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Aim The ocellated lizards of the genus Timon, comprising six species that are distributed across continental Asia, Europe and Africa, offer an interesting model to study the evolution of ecological niches through comparative phyloclimatic analysis. Our study provides insights into the evolutionary history of the ocellated lizards and helps to understand the role of climatic niche evolution during the speciation process. Location Eastern and western margins of the Mediterranean basin. Methods A dated molecular phylogeny was estimated based on three mitochondrial and two nuclear genes. Using multivariate statistics, species distribution models were developed to characterize the Grinnellian niches of all species. Subsequently, ancestral environmental niche occupancy of each taxon was reconstructed using niche occupancy profiles. Niche divergence among species was quantified by computing multivariate niche overlaps via twodimensional and n-dimensional approaches. Results Phylogenetic analysis supports that the ancestor of Timon diverged into the eastern and western groups following multiple vicariance events that shaped the current distribution pattern of Palaearctic lizards. High complexity in the ecological niche evolution between the Mediterranean and non-Mediterranean climatic regions was detected. The generally low niche divergence among members of the eastern group and the remarkable climatic divergence within the western group highlight an important role of temperature seasonality in a Mediterranean and Atlantic climate context. The results also suggest niche conservatism in terms of microhabitats described by vegetation cover. Main conclusions The ocellated lizards provide an interesting example of a vertebrate radiation where niche shift (with or without vicariance) and niche conservatism alternate in different niche axes shaping current biogeographical patterns.

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Die landgebundenen Wirbeltiere des festländischen Iberiens (Süßwasserfische, Amphibien, Reptilien mit Ausnahme der Meeresschildkröten, Säugetiere mit Ausnahme der Chiroptera, Pinnipedia und Cetacea) bilden, wie im einzelnen gezeigt wird, in ihrer Gesamtheit eine besondere faunistische Einheit: ungefähr die Hälfte der Formen (Spezies oder Subspezies) ist endemisch; das gilt für 73 % der echten Süßwasserfische, 47 % der Amphibien, 36 % der Reptilien und 53 % der Säugetiere. Die geographische Isolierung durch die Pyrenäen und das Mittelmeer ist gewiß die Hauptursache für die Entwicklung einer so eigenartigen Fauna, aber auch andere Faktoren (die sehr verschiedenen Klimate und Biotope innerhalb Iberiens, der im Vergleich mit anderen Gebieten Europas geringere Einfluß der Vereisungen usw. haben sicher ebenfalls eine sehr wichtige Rolle gespielt. Acanthodactylus erythrurus erythrurus, Algyroides hidalgoi, Algyroides marchi marchi, Algyroides marchi niethammeri, Lacerta agilis agilis, Lacerta hispanica hispanica, Lacerta hispanica bocagei, Lacerta hispanica vaucheri, Lacerta lepida lepida, Lacerta lepida nevadensis, Lacerta monticola monticola, Lacerta monticola cantabrica, Lacerta monticola cyreni, Lacerta schreiberi, Lacerta viridis viridis, Lacerta vivipara, Psammodromus algirus algirus, Psammodromus hispanicus hispanicus, Psammodromus hispanicus edwardsianus.

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The genus Timon currently comprises six species and is distributed in Europe, Asia and North Africa. The two European species are Timon nevadensis in south-eastern Spain and Timon lepidus in the rest of the Iberian Peninsula and parts of southern France (as well as an adjacent small area in Italy). There seems to be some intergrading between those two species. Two subspecies are recognized beside the nominate form Timon lepidus lepidus: Timon lepidus ibericus in north-eastern Iberia and Timon lepidus oteroi on Illa de Sálvora. Molecular research showed recently that five different lineages can be distinguished in Timon lepidus (MIRALDO et al. 2011). The various lineages differ to some extent in colour and pattern. Recently, PEEK (2011, 2017) reported a small-bodied form of T. lepidus from the north side of the Sierra de Gredos. Moreover there are two North African species of this genus, Timon pater and Timon tangitanus. Timon princeps and T. kurdistanicus inhabit regions in Asia, the former is found in Iran and the latter in Turkey, Iran and Iraq.

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I am passionate on keeping and breeding Ocellated Lizards for many years. Here, I report on the construction of an outdoor terrarium and my first experiences of keeping lizards outdoors!

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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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Insular biota represents a singular fraction of biodiversity in both evolutionary and ecological terms. However, it is especially sensitive to disturbance due to minimal interaction with competitor, predators and parasites, as well as low area and population size, which may lead to local extinction. Conserving such singularity involves monitoring organism quality and disturbance intensity to understand, anticipate and correct eventual declining trends. Reptiles are more likely to maintain populations in small islands than other vertebrates because of their low metabolic rates, resistance de dehydration and sedentariness. Some, particularly lizards, also display fast responses to insular conditions such as diet shifts. Timon nevadensis is a large lizard restricted to SE Iberia where it occupies a wide variety of open habitats, including some insular environments. The melanistic population of the Escombreras islet (off the coast of Cartagena, 4 ha) became threatened due to the accidental arrival of rabbits (Oryctolagus cuniculus), which altered the habitat structure and competed for the food. Annual campaigns of rabbit control are being conducted since 2018. Furthermore, in 2018 and 2019 Cartagena harbour authority promoted works to estimate the body condition, demography and population genetics parameters to assess the viability of the islet population. In 2018, four sampling sessions were carried out using 12 in vivo traps baited with fruit. A total of 62 captures were obtained with some individuals recaptured up to there times. Sexratio was 1.1 and only 1.6% of captures were juveniles. Abundance, estimated by the method of Schnabel, was 53 adults while, estimated by the method of Petersen, was 52. Samples from 25 lizards from Escobreras and 15 from the adjacent continent were genotyped with a battery of 16 microsatellites. The values of genetic diversity were by far lower (~50%) in the islet than in the continent, inbreeding was much higher (~300%) and effective population size was extremely small (Ne=41.7). Remarkably, relatedness between 6.1% of pairs in Escombreras attained > 0.5. Overall, these results indicate that the islet harbours a very small population with signs of inbreeding that may affect fitness. This, in combination with disturbance associated with rabbit introduction and the pensence of rats (Rattus norvegicus), makes it highly vulnerable extinction.

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South-western Europe has a rich diversity of lacertid lizards. In this study, we evaluated the occupancy patterns and niche segregation of five species of lacertids, focusing on large-bodied species (i.e., adults having >75 mm snout-vent length) that occur in south-western Europe (Italian to the Iberian Peninsula). We characterized the niches occupied by these species based on climate and vegetation cover properties. We expected some commonality among phylogenetically related species, but also patterns of habitat segregation mitigating competition between ecologically equivalent species. We used multivariate ordination and probabilistic methods to describe the occupancy patterns and evaluated niche evolution through phylogenetic analyses. Our results showed climate niche partitioning, but with a wide overlap in transitional zones, where segregation is maintained by species-specific responses to the vegetation cover. The analyses also showed that phylogenetically related species tend to share large parts of their habitat niches. The occurrence of independent evolutionary lineages contributed to the regional species richness favored by a long history of niche divergence.

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The Iberian Peninsula was one of the most important Pleistocene glacial refugia in Europe. A number of recent studies have documented the phylogeography of Iberian taxa and their relationship to more widely distributed species that expanded from this southern European refugium. We use a comparative approach to review the literature that challenges the paradigm of Iberia as a single refuge during Pleistocene glacial maxima and instead supports the occurrence of several Iberian refugia for a range of flora and fauna. Some patterns of phylogeographic concordance were found between the refugial areas identified by different case studies and these broadly overlapped with previously recognized areas of high endemism in the Iberian Peninsula. Such patterns help to illustrate the internal complexity of the Iberian Peninsula as a glacial refugium, and show that for many species, populations with a high degree of genetic structure have existed throughout the Pleistocene. Importantly, the occurrence of these ‘refugia within-refugia’ may confound the interpretation of phylogeographic patterns of European species, and can misleadingly support the occurrence of northern refugia. We discuss these and other consequences, especially when a limited number of samples from the southern European refugia are used.

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Ecological factors determining the geographic distribution of Timon lepida are poorly known. This work modelled the potential geographic distribution of Timon lepida at two spatial scales: 1. Landscape (Andalucía) 2. Regional (Spain) and analyzed the degree to which this distribution is associated with different predictor variables (e.g. temperature, solar radiation, topography, vegetation etc). The objectives of this study are to: (1) determine the most important predictor variables influencing the spatial distribution of Timon lepida; (2) generate potential geographic distribution maps for this species and (3) compare the predictive powers of environmental variables and hyper temporal NDVI to predict this distribution. Maxent, a presence-only distribution modelling technique was used to model predicted ranges for Timon lepida, using a large dataset of 10*10 km UTM presence only records collected between 1998-2002 period over Europe and a set of biophysical variable of 1*1 km resolution. To test the average behavior of the algorithm, 10 iterative models were produced by dividing all the presence records into 70% for training and 30% for testing. Three sets of model scenarios were generated: (i) models including environmental variables, (ii) models including environmental variables and vegetation and (iii) models including vegetation indices. Model accuracy was measured with binomial tests of omission rates and the area under the curve (AUC). All models were significantly better than random by the binomial test and AUC measure. The AUC score for models built using environmental variables was always higher indicating better discrimination of suitable and unsuitable areas for the species. For the two spatial scales, environmental variables models had a superior predictive ability than vegetation models. These findings did not support our hypothesis. The results indicate that at a landscape level, topographic variables (aspect and slope) are the most important whereas at a regional scale, climatic variables (temperature seasonality, solar radiation, altitude) and hyper temporal NDVI appear to have a significant effect on this distribution pattern. The results of the present study can be an important contribution to a better understanding of the ecological requirements of the species. The conclusions would be more precise if the adequate precise high resolution environmental data is included in the future application and reliable datasets of current conditions are identified to improve results.

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Measuring the diffusion of genes between diverging taxa through zones of secondary contact is an essential step to understand the extent and nature of the reproductive isolation that has been achieved. Previous studies have shown that the ocellated lizard (Lacerta lepida Daudin, 1802) has endured repeated range fragmentation associated with the climatic oscillations of the Plio-Pleistocene that promoted diversification of many different evolutionary units within the species. However, the oldest divergence within the group is estimated to have occurred much earlier, during the Miocene, around 9 Ma and corresponds to the split between the subspecies Lacerta lepida nevadensis Buchholz (1963) and Lacerta lepida lepida Daudin (1802). Although these two evolutionary units have documented genetic and morphological differentiation, most probably accumulated during periods of allopatry, little is known about patterns of gene flow between them. In this study, we performed a population genetic analysis of a putative area of secondary contact between these two taxa, using mtDNA and microsatellite data. We assessed levels of gene flow across the contact zone to clarify to what extent gene flow may be occurring. Hybridization between the subspecies was observed by the presence of genetically introgressed individuals. However, the overall coincidence of mitochondrial and multilocus nuclear clines and generally steep clines support the idea that this contact zone is acting as a barrier to gene flow. Taken together, these results suggest that L. l. lepida and L. l. nevadensis are in independent evolutionary trajectories and should be considered as two different species.

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The determination of the genetic basis of adaptive traits in natural populations is fundamental to better understand how populations adaptively diverge in heterogeneous environments and eventually give rise to new species. This work describes the efforts to identify candidate loci influenced by selection in ocellated lizards (Lacerta lepida) along an environmental gradient in the Iberian Peninsula, strongly affected by climatic variables. Two subspecies are recognized at the opposite extremes of the gradient, L. l. iberica in the northwest and L. l. nevadensis in the southeast, with morphological differences that suggest their local adaptation. Candidate loci were detected through an AFLP genome scan. Detection of candidate loci with a frequentist method and with Bayesian method resulted in a similar proportion of outliers (3-4%), but only a few loci were detected by both methods, denoting differences in methods’ sensitivity. Several outliers were associated with variation in temperature, insolation or precipitation along the gradient, suggesting the importance of these variables as selective pressures for local adaptation. Seven outliers were successfully characterized, being noncoding, with internal indels or repetitive elements as causes of length polymorphism, indicating that they might act as regulatory elements or are in linkage with the actual target of selection. The analysis of Mc1r, a candidate gene for coloration, revealed a nonconserved and derived substitution (T162I) associated with the brownish colour phenotype of L. l. nevadensis, suggesting a putative partial loss of function. Another substitution (S172C) was associated with the presence of black scales in both L. l. lepida and L. l. iberica, but no mutations were associated with the higher melanization of L. l. iberica. Analysis of genetic structure showed that L. l. nevadensis divergence is well supported by both neutral and non-neutral loci, confirming that the subspecies is at the final stages of its speciation process. L. l. iberica divergence from the nominal subspecies is mostly explained by a few adaptive loci, indicating that L. l. iberica might be at the early stages of ecological speciation.

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A comprehensive knowledge on the genetic basis of coloration is crucial to understand how new colour phenotypes arise and how they contribute to the emergence of new species. Variation in melanocortin-1 receptor (Mc1r), a gene that has been reported as a target for repeated evolution in a wide range of vertebrate taxa, was assessed in European ocellated lizards (Lacerta lepida) to search for associations with melanin-based colour phenotypes. Lacerta lepida subspecies’ distribution is associated with the three major bio-climatic regions in the Iberian Peninsula. A nonconserved and derived substitution (T162I) was associated with the L. l. nevadensis phenotype (prevalence of brown scales). Another substitution (S172C) was associated with the presence of black scales in both L. l. lepida and L. l. iberica, but no mutations were found to be associated with the higher proportion of black in L. l. iberica. Extensive genotyping of Mc1r along the contact zone between L. l. nevadensis and L. l. lepida revealed low gene flow (only two hybrids detected). The implications of these findings are discussed in the context of previous knowledge about the evolutionary history of ocellated lizards.

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This report describes the discovery and detailed description of a new small sized morph of ocellated lizard from the northern slopes of the Sierra de Gredos (Castilla y Leon, Spain). Morphological and molecular analysis showed that this new morph is different from the common species of ocellated lizard (Timon lepidus) on the Iberian peninsula in body size, sexual size dimorphism and cytochrome b haplotype. DNA sequencing revealed a unique and highly divergent cytochrome b haplotype indicating a long-term separation from other species of ocellated lizards. Evolutionary mechanisms that may have contributed to the formation of the small sized morph are discussed. The results presented in this paper suggest that this new morph is likely to be recognized as a full species in the near future.

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The Pyrenees, Sierra Nevada, Sierra de Peña de Francia and the Cantabrian Mountains were visited during the second half of May, 1994. In northern Spain Podarcis hispanica and P. muralis are living in syntopy. On all sites Lacerta lepida was common. North of Murcia the only typical L. lepida nevadensis have been seen. In the Sierra Nevada this subspecies had a large amount of green colouration. South of Madrid Acanthodactylus erythrurus and Psammodromus algirus were found in nearly every habitat. In the vicinity of Béjar Podarcis bocagei and Lacerta schreiberi were seen. In the Sierra Covadonga a large population of L. schreiberi was found in a very dry mountain slope.

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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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During two journeys from 27.04. – 07.05.2008 and from 21.04. – 27.04.2012, I visited several parts pf Spain together with some friends of mine. We focused on the lizards of the Iberiasn Peninsula. Specifically, we wanted to make some beautiful pictures of the Ocellated Lizatrds Timon lepidus and Timon nevadensis, which unfortunately did not always succeed on the first trip. In the following article, I briefly present the lizards of the family Lacertidae that were found during the two journeys together with some of their localities.