Lizard host abundances and climatic factors explain phylogenetic diversity and prevalence of blood parasites on an oceanic island.

Host abundance might favour the maintenance of a high phylogenetic diversity of some parasites via rapid transmission rates. Blood parasites of insular lizards represent a good model to test this hypothesis because these parasites can be particularly prevalent in islands and host lizards highly abundant. We applied deep amplicon sequencing and analysed environmental predictors of blood parasite prevalence and phylogenetic diversity in the endemic lizard Gallotia galloti across 24 localities on Tenerife, an island in the Canary archipelago that has experienced increasing warming and drought in recent years. Parasite prevalence assessed by microscopy was over 94%, and a higher proportion of infected lizards was found in warmer and drier locations. A total of 33 different 18s rRNA parasite haplotypes were identified, and the phylogenetic analyses indicated that they belong to two genera of Adeleorina (Apicomplexa: Coccidia), with Karyolysus as the dominant genus. The most important predictor of between-locality variation in parasite phylogenetic diversity was the abundance of lizard hosts. We conclude that a combination of climatic and host demographic factors associated with an insular syndrome may be favouring a rapid transmission of blood parasites among lizards on Tenerife, which may favour the maintenance of a high phylogenetic diversity of parasites.

Population Genomics of Wall Lizards Reflects the Dynamic History of the Mediterranean Basin.

The Mediterranean Basin has experienced extensive change in geology and climate over the past six million years. Yet, the relative importance of key geological events for the distribution and genetic structure of the Mediterranean fauna remains poorly understood. Here, we use population genomic and phylogenomic analyses to establish the evolutionary history and genetic structure of common wall lizards (Podarcis muralis). This species is particularly informative because, in contrast to other Mediterranean lizards, it is widespread across the Iberian, Italian, and Balkan Peninsulas, and in extra-Mediterranean regions. We found strong support for six major lineages within P. muralis, which were largely discordant with the phylogenetic relationship of mitochondrial DNA. The most recent common ancestor of extant P. muralis was likely distributed in the Italian Peninsula, and experienced an "Out-of-Italy" expansion following the Messinian salinity crisis (∼5 Mya), resulting in the differentiation into the extant lineages on the Iberian, Italian, and Balkan Peninsulas. Introgression analysis revealed that both inter- and intraspecific gene flows have been pervasive throughout the evolutionary history of P. muralis. For example, the Southern Italy lineage has a hybrid origin, formed through admixture between the Central Italy lineage and an ancient lineage that was the sister to all other P. muralis. More recent genetic differentiation is associated with the onset of the Quaternary glaciations, which influenced population dynamics and genetic diversity of contemporary lineages. These results demonstrate the pervasive role of Mediterranean geology and climate for the evolutionary history and population genetic structure of extant species.

No genomic repercussions of assortative pairing in a colour polymorphic lizard.

Long-term maintenance of colour polymorphisms often depends on the interplay of multiple selective forces. In the common wall lizard (Podarcis muralis), up to three pure and two mosaic ventral colour morphs co-exist across most of its range. Available evidence suggests that colour morphs in this species are maintained through the interaction between sexual and environment-dependent selection. In particular, colour-assortative pairing has been recorded, suggesting some degree of assortative mating. Here, we combined reduced-representation sequencing (ddRADseq) and fine-scale distribution data to explore the effects of assortative pairing on the common wall lizard. Overall, our results do not support any population structure (FST  = 0 and K = 1) nor a significant effect of colour morph or geographic location on genomic differentiation. Therefore, we argue that assortative pairing may not fully translate into assortative mating and genomic differentiation between colour morphs and discuss possible explanations. Nonetheless, we find potential support for an elevated population size and/or source-sink dynamics and debate the potential contribution of other forms of selection to the maintenance of colour polymorphisms in lacertids.

Time changes everything: a multispecies analyses of temporal patterns in evaporative water loss.

Increasing trends in environmental temperatures and changes in the hydrological cycle caused by climate change are affecting organisms around the world. Water balance is an important feature of organisms as warmer and drier environments increase water vapour pressure deficit and cutaneous water loss. The aim of our study was to understand the hourly kinetics of losses under water-restricted conditions. In general, these kinetics are thought to be negligible, and values from water loss studies have often been summed or averaged for population and species comparisons. We compiled a cross-species data set of instantaneous water loss (EWLi) of lizards in which weight was recorded every hour for 12 h (a period considered ecologically relevant for lizards) under standardized conditions with minimal disturbance. We statistically tested whether the duration of the experiment affected the values and found that the duration affected 83% of the data sets. In most cases, water loss was significantly higher during the first hour. This is likely a result of increased activity due to acclimation to the experimental conditions. However, as the experiment progressed, other temporal patterns of EWLi occurred that were not consistent between species. In addition, the average value and the first and last values of EWLi changed significantly for most species as the experiment progressed. These results suggest that the hourly kinetics of water loss should not be ignored, while experimental procedures in which EWL is recorded continuously rather than only at the beginning and end of the experiment should be used to find finer temporal variations.

Shared haemogregarine infections in competing lacertids.

In parasite­host interactions host species may differ in their ability to fight parasitic infections, while other ecological interactions, including competition, may differentially alter their physiological state, making them even more susceptible to parasites. In this study, we analyse the haemogregarine blood parasites infecting two competing lizard species, Iberolacerta horvathi and Podarcis muralis, and explore host­parasite relationships under different host competition scenarios. Both species were infected with haemogregarine parasites belonging to the genus Karyolysus. Using the 18S rRNA gene, six new Karyolysus haplotypes were identified clustering with other Central and Eastern European samples, and widely shared between both lizard hosts. Haemogregarine infections were detected at all sampled sites with over 50% of individuals parasitized. Overall, I. horvathi was more frequently and also more intensely parasitized than P. muralis, with higher infection rates observed in syntopy. Males of both species tended to be more frequently infected and showed a higher infection intensity than conspecific females. The results suggest that parasitisation by haemogregarines may be relevant in the dynamics of the competitive relationship between these lizard species. More studies, including immunological response analysis, and the identification of the vectors are needed to better understand host­parasite relationships and competition.

Regulatory changes in pterin and carotenoid genes underlie balanced color polymorphisms in the wall lizard.

Reptiles use pterin and carotenoid pigments to produce yellow, orange, and red colors. These conspicuous colors serve a diversity of signaling functions, but their molecular basis remains unresolved. Here, we show that the genomes of sympatric color morphs of the European common wall lizard (Podarcis muralis), which differ in orange and yellow pigmentation and in their ecology and behavior, are virtually undifferentiated. Genetic differences are restricted to two small regulatory regions near genes associated with pterin [sepiapterin reductase (SPR)] and carotenoid [beta-carotene oxygenase 2 (BCO2)] metabolism, demonstrating that a core gene in the housekeeping pathway of pterin biosynthesis has been coopted for bright coloration in reptiles and indicating that these loci exert pleiotropic effects on other aspects of physiology. Pigmentation differences are explained by extremely divergent alleles, and haplotype analysis revealed abundant transspecific allele sharing with other lacertids exhibiting color polymorphisms. The evolution of these conspicuous color ornaments is the result of ancient genetic variation and cross-species hybridization.

Translation and validation of the “personal evaluation of transitions in treatment (PETIT)” scale for people with schizophrenia.

To adapt the ‘Personal Evaluation of Transitions in Treatment (PETIT)’ scale into Spanish and analyse its psychometric properties on schizophrenic population.

An integrative analysis of the short-term effects of tail autotomy on thermoregulation and dehydration rates in wall lizards.

Maintaining body temperature is essential for the optimal performance of physiological functions. Ectotherms depend on external heat sources to thermoregulate. However, thermoregulation may be constrained by body condition and hydration state. Autotomy (i.e., the voluntary shed of a body part) evolved in various animal lineages and allowed surviving certain events (such as predator attacks), but it may affect body condition and volume/surface ratios, increase dehydration and constrain thermoregulation. In the framework of a general analysis of the evolution of autotomy, here we assessed the effects of tail loss on the thermal preferences and evaporative water loss rates (EWL) in the lizard Podarcis bocagei, integrating the thermal and hydric factors. We did not observe shifts in the thermal preferences of experimentally autotomized lizards when compared to the controls, which contradicted the hypothesis that they would raise preferred temperature to increase metabolic rates and accelerate regeneration. Evaporative water loss rates were also similar for tailed and tailless individuals, suggesting negligible increase of water loss through the injury and no specific ecophysiological responses after autotomy. Therefore, the changes observed in autotomized lizards in the field are to be considered primarily behavioural, rather than physiological, and thermoregulation could be secondarily affected by behavioural compensations for an increased predation risk after autotomy. Functional studies are necessary to understand how lizards' interaction with the environment is altered after autotomy, and further studies including different dehydration levels would be useful to fully understand the effect of water shortage on lizards' performance after caudal autotomy.

Evidence from Tarentola mauritanica (Gekkota: Phyllodactylidae) helps validate thermography as a tool to infer internal body temperatures of lizards.

Infrared (IR) thermal imaging has become an increasingly popular tool to measure body temperature of animals. The high-resolution data it provides with short lag and minimum disturbance makes it an appealing tool when studying reptile thermal ecology. However, due to the common phenomenon of regional heterothermy and surface-to-core temperature gradients, it is essential to select the appropriate body part to measure and provide calibrations to accurately infer internal body temperatures. This work follows from a previous study on lacertid lizards to assess the reliability of thermography-measured body temperatures, from several body locations, as a proxy for internal body temperature in lizards. This study focuses on the Moorish gecko, Tarentola mauritanica, due to its distant phylogenetic relationship and its different ecology and morphology from the previously tested species. A total of 60 adult geckos of both sexes and of a range of sizes were tested in thermal gradients and subjected to a sequence of randomly assorted treatments of heating and cooling. The temperatures of the animals were periodically measured with a thermal camera at six different body parts and, immediately after, the cloacal temperature was then measured with a thermocouple probe. Body parts' temperatures, obtained thermographically, were regressed against cloacal temperature using OLS regression and the pairwise correlations were tested using Spearman coefficients. Relationships among all body parts and between all body parts and the cloaca were strong in all cases (R2 > 0.87, Spearman Correlation > 0.95). The observed pattern was very similar to those previously obtained from lacertid lizards. Ultimately, the eye proved to provide the best overall proxy for internal temperature, when accounting for both the slope and intercept of the regression. Hence, this study provides further support for the establishment of the eye as the standard location to infer internal body temperatures of lizards through thermography.

Ecophysiology of a lacertid community in the high Moroccan mountains suggests conservation guidelines.

Lizard species may differ in their ecophysiology due to adaptation, plasticity and/or phylogeny. In restrictive environments, ecophysiological differences of species living in sympatry are expected to reveal long-term evolutionary responses to the abiotic environment while competitive interactions should be limited. These influences can be disentangled by combining field monitoring with experimental tests. Here, three lacertid lizard species, Atlantolacerta andreanskyi, Scelarcis perspicillata and Podarcis vaucheri sharing high mountain habitats in Oukaimeden (High Atlas, Morocco), were studied. In the field, spatiotemporal variation of the thermal and hydric environment used by the lizards was monitored using data-loggers. In the lab, thermal and hydric ecophysiology was estimated through assessments of preferred temperatures (Tp) and water loss (WL) rates. Species differed in microhabitat use and, hence, in their exposure to variations in temperature and humidity. However, they only differed in their WL (A. andreanskyi > S. perspicillata > P. vaucheri) while their Tp were similar. Such partial differences of species in in the fundamental niche, likely derived from their long-term independent phylogenetic trajectories, can be used to predict their responses to climate and habitat shifts in this and other parts of their respective ranges. Results also confirm previous suggestions that, together with thermal physiology, hydric physiology plays a prominent role in the organisation of lizard communities in the temperate region.

Habitat preference modulates trans-oceanic dispersal in a terrestrial vertebrate.

The importance of long-distance dispersal (LDD) in shaping geographical distributions has been debated since the nineteenth century. In terrestrial vertebrates, LDD events across large water bodies are considered highly improbable, but organismal traits affecting dispersal capacity are generally not taken into account. Here, we focus on a recent lizard radiation and combine a summary-coalescent species tree based on 1225 exons with a probabilistic model that links dispersal capacity to an evolving trait, to investigate whether ecological specialization has influenced the probability of trans-oceanic dispersal. Cryptoblepharus species that occur in coastal habitats have on average dispersed 13 to 14 times more frequently than non-coastal species and coastal specialization has, therefore, led to an extraordinarily widespread distribution that includes multiple continents and distant island archipelagoes. Furthermore, their presence across the Pacific substantially predates the age of human colonization and we can explicitly reject the possibility that these patterns are solely shaped by human-mediated dispersal. Overall, by combining new analytical methods with a comprehensive phylogenomic dataset, we use a quantitative framework to show how coastal specialization can influence dispersal capacity and eventually shape geographical distributions at a macroevolutionary scale.

Correction to: The role of hybridisation in the origin and evolutionary persistence of vertebrate parthenogens: a case study of Darevskia lizards.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

The role of hybridisation in the origin and evolutionary persistence of vertebrate parthenogens: a case study of Darevskia lizards.

Obligate parthenogenesis is found in only 0.1% of the vertebrate species, is thought to be relatively short lived and is typically of hybrid origin. However, neither the evolutionary persistence of asexuality in vertebrates, nor the conditions that allow the generation of new parthenogenetic lineages are currently well understood. It has been proposed that vertebrate parthenogenetic lineages arise from hybridisation between two divergent taxa within a specific range of phylogenetic distances (the 'Balance Hypothesis'). Moreover, parthenogenetic species often maintain a certain level of hybridisation with their closest sexual relatives, potentially generating new polyploid hybrid lineages. Here we address the role of hybridisation in the origin and evolutionary lifespan of vertebrate parthenogens. We use a set of microsatellite markers to characterise the origins of parthenogens in the lizard genus Darevskia, to study the distinctiveness of sexual and asexual taxa currently in sympatry, and to analyse the evolutionary consequences of interspecific hybridisation between asexual females and sexual males. We find that parthenogens result from multiple past hybridisation events between species from specific lineages over a range of phylogenetic distances. This suggests that the Balance Hypothesis needs to allow for lineage-specific effects, as envisaged in the Phylogenetic Constraint Hypothesis. Our results show recurrent backcrossing between sexual and parthenogenic Darevskia but neither gene flow nor formation of new asexual lineages. We suggest that, along with their demographic advantage, parthenogens gain additional leverage to outcompete sexuals in nature when the retention of sexual reproductive machinery allows backcrossing with their sexual ancestors.

A paradise for parasites? Seven new haemogregarine species infecting lizards from the Canary Islands.

Oceanic islands are hotspots of biodiversity due to their high levels of endemism, with the Canary Islands being a notable example. A previous molecular study on the biogeography and host associations of haemogregarines (Apicomplexa: Adeleorina) infecting lizards from this archipelago detected seven parasite haplogroups. These haplogroups exhibited high host-specificity and geographical structure, suggesting that they might correspond to distinct biological identities. In this study, along with sequencing a longer fragment of the 18S rRNA, we further explore the distinctiveness of these parasites by analysing their morphology, effects on host erythrocytes and parasitaemia levels. These lines of evidence together with their genetics, host associations, frequency of occurrence and geographical distribution support them as different biological entities. As such, we describe seven new species: Karyolysus canariensis sp. nov., Karyolysus galloti sp. nov., Karyolysus stehlini sp. nov., Karyolysus gomerensis sp. nov., Karyolysus atlanticus sp. nov., Karyolysus tinerfensis sp. nov. and Karyolysus makariogeckonis sp. nov. These new taxa are further examples of endemic diversity in the Canarian archipelago. They also contribute to clarify the taxonomy within the Apicomplexa, a phylum estimated to have one of the lowest percentages of described species.

Waitin' on a sunny day: Factors affecting lizard body temperature while hiding from predators.

Most animals face predators in their daily life and have evolved antipredator strategies that promote survival while minimizing escaping costs. For example, many animals often hide into refuges when chased. Ectotherms rely on external sources of heat to raise their body temperature, and thermoregulate to keep their body temperature close to the optimal for performance. For many ectotherms living in temperate areas, it can be expected that they pay a cost in terms of heat loss while staying hidden. Indeed, refuges are often more thermally unsuitable than the external environment. Hence, the aim of this study was to assess if and to what extent hiding may result in a decrease of body temperature in a temperate lizard. We used infrared technology to measure the body temperature of a large-sized lizard (Timon lepidus) before individuals escaped from a simulated predation attempt to hide inside a refuge, and after they emerged back from the refuge. We quantified the change of body temperature that lizards experienced while hiding. Results show that while the decrease in body temperature covaried with the time spent hidden, it was also affected by the initial body temperature. Our key finding is that the time spent hidden depends mostly on the temperature inside the refuge. Indeed, lizards hiding in warmer refuges spent more time hidden, likely benefitting from a reduced cooling rate. This suggests that lizards perceive and evaluate the thermal quality of their refuges and integrate this information to react to predation attempts and minimize the potential thermal consequences of hiding.

Getting there and around: Host range oscillations during colonization of the Canary Islands by the parasitic nematode Spauligodon.

Episodes of expansion and isolation in geographic range over space and time, during which parasites have the opportunity to expand their host range, are linked to the development of host-parasite mosaic assemblages and parasite diversification. In this study, we investigated whether island colonization events lead to host range oscillations in a taxon of host-specific parasitic nematodes of the genus Spauligodon in the Canary Islands. We further investigated whether range oscillations also resulted in shifts in host breadth (i.e., specialization), as expected for parasites on islands. Parasite phylogeny and divergence time estimates were inferred from molecular data with Bayesian methods. Host divergence times were set as calibration priors after a priori evaluation with a global-fit method of which individual host-parasite associations likely represent cospeciation links. Parasite colonization history was reconstructed, followed by an estimation of oscillation events and specificity level. The results indicate the presence of four Spauligodon clades in the Canary Islands, which originated from at least three different colonization events. We found evidence of host range oscillations to truly novel hosts, which in one case led to higher diversification. Contemporary host-parasite associations show strong host specificity, suggesting that changes in host breadth were limited to the shift period. Lineages with more frequent and wider taxonomic host range oscillations prior to the initial colonization event showed wider range oscillations during colonization and diversification within the archipelago. Our results suggest that a lineage's evolutionary past may be the best indicator of a parasite's potential for future range expansions.

Run for your life, but bite for your rights? How interactions between natural and sexual selection shape functional morphology across habitats.

A central issue in evolutionary biology is how morphology, performance, and habitat use coevolve. If morphological variation is tightly associated with habitat use, then differences in morphology should affect fitness through their effect on performance within specific habitats. In this study, we investigate how evolutionary forces mold morphological traits and performance differently given the surrounding environment, at the intraspecific level. For this purpose, we selected populations of the lizard Podarcis bocagei from two different habitat types, agricultural walls and dunes, which we expected to reflect saxicolous vs ground-dwelling habits. In the laboratory, we recorded morphological traits as well as performance traits by measuring sprint speed, climbing capacity, maneuverability, and bite force. Our results revealed fast-evolving ecomorphological variation among populations of P. bocagei, where a direct association existed between head morphology and bite performance. However, we could not establish links between limb morphology and locomotor performance at the individual level. Lizards from walls were better climbers than those from dunes, suggesting a very fast evolutionary response. Interestingly, a significant interaction between habitat and sex was detected in climbing performance. In addition, lizards from dunes bit harder than those from walls, although sexual differentiation was definitely the main factor driving variation in head functional morphology. Taking into account all the results, we found a complex interaction between natural and sexual selection on whole-organism performance, which are, in some cases, reflected in morphological variation.

Psychopathological profile and prevalence of dual pathology on patients with alcoholic dependence undergoing outpatient treatment.

AIMS: Assess the prevalence of dual pathology in patients with alcohol dependence and describe the psychopathological profile of mental disorders, impulsiveness, ADHD presence and craving. METHOD: It is a cross-sectional study about dual pathology, carried out on 102 patients undergoing outpatient treatment. The presence of dual pathology is established by means of the MINI-5 interview and the MCMI-III test; DSM-IV being used as the alcohol abuse criteria. Impulsiveness, ADHD presence, craving and quality of life were measured through SIS, ASRSv1, MACS and SF-36. RESULTS: The prevalence of dual pathology ranges from 45.1% to 80.4% according to MCMI-III and MINI-5, respectively. The most frequent pathologies are current major depressive episodes, followed by current generalized anxiety disorders, suicide risk and current dysthymia disorders; 73.2% of dual patients present a moderate and intense global score according to MACS, 56.1% got a meaningful score in impulsiveness according to SIS and 41.5% has highly consistent symptoms with ADHD. As regards quality of life, 53.7% of the sample had bad mental health. In the case of dual patients consuming other substances, 30% had a history of bipolar disorders and 10% had a high suicide risk. CONCLUSIONS: The prevalence of psychiatric comorbidity in patients with alcohol dependence undergoing outpatient treatment varies depending on the detection method, MINI being the one identifying a greater number of cases. More than half of dual patients present impulsive behavior, a bad mental health state and high craving levels. Special attention should be paid to dual patients consuming other substances.

Parthenogenesis through the ice ages: A biogeographic analysis of Caucasian rock lizards (genus Darevskia).

Darevskia rock lizards include both sexual and parthenogenetic species, mostly distributed in the heterogeneous and ecologically diverse Caucasus. The parthenogenetic species originated via directional hybridogenesis, with only some of the sexual species known to serve as parentals. However, it remains unclear when and where these events happened and how many parental lineages were involved. A multilocus phylogeographic analysis was performed on the parthenogens D. unisexualis, D. bendimahiensis and D. uzzeli, and their putative maternal species D. raddei. Results show the parthenogenetic species all have relatively recent origins, approximately 200-70kyr ago, and at least three hybridization events were involved in their formation. Ecological niche models identify the region where hybridization events leading to the formation of D. unisexualis took place, namely in the northeast of the current distribution. Models also suggest that the sexual D. raddei might have undergone a habitat shift between the Last Interglacial and the Last Glacial Maximum.

An ecophysiological background for biogeographic patterns of two island lizards?

Distributions of sedentary ectotherms are dependent on temperature and humidity due to their low homeostatic and dispersal abilities. Lizards are strongly conditioned by temperature, but hydric environment may be also important, at least in arid environments. Biotic interactions may also play a role in range patterns, but they are of minor importance in islands where native species monopolize well-delimited niche spaces. On the arid island of São Vicente (Cabo Verde), two endemic lizards display different spatial patterns. While the gecko Tarentola substituta is widely distributed across the island, the skink Chioninia stangeri is restricted to the NE, which is cooler, more humid, and vegetated. We hypothesized that this is due to differences in the fundamental niche, specifically in ecophysiology. We predict that C. stangeri should select for lower temperatures and lose more water by evaporation than T. substituta. We submitted adults of each species to standard experiments to assess preferred body temperatures (Tp) and evaporative water loss (EWL) rates, and examined the variation between species and through time using repeated-measures AN(C)OVAs. Results only partially supported our expectations. Contrary to the prediction, skinks attained higher Tp than geckos but in the long term showed a trend for higher EWL as predicted. Thus, while ecophysiology certainly contributes to functional interpretation of species distributions, it needs to be combined with other evidence such as habitat use and evolutionary history. These findings will be useful to perform mechanistic models to better understand the impact of climate change and habitat disturbance on these endemic species.