×

Twenty-four species of lacertid lizards were examined at 31 sites in western Yogoslavia, Greece and Iberia. Comparative observations were made on over 4500 individual lizards, noting such features as times of activitiy, hunting methods, diet, micro- and macrohabitat, refuges used and body temperature. These data are used as a basis for assessing resource partition in related sympatric species, for find out whether different systematic groups have characteristic types of niche, and for comparing cumminity structures in the peninsulas of southern Europa. Acanthodactylus erythrurus, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta graeca, Lacerta horvathi, Lacerta lepida, Lacerta monticola, Lacerta oxycephala, Lacerta mosorensis, Lacerta schreiberi, Lacerta trilineata, Lacerta viridis, Lacerta vivipara, Podarcis bocagei, Podarcis erhardii, Podarcis hispanica, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis sicula, Podarcis taurica, Psammodromus algirus, Psammodromus hispanicus.

×

Relationships of lacertid lizards were assessed on the basis of 84 primary and 112 binary characters drawn mainly from morphology, including features of the skeleton, external anatomy, various internal soft part systems and two aspects of behaviour. Among features not previously used, or not fully investigated before, are structure of the septomaxilla and nasal passages, arranged of the xiphisternal cartilages, mite pockets, kidney position, ulnar nerve arragement, thoracic fascia, aspects of the hemipenis and its associated muscles, female genitalia and jaw muscles. On the basis of parsimony analysis and compatibilty treatment of this character set, the Lacertidae fall into two main portions: A paraphyletic Palaearctic and Oriental group of primitive forms, from which is derived a holophyletic assemblage of Ethiopian and advanced Saharan and Eurasian taxa. The former group ist not fully resolvable, but Psammodromus and Gallotia appear to be sister groups and are probably related to Lacerta parva and L. fraasi and then L. brandtii, Podarcis appears to be related successively to L. andreanszkyi, the sister species L. dugesii and L. perspicillata, and perhaps L. danfordi and L. laevis. This assemblage may be related to archaeolacertas and Algyroides. The separation of Lacerta lepida, L. pater and L. princeps from the agilis group, based on chemical evidence, is weakly contradicted by morphology. Takydromus may be most closely related to L. vivipara, and L. jayakari and L. cyanura constitute the most likely sister group of the Ethiopian and advanced Saharo-Eurasian assemblage. Taxe in the Ethiopian and advanced SaharoEuroasian assemblage form a long essentially pectinate tree with relatively change between the side branches, except for a strong disjunction separating the more primitive from the more advanced taxa. Most of the former fall on two main branches, with ´Lacerta` australis and ´L.` rupicola possibly basal to them. 1. the Equatorial forest group containing Gastropholis, Bedriagaia, ´Lacerta` echinata, Adolfus, ´Lacerta` jacksoni and Holaspis. The first three of these constitute a holophyletic group and the same is probably true of the remainder. 2. Tropidosaura, Poromera and Nucras, the latter being the sister group of the more advanced forms. These include successively the Ethiopian Philochortus, Latastia, Ichnotropis and Heliobolus, Pseuderemias, Meroles and Aporosaura, and Pedioplanis, and then the Saharo-Eurasian Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita. It seems probable that the ancestors of modern Lacertidae arose in western Eurasia, where the family is known since the Palaeocene and is still represented there largely by quite primitive forms (89 species and seven nominal genera). The family later invaded Africa, perhaps first in the early or middle Miocene. Relatively primitive lacertids spread widely in largely mesic situations in the Ethiopian region, radiating to some extent (six present genera and 16 species) and producing Nucras and the related series of advaned groups (eight genera and 54 species) whoich show increasing adaptation to xeric environments. These genera tend to have heir most primitive species in the northeast and north of the Ethiopian region. The most advaned gave rise to the Saharo-Eurasian clade, now made up to Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita. This invaded the arid areas of North Africa and Eurasia, where it is presently represented by 70 species. Many morphological changes in increasingly advanced lacertids may be functionally related to the problems of survival in arid, hot, open environments. Considerable ecological parallelism exists in lacertids, with members of separate stocks occupying similar niches in different geographical areas. Morphological adaptations associated with these niches contribute significantly to the high levels of character homoplasy found in the family. There is also some correlation between the degree of niche differentiation in various groups and the quality of the phylogenies that can be produced from their physical characters. A number of morphological parallels exist between advaned lacertids and New World macroteiids. In the skull at least, advaned lacertids show a complex mixture of paedomorphosis and acceleration. Nomenclatorial changes are as follows: Cabrita is synonymised with Ophisops, necessitating a new name, Ophisops nictans, for Cabrita jerdonii. Aporosaura is synonymised with Meroles, Platyplacopus with Takydromus, and Bedriagaia with Gastropholis. ´Lacerta` (or Centromastyx) echinata is also transferred to the latter genus and Lacerta jacksoni to Adolfus. ´Lacerta` australis and ´L.` rupicola are put in a new genus, Australolacerta. It is recommended that Lacerta dugesii and L. perspicillata should not be placed in the otherwise very uniform genus Podarcis. Although clearly paraphyletic, Lacerta s. lat. Should be retained at least for the present and, if necessary putative relationships within it indicated by informal groups or subgenera.

×

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.

×

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.

×

The microornamentation of the scales (oberhautchen) has been studied by means of enamel casts (in negative), or with detached skin pieces (positive cast in the upper face or negative in their lower face of the scales) and its observation by transparency with optical microscopy at 40x, 400x and 1000x. In this study we have found a remarkable uniformity in all genera of Lacertini studied (already known, although significantly expanding the number of species previously studied). In light of the results obtained, it seems that the use of the microornamentation of the scales in taxonomy is limited. There is a remarkable degree of variation and probably of homoplasy, as it happens in so many characters of Lacertini. Nor is it clearly linked to certain conditions of the environment, although it may have a certain importance in mechanisms such as evapotranspiration and perhaps in the crypsis through its influence on the reflectance of the scales. These scales can present protuberances (like small tubercles) on their surface (derived character) in Algyroides (the four species), and in the case of Algyroides marchi also seems to present small pits on its scale surface. In most lacertids the cells of the “oberhautchen” are usually narrow (or short anteroposteriorly, if preferred), between 1 and 4 μm, having the appearance of ribbons or transverse straps. Smaller sizes (derivated) appear more or less marked in Algyroides spp, Darevskia praticola complex, Parvilacerta spp and in Dinarolacerta mosorensis (although not in D. montenegrina) and is difficult to interpret. Except in Dinarolacerta, in the other species it coincides that they are ground or ground-rock interface species, and the coarser stripes could have to do with reflectance (imitating the vegetation or the medium between which they move, according to the hypothesis of Allam et al.). In the case of Dinarolacerta, very saxicolous, it is probably the sister group of Algyroides and the common morphology could have a phylogenetic origin linked to its relationship with Algyroides. There is also no relationship with moisture since some come from dry places and others are from very wet areas. The posterior flange of the cells may be slightly mounted in the next cell (primitive character) but appears quite raised (derived character) in Algyroides nigropunctatus (much less in A. moreoticus and A. marchi, and apparently little or nothing in A. fitzingeri). The rear flange of the cells can be smooth, corrugated or notched. In our samples, irregular (and even dentate) posterior ridges appear in numerous Darevskia such as D. derjugini, D. parvula, D. raddei complex [D. raddei raddei, D. r. vanensis and D. nairensis], in part of the group of D. rudis [D. rudis sspp. and D. bithynica sspp.], several Iberolacerta (I. bonnali, I. horvathi and I. monticola astur), as well as in Teira dugesii. The character appears, although less marked in Algyroides (more in A. moreoticus, but less marked or barely visible in the others), in Hellenolacerta graeca, and other Iberolacerta (I. aurelioi, I. martinezricai, and scarcely marked in I. cyreni). Very little marked in Darevskia mixta, somewhat more in D. dryada and none in D. clarkorum, and also appears in the group of D. saxicola (D. saxicola, D. brauneri brauneri, D. b. darevskii, D. lindholdmi, D. szczerbaki), Dalmatolacerta oxycephala, Dinarolacerta mosorensis (only in the Biokovo sample, and not in Durmitor or D. montenegrina), Apathya cappadocica, in some places of the scales of Parvilacerta fraasii (but not in P. parva) and very little in D. (p) hungarica; also in the remaining group of D. rudis (D. valentini valentini, D. v. lantzicyreni, D. rudis bolkardaghica, D. portschinskii), and some parthenogenetic species (D. armeniaca, D. dahli, D. rostombekowi) but not in D. unisexualis, D. uzzelli, D. sapphirina, D. bendimahiensis or in the triploid hybrid between D. unisexualis and D. nairensis. In these parthenogenetic species it seems a general rule that the character of the crenulation or irregularity of the posterior flange of the cells resembles that of the paternal rather than the maternal species. The exception is the triploid hybrid [arisen from the hybridization of D. unisexualis (parthenogenetic female, without the character) and a male of D. nairensis (with fairly developed character)] that lacks character, and in this case is more similar to the maternal species of which has two genetic sets compared to only one of the paternal one.

×

First week of October 2004 a group of nature lovers visited the southern continental part of Hellas (Greece): the Peloponnesos. Nesos means island. According to mythology Pelops, son of Tantalos, was awarded with an island, in his case a pseudo-island. The Island of Pelops or Pelopo-nesos is almost an island: it is THE original peninsula connected to the Balkans via the original Isthmos! From a biogeographical point of view this peninsula is interesting, with plants and animals found elsewhere in Europe, but also with island characteristics and with some endemical forms. Cyclamens, Peloponnesos Colchicum and several other plants were encountered. The author gives a short travelogue. He describes batrachological and herpetological results of the excur- sions made. The situation of chameleons and slowworms on the peninsula receives some extra concern. Also a list of herpspecies found and photographed is added.

×

Acanthodactylus arabicus, Acanthodactylus aureus, Acanthodactylus bedriagai, Acanthodactylus blanci, Acanthodactylus blanfordii, Acanthodactylus boskianus boskianus, Acanthodactylus boskianus asper, Acanthodactylus boskianus euphraticus, Acanthodactylus boueti, Acanthodactylus busacki, Acanthodactylus cantoris, Acanthodactylus dumerili dumerili, Acanthodactylus dumerili exiguus, Acanthodactylus erythrurus erythrurus, Acanthodactylus erythrurus belli, Acanthodactylus erythrurus lineomaculatus, Acanthodactylus felicis, Acanthodactylus gongrorhynchatus, Acanthodactylus grandis, Acanthodactylus guineensis, Acanthodactylus haasi, Acanthodactylus longipes, Acanthodactylus maculatus, Acanthodactylus masirae, Acanthodactylus micropholis, Acanthodactylus opheodurus, Acanthodactylus orientalis, Acanthodactylus pardalis, Acanthodactylus robustus, Acanthodactylus savignyi, Acanthodactylus schmidti, Acanthodactylus schreiberi schreiberi, Acanthodactylus schreiberi syriacus, Acanthodactylus scutellatus scutellatus, Acanthodactylus scutellatus hardyi, Acanthodactylus spinicauda, Acanthodactylus tilburyi, Acanthodactylus tristrami, Acanthodactylus yemenicus, Adolfus africanus, Adolfus alleni, Adolfus jacksoni, Adolfus vauereselli, Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus nigropunctatus, Algyroides nigropunctatus kephallithacius, Australolacerta australis, Australolacerta rupicola

×

Courtship and aggressive behaviour of Algyroides moreoticus in captivity are described for the first time. The breeding season commences in March. A female produces five to seven clutches per year, each containing one to four eggs. These hatch in about 36 days at 29°C and 99 % relative humidity. The young reproduce for the first time in the following spring. In comparison with available behavioural data of other European lacertids Algyroides moreoticus displays two unusual traits. For up to three hours after copulation the mouth grip by the male on the female´s flank is maintained. During fighting between males one of the pair is suddenly flipped over on its back where it is immobilised by the neck for some time.

×

Note are given on the herpetofauna of Greece, with special reference to some of the (sub-) species. The observations were made during a spring trip.

×

The ecology of the lacertids genus Algyroides is still poorly known. Nevertheless, all four species recognized, A. nigropunctatus, A. moreoticus, A. fitzingeri and A. marchi, display some degree of restriction in distribution and habitat use when compared to other Mediterranean genera. As for other ectotherms with low dispersal abilities, in absence of other forces, such spatial restrictions are expected to be linked to ecophysiological constraints. Here, we test such prediction by focusing on two crucial features of reptilian life, thermal preferences and water loss. We performed two consecutive experiments in order to determine preferred body temperatures (Tp) and water loss rates (Wl). Tp of 5-10 adult males by species was hourly monitored in a photothermal gradient for a ten-hour period. Subsequently, Wl was estimated by placing the same specimens in sealed chambers with silica gel and weighing them hourly during a 12-hour period. Both Tp and Wl differed in mean and time profiles across species, which were arranged into two groups regarding their ecophysiology. Namely, under standard conditions, A. nigropunctatus and A. fitzingeri selected for lower temperatures but lost less water than A. moreoticus and A. marchi, although the existence a trade-off between Tp and Wl was unclear. While significant size variation between all species exists (A. fitzingeri and A. marchi being dwarfed), neither SVL nor body mass accounted for the differences in ecophysiology recorded. It is, hence, concluded that intrinsic (size/shape-free) differences in thermal and hydric physiology (high Tp and Wl) may be responsible for the decrease of the ranges and habitats available suffered by A. moreoticus and, mainly, by A. marchi. Such process is likely to be linked to the spread of Mediterranean climate during the Pliocene.

×

We assessed whether ecophysiological requirements are consistent with the environmental traits within the current ranges in a relictual lizard group, Algyroides, composed of four species with restricted and disjunctive distributions. We considered temperature, precipitation and their seasonal profiles, and focused on the preferred body temperature (Tp) and the evaporative water loss (EWL). The ranges of all four species differed in environmental traits. The two geographically more restricted species followed divergent patterns: A. moreoticus inhabits hot and climatically buffered areas, and A. marchi occupies cold seasonal environments. Tp and EWL also differed among species following a geographical grouping: A. nigropunctatus (Slovenia) and A. fitzingeri (Sardinia) selected for lower Tp and lost less water than the southern species A. moreoticus (Peloponnese) and A. marchi (Southern Spain). Tp and EWL were correlated at species level but not at individual level within species. Results suggest that the current distribution of Algyroides species partly reflects their ecophysiology, with water ecology taking precedence over thermal ecology as constraining factor. By unravelling the environmental factors limiting the distribution of species, ecophysiology may provide directions for conservation, predicting the degree of vulnerability to climate change.

×

The Greek lizard fauna consists of 26 species from which 5 are monotypic and the other ones are represented by a total of 86 subspecies. Five species and 61 subspecies are endemic of Greece. A checklist including the geographical distribution of each taxon in the Greek region is presented.

×

The present study examines the genetic variability and the taxonomic relationships among five lacertid species, i.e. Podarcis taurica, P. milensis, P. peloponnesiaca, Lacerta graeca and Algyroides moreoti- cus, representing the three main genera of this family in Europe. The last four of the above species are endemic to Greece and three of them live sympatrically in Peloponnisos. These relationships were studied by allozyme analysis. Of the loci analyzed, the Mpi-1 locus was found to be a convenient molecular marker for discrimi- nation of the genera Podarcis(allele a), Lacerta(allele b) and Algyroides(allele c). The values of Nei’s genetic distances between the examined species ranged from 0.025 to 0.484. According to the UPGMA-dendrogram plotted using the Nei’s genetic distances, two species groups are formed indicating that the genera Lacertaand Algyroidesshow a stronger relationship to one another than either does to Podarcis. These results are in agree- ment with DNAsequence data but are not in accordance with previous electrophoretic and immunological studies, which suggest that Lacertais more closely related to Podarcis than to Algyroides. The studied Podarcistaxa were found to be close relatives (Nei’s distances <0.18), especially P. taurica and P. milensis. These values are lower than those usually given in the literature for the distinction of lacertid species.

×

1. The herpetofauna of the Argo-Saronic islands includes two amphibians, one tortoise, six lizards, and six snakes. There are no indigenous species, 2. Owing to the lack of previous reports for the area most species are recorded for the first time. However, of particular interest are Chalcides o. ocellatus, Coluber najadum dahlii and Malpolon monspessulanus insignitus, as these are basically mainland forms and absent from the Cycladcs, although the first two are found on Kea, close to the Attic peninsula . The record for Eryx jaculus turcicus is valuable as knowledge of its range in Greece is very incomplete.

×

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.

×

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.

×

The four species of Algyroides Bibron & Bory, 1833 form part of the relatively plesiomorphic Palaearctic clade of lacertids comprising Lacerta and its allies. An estimate of phylogeny based on DNA sequence from parts of the 12S and 16S rRNA mitochondrial genes confirms the monophyly of the genus already suggested by several morphological features. The molecular data also indicates that relationships within the clade are: (A. nigropunctatus (A. moreoticus (A. fitzingeri, A. marchi))); this agrees with an estimate of phylogeny based on morphology that assumes the taxon ancestral to Alygroides was relatively robust in body form, and not strongly adapted to using crevices. Initial morphological evolution within Algyroides appears to involve adaptation to crypsis in woodland habitats. The most plesiomorphic form (A. nigropunctatus) is likely to have originally climbed extensively on tree boles and branches and there may have been two subsequent independent shifts to increased use of litter and vegetation matrices with related anatomical changes (A. moreoticus, A. fitzingeri), and one to increased use of crevices (A. marchi). Some members of Algyroides are strikingly similar in superficial morphology to particular species of the equatorial African genus Adolfus. This resemblance results from a combination of many shared primitive features plus a few independently acquired derived ones that are likely to give performance advantage in the relatively similar structural niches that these forms occupy. This study provides evidence that: (1) the use of a combination of molecular and morphological data may sometimes allow the estimation of ancestral anatomical features when these are otherwise unknown; (2) process considerations may permit a choice to be made in cases of character evolution where tree topology means that equally parsimonious alternatives exist; such decisions about character evolution may allow ecological shifts to be similarly assessed; (3) parallel evolution in ecological analogues may involve relatively few characters.

×

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.

×

A collecting trip to Zakynthos (Ionian Islands) has produced first evidence of Testudo h. hermanni and Elaphe qu. quatuorlineata. The occurence of Emys orbicularis, Hemidactylus t. turcicus and Coluber gemonensis has been confirmed. Algyroides nigropunctatus could not be found on the island, an old record must therefore be doubted. As to the small island Marathonisi, situated in the Bay of Lagana, Algyroides moreoticus and Tarentola mauritanica are recorded for the first time. The rookeries and nesting habits of the Loggerhead sea turtle (Caretta e. caretta) are briefly described.

×

All seven lacertid species recorded from the Peloponnese can be found in a very small area in the Arkadian highlands on the Peloponnese peninsula. Up to now only one example for such unique diversity in such a small area has been described in the Mediterranean region. This density requires the occupation of different restricted ecological and ethological niches for all seven species and therefore the habitats should be diverse. This niche segregation as well as the ecological demands and the ethological differences between the species are discussed in relation to this habitat diversity.

×

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.

×

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.

×

Aim Temperature influences most components of animal ecology and life history – but what kind of temperature? Physiologists usually examine the influence of body temperatures, while biogeographers and macroecologists tend to focus on environmental temperatures. We aim to examine the relationship between these two measures, to determine the factors that affect lizard body temperatures and to test the effect of both temperature measures on lizard life history. Location World-wide. Methods We used a large (861 species) global dataset of lizard body temperatures, and the mean annual temperatures across their geographic ranges to examine the relationships between body and mean annual temperatures.We then examined factors influencing body temperatures, and tested for the influence of both on ecological and life-history traits while accounting for the influence of shared ancestry. Results Body temperatures and mean annual temperatures are uncorrelated. However, accounting for activity time (nocturnal species have low body temperatures), use of space (fossorial and semi-aquatic species are ‘colder’), insularity (mainland species are ‘hotter’) and phylogeny, the two temperatures are positively correlated. High body temperatures are only associated with larger hatchlings and increased rates of biomass production. Annual temperatures are positively correlated with clutch frequency and annual longevity, and negatively correlated with clutch size, age at first reproduction and longevity. Main conclusions Lizards with low body temperatures do not seem to have ‘slower’ life-history attributes than species with high body temperatures. The longer seasons prevalent in warm regions, and physiological processes that operate while lizards are inactive (but warm enough), make environmental temperatures better predictors of lizard life-history variation than body temperatures. This surprisingly greater effect of environmental temperatures on lizard life histories hints that global warming may have a profound influence on lizard ecology and evolution.

×

Sex chromosomes were studied in eight species of lacertid lizards using C-banding, G-banding and restriction enzyme treatment. All of the species showed female heterogamety. The W chromosome was a microchromosome in Lacerta graeca and Ophisops elegans. Two types of W were found in Lacerta vivipara; in specimens from The Netherlands it was metacentric, whereas in specimens from Russia it was acrocentric or subtelocentric. The W chromosome was homomorphic or nearly homomorphic but completely C-banded and heterochromatic in Lacerta agilis, Podarcis hispanica, Algyroides moreoticus and A. nigropunctatus. In was only possible to find sex chromosomes using the G-banding method in Podarcis sicula. The results obtained, together with data in the literature, suggest that sex chromosomes are likely to be present in all Lacertidae and that their differentiation took place repeatedly and independently in different taxa within the family. A model for sex chromosome evolution in the family, in which the starting point was the heterochromatization of the W chromosome, is proposed.

×

The ability of an animal to shed its tail is a widespread antipredator strategy among lizards. The degree of expression of this defense is expected to be shaped by prevailing environmental conditions including local predation pressure. We test these hypotheses by comparing several aspects of caudal autotomy in 15 Mediterranean lizard taxa existing across a swath of mainland and island localities that differ in the number and identity of predator species present. Autotomic ease varied substantially among the study populations, in a pattern that is best explained by the presence of vipers. Neither insularity nor the presence of other types of predators explain the observed autotomy rates. Final concentration of accumulated tail muscle lactate and duration of movement of a shed tail, two traits that were previously thought to relate to predation pressure, are in general not shaped by either predator diversity or insularity. Under conditions of relaxed predation selection, an uncoupling of different aspects of caudal autotomy exists, with some elements (ease of autotomy) declining faster than others (duration of movement, lactate concentration). We compared rates of shed tails in the field against rates of laboratory autotomies conducted under standardized conditions and found very high correlation values (r > 0.96). This suggests that field autotomy rates, rather than being a metric of predatory attacks, merely reflect the innate predisposition of a taxon to shed its tail.

×

Caudal autotomy is one of the most effective and widespread defensive mechanisms among lizards. When predators grasp the tail, lizards are able to shed it from the point of the attack and further. Numerous factors have been reported to affect tail-shedding performance such as temperature, age, predation pressure, intraspecific competition etc. Interestingly, the impact of sex on tail loss remains greatly understudied. Here, we analyzed tail autotomy performance, simulated in the lab, in 12 species of lacertid lizards belonging to five genera (Algyroides, Anatololacerta, Hellenolacerta, Ophisops, Podarcis). Our aim was to investigate whether sex affects caudal autotomy and/or the duration of post-autotomic tail movement. We failed to detect any effect of sex on tail loss in the species examined. Also, we did not find any sexual impact on the duration of tail movement after autotomy, with a single exception. Our findings suggest that autotomy serves as a defensive tactic equally in both sexes and is used in the same extent.

×

Lacertinae is one of the three lacertid lizard subfamilies with a geographical distribution confined to the Palaearctic. Several past attempts to reconstruct its phylogeny resulted in unresolved bush-like topologies. We address the question of whether the lack of resolution is due to insufficient data or whether this lack reflects a rapid succession of speciation events. We analyzed four partial and one complete gene sequences from mitochondrial and nuclear genomes, totalling roughly 3600 bp. We included 29 species representing all 19 genera suggested in recent revision of Lacertinae [Arnold, E.N., Arribas, O., Carranza, S., 2007. Systematics of the palaearctic and oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430, 1–86]. The resulting phylogeny, first, corroborates monophyly at the genus level for the suggested genera, as well as the finding that Atlantolacerta andreanskyi, until recently part of Lacertinae, belongs to the subfamily Eremiadinae. Second, we find that increasing the sequence length and combining multiple nuclear and mitochondrial sequences did not resolve the polytomy, suggesting that the inferred topology indicates a multiple cladogenesis within a short geological period, rather than a methodical artefact. Divergence time estimates, based on previous estimates of several node ages, range from 13.9 to 14.9 million years for the radiation event, however with very broad confidence interval. To associate the radiation with a narrower geological time we consider palaeogeographic and palaeoclimatic data, assuming that the Lacertinae probably evolved in Central Europe andWAsia after the collision of Africa and Eurasia. We suggest that this radiation may date to the late Langhian (ca. 14–13.5 million years) when geological events caused abrupt changes in regional water–land distribution and climate, offering a window of distinct conditions.

×

The Dalmatian Algyroides displays only two subspecies, this despite its relative large range of distribution. The subspecies Algyroides nigropunctatus nigropunctatus is found from north-east Italy through the Dalmatian coast up to northern Greece, including several islands in the Adriatic and Ionian seas. The distribution of the other subspecies, Algyroides nigropunctatus kephallithacius, is limited to the Ionian islands of Kefallonia/Ithaca, and possibly Lefkada. Information on the latter subspecies is almost absent from current literature. The author was able to locate and photograph A. nigropunctatus kephallithacius in its natural habitat of mountaineous regions of Kefallonia. The bright red/orange belly and blue throat colors typical for A. n. nigropunctatus were absent. Male A. n. kephallithacius had yellow bellies and greenish throats, while females and juveniles were uniformly yellow at the ventral side. In addition a theory is presented which may explain the process that has lead to the formation of the subspecies A. n. kephallithacius on Kefallonia.

×

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.

×

New records and ecological observations are given for some greek reptiles: Coluber caspius was found in Kefallinia and Podarcis muralis in Evvia; the occurrence of Tarentola mauriianica is confirmed for Kefallinia. Ecological data are presented concerning Algyroides nigropunctatus from Kefallinia.

×

Sex chromosomes are believed to be stable in endotherms, but young and evolutionary unstable in most ectothermic vertebrates. Within lacertids, the widely radiated lizard group, sex chromosomes have been reported to vary in morphology and heterochroma- tinization, which may suggest turnovers during the evolution of the group. We compared the partial gene content of the Z-specific part of sex chromosomes across major lineages of lacertids and discovered a strong evolutionary stability of sex chromosomes. We can conclude that the common ancestor of lacertids, living around 70million years ago (Mya), already had the same highly differentiated sex chromosomes. Molecular data demonstrating an evolutionary conservation of sex chro- mosomes have also been documented for iguanas and caenophidian snakes. It seems that differences in the evolutionary conservation of sex chromosomes in vertebrates do not reflect the distinction between endotherms and ectotherms, but rather between amniotes and anamniotes, or generally, the differences in the life history of particular lineages.

×

This book shows a comprehensive picture of the reptiles, excluding snakes, living in Europe, North Africa, Middle and Near East and Central Asia. The bulk of the book is an annotated checklist and a distributional atlas of approx 500 species of lizards, crocodiles, turtles and terrapins, and amphisbaenians, living in the Western Palearctic. Information on distribution, subspecies, chorotype and main references are provided for each species, as well as 226 colour maps showing the global distribution of the species on a grid of one degree. For each species map the bibliographic and original sources (more than 850 sources are listed) are indicated. A further 83 maps show the distribution of all genera and main species groups of Palearctic reptiles. Beside the checklist and the atlas, in some additional chapters the following arguments are treated: Materials and methods; status of knowledge on the Palearctic fauna; composition of the Western Palearctic reptile fauna (including a statement on biodiversity and species richness); descriptive biogeography (including a discussion on the Palearctic region boundaries based on reptile distribution and the herpeto-geographic sectors of the Western Palearctic); conservation status. A list of more than 730 references quoted in the text and high quality colour plates including photos of the most of the supraspecific taxa living in the study area completes the book.

×

Research on the taxonomy of European amphibians and reptiles has increased noticeably over the last few decades, indicating the need for recognition of new species and the cancellation of others. This paper provides a critical review of recent changes and draws up a tentative species list.

×

The last species list of the European herpetofauna was published by Speybroeck, Beukema and Crochet (2010). In the meantime, ongoing research led to numerous taxonomic changes, including the discovery of new species-level lineages as well as reclassifications at genus level, requiring significant changes to this list. As of 2019, a new Taxonomic Committee was established as an official entity within the European Herpetological Society, Societas Europaea Herpetologica (SEH). Twelve members from nine European countries reviewed, discussed and voted on recent taxonomic research on a case-by-case basis. Accepted changes led to critical compilation of a new species list, which is hereby presented and discussed. According to our list, 301 species (95 amphibians, 15 chelonians, including six species of sea turtles, and 191 squamates) occur within our expanded geographical definition of Europe. The list includes 14 non-native species (three amphibians, one chelonian, and ten squamates).

×

In den Jahren 2005 bis 2007 fanden im Becken von Feneos (Peloponnes, Griechenland) umfangreiche Untersuchungen zur Avifauna und Herpetofauna statt. Bei der erstmaligen Erfassung der Vogelwelt im Bergtal konnten 98 Vogelarten nachgewiesen werden, davon 81 Brutvögel, 14 Durchzügler und drei Nahrungsgäste. Für die Herpetofauna gelang der Nachweis von sieben Amphibien- und 26 Reptilienarten.

×

The lizards of the genus Algyroides belong to the family of Lacertidae and currently comprise four morphological species. Two of them (Algyroides nigropunctatus and A. moreoticus) are distributed in Greece, whereas the second one is a Greek endemic. Algyroides nigropunctatus interestingly presents two different location-specific color phenotypes. Hitherto, there hasn’t been any assiduous molecular study on the phylogenetic relationships between the different insular and continental populations of the two species, which is the aim of the present work. Fresh tissue samples, as well as voucher samples from the Natural History Museums of Crete and Bonn, were obtained in order to represent the two species’ range sufficiently. Both mitochondrial and nuclear genes were sequenced in order to reveal the intra- and inter-specific genetic differentiation. Mitochondrial data reveal the presence of several clades of A. nigropunctatus with distinct geographic distribution in southwestern Greece. Combining molecular data and field observations it seems that there is reproductive isolation between the two color morphotypes, which are genetically distinct, despite the absence of any geographic barrier in western mainland Greece. No significant divergence was revealed within A. moreoticus. Current data indicate a possible need for a thorough morphological survey that may contribute to a reevaluation of A. nigropunctatus taxonomy.

×

In pristine conditions, Mediterranean Europe was covered by dense forests during all Pleistocene and in more recent warm periods. Combined with its rather northern position this means that at that time, apart from in coastal environments, lizard habitats would have been present only where open places were formed by the action of storms, natural fires and the erosive forces of rivers. The latter will have been the most reliable and also the most important force, especially combined with the influence of large grazing and browsing animals. Variations in habitat quality in the resulting open spaces are determined by longitudinal gradients in temperature, humidity, steepness and trophic conditions. Present day ecological differentiation and some special characteristics of the lizards will be discussed in the light of these hypotheses.

×

Elements of the herpetofauna are in the author`s opinion of great importance for the zoogeographical division of the Palaearctic region. It is suggested that this region be divided into the following four subregions: 1/ arctic, 2/ boreal Euro-Siberian, 3/ bo- real Himalayan-Manchurian and 4/ Mediterranean-Central-Asiatic.

×

Benny Trapp, Fotograf, Griechenlandkenner und Leiter des Schutzprojektes `Afrikanisches Chamäleon` auf dem Peloponnes, stellt in diesem Buch alle 62 Reptilien- und Amphibienarten des griechischen Festlandes in faszinierenden Bildern vor. Die Informationen zu Beschreibung, Verbreitung, Lebensraum, Fortpflanzung und Verhalten jeder Art werden durch praktische Tipps zu Beobachtungsmöglichkeiten der oftmals scheuen und heimlich lebenden Tiere ergänzt. Neben Informationen über Land und Leute sowie Naturschutz in Griechenland erfährt man Nützliches über Giftschlangen und zur Naturfotografie. Mit seinen bestechenden, zahlreichen Fotos sicherlich eines der prächtigsten Bücher über europäische Amphibien und Reptilien.

×

Observations of lacertids and their habitats from two trips in 2006 and 2023 in the west and south of the Peloponnese peninsula are summarized. Greek Algyroides Algyroides moreoticus Bibron & Bory de Saint-Vincent, 1833, Greek Rock Lizard Hellenolacerta graeca (Bedriaga, 1886), Balkan Green Lizard Lacerta trilineata trilineata Bedriaga, 1886, Ionic Wall Lizard Podarcis ionicus (Lehrs, 1902) and Peloponnese Wall Lizard Podarcis peloponnesiacus (Bibron & Bory de Saint-Vincent, 1833) were found at various locations. In some cases, four of the mentioned species were found at one location at the same time.