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

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The hemipenis of all the Iberolacerta species is studied. Overall morphology is very similar among the different species. Hemipenes are symmetrically bilobed, with the apical section shorter than the basal one (I. galani, I. cyreni, I. monticola monticola, I. horvathi, I. bonnali, I. aranica and I. aurelioi) or subequal or with slightly inverted proportions (I. m. astur and I. martinezricai). All are similarly sized and proportional to the animal’s SVL, except in I. aurelioi (and perhaps to a lesser extent in I. horvathi) that have notably small hemipenes. Concerning the hemipenial microornamentation, all the Iberolacerta s. str. (I. monticola, I. galani, I. martinezricai, I. cyreni and I. horvathi) have hemipenial microornamen-tation of crown-shaped tubercles, more or less elongated with a series or a crown of small spinules in their apex. Among these, isolate odd shaped tubercles can appear, probably corresponding to immature states that do not reach their complete development. The Pyrenean species (subgenus Pyrenesaura) are variable. Iberolacerta bonnali and I. aranica have mainly spiny-like tubercles (among which some crown shaped can appear), whereas I. aurelioi presents crown-shaped microornamentation (with scattered spiny-like, as occur in other Iberolacerta species). Our hypothesis is that crown shaped models are derived (or more mature, following an ontogenetic criterion) in respect to the spiniform ones, but some spiny (or finger-shaped) can be secondary reversals by incomplete maturation. These secondary derived and the original primitive spiny-shaped ones are very difficult if not impossible to distinguish. Phenomena as the heterochronies could be beneath these reversals in maturation.

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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.

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A new species of Iberolacerta is described from the Montes de León (northwest Iberia). This new species, Iberolacerta galani sp. nov., is characterized by its relatively large size, high number of blue ocelli on the shoulders and the relatively frequent contact or near-contact between the supranasal and the first loreal scale, the fairly straight squamosal bone (only curved on its posterior part), a unique karyotype in Iberolacerta combining 2n=36 chromosomes, an L-type NOR and differentiated W and Z sex chromosomes, and unique mitochondrial DNA sequences for the cytochrome b and 12S rRNA genes. The correlation analyses show that morphology in general, but especially scalation, is strongly correlated with the amount of precipitation during the months of lizard activity, which suggests that these are not good taxonomic characters, and that other characters apparently independent of the climate like for instance osteological, karyological and DNA features are much more reliable in delimiting species boundaries in Iberolacerta. According to our phylogenetic analyses, I. galani nov. is part of a very well supported clade that originated around 2.5 mya and also includes I. monticola and I. martinezricai. Phylogeny suggests I.martinezricai might be the sister taxon to I. galani nov. from which it split approximately 2 mya, at the beginning of the Pleistocene. The clade containing I. galani nov., I. martinezricai and I. monticola was probably widely distributed across western Iberia during moderately cool and moist phases of the Pleistocene, but it was probably restricted to its present range as a result of the general temperature increase during the Holocene and competition with other lacertid lizards. Iberolacerta galani nov. is endemic to the Montes de León, where it is isolated from the other species of the “monticola-group” by the Duero and Miño-Sil Rivers, but particularly by the Bibei river valley.

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This paper analyses the literature published in the last 17 years regarding records of albinism, axanthism, erythrism, leucism, melanism, nigrism or piebaldism, among other chromatic anomalies in Iberian herps.Whereas hipomelanistic disorders showed to be more widespread, melanism cases had a high occurrence pattern in the northern side of the Iberian Peninsula.We suggest that thermal melanism could be an adaptive advantage for ectotherms. Notwithstanding this, we also discuss the possible negative effect of the color aberrations taking into account differences in pre-dation pressure.

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West European Rock lizards, Lacerta (Iberolacerta) have small widely separated ranges in highland areas. Mitochondrial and nuclear DNA sequences corroborate the monophyly of the group and show it is not closely related to any of the other Rock lizards with which it was formerly placed in Archaeolacerta, an assemblage for which there is no evidence of clade status. L. (Iberolacerta) consists of four main units: L. (I.) horvathi of NW Croatia and neighbouring regions; the Pyrenees species, L. (I.) bonnali, L. (I.) aranica and L. (I.) aurelioi; L. (I.) cyreni of the Iberian Sistema Central, with distinctive populations in the Sierras de Béjar,Gredos and Guadarrama; and L. (I.) monticola of the Serra da Estrela of Central Portugal and NWSpain, this unit also contains L. (I.) cyreni martinezricai of La Peña de Francia, W. Spain and a distinctive population in the Montañas de Sanabria. L. (Iberolacerta) has persisted in some mountain ranges for at least 4.2±1.4Ma and may have been restricted tomountains by competition from Wall lizards (Podarcis). Its clade status shows it has lost range extensively and has produced few external branches since its initial fragmentation. In contrast, Podarcis diversified about the time L. (Iberolacerta) fragmented, producing a series of widespread lineages that have persisted until the present time. The mainly European subfamily Lacertinae, to which both L. (Iberolacerta) and Podarcis belong, diversified rapidly 13–9 Ma ago, probably largely replacing other lacertid lizards of earlier origin. As another round of replacement started at approximately 9 Ma ago with the spread of Podarcis, this may be a recurrent phenomenon in the evolution of some lizard communities.

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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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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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We conducted an experiment on chemical discrimination of two saurophagous snakes (the smooth snake, Coronella austriaca and the Seoane`s viper, Vipera seoanei) as well as the aquatic Natrix maura, by the mountain lizard Iberolacerta galani. Using terraria, 24 lizards were exposed to scents by the three snakes as well as an odourless control. We quantified fourteen behavioural variables, twelve of which significantly differed among treatments. Lizards are able to recognise the scents of both predatory snakes, and react to them with intense antipredatory responses. The antipredatory behaviour found in I. galani was similar for the scents of the two different predatory snakes, despite differences in their foraging behaviour. The behaviour displayed by lizards confronted with chemical cues suggests an adaptation to minimise the likelihood of being attacked.

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Mountain lizards are highly vulnerable to climate change, and the continuous warming of their habitats could be seriously threatening their survival. We aim to compare the thermal ecology and microhabitat selection of a mountain lizard, Iberolacerta galani, and a widely distributed lizard, Podarcis bocagei, in a montane area. Both species are currently in close syntopy in the study area, at 1,400 m above the sea level. We determined the precision, accuracy and effectiveness of thermoregulation, and the thermal quality of habitat for both species. We also compared the selection of thermal microhabitats between both species. Results show that I. galani is a cold-adapted thermal specialist with a preferred temperature range of 27.929.7 C, while P. bocagei would be a thermal generalist, with a broader and higher preferred temperature range (30.134.5 C). In addition, I. galani selects rocky substrates while P. bocagei selects warmer soil and leaf litter substrates. The thermal quality of the habitat is higher for P. bocagei than for I. galani. Finally, P. bocagei achieves a significantly higher effectiveness of thermoregulation (0.87) than I. galani (0.80). Therefore, these mountain habitat conditions seem currently more suitable for performance of thermophilic generalist lizards than for cold-specialist lizards.

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The León rock lizard, Iberolacerta galani, lives in isolated mountains of Spain.We studied the seasonal changes in the thermal biology of I. galani between spring and summer. We calculated precision, accuracy andeffectiveness of thermoregulation and the habitat thermal quality for spring, and compared with the values of summer. In addition, we studied how the shift in the thermal preferences of lizards would contribute to achieve a higher effectiveness of thermoregulation. Thermal preferences of León rock lizards are among the lowest in lacertids, and are also very narrow, maintaining the narrowness among seasons. As for summer (27.90–29.70 °C, mean value =28.76 °C), the thermal preferences of I. galani are also low in spring (29.60–31.10 °C, mean value =30.38 °C), supporting the idea that this species is adapted to cold environments. The habitat thermal quality is lower in spring (10.99 °C) than in summer (9.36 °C), while the effectiveness of thermoregulation is higher in spring (0.92) than in summer (0.80). We found that the seasonal shift in thermal preferences contributes significantly to enhance the effectiveness of thermoregulation in both seasons, more in spring (0.45 °C) than in summer (0.16 °C). Because I. galani inhabits isolated mountains, where the activity period is reduced from April to October, we hypothesize that the observed adaptation of the thermal preferences, which enhance thermoregulation to a larger extent in spring, may evolved to maximize performance during the reproductive season.

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Giardia duodenalis is a zoonotic parasite that infects the gut of a wide range of vertebrates, including numerous wildlife species. However, little is known about this protozoan parasite in reptiles. Fecal samples from 31 wild lizards were collected in Galicia (northwest Spain) and screened for the presence of Giardia by PCR amplification and sequencing of the ITS1-5.8S-ITS2 region in the ribosomal unit. This allowed detection of the parasite in 5 samples (16.1%), and enabled identification of G. duodenalis assemblage A2 in two samples of Iberian rock lizard (Iberolacerta monticola), G. duodenalis assemblage B in other two samples of I. monticola, and G. duodenalis assemblage E in one sample of Bocage`s wall lizard (Podarcis bocagei). The results obtained after PCR amplification and sequencing of the SSU-rDNA gene confirmed the presence of G. duodenalis assemblage A in two samples of I. monticola. This is the first report of G. duodenalis in free-living lizards, although further studies are needed to distinguish between actual infection and mechanical dissemination of cysts. The detection of zoonotic and livestock-specific assemblages of G. duodenalis demonstrates the wide environmental contamination by this parasite, possibly due to human activities.

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Fourteen polymorphic microsatellite loci are described for the Iberian rock lizard, Iberol- acerta monticola. Genetic variation in a sample of 20 individuals from Piornedo (north- western Spain) was quantified both by the number of alleles per locus, which ranged from six to 13, and by the expected frequency of heterozygotes under random mating (heterozy- gosity), which ranged from 0.761 to 0.902. Single locus and global exclusion probabilities were also computed, and indicate a high power of these markers for paternity assignments and mating system studies of I. monticola. All the analysed loci were also polymorphic in Iberolacerta galani, but only seven in Zootoca vivipara.

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Satellite DNAs compose a large portion of all higher eukaryotic genomes. The turnover of these highly repetitive sequences is an important element in genome organization and evolution. However, information about the structure and dynamics of reptilian satellite DNA is still scarce. Two satellite DNA families, HindIII and TaqI, have been previously characterized in four species of the genus Iberolacerta. These families showed different chromosomal locations, abundances, and evolutionary rates. Here, we extend the study of both satellite DNAs (satDNAs) to the remaining Iberolacerta species, with the aim to investigate the patterns of variability and factors influencing the evolution of these repetitive sequences. Our results revealed disparate patterns but also common traits in the evolutionary histories of these satellite families: (i) each satellite DNA is made up of a library of monomer variants or subfamilies shared by related species; (ii) species-specific profiles of satellite repeats are shaped by expansions and/or contractions of different variants from the library; (iii) different turnover rates, even among closely related species, result in great differences in overall sequence homogeneity and in concerted or non-concerted evolution patterns, which may not reflect the phylogenetic relationships among taxa. Contrasting turnover rates are possibly related to genomic constraints such as karyotype architecture and the interspersed organization of diverging repeat variants in satellite arrays. Moreover, rapid changes in copy number, especially in the centromeric HindIII satDNA, may have been associated with chromosomal rearrangements and even contributed to speciation within Iberolacerta.

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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.

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