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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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We present the results of several field campaigns, realized from June 2007 to September 2008, and from March 2012 to May 2014 in different Algerian localities (Cap Djinet, Tigzirt, Boukhalfa, Ouadias, El Misser, Darna and Talaguilef). During this field work we have collected 488 specimens belonging to 35 species: 328 small mammals (11 species), 53 amphibians (6 species), 106 squamates (17 species) and one chelonian. The trapping success was highly variable (0.1 to 13.34 %). Genetic and morphometric analyses in progress will improve our knowledge of the evolutionary history of these small vertebrates at population and community scales.

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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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DNA sequence indicates the Lacertidae contain two subfamilies, Gallotiinae and Lacertinae, the latter comprising two monophyletic tribes, the Eremiadini of Africa and arid southwest and central Asia, and the Lacertini of Europe, northwest Africa and southwest and east Asia. Relationships within the 108 species of Lacertini are explored using mtDNA (291 bp cytochrome b; 329 bp 12S rRNA for 59 nominal species, and reanalysis of the data of Harris et al. 1998, and Fu 2000). The morphology of the tribe is reviewed and 64 of its characters (equivalent to 83 binary ones) also used to assess relationships. The Lacertini are assigned to 19 monophyletic units of 1 to 27 species, recognised here as the following genera (contents are indicated in brackets): Algyroides, Anatololacerta gen. nov. (L. danfordi group), Apathya (L. cappadocica group), Archaeolacerta (L. bedriagae), Dalmatolacerta gen. nov. (L. oxycephala), Darevskia (L. saxicola group), Dinarolacerta gen. nov. (L. mosorensis), Hellenolacerta gen. nov. (L. graeca), Iberolacerta (L. monticola group), Iranolacerta gen. nov. (L. brandtii and L. zagrosica), Lacerta s. str. (sand and green lizards, L. agilis group), Parvilacerta gen. nov. (L. parva and L. fraasii), Phoenicolacerta gen. nov. (L. laevis group), Podarcis (wall lizards), Scelarcis (L. perspicillata), Takydromus (Asian grass lizards), Teira (L. dugesii), Timon (ocellated lizards, L. lepida group) and Zootoca (L. vivipara). Both mtDNA and morphology indicate that Lacerta and Timon are sister taxa, and DNA suggests further possible relationships among genera (Fig. 1, p. 6). Neither DNA nor morphology indicates that the archaeolacertas (sometimes formalised as Archaeolacerta sens. lat.) form a clade. Instead, they are representatives of an ecomorph associated with living on rock exposures and using the narrow crevices that these contain. The Lacertidae probably arose in the European area, with the Gallotiinae later reaching Northwest Africa and the Canary Islands, and the ancestor of the Eremiadini invading Africa in the mid-Miocene. The Lacertini spread through much of their present European range and diversified, perhaps largely by repeated vicariance, around 12–16 My ago, producing the ancestors of the present mainly small-bodied genera, which then underwent often modest speciation. Three units spread more widely: the Lacerta-Timon clade of large-bodied lizards probably dispersed earliest, followed by Algyroides and then Podarcis. Overall, European Lacertidae show a pattern of repeated spread, often accompanied by restriction of previous groups. Expansion of Lacertini may have displaced earlier lacertid lineages from all or much of Europe; while spread of Podarcis may have restricted many other genera of Lacertini. The earlier expansion of the Lacerta-Timon clade probably did not have this effect, as difference in adult body size restricted competitive interaction with other forms. Several invasions of more distant areas also occurred: of East Asia by Takydromus over 10 My ago, and more recently of northwest Africa by Podarcis, Scelarcis and Timon, and Madeira by Teira. Relationships within the Eremiadini estimated from both mtDNA, and nDNA differ considerably from those based on morphology. They indicate relatively mesic forms may have diversified widely across Africa and given rise to at least three independent invasions of arid habitats. MtDNA also indicates that Lacerta andreanskyi belongs in the Eremiadini and may occupy a basal position there. It is assigned to a further new genus, Atlantolacerta gen. nov.

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Archaeolacerta s. l., in the currently utilized sense results to be a polyphyletic taxon, composed of three monophyletic and phylogenetically well differentiated taxa: a)-Archaeolacerta (s. str.) Mertens, 1921, which includes three species: A. bedriagae, A. oxycephala, and A. mosorensis, distributed by the northern part of the Central Mediterranean (Corsica, Sardinia, and west Balcanic Peninsula). His most related genera seem to be Teira (his adelphotaxon in our study), Apathya, and Omanosaura. b)-Iberolacerta gen. nov., with two subgenera: Iberolacerta s. str. and Pyrenesaura subgen. nov. includes six species: I. monticola, I. cyreni, I. bonnali, I. aranica, I. aurelioi, and I. horvathi. They are distributed by the mountains of Western Europe (Iberian Peninsula, Pyrenees, Central and Eastern Alps and the north of the Dinaric Chains). His adelphotaxon are the Caucasian and Near East species of Darevskia gen. nov. c)-The third taxa is Darevskia gen. nov., which includes the species of the `L.` saxicola complex besides `L.` derjugini, `L.` praticola, and `L.` chlorogaster. They are distributed by Eastern Europe, Caucasus, and Near East. Other clear relationships among some taxa of the Eurasian Radiation [= `Eurasische linie` from (Mayer and Benyr, 1994)] have been found: Algyroides seems to be the sister group of the species considered of uncertain phylogenetic relationships belonging to the `L.` danfordi-laevis group and of Podarcis. `L.` brandtii seems to be a very primitive species within the Eurasian Radiation. Omanosaura and Apathya appear as sister taxa. `L.` graeca occupies a very basal position in the sister group of Podarcis and relatives (see above). `L.` andreanskyi is the sister species of the Darevskia nov. and Iberolacerta nov. clade. The assimilation of `L.` andreanskyi to Teira is very problematic. Also the `L.` parva group seems to be related to Timon. Some groups of species like `L.` brandtii, `L.` parva-fraasii and `L.` danfordi-laevis probably merit generic rank, but it is necessary a more deep study before to take a decision.

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Algeria is a case study of biodiversity ignorance. To date, there are neither complete inventories for any taxa nor atlas across the country! Yet, Algeria is of great importance for global biodiversity. The coastal area of Algeria is part of the global biodiversity hostpot which is the Mediterranean perimeter and has several regional red spots of biodiversity. Moreover, the Saharan part encompasses a diversity of endemic organisms adapted to the strong conditions of drought. The Algerian authorities are deploying an ambitious project for conservation aiming to reach 50% of the country`s surface as protected areas! However, the location of these protected areas and their management don’t obey to a scientific evidence based, but are rather based on experts opinion. The main aim of this thesis is the study of the biodiversity ignorance distribution in Algeria by focusing on the `Linnean, Wallacean, and Darwinian Shortfalls` and to contribute to reduce them. The first chapter aims at reducing the Linnean Shortfall by proposing 1) a first checklist of reptiles and amphibians of Algeria which is the result of an accurate review of scientific publications on these taxa in North Africa. 2) a first pre-assessment of the conservation status of reptiles and amphibians of Algeria for the production of the first national red list. The second part of this manuscript deals with the ignorance of the geographical distribution of species. The purpose of this part is to map the ignorance which is defined as the difference between the expected specific richness diversity and the observed specific richness. The predicted species richness was modelled with two approaches: 1 ecological niche modelling with the maximum entropy method (MaxEnt) to predict the suitable habitats for each species and then add the different binary layers of species presence to calculate the specific richness. 2) Minimum convex polygon method was used to create range maps of each species and were auditioned to obtain predicted species richness. Both approaches have shown the same trend at a national scale: the concentration of the richest areas along the Saharan Atlas and the High Plateaus which are the transition zone between the Sahara and the Mediterranean part of Algeria. The opposition of the Saharan with a low species richness and the northern part with a higher species richness. Finally, the Saharan massifs of the Hoggar and Tassili form a very distinct zone with a much greater wealth compared to the rest of the Sahara. In terms of gaps, in the northern part, Kabylie, El Kala National Park and the Oran region are fairly well surveyed. In the Sahara, only a few points are clearly well prospected as Biskra, Bechar and some areas of the Hoggar and Tassili. The third part deals with the distribution of genetic llineages in the Maghreb. The aim of this section is to locate the phylogeographic breaks between the divergent eastern and western populations of the Maghreb and to try to understand the mechanisms that led to this pattern of genetic diversity. To answer these questions, I carried out a comparative phylogeography on 11 species of reptiles and amphibians with continuous and wide distribution in the Maghreb. Two suture zones have been identified: one zone on the border of Algeria and Morocco, the other in Kabylia west of the Soummam valley. The divergences between the eastern clades and the western clades occurred between the pre-Messinian periods up to the pliopleistocene boundary and were maintained in climatic refugia in the East and West of the Maghreb.

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A series of phylogeographic studies in the Maghreb identified a repeated pattern of deep genetic divergence between an eastern (Tunisia) and western (Morocco) lineage for several taxa but lack of sampling in Algeria made it difficult to know if the range limits between the eastern and western lineages were shared among taxa or not. To address this question, we designed a comparative phylogeographic study using 8 reptile and 3 amphibian species with wide distribution in the Maghreb as models.We selected species where previous studies had identified an East-West phylogeographic divide and collected sampled in Algeria to 1) examine whether the simple East-West divergence pattern still holds after filling the sampling gap in Algeria or if more complex diversity patterns emerge; 2) if the E-W pattern still holds, test whether the limits between the E and W clades are shared between species, suggesting that common historical process caused the E-W divergences; 3) if E-W limits are shared between species, use information on the age of the divergence to identify possible geological or climatic events that could have triggered these E-W differentiations. We found that the E-W pattern was generally maintained after additional sampling in Algeria and identified two common disjunction areas, one around the Algeria-Morocco border, the other one in Kabylia (central Algeria), suggesting that common historical mechanisms caused the E-W divergences in the Maghreb. Our estimates for the times to most common recent ancestors to the E and W clades span a wide range between the Messinian salinity crisis and the Plio-Pleistocene limit (except for one older split), suggesting different origins for the initial divergences and subsequent preservation of the E and W lineages in common climatic refugia in the west and the east of the Maghreb.

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This study aims at providing the first faunistic and ecological data on the herpetofauna of the Oum El Bouaghi region. The observation period was spread over two seasons (2018-2019), from mid-March to the end of October. A total of 18 species of reptiles and 4 species of amphibians have been recorded. The data collected were analyzed in terms of the biogeographical and trophic aspects of the herpetofauna of the study area. The region of Oum El Bouaghi has a great herpetofaunistic diversity, which shows how important it is to ensure its protection.

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Ocellated lizards from Tunisia and Spain respectively were hybridized. The hybrids proved to be much more aggressive than normal specimens but did not show any sexual activity among cach other Back crossing experiments revealed a strongly reduced fertility of the hybrids and demonstrated that the degree of genetic incompatibility by far exceeds the subspecific, i. c. infraspecific Ievel. The two forms are, thus, ranked here as allospecies: Lacerta pater LATASTE, 1880, and Lacerta lepida DAUDIN, 1802.

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New informations concerning systematical questions within the family Lacertidae, which were published during the last year, are presented.

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Une prospection extensive a permis de préciser les aires de répartition et les habitats en Tunisie de L. hispanica et de L. lepida. Un examen des caractères d`écaillure et de coloration les plus utilisés en taxonomie a été réalisé sur un échantillon de 17 L. hispanica en raison des imperfections de la systématique de cette espèce.

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Cette note est destinée à faire le point sur le peuplement herpétologique du Maroc. Elle comprend: — Une liste des formes effectivement présentes au Maroc, avec les affinités biogéographiques des espèces, et éventuellement les travaux récents où ces formes sont étudiées. — Des commentaires sur les espèces africaines douteuses en Europe et européennes douteuses en Afrique. — Une liste des espèces à rechercher au Maroc. — Une discussion sommaire sur la systématique de certaines espèces marocaines.

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Study of the system Tick-Lizard in the national park of El Kala (North-East Algeria). -- Among the emerging diseases rickettsioses would hold a prominent place in Algeria. The news was a reminder that some diseases can easily cross the species barrier and affect humans especially those related to wildlife. The pathogenicity of ticks `Ixodes ricinus is located on several levels. Ixodes ricinus in Algeria remains among the most infected ticks in North Africa, however, which represent the highest rate of infection observed in southern Europe. Many species of ticks parasitize reptiles especially lizards. Those when they are parasitized by ticks (Ixodes) are regarded as competent reservoirs of Borrelia and in a few exceptions a reservoir incompetent. For most of Ixodes (immature) Ixodes ricinus, Ixodes scapularis, ticks feed on the blood of lizards. This has served to emphasize the importance of Ixodes in epidemiology as primary vector of Lyme disease and the important role that play the lizard in the maintenance and movement of Borreliae. In this context, we are interested to ectoparasites lizards and role in the epidemiological cycle of the Rickettsioses. Our results are quite important because after PCR ticks we could detect the presence of Rickettsia sp. in 35% of all ticks collected. Given the number of rickettsioses transmitted by ticks (Rickettsia conori, `spotty mediterranean fever”, Coxiella burneti `Q fever `, R. africae `The african tick fever `), our results could not be ignored for since this is the first time that rickettsiae were detected in ticks from lizards.

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A phylogenetic-comparative approach was used to assess and refine existing secondary structure models for a frequently studied region of the mitochondrial encoded large subunit (16S) rRNA in two large lizard lineages within the Scincomorpha, namely the Scincidae and the Lacertidae. Potential pairings and mutual information were analyzed to identify site interactions present within each lineage and provide consensus secondary structures. Many of the interactions proposed by previous models were supported, but several refinements were possible. The consensus structures allowed a detailed analysis of rRNA sequence evolution. Phylogenetic trees were inferred from Bayesian analyses of all sites, and the topologies used for maximum likelihood estimation of sequence evolution parameters. Assigning gamma-distributed relative rate categories to all interacting sites that were homologous between lineages revealed substantial differences between helices. In both lineages, sites within helix G2 were mostly conserved, while those within helix E18 evolved rapidly. Clear evidence of substantial site-specific rate variation (covarion-like evolution) was also detected, although this was not strongly associated with specific helices. This study, in conjunction with comparable findings on different, higher-level taxa, supports the ubiquitous nature of site-specific rate variation in this gene and justifies the incorporation of covarion models in phylogenetic inference.

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Morphological and biochemical divergence between Spanish and Moroccan populations of Lacerta lepida demonstrate that these populations are not conspecific; Iberian populations are referred to Lacerta lepida, Moroccan populations to L. pater. An evolutionary scenario, consistent with the biochemical and morphological data, is inferred from the physiographic history of the region.

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The study that we present has two objectives: to expand our knowledge of the herpetofauna in the region of Tiaret and bordering areas, with a primary focus on distribution and update the threats that will affect this group, to propose solutions to protect and conserve the species of these areas. In spring 2014, we identified in this study, 6 species of amphibians and 22 species of reptiles. In some cases, we collected biological samples for later analysis. The majority of locations where prospected between May 22 and June 7, has a high biodiversity of herpetofauna, which shows how important it is, to preserve this species.

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An analysis of geographic distribution areas of the 41 taxa of Reptiles sampled or observed in the Aures massif is provided at two levels. At the overall level, eight groups of biogeographic affinities have been set apart; the number of mediterranean species (49 %) about equals that of saharian ones (46 %). 95 % of the total number of Reptiles species are of palearctic origin. At the regional level, we have distinguished three distribution types: (1) insular areas, isolated from populations of the neighbouring regions, with a continuous or a fragmented pattern in the Aurcs massif ; (2) parapatric areas with juxtaposed or overlapped ecological habitat ranges; (3) sympatric areas of allotopic or syntopic species. Hypothesis about origin and settlement of the Aures herpetofauna are infered.

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Studies on the herpetofauna of Tunisia’s numerous islands and islets began in 2000 thanks to targeted research programs. The data collected were compared and merged with others reported in the literature, thus obtaining an updated and detailed checklist of the Amphibians and Reptiles present on the Tunisian islands.

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La présente étude vient apporter les premières données faunistiques et écologiques sur les amphibiens et reptiles en Kabylie de la Soummam, région dont, à notre connaissance, aucune étude n’a été consacrée à ce sujet auparavant. Le travail de terrain s’est étalé du début avril à fin juin 2006, en suivant un protocole d’échantillonnage par stations au niveau des principales zones humides de la région (cours d’eau permanents et mares temporaires) ainsi que de quelques milieux terrestres. Un total de 6 espèces d’amphibiens et 17 espèces de reptiles a été identifié dans la région. Les données recueillies sur le peuplement herpétologique et le milieu ont fait l’objet d’une discussion autour des aspects faunistiques, biogéographiques et écologiques de l’herpétofaune de la zone étudiée et dégager quelques pistes de recherche à développer dans l’avenir.

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Reptiles are a successful group in subtropical arid regions. However, little is known about the drivers that influence the diversity of arid reptile communities. One approach is to investigate the variation in phylogenetic structure of the communities along a broad environmental gradient. In this study, I investigated these patterns in the communities of lacertids at the periphery of the Sahara Desert in Tunisia. The effect of the environment was assessed based on a set of variables describing the climate, topography, substrate and cover of perennial vegetation. The phylogenetic alpha diversity was described using the mean pairwise distance (MPD) and the mean nearest taxon distance (NTD) metrics. The analyses showed that the lacertid species could be clustered into three main ecological groups: mesic (Podarcis vaucheri, Psammodromus algirus, Psammodromus blanci, Timon pater), xeric (Acanthodactylus blanci, Acanthodactylus boskianus, Acanthodactylus maculatus, Mesalina olivieri, Ophisops occidentalis), and hyperxeric (Acanthodactylus dumerilii, Acanthodactylus longipes, Acanthodactylus scutellatus, Mesalina guttulata). The analyses also indicated that environment had a weak influence on species richness (15.7% of explained variance), but a strong effect on the phylogenetic (76.5% NTD, 89.5% MPD) structure. The lacertid communities tended to be phylogenetically clustered on sandy substrates under arid climate conditions, and overdispersed under more humid climatic conditions.

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In Tunesia during the second half of the month October six species of amphibians and reptiles were seen: Rana ridibunda perezi (Sousse), Bufo v. viridis (Sousse, Enfida), Tarentola mauritanica (Sousse), Psammodromus algirus (Sousse, El Djem, Hammamet). Arab names for amphibian (Hammamet), Acanthodactylus sp. (Hammamet) and Lacerta lepida pater (El Fahs), and reptile species, given by boys from Enfida, differ on many points from the names given by Doumergue (1901) for the region of Oran.

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Climatic conditions changing over time and space shape the evolution of organisms at multiple levels, including temperate lizards in the family Lacertidae. Here we reconstruct a dated phylogenetic tree of 262 lacertid species based on a supermatrix relying on novel phylogenomic datasets and fossil calibrations. Diversification of lacertids was accompanied by an increasing disparity among occupied bioclimatic niches, especially in the last 10 Ma, during a period of progressive global cooling. Temperate species also underwent a genome- wide slowdown in molecular substitution rates compared to tropical and desert-adapted lacertids. Evaporative water loss and preferred temperature are correlated with bioclimatic parameters, indicating physiological adaptations to climate. Tropical, but also some popu- lations of cool-adapted species experience maximum temperatures close to their preferred temperatures. We hypothesize these species-specific physiological preferences may con- stitute a handicap to prevail under rapid global warming, and contribute to explaining local lizard extinctions in cool and humid climates.

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Extensive fight wounds are common in co-habiting reptiles. This case study describes the management of a large fight wound in a 1-year-old entire female Tunisian eyed lizard. Primary closure was initially attempted but subsequent postoperative infection and wound breakdown led to successful management by secondary intention healing. This case demonstrates the amazing capacity for healing of large integument defects in lizards that receive appropriate medical support.

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Plusieurs séjours au Maroc entre 1974 et 1978 effectués soit dans le cadre de la RCP 249 du C.N.R.S. soit à l`occasion de voyages touristiques ont permis de rassembler des informations complémentaires sur l`herpétofaune marocaine. Captures, observations de spécimens écrasés sur les routes, mues et déterminations visuelles ont permis d`établir le texte qui suit. Il nous a paru intéressant de rassembler les observations ainsi obtenues avant leur Intégration dans un atlas des Reptiles du Maroc qui devrait voir le jour sous peu. Le découpage régional qui a été retenu est celui qui a été proposé par J. BONS (1967). Les observations les plus originales sont marquées d`un astérisque (*) et certaines sont commentées dans la conclusion.

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DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.

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

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Several recent papers have reviewed the life and work of French herpetologist Louis Amédée Lantz. They have detailed the composition of his collections deposited in several museums. However, since then, several other important specimens from his collections deposited at the Natural History Museum (NHM, UK) have come to light and we here present all of them in detail. We discovered paralectotypes of Lacerta saxicola obscura Lantz & Cyrén (BMNH 1918.11.21.5–7), syntypes of Lacerta boemica Sukhov (BMNH 1960.1.4.26–30, BMNH 1965.337–342) and Lacerta viridis media Lantz & Cyrén (BMNH 1960.1.4.25, 1966.512). We also identified numerous specimens from the French Mediterranean islands in Lantz’s collection deposited at the NHM, some of which represent the first reported specimens of their species from certain islands. We here provide data on all these specimens. We also place the Mediterranean island specimens from the Lantz collection in their historical context and emphasize the historical and taxonomic value of these collections.

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The F2 generation of Lacerra lepida parer was bred from parents under one year of age. The young thrived on a diet of small insects, some of wich were enrichened by feeding them one bone-meal.

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Five Lacerta /epida pater (2,3) from Mo- rocco were kept in a terrarium with an exceptionally large surface (50m2, later 30 m2). During summer direct sunlight was pro- vided. The animals came to reproduction, each female depositing about ten eggs three to four iimes per season. The eggs develo- ped well, but dit not hatch, unless the preg- nant females were fed on new-born rats instead of the hitherto offered menu of in- sekts, snails and occasionally small verte- brates. The incubation time of the eggs was 3 months at a temperature of 25 - 30°C.

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Aim To map and assess the richness patterns of reptiles (and included groups: amphisbaenians, crocodiles, lizards, snakes and turtles) in Africa, quantify the overlap in species richness of reptiles (and included groups) with the other terrestrial vertebrate classes, investigate the environmental correlates underlying these patterns, and evaluate the role of range size on richness patterns. Location Africa. Methods We assembled a data set of distributions of all African reptile species. We tested the spatial congruence of reptile richness with that of amphibians, birds and mammals. We further tested the relative importance of temperature, precipitation, elevation range and net primary productivity for species richness over two spatial scales (ecoregions and 1° grids). We arranged reptile and vertebrate groups into range-size quartiles in order to evaluate the role of range size in producing richness patterns. Results Reptile, amphibian, bird and mammal richness are largely congruent (r = 0.79–0.86) and respond similarly to environmental variables (mainly productivity and precipitation). Ecoregion size accounts for more variation in the richness of reptiles than in that of other groups. Lizard distributions are distinct with several areas of high species richness where other vertebrate groups (including snakes) are species-poor, especially in arid ecoregions. Habitat heterogeneity is the best predictor of narrow-ranging species, but remains relatively important in explaining lizard richness even for species with large range sizes. Main conclusions Reptile richness varies with similar environmental variables as the other vertebrates in Africa, reflecting the disproportionate influence of snakes on reptile richness, a result of their large ranges. Richness gradients of narrow-ranged vertebrates differ from those of widespread taxa, which may demonstrate different centres of endemism for reptile subclades in Africa. Lizard richness varies mostly with habitat heterogeneity independent of range size, which suggests that the difference in response of lizards is due to their ecological characteristics. These results, over two spatial scales and multiple range-size quartiles, allow us to reliably interpret the influence of environmental variables on patterns of reptile richness and congruency.

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In order to study the composition of the south Kabylie’s herpétofauna (Bouira’s and Bordj Bou Arreridj’s wilaya), we have chosen six different ecological areas. 19 species (1 Amphibian and 18 Reptiles) have been inventoried. They are mainly of mediterranean origin (12 species of 19) with 3 species typically North Africans (Timon pater, Chalcides chalcides and Mesalina olivieri). But in this herpétofauna we have found two saharian species (Psammophis schokari and Mesalina olivieri). The field analysis of the species distribution showed that the landscape composition played an important role. Some species such as P. algirus frequented various areas, others such as Natrix natrix and Natrix maura preferred humid areas, Acanthodactylus erythrurus localized in biotopes with sandy substratum, also Timon. pater who frequents forests, on the six community studied none presents q high diversity. The main important specific riches were observed in the low scrub (10 species). The intermediate situation that corresponds to the average of the diversity and the regularity was observed in the rocky open ground and the low scrub.

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Nous avons étudié la variabilité protéinique et morphologique de lézards ocellés appartenant à différentes populations européennes et africaines de ce complexe d`espèces de la Méditerranée occidentale. Les lézards ocellés d`Europe (Lacerta lepida ssp. lepida, iberica, nevadensis) sont très variables pour la forme du corps, la pholidose, la couleur et la dentition dans l`ensemble de leur aire de distribution étendue latitudinalement, mais ils sont relativement peu différenciés par l`électrophorèse. A l`inverse, les lézards ocellés africains ne sont pas variables, quant à leur pholidose, le long de leur aire de distribution - une étroite bande entre la mer Méditerranée et le Sahara, coupée en 2 par la très aride vallée de la Moulouya -, mais l`électrophorèse révèle des distances importantes entre les populations algéro-tunisienne (Lacerta pater) et celles du Maroc (Lacerta tangitana [combinaison nouvelle]). Ces différences sont du même ordre de grandeur que celles existant entre les populations marocaines et européennes, taxons spécifiques bien différenciés. Ce type de répartition, avec des espèces séparées en péninsule Ibérique, dans les montagnes marocaines, et dans le Tell algéro-tunisien, se retrouve dans d`autres complexes et suggère l`existence d`une barrière géographique importante entre l`est et l`ouest du Maghreb à l`époque de l`ouverture du détroit de Gibraltar.

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

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Der Verfasser besuchte Tunesien Anfang April 1978 und berichtet von den Arten, die er da vorfand. Chalcides ocellatus und Tarentola mauritanica wurden häufig beobachtet, besonders in Sousse. Ertaunliche Mengen von mMauremys caspica leprosa wurden in dem kleinen Fluss ´Oued Hamdoun`bei Sousse gesehen, sie wurden aber auch in Kairouan und Gabès anagetroffen. Natrix maura wurde in Sousse und Kairouan immer im oder beim Wasser gesehen, oft ist dieses Wasser hier brackig. Zum Schluss gibt der Verfasser einen Determinationsschlüssel zu den tunesischen Amphibien-, Eidechsen- und Schildkrötenarten mit wissenscjaftlichen, französischen, niederländischen und manchmal auch arabischen Namen.

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A male specimen of Lacerta lepida pater (Lata s t e) from Tizi `n Tichka (near Marrakesj, Morocco), kept in a terrarium, displayed a form of threatening behaviour worth mentioning. The lizard, for a short time placed in a small terrarium where it could neither flee nor hide, would first threaten the author upon his aproach with an opened mouth, and then by throwing small stones into his direction, by means of scratching movements with one of the forelegs (photo). Finally it would bite, just to run away afterwards.

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Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk. Location World-wide. Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions. Results I obtained body size data for 99% of the world`s lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non-aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed. Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.

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The authors present an actualized and commented list of the amphibia and reptilia species noted in Morocco. The changes witch happened during the 16years after the last checklist established by BoNS(1972) are discussed. These are new additions (species recently discovered in Morocco and new taxa for Science, sorne of witch resulting from systematic modifications of other species or subspecies) and changes of names (new combinations) or status. Problems concerning the exact delimitation of several species are discussed, insisting particularly on the most conflictual groups (e.g. Chalcides, Aoenthodactylus). Thus we think that the number of species in the present checklist is probably surestimated. Taking in account these problems, we must consider this inventory as provisional, till clearing up the several questions still in suspense.

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Un découpage hiérarchisé du territoire tunisien en quatre domaines zoogéographiques a été réalisé à partir de la composition spécifique en Lacertidés de 25 secteurs correspondant aux différentes régions naturelles du pays. La délimitation, les caractéristiques faunistiques, les affinités et l`origine biogéographique du peuplement en Lacertidés de ces quatre entités, sont analysées et discutées.

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Each of die 25 natural regions in Tunisia harbour from I to 9 Lacerlids among the 14 species surveyed in this country. The richest regions are the South-western pari of the Tunisian Central Range and the Dhahars mountains in Southern Tunisia. They are ranging between lower semi-arid and upper saharian. In the sandy, lower saharian Far-south, only one species occurs. Geographie variation of the specific diversity and structure of Lacertid communities arc related to bioelimatical affinities and zoogeographical native areas of the species. Main steps in the immigration or local differentiation of Lacertids are reported. Impacts of climatical and biological factors on Lacertid communities arc discussed.

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A new taxonomic checklist is established for the Amphibians and “Reptiles” of Tunisia. In addition to international zoological scientific names, a French scientific name is attributed to each taxon. The Tunisian herpetofauna presently contains seven species of Amphibians and 62 of “Reptiles”.

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The schism between North Africa and Southern Europe caused by the opening of the Strait of Gibraltar and the consequent refilling of the Mediterranean basin at the end of Messinian salinity crisis (MSC), 5.33 million years ago, has been advocated as the main event shaping biogeographical patterns in the western Mediterranean as exemplified by the distribution of species and subspecies and genetic variation within the ocellated lizard group. To reassess the role of the MSC, partial sequences of three mitochondrial DNA genes (cytochrome b, 12S and 16S ribosomal RNA) and two nuclear genes (β-fibrinogen and C-mos) from species of the ocellated lizard group were analysed. Three alternative hypotheses were tested: that divergence was initiated (i) by post-MSC vicariance as the basin filled, (ii) when separate populations established either side of the strait by pre-MSC overseas dispersal, and (iii) by post-MSC overseas dispersal. The pattern and level of divergence detected clearly refute the post-MSC vicariance hypothesis, and support a model of divergence initiated by earlier overseas dispersal. Indeed, our best estimate is that the basal Euro-African divergence predates the MSC event by several million years. The estimated divergence times among the populations in former Miocene Mediterranean islands, the current Betic and Rifian mountains, from adjacent mainland populations suggest overseas dispersal for the former and overland dispersal, or perhaps vicariance, for the latter. These results suggest that the MSC may have played a much less important role in shaping the current western Mediterranean biogeographical patterns than might have been anticipated from the dramatic nature of the episode.

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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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Many animals display complex colour patterns that comprise several adjacent, often contrasting colour patches. Combining patches of complementary colours increases the overall conspicuousness of the complex pattern, enhancing signal detection. Therefore, selection for conspicuousness may act not only on the design of single colour patches, but also on their combination. Contrasting long- and short-wavelength colour patches are located on the ventral and lateral surfaces of many lacertid lizards. As the combination of long- and short-wavelength-based colours generates local chromatic contrast, we hypothesized that selection may favour the co-occurrence of lateral and ventral contrasting patches, resulting in complex colour patterns that maximize the overall conspicuousness of the signal. To test this hypothesis we performed a comparative phylogenetic study using a categorical colour classification based on spectral data and descriptive information on lacertid coloration collected from the literature. Our results demonstrate that conspicuous ventral (long wavelength-based) and lateral (short wavelength-based) colour patches co-occur throughout the lacertid phylogeny more often than expected by chance, especially in the subfamily Lacertini. These results suggest that selection promotes the evolution of the complex pattern rather than the acquisition of a single conspicuous colour patch, possibly due to the increased conspicuousness caused by the combination of colours with contrasting spectral properties.

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The integration of spatial area prioritization algorithms and species distribution modelling has shown great promise in conservation planning in recent years. However, despite the fact that reptiles and amphibians have the highest threat status of all terrestrial vertebrates, these species are often under-represented in conservation planning. The Kingdom of Morocco possesses the richest and most varied herpetofauna in the Maghreb and the western Mediterranean, and is characterized by high species richness, endemism and number of European relict species. Despite the fact that Moroccan reptiles and amphibians have been the subject of numerous studies by a large number of international herpetologists since the beginning of the 20th century, few or none of these concerned their conservation. This study had three main objectives: (1) to identify those areas that harbour the highest species richness; (2) to evaluate the existing and proposed future ‘important biological and ecological sites’ (SIBES) conservation area network (CAN) with respect to their ability to protect the herpetofauna adequately; and (3) to identify priority areas into which the existing protected areas can be augmented. We used maximum-entropy species distribution modelling to run distribution models for 11 amphibian and 86 reptile species (27.6% endemics and 12.4% threatened) for which we had 2,170 single geographic records. A total of 97 models were used to create a richness map of the Moroccan her- petofauna and thereby detect both areas of high species richness and the distribution patterns of individual species. This map was subsequently used as a basis for performance evaluation of the CAN and area prioritization using the ConsNet conservation planning software initialized by ‘‘Rarity’’ first, while using representation targets of 5% and 10%. Additionally, the proposed future Moroccan CAN (SIBES) was evaluated in terms of its overlay and proximity with ConsNet solutions using visual interpretation and distance measurements in a GIS. Our results show that Moroccan herpetofauna is poorly protected under the existing and future CAN. Prioritization of areas shows that a major increase in conservation area is required to guarantee the persistence of individual herpetofauna species even with a global minimum representation target of only 10%. An increase of the existing CAN is especially needed along parts of the Atlantic coast, in the north-western Mediterranean region, on the north-eastern Moroccan coast, as well as in several areas in the Sahara, notably vast proportions of the Valle ́e du Haut and Bas Draˆa.

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The large-sized lizard Timon pater (Lacertidae) has been recently recognised as a distinct species from the European ocellated lizard (Timon lepidus), and is endemic to Morocco, Algeria and Tunisia. The ecology of this lizard is entirely unknown, and in this study detailed data on the diet composition and on the daily activity patterns, in spring and summer, are presented. The field study was carried out at the Parc National d`El Kala, northeastern Algeria. Food data were recovered by faecal pellet analysis, and daily activity patterns by standardised surveys along 1000-m-long line transects. In total, 164 T. pater pellets were collected and analysed, and 428 prey items were identified. The mean volume of the pellets collected in July and August was significantly less than that of the pellets collected in all the other months. The diet of Algerian T. pater was dominated by three prey categories (Coleoptera, Gastropoda, and Formicidae); Coleoptera and Formicidae were the most important prey types through the whole study period, whereas other prey types were important only in some periods (i.e. Gastropoda in August and Orthoptera in September). The diet composition did not change noticeably between spring and summer, and the same was true for the daily activity patterns. Overall, T. pater appeared remarkably similar to the European T. lepidus populations as far as dietary habits and daily activity patterns are concerned.

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Algeria is situated at the crossroads between Europe and Africa. The northern part of the country is listed as an area of high biodiversity. Currently, the ecosystems (rivers, lakes, deserts, forests etc.) and the species are under more pressure than ever. The impact of humans is significant and many factors constitute a strong threat to this fauna, especially reptiles, which are the most vulnerable because of their low mobility. Thus, pollution, the drying up of wetlands and their conversion to agriculture have clearly affected the existence of many species. The herpetofauna of Algeria is one of the most diversified in the Mediterranean Basin, consisting of 104 species of which 16.98% are endemic. We suppose that the present list of reptilian fauna provided in this paper is not exhaustive and it is expected to include more species given the lack of research on reptiles in Algeria and its large area. Our dataset contains information on reptile occurrences in Algeria. The dataset is based on original research by the staff of the Laboratory of Environmental Sciences and Agroecology of Chadli Bendjedid University in Algeria. The conservation status of all recorded species is given.

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Although Timon pater is among the largest species of lacertids found throughout North Africa, mainly in Morocco, Algeria and Tunisia, studies examining the ecology of this lizard remain rare. In this paper we describe the daily activity patterns, biometry and diet of T. pater in a mountain area during its period of maximum activity. We carried out the field study in the southern slope of Mount Chélia in the middle of the Aurès Massif, north-eastern Algeria. In order to analyse the daily activity patterns we performed surveys along 4 000 m long random transects. To obtain biometrical measurements we captured and measured 20 adults with a digital calliper. We obtained diet data by analysing 17 extracted stomach contents. The daily activity patterns of T. pater were similar during the summer periods (early and late summer), and no variation in activity was detected. Adult males were slightly larger and heavier than adult females and their heads were longer, larger and thicker than females’ heads. T. pater primarily consumed Coleopterans, but diet composition was noticeably different between the two summer periods. There were no sexual differences in dietary preferences.

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Observations on three species of lacertid lizards in the Ain Draham region in the Kroumir mountains of northern Tunisia are reported. Timon pater lives on big branches of cork oak trees and boulders. Podarcis (hispanica) vaucheri on tree trunks, walls and rock faces and Psammodromus algirus mainly on the ground in open situations (pathes, clearings) in the forest.

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Six localities within a radius of about 35 km around Ouarzazate (Central Morocco) were visited by the author five times between 1977 and 1989 to investigate their herpetofauna. 4 anuran and 20 reptile species were observed which raises the total number of herp species known from the area to 5 and 26 respectively. The threats to the local herpetofauna are discussed.

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Parasitism of Ixodes ricinus on three species of lizards (Psammodromus algirus, Podarcis vaucheri and Timon pater) in northeastern Algeria was studied. Psammodromus algirus was the most preferred host, T. pater was least preferred. Nymphs of I. ricinus were found attached to lizards from March to August in various numbers. Most nymphs were collected during March, April and June for P. algirus, and most larvae from April until August. Larval stages were found to be mostly associated with P. vaucheri during the study period, with the highest number of recovered larvae in June.

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