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A complete catalogue is provided for the type specimens of lacertid lizards in the herpetological collection of the Zoological Museum, Berlin, as of December, 1993. The collection contains a total of 514 type specimens, representing 63 taxa, of which 42 are currently regarded as valid at the specific or subspecific level. Types representing an additional four taxa appear to have been lost from the collection. The collections are especially rich in African lacertid types and in historically significant specimens collected early in the Nineteenth Century by Pallas, Eversmann, and Hemprich & Ehrenberg. Also present are the type series of many subspecies of Podarcis lilfordi and P. pityusensis described by Eisentraut.

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In this article an introduction is given on the geological history leading to the separation of Podarcis lilfordi (GÜNTHER, 1874) and Podarcis pityusensis (BOSCÁ, 1883) as separate species, as well as a Holocene sea level rise model which combined with bathymetric data leads to an estimation of recent divergence time in populations of the Balearic lizards.

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New data on estimated divergence times of the populations of lacertid lizards in the Balearic Islands are provided in this second update of the October 2011 article: Estimating recent divergence time in populations of Podarcis lilfordi (GÜNTHER, 1874) and Podarcis pityusensis (BOSCÁ, 1883) (VAN DEN BERG 2011), which received its first update May 2012. In most cases better estimations of divergence times were available by using the NAVIONICS SonarCharts™webapp.

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We report on our observations on Podarcis lilfordi (GÜNTHER, 1874) and other Menorcan herpetofauna during our 12 day trip to Menorca (Balearics, Spain). 13 populations of Podarcis lilfordi were surveyed, 101 specimens were measured and photographed in detail, from which 84 specimens were “buccal-swabbed” for DNA samples. The subspecific status of Hemidactylus turcicus spinalis from Addaia Gran is discussed and rejected.

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Phylogenetic relationships and timings of major cladogenesis events are investigated in the Balearic Island lizards Podarcis lilfordi and Podarcis pityusensis using 2675 bp of mitochondrial and nuclear DNA sequences. Partitioned Bayesian and Maximum Parsimony analyses provided a well-resolved phylogeny with high node-support values. Bayesian MCMC estimation of node dates was investigated by comparing means of posterior distributions from different subsets of the sequence against the most robust analysis which used multiple partitions and allowed for rate heterogeneity among branches under a rate-drift model. Evolutionary rates were systematically underestimated and thus divergence times overestimated when sequences containing lower numbers of variable sites were used (based on ingroup node constraints). The following analyses allowed the best recovery of node times under the constant-rate (i.e., perfect clock) model: (i) all cytochrome b sequence (partitioned by codon position), (ii) cytochrome b (codon position 3 alone), (iii) NADH dehydrogenase (subunits 1 and 2; partitioned by codon position), (iv) cytochrome b and NADH dehydrogenase sequence together (six gene–codon partitions), (v) all unpartitioned sequence, (vi) a full multipartition analysis (nine partitions). Of these, only (iv) and (vi) performed well under the rate-drift model. These findings have significant implications for dating of recent divergence times in other taxa. The earliest P. lilfordi cladogenesis event (divergence of Menorcan populations), occurred before the end of the Pliocene, some 2.6 Ma. Subsequent events led to a West Mallorcan lineage (2.0 Ma ago), followed 1.2 Ma ago by divergence of populations from the southern part of the Cabrera archipelago from a widely-distributed group from north Cabrera, northern and southern Mallorcan islets. Divergence within P. pityusensis is more recent with the main Ibiza and Formentera clades sharing a common ancestor at about 1.0 Ma ago. Climatic and sea level changes are likely to have initiated cladogenesis, with lineages making secondary contact during periodic landbridge formation. This oscillating cross-archipelago pattern in which ancient divergence is followed by repeated contact resembles that seen between East-West refugia populations from mainland Europe.

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Actively foraging lizards are capable of identifying prey using only chemical cues sampled by tongue-flicking, and the relatively few omnivorous and herbivorous lizards tested similarly can identify both animal and plant foods from chemical cues. Whether lizards that eat plants respond to cues specific to preferred plant types and whether there is geographic variability in responses to cues from various plants correlated with the importance of those plants in local diets is unknown. In three populations of an omnivorous lacertid, the Balearic lizard Podarcis lilfordi, we studied chemosensory sampling and feeding responses to chemical cues from plant and animal foods presented on cotton swabs. Each lizard population is endemic to one islet off the coast of Menorca, Balearic Islands, Spain. Lizards in all three populations discriminated chemical cues from plant and animal foods from control substances. Our results extend findings of prey chemical discrimination and plant chemical discrimination in omnivores, increasing confidence that correlated evolution has occurred between plant diet and chemosensory response to palatable plants. There were no consistent differences among populations in tongue-flicking and biting responses to stimuli from flowers of syntopic and allopatric plant species. The lizards may respond to cues indicative of palatability in a wide range of plant species rather than exhibiting strong responses only to locally available plant species. Nevertheless, tongue-flicking and biting frequencies varied among plant species, perhaps indicating food preferences. In addition, there were differences among populations in tongue-flick rates, latency to bite, and licking behavior. Licking was observed in only one lizard population as a response to floral chemicals from only one of the plants species tested, raising the possibility of a population-specific linkage between identification of a particular plant species and performance of an appropriate feeding response.

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The aim of this study is to assess the health status of different populations of P. lilfordi, an endemic lizard of the Balearic Islands. The health status and potential environmental stress to which these populations are subject estimaatendiendo four independent indicators, putting both to test its reliability and searching for the possible relationship between them. These four indicators are: the so-called state of condition of individuals, the parasite load, immune response and the degree of fluctuating asymmetry observed in bilateral characters. Also, relationships are established with adaptive responses in terms of features of behavioral ecology, including: strategies to secure food, defense capabilities and tactics of escape from predators.

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A survey about the helminthfauna of Podarcis lilfordi (Günther, 1874) (Sauria, Lacertidae) in the Gimnesic Islands (Balearic Islands, Spain) has been carried out. Eleven species of helminths were found, two Digenea, eight Nematoda and one Acanthocephala. Data of prevalence of each helminth species are indicated. The nematodes Skrjabinodon medinae and Spauligodon cabrerae show the highest values of prevalence and abundance. Host-parasite relationships are discussed, and the absence of cestodes is pointed out.

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Caudal autotomy is an effective antipredator strategy widespread among lizards. The shed tail thrashes vigorously for long periods to distract the predator and facilitate the lizard`s escape. This movement is maintained by energy supplied by the anaerobic conversion of glycogen into lactate. It has been suggested that lactate accumulation serves as an index for the vigor of tail thrashing. We made three predictions: (1) tail loss frequency should be higher under heavier predation regime, (2) the duration of postautotomy tail movement should be extended in populations under heavy predation pressure as an adaptation to the higher risk and the increased need for defense, and (3) as result, lactate in these tail tissues should be concentrated at higher levels. To eliminate the impact of phylogeny and environmental factors on the interpretation of our result, we focused exclusively on one species, the Balearic lizard (Podarcis lilfordi). We studied three populations, Colom (Illa d`en Colom), Aire (Illa de l`Aire), and Sargantana (Illa Sargantana), under different predation pressure but sharing the same climatic conditions. We found no differences among the studied populations either in postautotomy duration of tail movement or in levels of final lactate accumulation while autotomy frequency was higher where predation pressure was more intense. Tail loss effectiveness is directly influenced by the level of predation, while secondary features of the trait appear to remain independent from the impact of environment.

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Aims: To characterize the genetic and morphological diversification of the endan- gered Balearic lizard Podarcis lilfordi and to assess the relevance of this diversity, and how it is described, to conservation measures. Location: This study covers all the populations of the Balearic lizard, Podarcis lilfordi, present in its range of distribution at coastal islets of Menorca, Mallorca and Cabrera Archipelago. Methods: We analysed genetic and morphological variation across the 43 known extant populations of the Balearic lizard, using mitochondrial and nuclear markers. We examined morphometric and scalation characters using, in some cases, phyloge- netically independent contrasts. We also incorporated the study of dorsal coloration and dorsal colour pattern including the analysis of melanism in several populations. Results: We detected clear genetic divergence between Menorcan populations and populations from Mallorca and Cabrera, in both nuclear and mtDNA markers, but genetic divergence is relatively low among different insular populations within these groups. In contrast, morphological divergence was substantial both between Menorcan and remaining populations and within these groups. Morphological traits, such as dorsal coloration, body size and the number and size of scales, seemed to be linked with differences in climatic conditions between populations. In addition, some traits, as melanism, showed a strong phylogenetic signal. Main conclusions: The morphological and genetic diversity of the Balearic lizard is incongruent with the subspecies described in the classical taxonomic literature. Moreover, current populations differ not only in some genetic and morphological features, but also in several ecological and ethological characteristics, in many cases unique to one population. Based on our results, we propose abandoning the use of subspecies to describe the extraordinary morphological diversity of the Balearic liz- ard and its replacement with the concept of evolutionary significant units (ESUs). ESUs are particularly suitable to describe and recognize such diversity and, especially, to ensure the continuity of the evolutionary process.

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New information on the geographical location of the population of the Balearic lizard described as Podarcis lilfordi carbonerae Pérez-Mellado & Salvador, 1988 is given. The records of the herpetological collection of the Alexander Koenig Museum of Bonn, the capture dates of specimens and three recent surveys to coastal islets of Menorca (Balearic Islands) showed that the capture locality so-called Isla Carbonera (Na Corberana) corresponds to En Carbó islet (Illot d`en Carbó), off northern Minorcan coast. New colour, design and morphometric data of lizards from En Carbó confirm the validity of the subspecies P. lilfordi carbonerae. The present day situation and conservation status of the lizard population from En Carbó are discussed.

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We examined the diets of several species of Podarcis from different western Mediterranean islands. We tested the prediction of more generalized diets in lizards from poor environments. We also examined the extert and mode of herbivory in relation to arthropod diversity in the diet. Lizards from the Tuscan Archipelago, P. sicula from Menorca, and P. tiliguerta from Corsica all show a rather generalized diet. In contrast, P. lilfordi and P. pilyusensis from the Balearic Islands and, to a lesser extent, P. hispanica from Benidorm Island, appear much more stenophagous, including a high proportion of clumped prey in their diets. Plant matter is a common food item in most of the population studied, specially in Balearic lizards. A marked seasonal shift in the degree of herbivory was observed in these species where high-energy plant parts are frequently consumed during summer. In one of the Balearic Islands (Nitge), the bulk of the summer diet was formed by pollen and nectar of one particular plant species. Herbivory, consumption of clumped prey and other dietary peculiarities of insular populations of Mediterranean lizards indicate a very flexible feeding behaviour compared to mainland congenerics. This could result from a lack of predation pressure allowing high population densities to be reached in islets with poor trophic resources. Differences found in dietary characteristics among island populations can be interpreted as a consequence of different current ecological conditions, as well as historical and biogeographical differences among the islands.

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Tail autotomy is one of the main anti-predator mechanisms of lacertid lizards, but it has been predicted that it is only retained in its full capacity when its benefits exceed its costs (Arnold, 1988). To test this hypothesis, ease of tail shedding was examined in a number of continental and insular lacertid lizard populations, each of which showed a different shedding capacity. Tails are shed more easily in those continental and insular populations where there is a greater probability of predation. In insular populations not subjected to strong predation, the tail tends to be retained. The relationship of these findings to insular Mediterranean lizard populations and to the extinction of the Balearic lizard, Podarcis lilfordi are discussed.

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he Balearic lizard, Podarcis lilfordi, is present in 43 insular populations in the Cabrera archipelago and around the coasts of Mallorca and Menorca islands (Spain). We studied lizard densities over the entire range of distribution, analyzing observed differences of density in relation to island area, habitat diversity, availability of resources, presence of predators, competitors and human disturbances. The density of the Balearic lizard varies from less than 35 to almost 8000 lizards ha-1, with an average of around 1500 lizards ha-1. In some very small islets we detected no more than 10 individuals. Using a subsample of nine coastal islets (Menorca) we did not find any significant correlation between ground arthropod biomass and lizard density. The combination of island area and its maximal altitude, its so-called biotic capacity, was also uncorrelated with lizard density. In addition, neither degree of island accessibility nor presence/absence of seagull breeding colonies, were able to explain lizard densities. Islands without ship rats (Rattus rattus) showed a significantly higher lizard density, but islands in which rat eradication programs were launched during the study period, showed lower densities than those with rats but no eradication actions. Genetic variability was significantly higher on bigger lizard populations, lacking a correlation with lizard densities. No single independent variable can explain density differences among populations under study. Our results are discussed in the light of available hypotheses on factors affecting population densities.

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While the use of faecal pellets is widely accepted as a primary methodological source of data for dietary studies, a recent paper advocated for the use of gut contents. This was due to the fact that faecal samples would give biased results of the diet of arthropod predators, due to a lower representation of soft-bodied prey in faecal pellets. To test this assumption, we compared the spring diet of several populations of two insular lizards from the Balearic Islands (Spain), Podarcis lilfordi and Podarcis pityusensis, using both faecal pellets and gut contents. Our results do not support the supposed bias of dietary analyses based on faecal pellets. Indeed, soft-bodied prey and particularly insect larvae are often equally represented in faecal pellets and gut contents. Alternatively, soft bodied prey are represented in different proportions in gut contents and faecal pellets, but in some cases with higher proportions being observed in the gut contents, and in other cases with higher proportions in faecal samples. We conclude that faecal pellets can be a reliable source of information for dietary studies.

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A studie was carried out on 17 populations of Podarcis lilfordi inhabiting the islets off the coast of Menorca (Balearic Islands). Pattern and colouring together with the statistical analysis performed suggested that the menorcan rassenkreis comprises a total of 9 subspecies. The evolutionary processes involved appear to be fundamentally governed by the age of the islets, their distance from the coast and by their ecological conditions. Though in certain cases it may be postulated that genetic drift may have acted as a differentiating mechanism in populations which were initially small. However, interpretation of the microevolutionary processin Menorca is more difficult thnin the case of other rassenkreis of the Balearic Islands due to extinction of the `mother` population on the main island.

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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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A 684 bp fragment from cytochrome-b allowed a coherent arrangement of several Podarcis populations from the Balearic archipelago, in a study focused on the phylogeography of the 15 Minorcan populations of Podarcis lilfordi. A reference site with Podarcis lilfordi from Mallorca clusters with the Minorcan population, although it shares a few variations with Podarcis pityusensis that are not present in the Minorcan clade. Concerning the Minorcan islets, a two steps model is drawn accounting for the observed molecular pattern. An ancient and a derived stock of populations can be discerned, seperated by a population event replacing the former populations by the modernones on the shallowest protoislets, placing it around 7000 to 5000 BP, based on bathymetric information and Holocene sea level data. Autapomorphies are present in several sites and could be indicative of seperated demes prevailing on penninsular areas of the island, as in Cavalleria, Mola de Fornells, or even on larger regions as in the southeast Menorca. Within the framework of a vicariant model, we suggest that islet`s inhabitantshave thus retained some of the history of the now extinct mainland population.

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The Balearic lizard P. lilfordi inhabits the Gymnesies Islands and have sexual dimorphism. By means of biometric studies and analyses such as the discriminant analysis, it has been determined the morphometric characters that determine this sexual dimorphism. The characters chosen by the analysis were: body length, pileus width, number of gular scales, length of the four finger of the posterior leg. The differential measures of these characters makes the males to have a burlness and robustness greater than the females of the same population. Nevertheless, it cannot be said that in all P. lilfordi populations, the males will be aways the most burliness and robustness,but this will depend upon th population of P. lilfordi we take into consideration.

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Comparisons among eight populations of Podarcis lilfordi, seven of Podarcis pityusensis (representative lizards of the Balearic Archipielago), and one of Podarcis hispanica atrata fro the Columbretes Islands, have been made from a morphological point of view, measuring 15 morphometric characters of the lizards. Cluster and discriminant analyses were obtained for males and females to show the relationships among the different populations and species. These analyses gave three groups: one of them with the most P. lilfordi populations, other with the most P. pityusensis populations, and a well seperated group with the P. hispanica atrata species. Some populations connect the groups of P. lilfordi and P. pityusensis. The possibility that P. lilfordi and P. pityusensis be two pattern of variation inside the same species instead of two well differentiated species is considered, altrough further studies are necessary.

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The effect of several abiotic and biotic factors (island area, distance from a potential colonizing source, sex, age and diet of the host) on helminth diversity of the endemic balearic lizards Podarcis pityusensis and Podarcis lilfordi caught on small islets of the Balearic islands, has been studied.

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The knowledge of the helminth communities of reptiles and their ecological relationships with their hosts are until the present not well known. Some general researches have been made only on American herps (AHO 1990). Recently, ROCA & HORNERO (1991a, 1991b) attempted similar kind of researches from Mediter- ranean insularlizards.

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A survey of the helminth communities of Podarcis pityusensis (Bosca, 1883) and Podarcis lilfordi (Gunther, 1874) (Sauria: Lacertidae) from the Pityusic and Gimnesic islands, respectively, in the Balearic Islands, Spain, was conducted. Both host species harbour similar though not identical helminth infracommunities. An analysis of the patterns of diversity for all helminths revealed that both lizards have depauperate isolationist infracommunities, possibly as a result of factors such as host ectothermy, the simplicity of their digestive canal, low vagility, and generalist feeding.

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Aim To describe and analyse phylogeographical patterns in the endangered endemic lizard Podarcis lilfordi from across its remaining range and thereby establish baseline information on genetic diversity that will help determine conservation priorities and assist future reintroduction programs. Location Balearic Islands, Spain. Methods We analysed mitochondrial DNA (2382 bp sequence from eight genes) from 118 individuals and characterized the relationships among haplotypes using parsimony networks, as well as phylogenetic inference. Analyses of historical gene flow and population growth were used to provide further insights into population histories. Results Four unconnected parsimony networks were obtained that mirrored the main clades in the phylogenetic tree: (I) all Menorcan populations, (II) Dragonera, Malgrats and Toro islands (Western Mallorca) (III and IV) and the remaining populations from Cabrera and Mallorca. Two major haplotype groups were detected in Menorca (I) and these provided signatures of a demographic expansion and asymmetrical historical gene flow, respectively, concordant with the expected direction of colonization from south to north of the island. Populations from western Mallorca (II) showed evidence of historical allopatric fragmentation events following isolation around the start of the Pleistocene. In networks III and IV, Cabreran populations appear to have become isolated from north and south Mallorca quite recently, with asymmetric gene flow indicating a northwards dispersal direction. Main conclusions P. lilfordi is a genetically diverse species that shows substantial mtDNA structuring both between regions and, at a finer scale, between some islet populations within regions. The precarious state of some islet populations shown here to be quite divergent (e.g. Toro island in western Mallorca) means that conservation of this intraspecific biodiversity requires urgent action.

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Presentation held at the annual meeting of the AG Lacertiden in Gersfeld/Rhön the 10th of April 2011. It is reported about the rediscovery of a population of the Balearic Lizard (Podarcis lilfordi) on the Illa de Ses Mones in the harbour of Port d´Addaia. This population has been considered to have become extinct during the 1990s due to the introduction of the Italian Wall Lizard (Podarcis siculus). Descriptions of the island and the lizards are given and for the first time it is reported about the sympatric occurrence of Podarcis lilfordi and Podarcis siculus. Because in quite a lot of publications the nearby Illot d´en Carbó has been confused with the Illa de Ses Mones the taxonomic status of the rediscovered Podarcis lilfordi population from the Illa de Ses Mones and the Podarcis lilfordi carbonerae population of Illot d`en Carbó are discussed.