Phylogenomic insights into the diversity and evolution of Palearctic vipers.

Despite decades of molecular research, phylogenetic relationships in Palearctic vipers (genus Vipera) still essentially rely on a few loci, such as mitochondrial barcoding genes. Here we examined the diversity and evolution of Vipera with ddRAD-seq data from 33 representative species and subspecies. Phylogenomic analyses of ∼ 1.1 Mb recovered nine major clades corresponding to known species/species complexes which are generally consistent with the mitochondrial phylogeny, albeit with a few deep discrepancies that highlight past hybridization events. The most spectacular case is the Italian-endemic V. walser, which is grouped with the alpine genetic diversity of V. berus in the nuclear tree despite carrying a divergent mitogenome related to the Caucasian V. kaznakovi complex. Clustering analyses of SNPs suggest potential admixture between diverged Iberian taxa (V. aspis zinnikeri and V. seoanei), and confirm that the Anatolian V. pontica corresponds to occasional hybrids between V. (ammodytes) meridionalis and V. kaznakovi. Finally, all analyzed lineages of the V. berus complex (including V. walser and V. barani) form vast areas of admixture and may be delimited as subspecies. Our study sets grounds for future taxonomic and phylogeographic surveys on Palearctic vipers, a group of prime interest for toxinological, ecological, biogeographic and conservation research.

Contribution of host species and pathogen clade to snake fungal disease hotspots in Europe.

Infectious diseases are influenced by interactions between host and pathogen, and the number of infected hosts is rarely homogenous across the landscape. Areas with elevated pathogen prevalence can maintain a high force of infection and may indicate areas with disease impacts on host populations. However, isolating the ecological processes that result in increases in infection prevalence and intensity remains a challenge. Here we elucidate the contribution of pathogen clade and host species in disease hotspots caused by Ophidiomyces ophidiicola, the pathogen responsible for snake fungal disease, in 21 species of snakes infected with multiple pathogen strains across 10 countries in Europe. We found isolated areas of disease hotspots in a landscape where infections were otherwise low. O. ophidiicola clade had important effects on transmission, and areas with multiple pathogen clades had higher host infection prevalence. Snake species further influenced infection, with most positive detections coming from species within the Natrix genus. Our results suggest that both host and pathogen identity are essential components contributing to increased pathogen prevalence.

Paranannizziopsis spp. Infection in Wild Vipers, Europe.

We describe the detection of Paranannizziopsis sp. fungus in a wild population of vipers in Europe. Fungal infections were severe, and 1 animal likely died from infection. Surveillance efforts are needed to better understand the threat of this pathogen to snake conservation.

Isolation and characterization of polymorphic microsatellite loci for the three Iberian vipers, Vipera aspis, V. Latastei and V. seoanei by Illumina MiSeq sequencing.

BACKGROUND: European vipers (genus Vipera) are a well-studied taxonomic group, but the low resolution of nuclear sanger-sequenced regions has precluded thorough studies at systematic, ecological, evolutionary and conservation levels. In this study, we developed novel microsatellite markers for the three Iberian vipers, Vipera aspis, V. latastei and V. seoanei, and assessed their polymorphism in north-central Iberian populations. METHODS AND RESULTS: Genomic libraries were developed for each species using an Illumina Miseq sequencing approach. From the 70 primer pairs initially tested, 48 amplified reliably and were polymorphic within species. Cross-species transferability was achieved for 31 microsatellites loci in the three target species and four additional loci that were transferable to one species only. The 48 loci amplified in average seven alleles, and detected average expected and observed heterozygosities of 0.7 and 0.55, in the three genotyped populations/species (26 V. aspis, 20 V. latastei and 10 V. seoanei). CONCLUSIONS: Our study provides a selection of 48 polymorphic microsatellite markers that will contribute significantly to current knowledge on genetic diversity, gene flow, population structure, demographic dynamics, systematics, reproduction and heritability in these species, and potentially in other congeneric taxa.

Hidden in the sand: Phylogenomics unravel an unexpected evolutionary history for the desert-adapted vipers of the genus Cerastes.

The desert vipers of the genus Cerastes are a small clade of medically important venomous snakes within the family Viperidae. According to published morphological and molecular studies, the group is comprised by four species: two morphologically similar and phylogenetically sister taxa, the African horned viper (Cerastes cerastes) and the Arabian horned viper (Cerastes gasperettii); a more distantly related species, the Saharan sand viper (Cerastes vipera), and the enigmatic Böhme's sand viper (Cerastes boehmei), only known from a single specimen in captivity allegedly captured in Central Tunisia. In this study, we sequenced one mitochondrial marker (COI) as well as genome-wide data (ddRAD sequencing) from 28 and 41 samples, respectively, covering the entire distribution range of the genus to explore the population genomics, phylogenomic relationships and introgression patterns within the genus Cerastes. Additionally, and to provide insights into the mode of diversification of the group, we carried out niche overlap analyses considering climatic and habitat variables. Both nuclear phylogenomic reconstructions and population structure analyses have unveiled an unexpected evolutionary history for the genus Cerastes, which sharply contradicts the morphological similarities and previously published mitochondrial approaches. Cerastes cerastes and C. vipera are recovered as sister taxa whilst C. gasperettii is a sister taxon to the clade formed by these two species. We found a relatively high niche overlap (OI > 0.7) in both climatic and habitat variables between C. cerastes and C. vipera, contradicting a potential scenario of sympatric speciation. These results are in line with the introgression found between the northwestern African populations of C. cerastes and C. vipera. Finally, our genomic data confirms the existence of a lineage of C. cerastes in Arabia. All these results highlight the importance of genome-wide data over few genetic markers to study the evolutionary history of species.

Local niches explain coexistence in environmentally-distinct contact zones between Western Mediterranean vipers.

Species' ecological niches are frequently analysed to gain insights into how anthropogenic changes affect biodiversity. Coping with these changes often involves shifts in niche expression, which can disrupt local biotic interactions. Secondary contact zones, where competition and ecological segregation commonly occur, are ideal for studying the ecological factors influencing species' niches. In this study, we investigated the effect of climate and landscape factors on the ecological niches of two viper species, Vipera aspis and Vipera latastei, across three contact zones in northern Iberia, characterized by varying levels of landscape alteration. Using niche overlap tests, ecological niche models and spatial analyses we observed local variation in the expression of the species' niches across the three contact zones, resulting from the different abiotic and biotic conditions of each area. Rather than spatial niche segregation, we observed high niche overlap, suggesting niche convergence. A pattern of asymmetrical niche variation was identified in all contact zones, driven by species' climatic tolerances and the environmental conditions of each area. V. aspis generally exhibited a wider niche, except in the southernmost zone where the pure Mediterranean climate favored V. latastei. Human-induced landscape changes intensified niche asymmetry, by favoring the most generalist V. aspis over the specialist V. latastei, increasing habitat overlap, and likely competition. This study presents a comprehensive analysis of niche expression at range margins, anticipating a heightened impact of landscape changes in V. latastei. The methodological framework implemented here, and our findings, hold significant relevance for biodiversity management and conservation in human-impacted areas.

Phylogeographic and Paleoclimatic Modelling Tools Improve Our Understanding of the Biogeographic History of Hierophis viridiflavus (Colubridae).

Phylogeographic and paleoclimatic modelling studies have been combined to infer the role of Pleistocene climatic oscillations as drivers of the genetic structure and distribution of Mediterranean taxa. For the European whip snake, Hierophis viridiflavus, previous studies based on paleoclimatic modelling have depicted a low reliability in the pattern of past climatic suitability across the central Mediterranean Basin, which barely fits the species' genetic structure. In this study, we combined phylogeographic and paleoclimatic modelling tools to improve our understanding of the biogeographic history of H. viridiflavus, particularly extending the sampling and phylogeographic inferences to previously under-sampled regions. Phylogeographic analyses recovered two major clades that diverged at the beginning of the Pleistocene and had diversified in different ways by the late Pleistocene: the east clade (composed of three subclades) and the west clade (with no further structure). Paleoclimatic models highlighted the temperate character of H. viridiflavus, indicating range contractions during both the last inter-glacial and last glacial maximum periods. Range expansions from southern-located climatic refugia likely occurred in the Bølling-Allerød and Middle Holocene periods, which are supported by signals of demographic growth in the west clade and South-East-North subclade. Overall, this work improves our understanding of the historical biogeography of H. viridiflavus, providing further insights into the evolutionary processes that occurred in the Mediterranean Basin hotspot.

One Size Fits All-Venomics of the Iberian Adder (Vipera seoanei, Lataste 1878) Reveals Low Levels of Venom Variation across Its Distributional Range.

European vipers (genus Vipera) are medically important snakes displaying considerable venom variation, occurring at different levels in this group. The presence of intraspecific venom variation, however, remains understudied in several Vipera species. Vipera seoanei is a venomous snake endemic to the northern Iberian Peninsula and south-western France, presenting notable phenotypic variation and inhabiting several diverse habitats across its range. We analysed the venoms of 49 adult specimens of V. seoanei from 20 localities across the species' Iberian distribution. We used a pool of all individual venoms to generate a V. seoanei venom reference proteome, produced SDS-PAGE profiles of all venom samples, and visualised patterns of variation using NMDS. By applying linear regression, we then assessed presence and nature of venom variation between localities, and investigated the effect of 14 predictors (biological, eco-geographic, genetic) on its occurrence. The venom comprised at least 12 different toxin families, of which five (i.e., PLA2, svSP, DI, snaclec, svMP) accounted for about 75% of the whole proteome. The comparative analyses of the SDS-PAGE venom profiles showed them to be remarkably similar across the sampled localities, suggesting low geographic variability. The regression analyses suggested significant effects of biological and habitat predictors on the little variation we detected across the analysed V. seoanei venoms. Other factors were also significantly associated with the presence/absence of individual bands in the SDS-PAGE profiles. The low levels of venom variability we detected within V. seoanei might be the result of a recent population expansion, or of processes other than directional positive selection.

Head Shape Heritability in the Hungarian Meadow Viper Vipera ursinii rakosiensis.

Understanding heritability patterns in functionally relevant traits is a cornerstone for evaluating their evolutionary potential and their role in local adaptation. In this study, we investigated patterns of heritability in the head shape of the Hungarian meadow viper (Vipera ursinii rakosiensis). To this end, we used geometric morphometric data from 12 families composed of 8 mothers, 6 fathers and 221 offspring, bred in captivity at the Hungarian Meadow Viper Conservation Centre (Hungary). We separately evaluated maternal and paternal contributions to the offspring phenotype, in addition to additive genetic effects, all determined using a mixed animal model. Our results indicate a strong genetic and maternal contribution to head shape variations. In contrast, the paternal effects-which are rarely evaluated in wild-ranging species-as well as residual environmental variance, were minimal. Overall, our results indicate a high evolutionary potential for head shape in the Hungarian meadow viper, which suggests a strong contribution of this ecologically important trait in shaping the ability of this endangered species to adapt to changing conditions and/or habitats. Furthermore, our results suggest that maternal phenotypes should be carefully considered when designing captive breeding parental pairs for reinforcing the adaptive capacity of threatened populations, whereas the paternal phenotypes seem less relevant.

Toxic Habits: An Analysis of General Trends and Biases in Snake Venom Research.

Biases in snake venom research have been partially identified but seldomly quantified. Using the Google Scholar web search engine, we collected a total of 267 articles published between 1964 and 2021, and reviewed them to assess the main trends in this field of study. We developed a 4-category classification of the harmful potential of each of the 298 snake species retrieved from the analysed publications, and tested whether taxonomy, realm of origin, and/or assigned hazard category could affect how often each of them appeared in the articles considered. Overall, viperids were significantly more represented than any other snake taxon retrieved. The Neotropics were the most represented biogeographic realm for number of studied species, whereas information about the country of origin of the analysed specimens was often incomplete. The vast majority of the publications focused on snake venom characterisation, whereas more ecology-related topics were rarely considered. Hazard category and biogeographic realm of origin of each species had a significant effect on the number of articles dedicated to it, suggesting that a snake's harmful potential and place of origin influence its popularity in venom studies. Our analysis showed an overall positive trend in the number of snake venom studies published yearly, but also underlined severe neglect of snake families of supposedly minor medical relevance (e.g., Atractaspididae), underrepresentation of some of the areas most impacted by snakebite (i.e., Indomalayan and Afrotropic realms), and limited interest in the ecological and functional context of snake venom.

DNA barcode reference library for the West Sahara-Sahel reptiles.

DNA barcode reference libraries are now continuously produced for the tree of life, which are essential pillars for the study of biological diversity. Yet, our knowledge about global diversity is largely limited in undersampled regions such as the largest warm desert, the Sahara-Sahel. This dataset provides a DNA barcode reference library for the reptiles of the Western Sahara-Sahel (WSS) and neighbouring countries across this region. It includes 760 barcodes from 133 reptile taxa, distributed in 23 families, and covering the intraspecific diversity of some species. A total of 84 species were collected in the WSS (83% of the total reptile species richness) over 18 overland field expeditions conducted since 2003. DNA barcodes resulted in a high success rate (95%) of species identification and barcoding gap analysis highlighted the effectiveness of the COI fragment as a barcode marker for the WSS reptiles. This dataset represents a comprehensive and reliable DNA reference library for the WSS, filling an important biodiversity gap across a remote and hard-to-sample region.

Diversity, distribution and conservation of land mammals in Mauritania, North-West Africa.

Detailed knowledge about biodiversity distribution is critical for monitoring the biological effects of global change processes. Biodiversity knowledge gaps hamper the monitoring of conservation trends and they are especially evident in the desert biome. Mauritania constitutes a remarkable example on how remoteness and regional insecurity affect current knowledge gaps. Mammals remain one of the least studied groups in this country, without a concerted species checklist, the mapping of regions concentrating mammal diversity, or a national assessment of their conservation status. This work assessed the diversity, distribution, and conservation of land mammals in Mauritania. A total of 6,718 published and original observations were assembled in a spatial database and used to update the occurrence status, distribution area, and conservation status. The updated taxonomic list comprises 107 species, including 93 extant, 12 Regionally Extinct, and 2 Extinct in the Wild. Mapping of species distributions allowed locating concentrations of extant mammal species richness in coastal areas, along the Senegal River valley, and in mountain plateaus. Recent regional extinction of large-sized Artiodactyla and Carnivora has been very high (11% extinct species). From the extant mammals, 11% are threatened, including flagship species (e.g., Addax nasomaculatus and Panthera pardus). Species richness is poorly represented by the current protected areas. Despite the strong advances made, 23% of species categorise as Data Deficient. Persisting systematics and distribution uncertainties require further research. Field surveys in currently unexplored areas (northern and south-eastern regions) are urgently needed to increase knowledge about threatened mammals. The long-term conservation of land mammals in Mauritania is embedded in a complex web of socioeconomic and environmental factors that call for collaborative action and investment in sustainable human development. The current work sets the baseline for the future development of detailed research studies and to address the general challenges faced by mammals and biodiversity in the country.

Interpopulational variation and ontogenetic shift in the venom composition of Lataste's viper (Vipera latastei, Boscá 1878) from northern Portugal.

Lataste's viper (Vipera latastei) is a venomous European viper endemic to the Iberian Peninsula, recognised as medically important by the World Health Organization. To date, no comprehensive characterisation of this species' venom has been reported. Here, we analysed the venoms of juvenile and adult specimens of V. latastei from two environmentally different populations from northern Portugal. Using bottom-up venomics, we produced six venom proteomes (three per population) from vipers belonging to both age classes (i.e., two juveniles and four adults), and RP-HPLC profiles of 54 venoms collected from wild specimens. Venoms from juveniles and adults differed in their chromatographic profiles and relative abundances of their toxins, suggesting the occurrence of ontogenetic changes in venom composition. Specifically, snake venom metalloproteinase (SVMP) was the most abundant toxin family in juvenile venoms, while snake venom serine proteinases (SVSPs), phospholipases A2 (PLA2s), and C-type lectin-like (CTLs) proteins were the main toxins comprising adult venoms. The RP-HPLC venom profiles were found to vary significantly between the two sampled localities, indicating geographic variability. Furthermore, the presence/absence of certain peaks in the venom chromatographic profiles appeared to be significantly correlated also to factors like body size and sex of the vipers. Our findings show that V. latastei venom is a variable phenotype. The intraspecific differences we detected in its composition likely mirror changes in the feeding ecology of this species, taking place during different life stages and under different environmental pressures. SIGNIFICANCE: Lataste's viper (Vipera latastei) is a medically important viper endemic to the Iberian Peninsula, inhabiting different habitats and undergoing a marked ontogenetic dietary shift. In the current study, we report the first proteomic analysis of V. latastei venom from two environmentally different localities in northern Portugal. Our bottom-up venomic analyses show that snake venom serine proteinases (SVSPs), phospholipases A2 (PLA2s), and C-type lectin-like (CTLs) proteins are the major components of adult V. latastei venom. The comparative analysis of young and adult venoms suggests the occurrence of ontogenetic shift in toxin abundances, with snake venom metalloproteinases (SVMPs) being the predominant toxins in juvenile venoms. Moreover, geographic venom variation between the two studied populations is also detected, with our statistical analyses suggesting that factors like body size and sex of the vipers are possibly at play in its determination. Our work represents the first assessment of the composition of V. latastei venom, and the first step towards a better understanding of the drivers behind its variability.

Species versus within-species niches: a multi-modelling approach to assess range size of a spring-dwelling amphibian.

Species Distribution Models (SDMs) can be used to estimate potential geographic ranges and derive indices to assess species conservation status. However, habitat-specialist species require fine-scale range estimates that reflect resource dependency. Furthermore, local adaptation of intraspecific lineages to distinct environmental conditions across ranges have frequently been neglected in SDMs. Here, we propose a multi-stage SDM approach to estimate the distributional range and potential area of occupancy (pAOO) of Neurergus kaiseri, a spring-dwelling amphibian with two climatically-divergent evolutionary lineages. We integrate both broad-scale climatic variables and fine-resolution environmental data to predict the species distribution while examining the performance of lineage-level versus species-level modelling on the estimated pAOO. Predictions of habitat suitability at the landscape scale differed considerably between evolutionary level models. At the landscape scale, spatial predictions derived from lineage-level models showed low overlap and recognised a larger amount of suitable habitats than species-level model. The variable dependency of lineages was different at the landscape scale, but similar at the local scale. Our results highlight the importance of considering fine-scale resolution approaches, as well as intraspecific genetic structure of taxa to estimate pAOO. The flexible procedure presented here can be used as a guideline for estimating pAOO of other similar species.

[Venomous Animals in the Portuguese Territory: Clinical Management of Bites and Stings]. / Animais Venenosos em Território Português: Abordagem Clínica de Picadas e Mordeduras.

There are many native species of animals with the potential to cause severe disease in Portugal. Of the four clinically relevant snake species, the vipers (Vipera latastei and Vipera seoanei) are the most concerning. They can cause severe disease and require in-hospital management, monitoring and specific treatment, including the administration of anti-snake venom serum. The Malpolon monspessulanus and Macroprotodon brevis snakes cause mostly localized clinical manifestations, which do not require specific treatment. Only a minority of the spider species in Portugal possess chelicerae (mouthparts of the Chelicerata, shaped as either articulated fangs or pincers, which may contain venom or be connected to the venom glands and are used for predation or capture of food) long enough to perforate the human skin. Nevertheless, Latrodectus tredecimguttatus and Loxosceles rufescens spider bites may require specialized treatment in a hospital setting, as well as careful active symptom surveillance. Diversely, the treatment for Scolopendra cingulata and Scolopendra oraniensis centipedes' bites is merely symptomatic. The only existing scorpion species in Portugal is the Buthus ibericus; its sting typically causes local symptoms with intense pain, and its treatment consists essentially of analgesia. The insects of the Hymenoptera order, such as bees and wasps, have the ability to inject venom into the skin. Most people present only with local or regional inflammatory response, and symptomatic treatment is usually effective. Even so, individuals with hypersensitivity to bee venom may develop anaphylaxis. Several marine species in the Portuguese coast are venomous to humans, including weeverfish (Trachinidae family), stingrays (Dasyatis pastinaca, Taeniura grabata, Myliobatis aquila), red scorpion fish (Scorpaena scrofa), cnidaria (Pelagia noctiluca, Chrysaora hysoscella, Physalia physalis) and bearded fireworm (Hermodice carunculata); treatment is symptomatic. Contact with the larvae or bristles (chitinous structures with locomotor or tactile functions) of Thaumetopoea pityocampa (pine processionary) can cause cutaneous, ocular and, rarely, respiratory reactions; its management is also symptomatic.

Em território português, existem várias espécies autóctones de animais capazes de provocar envenenamento e doença clinicamente significativa, com potencial de gravidade. Das quatro espécies de serpentes com relevância clínica, as víboras (Vipera latastei e Vipera seoanei) são as mais preocupantes; podem causar doença grave e necessitam de abordagem hospitalar, monitorização e tratamento específico, incluindo soro antiveneno. As serpentes Malpolon monspessulanus e Macroprotodon brevis causam maioritariamente manifestações clínicas locais, sem tratamento específico. Apenas uma minoria das espécies de aranha presentes em Portugal possui quelíceras (apêndices bucais dos Chelicerata, em forma de tenaz ou gancho, que podem conter veneno ou encontrar-se ligados às glândulas venenosas, usados para predação ou captura de alimentos) suficientemente compridas para perfurar a pele humana, mas as picadas de Latrodectus tredecimguttatus e Loxosceles rufescens podem implicar tratamento diferenciado em ambiente hospitalar, com necessidade de vigilância ativa. O tratamento da picada das centopeias Scolopendra cingulata e Scolopendra oraniensis é sintomático. A única espécie de escorpião presente em território nacional, Buthus ibericus, causa geralmente sintomas locais com dor intensa; o tratamento habitualmente baseia-se apenas na administração de analgesia. Os insetos da ordem Hymenoptera, como as abelhas e vespas, possuem capacidade de libertar veneno através de picadas; a maior parte dos indivíduos apresenta apenas reações inflamatórias locais ou regionais, para as quais a aplicação de medidas sintomáticas é suficiente, mas indivíduos que apresentem hipersensibilidade ao veneno podem desenvolver reações anafiláticas. Na costa portuguesa e em águas pouco profundas, encontram-se várias espécies marinhas venenosas para os humanos, como Echiichthys vipera (peixe-aranha comum), Dasyatis pastinaca, Taeniura grabata e Myliobatis aquila (ratões), Scorpaena scrofa (rascasso-vermelho), Pelagia noctiluca, Chrysaora hysoscella e Physalia physalis (cnidários) e Hermodice carunculata (verme-do-fogo), cujas picadas necessitam apenas de tratamento sintomático. O contacto com as larvas ou cerdas (estruturas quitinosas com funções locomotoras ou tácteis) de Thaumetopoea pityocampa (lagarta-do-pinheiro) pode provocar reações cutâneas, oculares e, raramente, respiratórias; a sua abordagem também é sintomática.

Using fire to enhance rewilding when agricultural policies fail.

Rewilding has been proposed as an opportunity for biodiversity conservation in abandoned landscapes. However, rewilding is challenged by the increasing fire risk associated with more flammable landscapes, and the loss of open-habitat specialist species. Contrastingly, supporting High Nature Value farmlands (HNVf) has been also highlighted as a valuable option, but the effective implementation of agricultural policies often fails leading to uncertain scenarios wherein the effects of wildfire management remain largely unexplored. Herein, we simulated fire-landscape dynamics to evaluate how fire suppression scenarios affect fire regime and biodiversity (102 species of vertebrates) under rewilding and HNVf policies in the future (2050), in a transnational biosphere reserve (Gerês-Xurés Mountains, Portugal-Spain). Rewilding and HNVf scenarios were modulated by three different levels of fire suppression effectiveness. Then, we quantified scenario effects on fire regime (burned and suppressed areas) and biodiversity (habitat suitability change for 2050). Simulations confirm HNVf as a long-term opportunity for fire suppression (up to 30,000 ha of additional suppressed areas between 2031 and 2050 in comparison to rewilding scenario) and for conservation (benefiting around 60% of species). Rewilding benefits some species (20%), including critically endangered, vulnerable and endemic taxa, while several species (33%) also profit from open habitats created by fire. Although HNVf remains the best scenario, rewilding reinforced by low fire suppression management may provide a nature-based solution when societal support through agricultural policies fails.

Thermal melanism explains macroevolutionary variation of dorsal pigmentation in Eurasian vipers.

Colouration may endorse thermoregulatory and antipredatory functions in snakes. The thermal melanism hypothesis predicts that dark-coloured individuals are ecologically favoured in cool climates. However, the loss of aposematic and cryptic colourations may imply high predation for melanistic snakes. Here, we used the monophyletic group of Eurasian vipers (subfamily Viperinae) to test whether an increase in the extent of dark area inside the characteristic zigzag dorsal pattern is associated to colder environments. We measured two colouration traits in zigzag-patterned individuals (number of dorsal marks and weighted pigmentation index) and used a phylogenetic comparative approach to explore macroevolutionary patterns of dorsal pigmentation and test whether its extent is associated to ecogeographic characteristics of lineages' ranges. Phylogenetically-naïve and phylogenetically-informed analyses yielded a significant association between the degree of pigmentation of the zigzag pattern and environmental variables such as solar radiation, elevation and latitude. The degree of pigmentation of the zigzag pattern is highlighted as an adaptive trait that matches range attributes mirroring cold environments irrespective of the phylogeny. These results constitute the first large-scale evidence supporting the thermal melanism hypothesis in snakes, opening new avenues of inquiry for the mechanisms that shape the evolution of colour phenotypes.

Evolutionary history of two cryptic species of northern African jerboas.

BACKGROUND: Climatic variation and geologic change both play significant roles in shaping species distributions, thus affecting their evolutionary history. In Sahara-Sahel, climatic oscillations shifted the desert extent during the Pliocene-Pleistocene interval, triggering the diversification of several species. Here, we investigated how these biogeographical and ecological events have shaped patterns of genetic diversity and divergence in African Jerboas, desert specialist rodents. We focused on two sister and cryptic species, Jaculus jaculus and J. hirtipes, where we (1) evaluated their genetic differentiation, (2) reconstructed their evolutionary and demographic history; (3) tested the level of gene flow between them, and (4) assessed their ecological niche divergence. RESULTS: The analyses based on 231 individuals sampled throughout North Africa, 8 sequence fragments (one mitochondrial and seven single copy nuclear DNA, including two candidate genes for fur coloration: MC1R and Agouti), 6 microsatellite markers and ecological modelling revealed: (1) two distinct genetic lineages with overlapping distributions, in agreement with their classification as different species, J. jaculus and J. hirtipes, with (2) low levels of gene flow and strong species divergence, (3) high haplotypic diversity without evident geographic structure within species, and (4) a low level of large-scale ecological divergence between the two taxa, suggesting species micro-habitat specialization. CONCLUSIONS: Overall, our results suggest a speciation event that occurred during the Pliocene-Pleistocene transition. The contemporary distribution of genetic variation suggests ongoing population expansions. Despite the largely overlapping distributions at a macrogeographic scale, our genetic results suggest that the two species remain reproductively isolated, as only negligible levels of gene flow were observed. The overlapping ecological preferences at a macro-geographic scale and the ecological divergence at the micro-habitat scale suggest that local adaptation may have played a crucial role in the speciation process of these species.

Allopatric diversification and evolutionary melting pot in a North African Palearctic relict: The biogeographic history of Salamandra algira.

North Africa is a climatically and topographically complex region with unique biotic assemblages resulting from the combination of multiple biogeographic realms. Here, we assess the role of climate in promoting intra-specific diversification in a Palearctic relict, the North African fire salamander, Salamandra algira, using a combination of phylogenetic and population genetic analyses, paleoclimatic modelling and niche overlap tests. We used mitochondrial DNA (Cyt-b), 9838 ddRADseq loci, and 14 microsatellite loci to characterize patterns of genetic diversity and population structure. Phylogenetic analyses recover two major clades, each including several lineages with mito-nuclear discordances suggesting introgressive patterns between lineages in the Middle Atlas, associated with a melting pot of genetic diversity. Paleoclimatic modelling identified putative climatic refugia, largely matching areas of high genetic diversity, and supports the role of aridity in promoting allopatric diversification associated with ecological niche conservatism. Overall, our results highlight the role of climatic microrefugia as drivers of populations' persistence and diversification in the face of climatic oscillations in North Africa, and stress the importance of accounting for different genomic regions when reconstructing biogeographic processes from molecular markers.