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Among vertebrates, obligate parthenogenesis occurs exclusively in squamate reptiles. Premeiotic endoreplication in a small subset of developing oocytes has been documented as the mechanism of production of unreduced eggs in minutely explored obligate parthenogenetic lineages, namely in teiids and geckos. The situation in the lacertid genus Darevskia has been discussed for decades. Certain observations suggested that the ploidy level is restored during egg formation through a fusion of egg and polar body nuclei in Darevskia unisexualis and D. armeniaca. In this study, we re-evaluated the fusion hypothesis by studying diplotene chromosomes in adult females of sexual species D. raddei nairensis and obligate parthenogens D. armeniaca, D. dahli and D. unisexualis. We revealed 19 bivalents in the sexual species and 38 bivalents in the diploid obligate parthenogens, which uncovers premeiotic endoreplication as the mechanism of the production of non-reduced eggs in parthenogenetic females. The earlier contradicting reports can likely be attributed to the difficulty in identifying mispairing of chromosomes in pachytene, and the fact that in parthenogenetic reptiles relying on premeiotic endoreplication only a small subset of developing oocytes undergo genome doubling and overcome the pachytene checkpoint. This study highlights co-option of premeiotic endoreplication for escape from sexual reproduction in all independent hybrid origins of obligate parthenogenesis in vertebrates studied to date.

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Female and male hosts may maximise their fitness by evolving different strategies to compensate for the costs of parasite infections. The resulting sexual dimorphism might be apparent in differential relationships between parasite load and body condition, potentially reflecting differences in energy allocation to anti-parasitic defences. For example, male lacertids with high body condition may produce many offspring while being intensely parasitised. In contrast, female lacertids may show a different outcome of the trade-offs between body condition and immunity, aiming to better protect themselves from the harm of parasites. We predicted that females would have fewer parasites than males and a lower body condition across parasitaemia levels because they would invest resources in parasite defence to mitigate the costs of infection. In contrast, the male strategy to maximise access to females would imply some level of parasite tolerance and, thus, higher parasitaemia. We analysed the relationship between the body condition of lizards and the parasitemias of Karyolysus and Schellackia, two genera of blood parasites with different phylogenetic origins, in 565 females and 899 males belonging to 10 species of the Lacertidae (Squamata). These lizards were sampled over a period of 12 years across 34 sampling sites in southwestern Europe. The results concerning the Karyolysus infections were consistent with the predictions, with males having similar body condition across parasitaemia levels even though they had higher infection intensities than females. On the other hand, females with higher levels of Karyolysus parasitaemia had lower body condition. This is consistent with the prediction that different life strategies of male and female lacertids can explain the infection patterns of Karyolysus. In contrast, the parasitaemia of Schellackia was consistently low in both male and female hosts, with no significant effect on the body condition of lizards. This suggests that lizards of both sexes maintain this parasite below a pathogenic threshold.

Los machos y hembras pueden maximizar su eficacia biológica mediante la evolución diferencial de estrategias que compensen los costes asociados con las infecciones parasitarias. Por ejemplo, los machos con una alta condición corporal pueden producir muchas crías aun estando altamente parasitados. Mientras que es común que las hembras inviertan más energía en protegerse frente a los parásitos, lo que podría comprometer el mantenimiento de su condición corporal. Nuestra hipótesis es que las diferencias sexuales en la asignación de energía pueden quedar de manifiesto al analizar la relación entre la intensidad de infección con la condición corporal. También esperamos que las hembras tengan menos parásitos que los machos y que su condición corporal esté correlacionada negativamente con su carga parasitaria. Por el contrario, esperamos que los machos tengan más parásitos en parte porque su estrategia implicaría cierto nivel de tolerancia si con ello pueden incrementar su inversión reproductiva. Hemos analizado la relación entre la condición corporal y las parasitemias de Schellackia y Karyolysus, dos parásitos sanguíneos, en 565 hembras y 899 machos de 10 especies de la familia Lacertidae (Squamata) durante 12 años en 34 localidades en el suroeste de Europa. El patrón para Karyolysus fue coherente con las predicciones, ya que la condición corporal de los machos no se correlacionó con su parasitemia, a pesar de que estas fueron mayores que en las hembras. Mientras que la relación fue negativa en las hembras. Por lo que el patrón de infección de Karyolysus concuerda con las diferentes estrategias de inversión energética en machos y hembras. Sin embargo, en el caso de Schellackia las parasitemias fueron más bajas en hospedadores de ambos sexos y su parasitemia no se correlacionó con la condición corporal. Este resultado sugiere que Schellackia es mantenido en umbrales subpatogénicos.

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During our collecting trip to Guangxi in 2016, we collected ten specimens of the genus Takydromus from the suburb of Guilin, northeastern Guangxi, South China, and found that they did not belong to any currently known species. Here, we described this new species, Takydromus guilinensis sp. nov., based on morphological and mitochondrial DNA (CO1 and cyt b) data. This new species is a sister taxon to T. intermedius with a p-distance of 0.070 in CO1 and 0.080 in cyt b. These two p-distances exceed not only the minimum value (0.067) between T. septentrionalis and T. stejnegeri but also the minimum value (0.079) between T. intermedius and T. yunkaiensis. Morphologically, this new species differs from other currently recognized Takydromus species from the same clade, more evidently in the longitudinal rows of dorsal scales, transverse rows of scales at the mid-body and mensural variables. The description of Takydromus guilinensis sp. nov. increases the total number of Takydromus species to 25, of which 16 can be found in China. Takydromus guilinensis sp. nov. is currently known only from Guilin, Guangxi, South China, where it is sympatric with the other four Takydromus species (T. septentrionalis, T. kuehnei, T. sexlineatus and T. intermedius).

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We designed a common garden design to collect data on female reproductive traits from three populations of the southern grass lizard Takydromus sexlineatus, testing the hypothesis that a species-specific pattern of seasonal shifts in reproductive allocation should be shared by geographically separated populations. Of the seven examined traits, six differed among populations, with four of the six also differing among successive clutches. Females grew longer during the breeding season and produced more eggs in the first clutch than in the subsequent clutches; egg size was unchanged throughout the breeding season. After removing the influence of female size or postpartum body mass we found the following. First, postpartum body mass, clutch mass, and relative clutch mass were greater in the Wuzhishan population than in the Shaoguan and Zhaoqing populations. Second, egg size was greatest in the Wuzhishan population and smallest in the Zhaoqing population. Third, clutch size was greatest in the Wuzhishan population and smallest in the Shaoguan population. Females did not trade-off egg size against number within each population × clutch combination. Our study validates the hypothesis tested, supports the conventional view that reproductive output is highly linked to maternal body size in lizards, and follows the classic prediction that females with different amounts of resources to invest in reproduction should give priority to adjusting the total number rather than size of their offspring.

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The mitochondrion, which is an intracellular organelle responsible for most of the energy-producing pathways, can have its genome targeted for climate-driven selection. However, climate-driven mitochondrial selection remains a sparsely studied area in reptiles. Here, we reported the complete mitochondrial genome sequence of a lacertid lizard (Takydromus intermedius) and used mitogenomes from 54 species of lacertid lizards to study their phylogenetic relationships and to identify the mitochondrial genes under positive selection by climate. The length of the complete mitochondrial genome sequence of T. intermedius was 17,713 bp, which was within the range of lengths (17,224-18,943) ever reported for Takydromus species. The arrangement of mitochondrial genes in T. intermedius was the same as in other congeneric species. The 54 lacertid species could be divided into three geographically and climatically different clades. We identified three mitochondrial genes (ATP6, ATP8, and ND3) under positive selection by climate, and found that isothermality, temperature seasonality, precipitation of wettest month, and precipitation seasonality were the most important climatic variables contributing to the gene selection.

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Ectotherms are vastly affected by climatic conditions as they rely on external sources of heat to regulate their body temperature, and changes in their habitat thermal quality could seriously affect their overall biology. To overcome the problems of a thermally unfavorable habitat, lizards need to either adjust their thermoregulatory behavior or respond to directional selection and shift their preferred body temperatures. To assess the impact of habitat thermal quality on the thermoregulatory profile, we studied multiple islet and 'mainland' populations of the Skyros wall lizard Podarcis gaigeae, an endemic lacertid to Skyros Archipelago, Greece. We evaluated the effectiveness of thermoregulation (E) using the three main thermal parameters: body (Tb), operative (Te), and preferred (Tpref) temperatures. We first hypothesized that the spatial homogeneity, the scarcity of thermal shelters, and the exposure to higher winds on islets would result in more demanding climate conditions. Second, we anticipated that islet lizards would achieve higher E in response to the lower thermal quality therein. As hypothesized, thermal parameters differed between populations but not in the expected manner. Skyros 'mainland' habitats reached higher temperatures, had more intense fluctuations, and were of lower thermal quality. As a result, lizards showed higher accuracy, precision, and effectiveness of thermoregulation. Noteworthy, we found that lizards from different populations have shifted their thermal profile and preferred body temperatures to cope with the particular conditions prevailing in their habitats. The latter supports the labile view on the evolution of thermoregulation.

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Among the extant populations of the critically endangered Aeolian wall lizard, the most vulnerable is the one surviving on La Canna, a columnar volcanic stack off the Filicudi Island. Here, I report the results of the first climbing expedition by a biologist on La Canna, that contributed direct observations and updated information on the size, morphology, and genetic variability of this population. Lizard density at the sampling site (a small terrace at 50 m of elevation) was 1.7 m-2, twice of a previous estimate. Standard methods for estimating population size are unsuitable for La Canna. An educated guess of about a hundred individuals can be drawn, considering the extent of habitat available on the stack and the number of observed lizards. Lizards on La Canna were not fearless, despite what was reported by alpinists, possibly because of aggressive intraspecific interactions or high environmental temperatures during sampling. Biometric data significantly extend the body size of La Canna's lizards and indicate that it is not smaller than other P. raffonei populations. A complete lack of genetic diversity was found at the mitochondrial nd4 gene, in line with previous allozyme data and with estimates on other microinsular Podarcis populations. The small size of the La Canna population implies severe genetic drift and an extremely high level of inbreeding, as supported by low heterozygosity found across the genome. Detrimental effects of inbreeding depression are evident as cephalic malformations observed in all captured lizards of La Canna and might represent the more immediate threat to the persistence of this population.

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In ecosystems threatened by the expansion of croplands, habitat fragmentation and climate change, two of the main extinction drivers, may have thermoregulation-mediated interacting effects on demographic trends of terrestrial ectotherms. We studied the thermal biology of a metapopulation of the widespread Mediterranean lacertid Psammodromus algirus in ten fragments of evergreen or deciduous oak forests interspersed among cereal fields. We obtained thermoregulation statistics (selected temperature range, body and operative temperatures, thermal quality of the habitat, and precision, accuracy, and effectiveness of thermoregulation) that could be compared among fragments and with conspecific populations living in unfragmented habitat. We also measured the selection (use vs. availability) and spatial distribution of sunlit and shaded patches used for behavioral thermoregulation in fragments, and we estimated operative temperatures and thermal habitat quality in the agricultural matrix surrounding the fragments. Variation of the thermal environment was much larger within fragments than among them, and thermoregulation was accurate, precise, and efficient throughout the fragmented landscape; its effectiveness was similar to that of previously studied unfragmented populations. The average distance between sunlit and shaded patches was shorter in deciduous than in evergreen fragments, producing a more clumped distribution of the mosaic of thermal resources. Consequently, in evergreen habitat the cost of thermoregulation was higher, because lizards were more selective in their choice of sunlit sites (i.e. they used sunlit patches closer to shade and refuge than expected at random, and the extent of such selection was larger than at deciduous habitat). Temperatures available in croplands were too high to allow lizard dispersal, at least in the post-breeding season. This result confirms the role of croplands as a thermal barrier that promotes inbreeding and associated fitness losses in isolated fragments, and it forecasts a dark future for populations of forest lizards in agricultural landscapes under the combined effects of habitat fragmentation and global warming.

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In organisms reproducing sexually, speciation occurs when increasing divergence results in pre- or post-zygotic reproductive isolation between lineages. Studies focusing on reproductive isolation origin in early stages of speciation are common and many rely on genomic scans to infer introgression providing limited information on the genomic architecture of reproductive isolation long-term maintenance. This study analyses a natural hybrid zone between two species in a late stage of speciation. We used ddRADseq genotyping in the contact between Podarcis bocagei and P. carbonelli to examine admixture extent, analyse hybrid zone stability and assess genome-wide variation in selection against introgression. We confirmed strong but incomplete reproductive isolation in a bimodal hybrid zone. New findings revealed population genetic structure within P. carbonelli in the contact zone; geographical and genomic clines analysis suggested strong selection against gene flow, but a relatively small proportion of the loci can introgress, mostly within the narrow contact zone. However, geographical clines revealed that a few introgressed loci show signs of potential positive selection, particularly into P. bocagei. Geographical clines also detected a signal of hybrid zone movement towards P. bocagei distribution. Genomic cline analysis revealed heterogeneous patterns of introgression among loci within the syntopy zone, but the majority maintain a strong association with the genomic background of origin. However, incongruences between both cline approaches were found, potentially driven by confounding effects on genomic clines. Last, an important role of the Z chromosome in reproductive isolation is suggested. Importantly, overall patterns of restricted introgression seem to result from numerous strong intrinsic barriers across the genome.

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During the last 24 years, the mutualistic interaction between the dead horse arum, Helicodiceros muscivorus, and the Balearic lizard, Podarcis lilfordi, was studied on Aire Island (Balearic Islands, Spain). From a small population of a hundred plants, the dead horse arum expanded extraordinarily throughout the island, reaching the highest known densities of the species and occupying areas of the island where it was not previously present. The current abundance of plants is a direct effect of the frugivorous activity of the Balearic lizard, which is the main, if not the only, effective seed disperser of the plant on Aire Island. However, abiotic factors predominated over biotic factors in driving abundance of plants. Over the years, plant densities varied significantly depending on the aridity of the island, with higher densities recorded in drier years. Lizards' frugivorous activity and dispersal intensity was inversely correlated with annual rainfall. We found higher dispersal intensity in years with lower rainfall. We propose that the years of lower rainfall are those in which there is a lower prey availability. In such years, lizards compensate the shortage of other trophic resources with a more intense consumption of dead horse arum fruits. The mutualistic interaction is therefore asymmetric, since there is a greater influence of the frugivorous activity of the lizards on the plants than of the plants on lizards. It is, in short, a system chronically out of balance.

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The genus Karyolysus was originally proposed to accommodate blood parasites of lacertid lizards in Western Europe. However, recent phylogenetic analyses suggested an inconclusive taxonomic position of these parasites of the order Adeleorina based on the available genetic information. Inconsistencies between molecular phylogeny, morphology, and/or life cycles can reflect lack of enough genetic information of the target group. We therefore surveyed 28 localities and collected blood samples from 828 lizards of 23 species including lacertids, skinks, and geckoes in the western Mediterranean, North Africa, and Macaronesia, where species of Karyolysus and other adeleorine parasites have been described. We combined molecular and microscopic methods to analyze the samples, including those from the host type species and the type locality of Karyolysus bicapsulatus. The phylogenetic relationship of these parasites was analyzed based on the 18S rRNA gene and the co-phylogenetic relationship with their vertebrate hosts was reconstructed. We molecularly detected adeleorine parasites in 37.9% of the blood samples and found 22 new parasite haplotypes. A phylogenetic reconstruction with 132 sequences indicated that 20 of the newly detected haplotypes clustered in a well-supported clade with another 18 sequences that included Karyolysus galloti and Karyolysus lacazei. Morphological evidence also supported that K. bicapsulatus clustered in this monophyletic clade. These results supported the taxonomic validity of the genus. In addition, we found some parasite haplotypes that infected different lizard host genera with ancient diverging histories, which suggested that Karyolysus is less host-specific than other blood parasites of lizards in the region. A co-phylogenetic analysis supported this interpretation because no significant co-speciation signal was shown between Karyolysus and lizard hosts.

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The diets of insular lizards are extremely varied, depending on the different environmental characteristics of each island population. This is particularly evident in the case of the populations of small coastal islets of the Balearic Islands, where the Balearic lizard, Podarcis lilfordi, is found. The study of trophic ecology carried out by means of traditional tools, such as morphological analysis of feces, has made it possible to detect numerous prey and nutritional elements. However, these methods are clearly insufficient, as some rare groups are not detected. It is also difficult to identify remains of marine subsidies or of foods contributed to these small islands by other predators, such as seabirds. The current study demonstrates the advantages of combining morphological diet analysis with the molecular study of individual feces samples obtained from the same populations. We obtained a greater diversity of prey groups using the combined methodologies, with each method identifying prey items that were not detected using the other method. Particularly, the study of diets at the molecular level identified plant species consumed by lizards that were, occasionally, not identified in morphological analyses. Conversely, the traditional morphological study of an equivalent number of fecal samples allowed for the identification of several prey groups that had not been detected in the molecular study. From this viewpoint, the advantages and disadvantages of each methodology are discussed.

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How does ecology influence cognitive evolution in lizards? Taking a comparative approach, De Meester et al. (2022) discovered that species living in temporally fluctuating environments tend to perform relatively poorly on cognitive tasks associated with behavioral flexibility compared to species living in more climatically stable environments. The negative association between environmental variability and cognitive performance suggests that stochastic environments can hamper, rather than stimulate, the evolution of cognitive ability.

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The extant reptiles are one of the most diverse clades among terrestrial vertebrates and one of a few groups with instances of parthenogenesis. Due to the hybrid origin of parthenogenetic species, reference genomes of the parental species as well as of the parthenogenetic progeny are indispensable to explore the genetic foundations of parthenogenetic reproduction. Here, we report on the first genome assembly of rock lizard Darevskia valentini, a paternal species for several parthenogenetic lineages. The novel genome was used in the reconstruction of the comprehensive phylogeny of Squamata inferred independently from 7369 trees of single-copy orthologs and a supermatrix of 378 conserved proteins. We also investigated Hox clusters, the loci that are often regarded as playing an important role in the speciation of animal groups with drastically diverse morphology. We demonstrated that Hox clusters of D. valentini are invaded with transposons and contain the HoxC1 gene that has been considered to be lost in the amniote ancestor. This study provides confirmation for previous works and releases new genomic data that will contribute to future discoveries on the mechanisms of parthenogenesis as well as support comparative studies among reptiles.

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Cognition is an essential tool for animals to deal with environmental challenges. Nonetheless, the ecological forces driving the evolution of cognition throughout the animal kingdom remain enigmatic. Large-scale comparative studies on multiple species and cognitive traits have been advanced as the best way to facilitate our understanding of cognitive evolution, but such studies are rare. Here, we tested 13 species of lacertid lizards (Reptilia: Lacertidae) using a battery of cognitive tests measuring inhibitory control, problem-solving, and spatial and reversal learning. Next, we tested the relationship between species' performance and (a) resource availability (temperature and precipitation), habitat complexity (Normalized Difference Vegetation Index), and habitat variability (seasonality) in their natural habitat and (b) their life history (size at hatching and maturity, clutch size, and frequency). Although species differed markedly in their cognitive abilities, such variation was mostly unrelated to their ecology and life history. Yet, species living in more variable environments exhibited lower behavioral flexibility, likely due to energetic constrains in such habitats. Our standardized protocols provide opportunities for collaborative research, allowing increased sample sizes and replication, essential for moving forward in the field of comparative cognition. Follow-up studies could include more detailed measures of habitat structure and look at other potential selective drivers such as predation.

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Morocco has a great diversity of reptiles; more than 26% of the herpetofauna species are considered to be endemic. Nevertheless, there is little information available on helminth parasites of Moroccan lizards. The purpose of this article is to establish a helminth list using non-invasive methods for natural populations of seven lizard species: Acanthodactylus maculatus, Chalcides mionecton, Chalcides montanus, Chalcides polylepis, Quedenfeldtia moerens, Quedenfeldtia trachyblepharus, and Tarentola mauritanica. For each species, prevalence and intensity of infection were given as well as their relationship to some determinant parameters (sex, age class, size, altitude, and seasons). Four species of nematodes, Parapharyngodon micipsae, Pharyngodon mamillatus, Spauligodon auziensis, Thelandros alatus, and unassigned Spauligodon were found. Mean helminth intensity for the seven lizard species was 7.3 ± 16.2 SD (range = 1-92 in infected lizards), with total infection prevalence of 5.6%. Mixed infections were observed in Chalcides montanus and Chalcides polylepis co-infected by Pharyngodon mamillatus and Spauligodon sp. Furthermore, the intensity of infection by nematodes were significantly different among the seven studied species. The nematode infection in Chalcides mionecton were related to sex, host size, and altitude; in Chalcides polylepis to the host age; in Quedenfeldtia trachyblepharus to elevation, age, and host size; and in Tarentola Mauritanica to age, body size, seasons, and altitude. In conclusion, our study provides data about nine new host and locality records and the effect of some determinant factors on host parasitism.

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The Kokshaal racerunner, Eremias kokshaaliensis Eremchenko et Panfilov, 1999, together with other central Asian racerunner species, is included in the Eremias multiocellata complex. In the present work, for the first time, the results of the analysis of historical mitochondrial DNA (barcode) are presented and the taxonomic status and preliminary phylogenetic relationships within the complex are specified. We present, for the first time, the results of the molecular analysis using historical DNA recovered from specimens of several species of this complex (paratypes of the Kokshaal racerunner and historical collections of the Kashgar racerunner E. buechneri from Kashgaria) using DNA barcoding.

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Studies which quantify the influence of abiotic factors on physiological variation are paramount to comprehend organismal responses to diverse environments. We studied three physiological aspects of metabolism in two sympatric and ecologically similar European lizard species, Podarcis muralis and Iberolacerta horvathi, across an 830-m elevational gradient. We collected blood samples and tail tips from adult lizards, which were analyzed for parasitemia, hemoglobin concentration, potential metabolic activity and catalase activity. Hemoglobin concentration was higher in males than females and it increased across elevation in one of the studied species - P. muralis. Parasitemia was not an important predictor of the variation in hemoglobin concentration, which suggests that blood parasites do not constraint the aerobic capacity of the lizards. On the other hand, catalase activity reflected increased antioxidant activity in the presence of higher parasitemia, possibly acting as an adaptive mechanism to reduce oxidative stress during immune activation. Potential metabolic activity, as a proxy for maximum respiratory enzymatic capacity, did not differ between species or sexes nor was it affected by elevation or levels of parasitemia. The results provide insight into the relationships between physiological, biotic, and environmental traits in sympatric lizards.

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Comparative studies on mitochondrial genomes (mitogenomes) as well as the structure and evolution of the mitochondrial control region are few in the Lacertidae family. Here, the complete mitogenomes of five individuals of Eremias scripta (2 individuals), Eremias nikolskii, Eremias szczerbaki, and Eremias yarkandensis were determined using next-generation sequencing and were compared with other lacertids available in GenBank. The circular mitogenomes comprised the standard set of 13 protein-coding genes (PCGs), 22 transfer RNA genes, 2 ribosomal RNA genes and a long non-coding control region (CR). The extent of purifying selection was less pronounced for the COIII and ND2 genes in comparison with the rest of the PCGs. The codons encoding Leucine (CUN), Threonine, and Isolecucine were the three most frequently present. The secondary structure of rRNA of Lacertidae (herein, E. scripta KZL15 as an example) comprised four domains and 28 helices for 12S rRNA, with six domains and 50 helices for 16S rRNA. Five types and twenty-one subtypes of CR in Lacertidae were described by following the criteria of the presence and position of tandem repeats (TR), termination-associated sequence 1 (TAS1), termination-associated sequence 2 (TAS2), conserved sequence block 1 (CBS1), conserved sequence block 2 (CSB2), and conserved sequence block 3 (CSB3). The compositions of conserved structural elements in four genera, Acanthodactylus, Darevskia, Eremias, and Takydromus, were further explored in detail. The base composition of TAS2 - TATACATTAT in Lacertidae was updated. In addition, the motif "TAGCGGCTTTTTTG" of tandem repeats in Eremias and the motif "GCGGCTT" in Takydromus were presented. Nucleotide lengths between CSB2 and CSB3 remained 35 bp in Eremias and Darevskia. The phylogenetic analyses of Lacertidae recovered the higher-level relationships among the three subfamilies and corroborated a hard polytomy in the Lacertinae phylogeny. The phylogenetic position of E. nikolskii challenged the monophyly of the subgenus Pareremias within Eremias. Some mismatches between the types of CR and their phylogeny demonstrated the complicated evolutionary signals of CR such as convergent evolution. These findings will promote research on the structure and evolution of the CR and highlight the need for more mitogenomes in Lacertidae.

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Proteins are ubiquitous macromolecules that display a vast repertoire of chemical and enzymatic functions, making them suitable candidates for chemosignals, used in intraspecific communication. Proteins are present in the skin gland secretions of vertebrates but their identity, and especially, their functions, remain largely unknown. Many lizard species possess femoral glands, i.e., epidermal organs primarily involved in the production and secretion of chemosignals, playing a pivotal role in mate choice and intrasexual communication. The lipophilic fraction of femoral glands has been well studied in lizards. In contrast, proteins have been the focus of only a handful of investigations. Here, we identify and describe inter-individual expression patterns and the functionality of proteins present in femoral glands of male sand lizards (Lacerta agilis) by applying mass spectrometry-based proteomics. Our results show that the total number of proteins varied substantially among individuals. None of the identified femoral gland proteins could be directly linked to chemical communication in lizards, although this result hinges on protein annotation in databases in which squamate semiochemicals are poorly represented. In contrast to our expectations, the proteins consistently expressed across individuals were related to the immune system, antioxidant activity and lipid metabolism as their main functions, showing that proteins in reptilian epidermal glands may have other functions besides chemical communication. Interestingly, we found expression of the Major Histocompatibility Complex (MHC) among the multiple and diverse biological processes enriched in FGs, tentatively supporting a previous hypothesis that MHC was coopted for semiochemical function in sand lizards, specifically in mate recognition. Our study shows that mass spectrometry-based proteomics are a powerful tool for characterizing and deciphering the role of proteins secreted by skin glands in non-model vertebrates.

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