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Oman material of four lizard species is described and discussed ; it comprises specimens of a previously undescribed gecko, Phyllodactylus gallagheri, two lacertids — the poorly known Lacerta jayakari Boulenger, 1887, and L. cyanura, also described here for the first time, and a skink, Ablepharus pannonicus Lichtenstein, 1823. These species have no strong affinities with any members of the mainly Saharo-Sindian fauna now occupying much of Arabia but are closely related to forms occurring to the north of the peninsula. It is probable that they are relicts of a more temperate fauna which once entered more extensively into southeastern Arabia. As might be expected, all four species are absent from the desert areas of Oman and are confined to the disjunct mesic habitats which exist in the mountains.

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

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The ability to shed (autotomize) all or part of the tail, usually in response to predator attack, and often to subsequently regenerate it is widespread in lizards and amphisbaenians and also occurs in a few snakes and in the tuatara. Most species possess a sophisticated intravertebral autotomy mechanism which seems to be primitive in the Squamata. This appears to have been independently lost in members of many groups, but some agamids and snakes have regained the ability to shed their tails by a simpler intervertebral means and a -number of agamids have also redeveloped tail regeneration as well. Breakable tails are used to evade capture in two main ways: by enabling reptiles to break away from predators that have grasped them by the tail and by providing a distraction which deflects the attention of the attacker away from the vulnerable head and body. It is argued that loss of caudal autotomy has occurred when the costs of tail shedding outweigh its benefits. Likely costs include the expense of regrowing the tail and the loss of a variety of possible tail functions that may cause partial incapacitation, at least until the tail regenerates. Benefits of autotomy are liable to be low if predation is rare, if the animal is able to protect itself effectively in other ways, if it is too slow to evade further pursuit after the tail is shed, or if the tail is small or unpalatable and consequently not likely to distract a predator. Benefit variation may well be greater than cost variation and therefore more important in initiating the loss of autotomy mechanisms. Many taxa that do not shed the tail appear to conform to the above interpretation, but in some cases, such as the Platynota, Agamidae and Chamaeleonidae, lack of intravertebral autotomy may reflect the history of these groups rather than being a direct result of present ecological pressures. The distribution of intervertebral autotomy in the Agamidae suggests that it may have evolved only in rather special circumstances where tail fragility is advantageous even in the absence of the ability to regenerate. Restriction ot autotomy planes to the tail-base, so that the whole organ is lost, a condition found in a number of relatively slow-moving geckoes, is interpreted as a means of ensuring that enough of the tail is shed to distract a predator from further pursuit. The stimulus necessary to induce autotomy can vary rapidly in individual lizards and at least some of these changes probably maximize the effectiveness of the tail-shedding mechanism. Differences in the readiness with which all or part of the tail is shed exist between species and are likely to reflect the balance of costs and benefits in particular cases. Variations in incidence of broken tails between species and populations may be due to such differences in fragility but many other factors may play a part, including the age structure of samples, incidence of unsuccessful attacks by predators and ability to evade predators after autotomy. There is a clear tendency for climbing lizards, especially those living on rock surfaces, to have higher incidences of broken tails than ground-dwelling species, perhaps because the tail is usually less important in locomotion in the first group. Many lizards possess conspicuously coloured tails and tail movements that seem likely to help distract attention from the head and body. Conspicuous tail colouring is more frequent and often better developed in young animals, which tend to be more vulnerable than adults, and in active species from open habitats where crypsis may not always be very effective. Conspicuous tails usually have contrasting light and dark areas in nocturnal forms but are often a single bright colour in diurnal ones, probably reflecting the visual capacities of their respective predators. The predominance of blue tails in day-active species may be because this colour is striking close to but not very arresting at a distance, so it may not attract predators from far away while still drawing their attention at close quarters.

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A key and check list is provided to the 96 species and six additional subspecies of lizards and two species of amphisbaenians known to occur in and around Arabia. Philochortus neumanni, Lacerta jayakari, Lacerta cyanura, Latastia longicaudata andersonii, Mesalina guttulata, Mesalina olivieri, Mesalina martini, Mesalina brevirostris, Mesalina ayunensis, Mesalina adramitana, Acanthodactylus orientalis, Acanthodactylus robustus, Acanthodactylus opheodurus, Acanthodactylus boskianus, Acanthodactylus felicis, Acanthodactylus yemenicus, Acanthodactylus masirae, Acanthodactylus opheodurus, Acanthodactylus gongrorhynchatus, Acanthodactylus haasi, Acanthodactylus schmidti, Acanthodactylus blanfordii, Acanthodactylus arabicus, Acanthodactylus grandis, Acanthodactylus tilburyi, Acanthodactylus scutellatus hardyi.

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

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Arabia, including the arid lowland regions of Jordan, Syria, Iraq and southwest Iran, has approximately 128 species of terrestrial reptiles and about 7 of amphibians. Endemicity is high, 55% of species being largely or entirely confined to the area. Abrupt faunal discontinuities are present between Arabia and neighbouring regions, namely the Horn of Africa, the Sahara desert and the relatively mesic areas to the north. However, there must have been substantial interchange with these areas in the past. Within Arabia, the main Zoogeographie division is between the arid regions occupying most of the peninsula and much smaller, often more mesic areas of endemicity in the south. The former are mainly occupied by Saharo-Sindian taxa and have rather more affinities with the Sahara than to arid regions east of Arabia. Southern areas of endemicity include: 1. The southwestern coast, with affinities to the Horn of Africa and the inland deserts of Arabia. 2. The southwest and southern highlands which share a number of forms with northeast African relationships but differ significantly, the former having more, often endemic species with African affinities and a number with relatives in the Mediterranean and other northern mesic regions. The drier southern mountains have their own endemics, some of which have representatives in the öiddat al-HaräsIs area to the east. 3. The north Oman mountains also possess distinctive species and populations, some of which are related to forms in the less arid parts of the Iranian area. Finally there are a number of mesic forms distributed around the periphery of Arabia, and populations at oasis sites in the arid region which are probably relicts of forms more widespread in Quaternary pluvial phases.

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Estimates of phylogeny may allow historical events to be reconstructed even without a fossil record. The reliability of such interpretations depends not only on the robustness of the phylogeny but also on its topology. Changes in individual features can be traced and general histories of groups developed and compared with each other. Results are ofter surprising, for Instance the sophisticated tail shedding mechanism of lizards turns out to be a primitive feature that has been lost many times. Similarly, ecological analogues may have developed their common characaters in quite different orders. Phylogenies also provide a way of recognizing constraints and the effects of history on present ecological and behavioural patterns. When using anatomical characters the quality of the apparent phylogenies produced may be related to ecological history: expansion of a group along an ecological continuum into increasingly demanding niches ina small geographical area tends to produce a robust phylogeny (for instance in Meroles), while this is often not so for widely distributed groups that occupy a more modest range of niches (such as Pedioplanis and Podarcis). Non-morphological data may not show this tendency, but can have their own problems. Lacertids can be referred to clade with many successive branches in Africa and the Saharo-Eurasian arid zones and a less resolved probably paraphyletic complex of more primitive forms in the Mediterranean and wider Palaearctic areas. The African-Eurasian clade shows a general trend towards ground-dwelling and increasingly arid habitats but is ecologically variied. These animals are important to the study of Mediterranean forms because they provide eco-morphological parallels to them (for instance to Algyroides, Psammodromus and the archaeolacertas) and help form a basis for testing hypotheses about function. Although it is possible to recognise a number of distinct clades among the Mediterranean and Eurasian forms, relationships within and between these are often much less well substantiated. If results are available in time, current work on mitochondrial DNA sequencing will be discussed.

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Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.

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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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In this paper we deal on the ultraviolet color (invisible to us): where we can find it, the capability of animals to see it and the advantages that this color perception offers to them. As the simplest way to detect it is the photography, we describe and review how to photograph the UV, as a result of 15 years of amateur experience, searching and testing nearly in complete blindness due to the lack of practical information about “how to do it”. We describe the different kinds of photography (chemical and digital); the cameras and objectives suitable (both astronomically expensive ones and cheap options); what are the best characteristics that the objectives should have for this purpose; the films suitable for their use in chemical photography; the different filters (current or discontinued) manufactured along the years; and the subtle combinations among the different materials to obtain pure UV photographs. This kind of scientific photography is mainly used in forensics, forgery detection, art dermatology and less in Natural History, despite the fact that a great part of animals see this color and use it in important questions of their biology as the social behavior, mate choice or the food search.

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Animals communicate via a variety of sensory channels and signals. Studies on acoustic and visual communication systems suggest that differences in the physical environment contribute to the variety of signalling behaviour, with species investing in those signals that are transmitted best under the local conditions. Whether or not environmental tuning also occurs in chemical communication systems has received much less attention. In the present study, we examined the effect of several aspects of the physical environment on the chemical communication system of lacertid lizards (family Lacertidae). The numbers of femoral pores are used as a proxy reflecting how much a particular species invests in and relies upon chemical signalling. Femoral pores are specialized epidermal structures that function as a secretion channel for the waxy substance produced by glands. In some lacertid species, the secretion carries infochemicals that play an important role in social communication. The number of femoral pores varies considerably among species. We have compiled data on femoral pore numbers for 162 species and tested for the effects of climate and substrate use. After correcting for body size and taking the phylogenetic relationships among the species into account, we found no effect of climate conditions or latitude on species pore numbers. Substrate use did affect pore numbers: shrub-climbing species tended to have fewer femoral pores than species inhabiting other substrates.

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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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Diskussion über die Darstellung von Lacerta jayakari auf der Tafel II in BOULENGER (1887). Es konnte keine Erklärung für die düstere Färbung des Lebensbildes gefunden werden.

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In this paper, we report on our failure to breed a lasting population of Omanosaura cyanura in a vivarium environment. In our vivaria, O. cyanura produced 1 – 5 (maximally 8) clutches per year with 2.5 – 3.5 eggs per clutch. The eggs grow to approximately 10.8 x 18.9 mm and weigh, on average, 1.29 g. The eggs normally hatch at 25 °C after 72 days and at 29 °C after 56 days. In this study, of the externally healthy looking, only approximately one third hatched. The average dimensions of the eggs immediately following oviposition were 6.0 x 10.8 mm with a mean weight of 0.25 g. We surmise that a potential reason for the lack of hatching success is a vitamin D3 deficiency. Raising the successful hatchlings (SVL 23.9 mm; TL 46.8 mm; weight 0.25 g) turned out to be quite difficult because they were very finicky about eating and drinking, and moreover turned out to be easily frightened. We recommend keeping the individual lizards separately most of the time because of a high level of interspecific aggression.

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In the present study we use an unprecedented database of 5,535 distributional records to infer the diversity, ecological preferences and spatial distribution of the 60 species of terrestrial reptiles of the United Arab Emirates (UAE), and use the 57 native species to test the effectiveness of the protected areas’ network in conserving this unique vertebrate fauna. We infer a time-calibrated phylogeny with 146 species of squamates and 15 genes including all UAE terrestrial reptile species to determine the phylogenetic diversity (PD) and evolutionary distinctiveness (ED) of the native species and to compare it with the distribution of the hotspots of native species richness. The results of this study indicate that the sampling effort is remarkable, covering 75% of the country’s territory representing nearly the entire climatic space of the UAE defined by the mean annual temperature and the total annual precipitation, as well as the multivariate climatic space defined by a principal component analysis (PCA). Species richness is highest in the northeast of the country, in a transitional area from sandy desert to the mountainous terrain of the Hajar Mountains. The highest PD of a single square cell of 10 arc-minutes grid is of 2,430 million years (my) of accumulated evolutionary history and the strong correlation between PD and species richness suggests that the raw number of species is a good surrogate to quantify the evolutionary history (i.e., PD). The species with the highest values of ED are those in families represented by only one species in the UAE. Finally, the assessment of the UAE protected areas shows that, despite their relevance in protecting the terrestrial reptiles, they do not offer adequate protection for some threatened species. Therefore, a reassessment of some of the protected areas or the creation of species specific conservation action plans are recommended in order to ensure the preservation of the unique diversity of UAE terrestrial reptiles.

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In the present work, we use an exceptional database including 5,359 records of 101 species of Oman’s terrestrial reptiles together with spatial tools to infer the spatial patterns of species richness and endemicity, to infer the habitat preference of each species and to better define conservation priorities, with especial focus on the effectiveness of the protected areas in preserving this unique arid fauna. Our results indicate that the sampling effort is not only remarkable from a taxonomic point of view, with multiple observations for most species, but also for the spatial coverage achieved. The observations are distributed almost continuously across the two-dimensional climatic space of Oman defined by the mean annual tempera- ture and the total annual precipitation and across the Principal Component Analysis (PCA) of the multivariate climatic space and are well represented within 17 out of the 20 climatic clusters grouping 10% of the explained climatic variance defined by PC1 and PC2. Species richness is highest in the Hajar and Dhofar Mountains, two of the most biodiverse areas of the Arabian Peninsula, and endemic species richness is greatest in the Jebel Akhdar, the highest part of the Hajar Mountains. Oman’s 22 protected areas cover only 3.91% of the country, including within their limits 63.37% of terrestrial reptiles and 50% of all endemics. Our analy- ses show that large areas of the climatic space of Oman lie outside protected areas and that seven of the 20 climatic clusters are not protected at all. The results of the gap analysis indi- cate that most of the species are below the conservation target of 17% or even the less restrictive 12% of their total area within a protected area in order to be considered adequately protected. Therefore, an evaluation of the coverage of the current network of protected areas and the identification of priority protected areas for reptiles using reserve design algorithms are urgently needed. Our study also shows that more than half of the species are still pending of a definitive evaluation by the International Union for Conservation of Nature (IUCN).

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Es wird über die langjährige und erfolgreiche Haltung und Nachzucht der Omaneidechse Omanosaura jayakari im Terrarium berichtet. Hierbei werden insbesondere Angaben zum Verhalten der Echsen sowie gewonnene Daten zur Fortpflanzungsbiologie mitgeteilt.

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The analysis of biogeographical structure and patterns of endemism are central topics of biogeography, but require exhaustive distribution data. A lack of accurate broad-scale information on the distribution of reptiles has so far limited the analyses of biogeographical structure. Here we analysed the distribution of reptiles within the broad-sense Western Palearctic to assess biogeographical regionalization using phylogenetic and non-phylogenetic approaches, identified areas of endemism and evaluated the environmental factors pro- moting community uniqueness and endemism. We gathered distributional records from the literature and from the field, mapping the distribution of all the Western Palearctic reptiles on a 1-degree resolution grid. bsim dissimilarity and hierarchical clustering was used to identify bioregions, analysing data both at the species and at the genus level, and considering phylogenetic dissimilarity. Consensus areas of endemism were identified on the basis of the optimality criterion. We then assessed whether biogeographical structure is related to present-day climate, insularity, orography and velocity of climate change during the Late Qua- ternary. The genus-level analysis identified five main biogeographical regions within the Western Palearctic, in partial agreement with previous proposals, while the species-level analysis identified more bioregions, largely by dividing the ones identified by genera. Phylogenetic bioregions were generally consistent with the non- phylogenetic ones. The strongest community uniqueness was observed in subtropical warm climates with seasonal precipitation and low productivity. We found nine consensus areas of endemism, mostly in regions with limited velocity of Quaternary climate change and warm subtropical climates. The biogeographical structure of Western Palearctic reptiles is comparable to what has been observed in other vertebrates, with a clear distinction between the Saharo-Arabian-Sindian and Euro-Mediterranean herpetofaunas. Unlike other vertebrates, in reptiles the highest uniqueness and endemism is observed in dry climates, but the velocity of climate change during the Quaternary remains a major driver of endemism across all the vertebrates.

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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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Th e terrestrial reptile fauna of Oman and the United Arab Emirates is rich, with at least 79 species of lizards and snakes and a single species of worm lizard. However, to date there have been no accurate maps published of their distribution ranges, and distribution data relies on scattered museum specimen localities and published accounts. Considerable numbers of locality data points do exist, collected by visting and resident herpetologists, and more recently, from ecologists working on surveys for environmental impact assessments and biodiversity action plans. Th ese data are invaluable, as amongst other uses, they can assist conservation planning and management, and will eventually document changes in distributions over time. Th is is especially true where there has been extensive habitat loss and degradation due to urbanisation and development activities. Data have been collected from museum records, published accounts and unpublished data from a variety of sources, including many records made by the author over the last 20 years, with the aim of producing an atlas of species distributions. Th e number of records is now approaching 5.000, giving suffi cient coverage to produce maps that are useful for a variety of applications. Examples are discussed, including endangered and endemic species, snakes of medical importance and species of potential interest in ecological and evolutionary studies.

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The terrestrial reptiles oft he UAE the lizards, snakes and a single species of amphisbaenian or worm lizard. There are no extant native crocodiles, tortoises or freshwater terrapins. Lizards, snakes and amphisbaenians are all classified under the reptilian order Squamata, in the suborders Sauria, Serpentes and Amphisbaenia respectively. Characterised by their dry, scaly skin, which protects them from drying out, and by their reliance on external heat sources to maintain their body temperatures, reptiles are well-suited to hot and arid areas. In Arabia, reptiles are a major component of the desert and mountain ecosystems and a total of at least 54 species of terrestrial reptile are found in the UAE. So far, a comprehensive handbook covering all of the UAE reptiles in detail is not available. The Handbook to Middle East Amphibians and Reptiles (Leviton et al. 1992) does not cover the species of the mountains or Northern Emirates and gives little information on natural history. Useful general sources of information are Wild about Reptiles (Jongbloed 2000) and Terrestrial Reptiles of Abu Dhabi (Baha El Din 1986).

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From the 26th of April until the 3rd of May I had a herpetological trip to Oman together with Javier Aznar, Javier Gallego and Gabriel Martinez. During these eight days we were able to find and photograph 48 reptile- and 2 amphibian species. The trip started in Maskat and took us by Al-Qabil, Thumrait to Salalah and back by Al-Qabil, Nizwa, Ibri, Barka to Maskat. The course of this journey is presented in chronological order.

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Die kürzlich erstellten Checklisten der Amphibien und Reptilien für den Jebel al-Akhdar und im Besonderen für das Saiq-Plateau (Kowalski et al. 2011, Grossmann et al. 2012) werden durch die Ergebnisse einer weiteren, im Juni 2013 auf dem Saiq-Plateau durchgeführten Exkursion ergänzt. Dieser Besuch erbrachte den Erstnachweis für Echis omanensis, sowie neue Erkenntnisse an anderen Taxa der Herpetofauna. Damit sind nun zwei Amphibien- und Reptilientaxa für das Saiq-Plateau bekannt.

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In recent years there has been an exponential increase in the amount of published DNA sequences, as it has become one of the most popular sources of data for reconstructing phylogenies. A number of studies have attempted to resolve the relationships of groups of lacertids using mitochondrial DNA sequence data, in particular Podarcis and Gallotia (e.g. THORPE et al., 1994; 1996; GONZALEZ et al., 1996; FU et al., 1997; HARRIS, 1997; HARRIS et al., 1998a; 1999; CASTILLA et al., 1999a,b). Reassessment of these data suggest some alternative phylogenies to those previously published, and also highlight possible problems. Two studies have attempted to unravel the phylogeny of the whole family using mitochondrial DNA sequences (FU, 1998; HARRIS et al., 1998b). By combining these essentially congruent data sets, and reanalysing levels of support for the phylogenies produced, a more complete estimate of relationships across the family is produced. This is then compared to alternative sources of evidence, primarily morphological (ARNOLD, 1989), biochemical (MAYER & BENYR, 1994) and karyological (OLMO et al., 1993). Reasons for discrepencies are discussed, as are the relative advantages and disadvantages of the different sources of phylogenetic inference.

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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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Background: Although current molecular clock methods offer greater flexibility in modelling evolutionary events, calibration of the clock with dates from the fossil record is still problematic for many groups. Here we implement several new approaches in molecular dating to estimate the evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and uncertain phylogeny. Four different models of rate variation are tested in a new program for Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by expressing multiple calibration dates as a range of probabilistic distributions. We also test the reliability of our proposed calibrations by exploring effects of individual priors on posterior estimates. Results: According to the most reliable model, as indicated by Bayes factor comparison, modern lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate much earlier origins for these clades than previously reported, and we discuss our results in light of paleogeographic trends during the Cenozoic. Conclusion: This study represents the first attempt to estimate evolutionary ages of a specific group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates from the fossil record as minimum node constraints. The bioinformatics program used here, TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with similar challenges.

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The Afrotropical element in the Arabian herpetofauna is concentrated in the southwestern mountainous region, whereas everywhere else in the peninsula the reptile fauna is Saharo-Sindian. There is a high degree of correspondence between the reptile faunas of the Yemens and Somalia. A remarkable faunistic resemblance also exists between the relict faunas of SW Arabia and Morocco. Three evolutionary levels can be distinguished among Ethiopian taxa in Arabia and are interpreted as evidence of different times of faunal contact: 1. Old relicts (genus level) of a Palaeogene (pre continental drift) Arabian/North African fauna (Pristurus, Trogonophidae); 2. Endemic species closely related to East African species (most probably Miocene); 3. Species distributed on both sides of the Bäb al-Mandab (Pleistocene). Contrary to the Saharo-Sindian faunal elements, there is good evidence that groups 2 and 3 used a landbridge across the Bäb al-Mandab for faunal exchange: the Miocene closure of the southern outlet of the Red Sea and a Pleistocene land connection due to eustatic sea level change.

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the Lacertidae is one of the most diverse and widespread lizard families throughout Eurasia and Africa. Several studies so far have attempted to unravel the phylogeny of Lacertidae using morphological and mole- cular data. However, the intra-family relationships remain unclear. in an effort to explore the phylogenetic relationships within the family Lacertidae, a concatenated dataset of 5727 bp from six genes (two nuclear and four mitochondrial) and 40 genera was assembled based on GenBank database. Phylogenetic inference analy- ses were conducted using Maximum Parsimony (MP), Bayesian inference (Bi) and Maximum Likelihood (ML), revealing that even a combined dataset of both mitochondrial and nuclear genes is not able to resolve the phylogenetic relationships of the Lacertidae family under the tribe level.

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In this third paper on the herpetofauna of Oman, three families are treated: eleven Lacertidae, six Scincidae and one varanid. New records, notes on ecology and localities are combined with data from literature.

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The herpetological results of three brief journeys to the saiq Plateau in the Sultanate of Oman are summarized. These yielded eight new locality records and numerous field observations. Existing distribution data are discussed relative to the new finds, and a checklist of recorded and expected species is presented.

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The dangers of helminth infestations for lizards, particularly infestations with nematodes und cestodes, are reported. Infection routes, diagnoses, und treatments are described.

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CAPTIVE MAINTENANCE AND BREEDING OF THE OMAN LIZARD LACERTA JAYAKARI The rare and beautiful Oman lizard proves to be an animal suitable for keeping in terraria. Since 1980 this lizard is bred in Europe, while only seven specimens of this species were known in 1971. The author succeeded in breeding 26 hatchlings since 1982. The lizard can be kept in a glass covered gardentenarium during summer. The reproductive cycle starts in autumn. This is probably linked with an estivation in the hot dry summerofOman. ]`hough hibernation in Oman is not a necessity. hibernation is triggered if unfavorable climatical conditions occur.

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The ecology, behaviour and first breeding of Lacerta cyanura are described. The breeding female laid three eggs every three weeks du- ring July to November. The first hatchlings were born after 59-61 days of artificial incuba- tion at 28°C. The colour pattern of hatchlings is entirely different from that of adults. The adults are dorsally light grey-brown, with a blue throat, white belly and a turquoise tail. The young have a black back and flanks with cream-yellow longitudinal stripes, a black and yellow marbled head, a very faintly yellowish or greenish underside, and a fluorescent blue tail. In contrast, the young of the presumed close relative Lacerta jayakari have almost the same colour pattern as the adults; this sug- gests a more distant relationship between the two species than has been assumed.

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The rare Southeast Arabian lizard Lacerta cyanura ARNOLD, 1972 has been found at two additional localities (seven being actually known: GRUBER & al. 1993), which are both situated on the Musandam Pen- insula, Oman. Two males and one female are kept in captivity. The female, associated with only one of the males, so far produced 6 clutches of 3 eggs each in intervals oj roughly 3 weeks. This reproductive pattern and the juvenile dress are discussed, also in respect to the systematic relationship of this species.

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

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

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According to their phylogenetic relationships, the subgenera Zootoca, Omanosaura, and Timon are excluded from the genus Lacerta, and Teira is excluded from the genus Podarcis. On the basis of their morphological and karyological pecularities, they are defined as full genera.

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The family Lacertidae encompasses more than 250 species distributed in the Palearctis, Ethiopis and Orientalis. Lacertids have been subjected in the past to several morphological and molecular studies to establish their phylogeny. However, the problems of convergent adaptation in morphology and of excessively variable molecular markers have hampered the establishment of well supported deeper phylogenetic relationships. Particularly the adaptations to xeric environments have often been used to establish a scenario for the origin and radiation of major lineages within lacertids. Here we present a molecular phylogenetic study based on two nuclear marker genes and representatives of 37 lacertid genera and distinct species groups (as in the case of the collective genus Lacerta). Roughly 1600 bp of the nuclear rag1 and c-mos genes were sequenced and analyzed. While the results provide good support to the hitherto suggested main subfamilies of Gallotiinae (Gallotia and Psammodromus), Eremiainae and Lacertinae [Harris, D.J., Arnold, E.N., Thomas, R.H., 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proc. R. Soc. Lond. B 265, 1939–1948], they also suggest unexpected relationships. In particular, the oriental genus Takydromus, previously considered the sister-group to the three subfamilies, is nested within Lacertinae. Moreover, the genera within the Eremiainae are further divided into two groups, roughly corresponding to their respective geographical distributions in the Ethiopian and the Saharo-Eurasian ranges. The results support an independent origin of adaptations to xeric conditions in different subfamilies. The relationships within the subfamily Lacertinae could not be resolved with the markers used. The species groups of the collective genus Lacerta show a bush-like topology in the inferred Bayesian tree, suggesting rapid radiation. The composition of the subfamilies Eremiainae and Lacertinae as well as their phylogeography are discussed.

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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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A total of 2729 base pairs from three mitochondrial (12S, ND4 and CYTB) and two nuclear (C‐MOS, MC1R) genes were used to assess the molecular diversity and phylogenetic structure of the two Omanosaura lacertid lizards endemic to the Hajar Mountains in Northern Oman. Sequence data from fifteen samples of O. jayakari and five of O. cyanura were analyzed using Maximum likelihood and Bayesian inference phylogenetic methods, haplotype networks, and molecular dating. Our divergence time estimates suggest that the two species diverged around 8 million years ago (Mya). Mitochondrial sequence data show that the genetic diversity within O. jayakari is very low. Phylogenetic analyses support a partition into two clades without a clear geographic association across the Hajar Mountains, their split dating back to approximately 0.31 Mya. On the other hand, O. cyanura presents three relatively deep lineages associated with three geographic regions of the mountains: lineage 1, restricted to the Musandam Peninsula in the extreme north of the mountain range; lineage 2, including only one sample located in the Western Hajars; and lineage 3, including two samples from the Jebel Akhdar, the highest part of the Mountains, situated between the Western and Eastern Hajars. According to our time estimates, lineage 1 diverged from the other two lineages around 3.95 Mya, and lineages 2 and 3 would have diverged approximately 2.44 Mya. Nuclear data confirm the genetic homogeneity of O. jayakari and the high genetic diversity within O. cyanura. More samples of O. cyanura from undersampled areas will be needed to confirm the consistency of this pattern.

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An increase in studies in the Hajar Mountains from the southeastern Arabian Peninsula has revealed a high richness of endemic evolutionary lineages with many cryptic taxa. Omanosaura is the only lacertid lizard genus endemic to the Hajar Mountains, with two species O. cyanura and O. jayakari distributed throughout this mountain range. The phylogenetic relationships and genetic diversity between and within these species have been poorly studied. In this study, we collected mitochondrial (12S, cytb, and nd4) and nuclear (cmos and mc1r) sequences for 25 specimens of Omanosaura, including 15 individuals of O. jayakari and 10 of O. cyanura. We performed phylogenetic analyses based on network reconstruction, maximum likelihood and Bayesian inference to estimate the relationships and intraspecific genetic diversity of these species. We estimated the time of divergence between the two species in the Miocene, around 8.5 million years ago. Omanosaura jayakari shows little genetic diversity, while O. cyanura presents two differentiated lineages. These are reciprocally monophyletic at mitochondrial and nuclear genes and present a high genetic distance between them. These two lineages are associated with the geographic features of the Hajar Mountains, with one lineage distributed in the northernmost part of the Hajar Mountains and the other in the rest of the western Hajars, the Jebel Akhdar, and the eastern Hajars. This geographic relationship has been recovered previously in other reptile taxa and is generally associated with high levels of local genetic diversity. Our results suggest the existence of cryptic diversity within O. cyanura and support a general biogeographic pattern of high diversity and endemism in the northern Hajar Mountains that certainly deserves additional research in the 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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While the stability of sex chromosomes is widely accepted in viviparous mammals and birds, ectothermic vertebrates are still largely viewed as having frequent turnovers in sex determining systems. Frequent changes in sex determining systems in ectotherms could be problematic for field ecological studies as well as for breeding programs, as molecular sexing across a phylogenetically widespread spectrum of ectothermic vertebrates would not be possible. However, we recently documented that sex determining systems in three important reptile lineages (caenophidian snakes, iguanas, and lacertid lizards) are in fact highly conserved. 2.We applied a new molecular procedure to identify sex within each of these three lineages (encompassing altogether around 4,000 species, i.e. nearly 50% of the recent species of reptiles). This technique uses quantitative PCR (qPCR) to compare copy numbers of genes specific for their respective Z (in caenophidian snakes and lacertids) and X (in iguanas) chromosomes between male and female genomes. The DNA samples required can be collected relatively non-invasively. Unlike molecular sexing based on repetitive elements, this technique can be easily applied to previously unstudied species of these lineages, as the number of copies of protein coding genes linked to the differentiated sex chromosomes is evolutionary highly conserved in each. 3.We suggest that qPCR-based molecular sexing using the comparison of gene copy number is a practical choice for non-model species of caenophidian snakes, iguanas and lacertids. Morever, it should also soon be available for other reptile lineages with differentiated sex chromosomes.

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The order Squamata is a species-rich group of reptiles including the groups Sauria, Serpentes and Amphisbaenia. Several members of these groups have been extensively used as model organisms for a great variety of studies of different fields including ecology, behaviour and medicine. Such studies require a well-established phylogenetic framework to trace the evolution and diversity of the studied traits in the squamate tree of life. However, the accuracy of phylogenetic inference in Squamata is unbalanced: while the position of the main squamate groups and high level relationships have been successfully and consistently estimated across phylogenetic studies, the phylogenetic relationships of some groups have provided inconsistent results over the years. The accurate estimation of relationships between these groups might have been hindered by methodological artefacts of the phylogenetic inference due to limited data and analytical tools. For the last couple of decades, and even presently, phylogenetic inference has relied heavily on mitochondrial DNA as the main molecular marker as a result of its advantages, such as the lack of recombination, easy amplification and high evolutionary rate. While its use is based on sound reasons, some drawbacks and limitations have been highlighted, such as the overall small size, and the fact that all genes are linked thus it represents a single locus. To overcome this, the addition of nuclear DNA complements the use of mitochondrial DNA in phylogenetic inference. Within the nuclear DNA, some markers may present so little variation that it is insufficient to recover phylogenetic relationships, yet slow-evolving nuclear genes have been the most widely used markers in phylogenetic studies. On the other hand, it has been shown that the use of highly informative fast evolving nuclear genes, in combination with mitochondrial DNA, can provide resolution at different parts and depths of the evolutionary history of the species. Phylogenetic analyses of multiple loci have historically been based on the concatenation approach, .i.e. the combination of the sequences into a single alignment that is then analysed to represent the species tree. This method, however, relies on the assumption that the most commonly occurring gene tree is equivalent to the true species tree of any taxa, which may not be true in all cases. Another limitation of the concatenation approach is the elevated percentage of missing data that it sometimes entails, especially in large taxon sets, meaning that the sequence representation is The order Squamata is a species-rich group of reptiles including the groups Sauria, Serpentes and Amphisbaenia. Several members of these groups have been extensively used as model organisms for a great variety of studies of different fields including ecology, behaviour and medicine. Such studies require a well-established phylogenetic framework to trace the evolution and diversity of the studied traits in the squamate tree of life. However, the accuracy of phylogenetic inference in Squamata is unbalanced: while the position of the main squamate groups and high level relationships have been successfully and consistently estimated across phylogenetic studies, the phylogenetic relationships of some groups have provided inconsistent results over the years. The accurate estimation of relationships between these groups might have been hindered by methodological artefacts of the phylogenetic inference due to limited data and analytical tools. For the last couple of decades, and even presently, phylogenetic inference has relied heavily on mitochondrial DNA as the main molecular marker as a result of its advantages, such as the lack of recombination, easy amplification and high evolutionary rate. While its use is based on sound reasons, some drawbacks and limitations have been highlighted, such as the overall small size, and the fact that all genes are linked thus it represents a single locus. To overcome this, the addition of nuclear DNA complements the use of mitochondrial DNA in phylogenetic inference. Within the nuclear DNA, some markers may present so little variation that it is insufficient to recover phylogenetic relationships, yet slow-evolving nuclear genes have been the most widely used markers in phylogenetic studies. On the other hand, it has been shown that the use of highly informative fast evolving nuclear genes, in combination with mitochondrial DNA, can provide resolution at different parts and depths of the evolutionary history of the species. Phylogenetic analyses of multiple loci have historically been based on the concatenation approach, .i.e. the combination of the sequences into a single alignment that is then analysed to represent the species tree. This method, however, relies on the assumption that the most commonly occurring gene tree is equivalent to the true species tree of any taxa, which may not be true in all cases. Another limitation of the concatenation approach is the elevated percentage of missing data that it sometimes entails, especially in large taxon sets, meaning that the sequence representation is Miocene and supported a scenario of east-to-west diversification. The little morphological and phylogenetic evidence for the distinctiveness between Rhinechis and Zamenis supports a classification lumping which better reflects their evolutionary history and, based on the priority rule, R. scalaris is moved into the genus Zamenis and designated as Zamenis scalaris comb. nov. Within the Gallotiinae subfamily, the combined use of fast nuclear markers and the species tree was fruitful for the inference of a robust phylogeny of the genus Psammodromus, distributed in the Iberian Peninsula and North Africa. In this study we analysed for the first time the six Psammodromus species to infer the association between the main cladogenetic events within this genus and the complex biogeographic dynamics across the Strait of Gibraltar. The inferred phylogeographical history suggests that Psammodromus probably originated in Iberia. The African species and the African lineage of P. algirus were a result of two over-sea dispersal events towards Africa that occurred 10 Ma and 1.5 Ma, while continental vicariance events might have shaped the diversification of the species within Iberia and Africa. These results, combined with previous literature, provide compelling evidence that major biotic exchanges occurred across the Strait of Gibraltar well before or long after the land connection during the Messinian Salinity Crisis (5.9 – 5.33 Ma). These findings suggest caution in the application of the relatively short event of the opening of the Strait of Gibraltar at the end of the Messinian Salinity Crisis as a cause for divergence in molecular clock calibrations, which is a common approach in literature. At the species level, the use of fast evolving nuclear markers proved to be fruitful to understand the geographical pattern of phyletic diversification within the Omanosaura lizards belonging to the Eremiadini tribe, endemic to the Hajar Mountains in the Arabian Peninsula. Multilocus phylogenetic analyses recovered two highly divergent lineages within O. cyanura which are geographically associated to the northernmost and to the south and eastern regions of the Hajar Mountains. These lineages are reciprocally monophyletic at both mitochondrial and nuclear loci suggesting a long history of independent evolution and the need of a comprehensive taxonomic assessment. Omanosaura cyanura represents an additional case of cryptic diversity in the north Hajar Mountains, an area that has been demonstrated to hide high levels of genetic diversity in other reptile groups. Overall, the work developed in this dissertation has demonstrated that the inclusion of fast evolving nuclear genes and the coalescent species tree approach was relevant for resolving challenging phylogenetic questions on selected squamate groups. This approach allowed the recovery of new clades and provided support for old and recent relationships. These methods have further allowed the comparison between contrasting phylogenetic hypotheses on the Lacertini and Colubrinae, demonstrating that the inference based on the supermatrix approach used by many recent studies may provide high support for incorrect nodes. Future research directions include (i) the application of the phylogenomic approach based on Next Generation Sequence data to further resolve basal polytomy within the Lacertini and Colubrinae radiations; (ii) a taxonomical assessment of Omanosaura; (iii) a re-evaluation of molecular clock calibrations based on the end of the Messinian Salinity Crisis as general cause for divergence, in order to account for biotic exchanges by over-sea dispersal across the Mediterranean well before or long after the land connection during this period, as suggested by emerging literature and the study case on Psammodromus.

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Aim: Deserts are generally perceived as areas of low diversity, and hence receive little attention from researchers and conservationists. Squamates are the dominant group of vertebrates in arid regions, and as such represent an ideal model to study biodiversity patterns in these areas. We examine spatial patterns of diversity, evolutionary history and endemism of terrestrial squamates of the Arabian Peninsula and test hypotheses on the role of topography and history of isolation so as to identify possible environmental drivers of diversification. Location: The Arabian Peninsula. Taxon: Squamate reptiles (Squamata; lizards and snakes). Methods: We generated distribution maps for all Arabian squamate species (including yet undescribed) and reconstructed their phylogenetic relationships using existing and newly produced genetic data for nearly all the species. We assessed patterns of the distribution of species richness, phylogenetic diversity and phylogenetic en demism across the peninsula to identify areas that could be considered evolutionary or endemicity hotspots for squamates. We evaluated community turnover across the peninsula and assessed the possible environmental drivers affecting the diversity of Arabian squamates in a regression framework. Results: The main hotspots of Arabian squamate diversity are mostly along the mountains that rim the peninsula while the most arid, central regions support a low diversity of species. The distribution of the phylogenetic diversity mirrors that of the species richness. Phylogenetic endemism is also highest in the mountains, especially when only endemic species are analysed. The deserts of northern Arabia are poor in terms of species richness and they show low connectivity to the peninsular communities. Topographic heterogeneity is the strongest predictor for Arabian squamates, followed by elevation. There is no correlation between richness and temperature. Main conclusions: The mountains of Arabia support rich and unique squamate communities that are dominated by local radiations of closely related and narrow-ranging species. In particular, the Asir Mountains of SW Arabia, Dhofar Province of Oman and the Hajar Mountains of northern Oman and UAE show unprecedented levels of squamate endemism and phylogenetic endemism. While many generalist species range across Arabia, a low number of species is shared between the peninsula and mainland Asia, indicating an effective isolation of the Arabian fauna. Squamate richness is highest in heterogeneous, topographically complex habitats.

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This paper provides an updated checklist of the United Arab Emirates (UAE) native and alien herpetofauna. The UAE, while largely a desert country with a hyper-arid climate, also has a range of more mesic habitats such as islands, mountains, and wadis. As such it has a diverse native herpetofauna of at least 72 species as follows: two amphibian species (Bufonidae), five marine turtle species (Cheloniidae [four] and Dermochelyidae [one]), 42 lizard species (Agamidae [six], Gekkonidae [19], Lacertidae [10], Scincidae [six], and Varanidae [one]), a single amphisbaenian, and 22 snake species (Leptotyphlopidae [one], Boidae [one], Colubridae [seven], Hydrophiidae [nine], and Viperidae [four]). Additionally, we recorded at least eight alien species, although only the Brahminy Blind Snake (Ramphotyplops braminus) appears to have become naturalized. We also list legislation and international conventions pertinent to the herpetofauna.

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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.