The two species have been found in syntopy in eight localities between 520 and 1800 metres a.s.l.. The Mount Dimon represents the highest locality where in Friuli Venezia Giulia Podarcis muralis has been found and the two lizards in Italy are in syntopy. Finally it has to be pointed out that the works for construction, maintenance and extension of walls, roads etcetera might drastically reduce the suitable habitat of the two species.

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There are limited studies on the Iskenderun Lizard, Acanthodactylus schreiberi Boulenger, 1878, which is known to distribute in a single area in Turkey, up to now. This study aimed to determine the distribution area, some ecological features, population size, species threatening factors and precautions to be taken for it. As a result of the field studies, it was determined that Acanthodactylus schreiberi distributes in the 7 local areas from Burnaz Coast in Erzin district to Dörtyol district, Hatay Province. Population size estimated as 13193 specimens.

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After a short synthesis of the present knowledge on the distribution of the Horvath’s rock lizard (Iberolacerta horvathi) in Italy, the Author refers about recent news and field verifications on the distribution of the lizard in Veneto and Friuli Venetia Giulia Regions (north-eastern Italy).

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Background The present study aimed to illustrate comparative morphological, histological, and functional variations of the hyoid apparatus of Acanthodactylus boskianus and Ptyodactylus guttatus. Result The hyoid apparatuses and musculature of the two investigated species are well-developed. The hyoid apparatuses exhibit high mobility with different articulation sites between their skeletal elements. The degree of ossification of the hyoid apparatus of gecko is more developed than that of the lacertid lizard. In P. guttatus, a well-developed synchondrosis articulation appears between the basihyoid and ceratobranchialis of the hyoid apparatus. Meanwhile, in A. boskianus, the articular ligament which appears between basihyoid and ceratobranchialis is less developed and also appears as a ligament connection between the two parts of hyoid cornua in which ligaments provide more mobility to the hyoid of lacertid lizard. Conclusion The results reflect a phylogenetically informative character about these clad and explain the different functional demands imposed on the hyoid apparatus as well as confirm the important role of hyoid apparatus in the movement of the tongue during the prey transport.

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Predation is one of the most important factors affecting biology, ecology and behaviour of the prey. We have studied predation of the common lizard (Zootoca vivipara) by the great grey shrike (Lanius excubitor) in farmland habitats in western Poland. Shrike caches were used as a source of information about preyed lizards. Shrikes hunt significantly more adult males than females, juveniles and sub-adults; the pattern was similar over all three study years. Male lizard had longer activity time than female; therefore, they seem to be under predation pressure for a longer time during breeding season. Capture and count transect data of common lizard populations living within and outside shrike territories showed significant seasonal differences: within shrike territories, there was lower proportion of males than females; moreover, in some territories, the number of males in the lizard population was negatively correlated with the number of males impaled by shrikes. Our findings suggest that in this particular predator-prey system, shrikes may be a strong selective force for lizards’ population dynamics. Male-biased predation could be caused by differences in the behaviour of adult male and female lizards, namely longer activity of males and differences in space and refuge use.

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Invasive species have been recognised as an important hazard to native communities. Amongst the mitigation measures that have been proposed to confront biological invasions, eradication projects are certainly the most drastic. In this short communication, a successful eradication project against a recently established population of the Italian wall lizard (Podarcis siculus) in Athens, Greece, is reported. To this aim, the Hellenic Herpetological Society received unforeseen aid from stray cats and, possibly, from the Οcellated skink (Chalcides ocellatus) and vegetation growth. Within three years, the initial thriving P. siculus population has shrunk to very few individuals.