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Biological invasions pose one of the most severe environmental challenges of the twenty-first century. A longstanding idea is that invasion risk is predictable based on the phylogenetic distance - and hence ecological resemblance - between non-native and native species. However, current evidence is contradictory. To explain these mixed results, it has been proposed that the effect is scale-dependent, with invasion inhibited by phylogenetic similarity at small spatial scales but enhanced at larger scales. Analyzing invasion outcomes in a global sample of bird communities, we find no evidence to support this hypothesis. Instead, our results suggest that invaders are locally more successful in the presence of closely related and ecologically similar species, at least in human-altered environments where the majority of invasions have occurred. Functional trait analyses further confirm that the ecological niches of invaders are phylogenetically conserved, supporting the notion that successful invasion in the presence of close relatives is driven by shared adaptations to the types of niches available in novel environments.

Understanding the genomic basis of adaptation to different abiotic environments is important in the context of climate change and resulting short-term environmental fluctuations. Using functional and comparative genomics approaches, we here investigated whether signatures of genomic adaptation to a set of environmental parameters are concentrated in specific subsets of genes and functions in lacertid lizards and other vertebrates. We first identify 200 genes with signatures of positive diversifying selection from transcriptomes of 24 species of lacertid lizards and demonstrate their involvement in physiological and morphological adaptations to climate. To understand how functionally similar these genes are to previously predicted candidate functions for climate adaptation and to compare them with other vertebrate species, we then performed a meta-analysis of 1, 100 genes under selection obtained from -omics studies in vertebrate species adapted to different abiotic factors. We found that the vertebrate gene set formed a tightly connected interactome, which was to 23% enriched in previously predicted functions of adaptation to climate, and to a large part (18%) involved in organismal stress response. We found a much higher degree of identical genes being repeatedly selected among different animal groups (43.6%), and of functional similarity and post-translational modifications than expected by chance, and no clear functional division between genes used for ectotherm and endotherm physiological strategies. In total, 171 out of 200 genes of Lacertidae were part of this network. These results highlight an important role of a comparatively small set of genes and their functions in environmental adaptation and narrow the set of candidate pathways and markers to be used in future research on adaptation and stress response related to climate change.

A longstanding issue in biology is whether the intelligence of animals can be predicted by absolute or relative brain size. However, progress has been hampered by an insufficient understanding of how neuron numbers shape internal brain organization and cognitive performance. On the basis of estimations of neuron numbers for 111 bird species, we show here that the number of neurons in the pallial telencephalon is positively associated with a major expression of intelligence: innovation propensity. The number of pallial neurons, in turn, is greater in brains that are larger in both absolute and relative terms and positively covaries with longer post-hatching development periods. Thus, our analyses show that neuron numbers link cognitive performance to both absolute and relative brain size through developmental adjustments. These findings help unify neuro-anatomical measures at multiple levels, reconciling contradictory views over the biological significance of brain expansion. The results also highlight the value of a life history perspective to advance our understanding of the evolutionary bases of the connections between brain and cognition.

Anthropogenic climate change is rapidly altering local environments and threatening biodiversity throughout the world. Although many wildlife responses to this phenomenon appear largely idiosyncratic, a wealth of basic research on this topic is enabling the identification of general patterns across taxa. Here, we expand those efforts by investigating how avian responses to climate change are affected by the ability to cope with ecological variation through behavioural flexibility (as measured by relative brain size). After accounting for the effects of phylogenetic uncertainty and interspecific variation in adaptive potential, we confirm that although climate warming is generally correlated with major body size reductions in North American migrants, these responses are significantly weaker in species with larger relative brain sizes. Our findings suggest that cognition can play an important role in organismal responses to global change by actively buffering individuals from the environmental effects of warming temperatures.

Despite biological invasions are one of the main environmental problems of the twenty-first century, there is still no theoretical or empirical agreement on whether a high phylogenetic relatedness between exotic and native species positively or negatively affect invasion success. To resolve this conundrum, it has been proposed that the effect might be scale-dependent, being negative at smaller spatial scales and positive at larger scales. Here we show that this scale-dependent pattern may be a sampling artefact associated with species-area effects and a non-random pattern of species introductions. We support this conclusion with simulations and empirical data on invaded and non-invaded avian communities in regions from five continents. We further show that at smaller-scales —where these artifacts are negligible— invasion success generally increases with the presence of closely-related species, but that predictive accuracy largely depends on considering the influence of human-related disturbances in facilitating invasions.

Ecological stoichiometry and studies of biogeochemical niches have mainly focused on plankton and vascular plants, but the phenotypically closest modern relatives of early plants, bryophytes, have been largely neglected. We analysed C:N:P stoichiometries and elemental compositions (K, Na, Mg, Ca, S, Fe) of 35 widely distributed bryophyte species inhabiting springs. We estimated bryophyte C:N:P ratios and their biogeochemical niches, investigated how elementomes respond to the environment and determined whether they tend to diverge more for coexisting than non-coexisting individuals and species. The median C:N:P was 145:8:1, intermediate between Redfield's ratio for marine plankton and those for vascular plants. Biogeochemical niches were differentiated amongst species and were phylogenetically conserved. Differences in individual and species-specific elementomes increased with coexistence between species. Our results provide an evolutionary bridge between the ecological stoichiometries of algae and vascular plants and suggest that differences in elementomes could be used to understand community assemblages and functional diversity.

Nutrient economy and plant nutrient concentrations have been suggested as important selective pressures of reproductive traits in vascular plants. However, evidence supporting that the reproduction of bryophytes and their reproductive traits have also been conditioned by nutrient availability is lacking. We here analysed the nutrient (N, P, K, and Fe) concentration of 35 aquatic and semi-aquatic bryophytes and the chemistry of the spring water in which they lived to determine whether bryophyte nutrient concentrations were correlated with the frequency of sexual reproduction, main reproductive mode (sexual vs. asexual), dioicy, and spore diameter, while controlling for climate and nutrient concentrations in the spring water. Water nutrient concentrations, except for K, had a small but positive influence on bryophyte nutrient concentrations. P-rich and K-poor species produced sporophytes more often and were more likely to reproduce sexually and be monoicous. Spore diameter was smaller in P- and Fe-rich mosses but larger in N-rich mosses living in humid climates. P-rich species thus produced more and smaller spores and were able to propagate more extensively. Overall, our results suggest that nutrient economy, especially P economy, has played an important role in shaping the reproductive traits of bryophytes.

The fossil record provides compelling examples of heterochrony at macroevolutionary scales such as the peramorphic giant antlers of the Irish elk. Heterochrony has also been invoked in the evolution of the distinctive cranial frill of ceratopsian dinosaurs such as Triceratops. Although ceratopsian frills vary in size, shape, and ornamentation, quantitative analyses that would allow for testing hypotheses of heterochrony are lacking. Here, we use geometric morphometrics to examine frill shape variation across ceratopsian diversity and within four species preserving growth series. We then test whether the frill constitutes an evolvable module both across and within species, and compare growth trajectories of taxa with ontogenetic growth series to identify heterochronic processes. Evolution of the ceratopsian frill consisted primarily of progressive expansion of its caudal and caudolateral margins, with morphospace occupation following taxonomic groups. Although taphonomic distortion represents a complicating factor, our data support modularity both across and within species. Peramorphosis played an important role in frill evolution, with acceleration operating early in neoceratopsian evolution followed by progenesis in later diverging cornosaurian ceratopsians. Peramorphic evolution of the ceratopsian frill may have been facilitated by the decoupling of this structure from the jaw musculature, an inference that predicts an expansion of morphospace occupation and higher evolutionary rates among ceratopsids as indeed borne out by our data. However, denser sampling of the meager record of early‐diverging taxa is required to test this further., We find support for peramorphic evolution of the frill of ceratopsian dinosaurs, likely facilitated by the decoupling of this structure from jaw musculature.

Anuran amphibians undergo major morphological transitions during development, but the contribution of their markedly different life-history phases to macroevolution has rarely been analysed. Here we generate testable predictions for coupling versus uncoupling of phenotypic evolution of tadpole and adult life-history phases, and for the underlying expression of genes related to morphological feature formation. We test these predictions by combining evidence from gene expression in two distantly related frogs, Xenopus laevis and Mantidactylus betsileanus, with patterns of morphological evolution in the entire radiation of Madagascan mantellid frogs. Genes linked to morphological structure formation are expressed in a highly phase-specific pattern, suggesting uncoupling of phenotypic evolution across life-history phases. This gene expression pattern agrees with uncoupled rates of trait evolution among life-history phases in the mantellids, which we show to have undergone an adaptive radiation. Our results validate a prevalence of uncoupling in the evolution of tadpole and adult phenotypes of frogs., In animals with complex life cycles, selection on one life phase may constrain adaptation in another phase. Here the authors find that, during the adaptive radiation of mantellid frogs, the evolution of tadpole and adult morphologies has been uncoupled through phase-specific gene expression.

Despite their miniature brains, insects exhibit substantial variation in brain size. Although the functional significance of this variation is increasingly recognized, research on whether differences in insect brain sizes are mainly the result of constraints or selective pressures has hardly been performed. Here, we address this gap by combining prospective and retrospective phylogenetic-based analyses of brain size for a major insect group, bees (superfamily Apoidea). Using a brain dataset of 93 species from North America and Europe, we found that body size was the single best predictor of brain size in bees. However, the analyses also revealed that substantial variation in brain size remained even when adjusting for body size. We consequently asked whether such variation in relative brain size might be explained by adaptive hypotheses. We found that ecologically specialized species with single generations have larger brains—relative to their body size—than generalist or multi-generation species, but we did not find an effect of sociality on relative brain size. Phylogenetic reconstruction further supported the existence of different adaptive optima for relative brain size in lineages differing in feeding specialization and reproductive strategy. Our findings shed new light on the evolution of the insect brain, highlighting the importance of ecological pressures over social factors and suggesting that these pressures are different from those previously found to influence brain evolution in other taxa.

Evolutionary innovations and ecological competition are factors often cited as drivers of adaptive diversification. Yet many innovations result in stabilizing rather than diversifying selection on morphology, and morphological disparity among coexisting species can reflect competitive exclusion (species sorting) rather than sympatric adaptive divergence (character displacement). We studied the innovation of gliding in dragons (Agamidae) and squirrels (Sciuridae) and its effect on subsequent body size diversification. We found that gliding either had no impact (squirrels) or resulted in strong stabilizing selection on body size (dragons). Despite this constraining effect in dragons, sympatric gliders exhibit greater size disparity compared with allopatric gliders, a pattern consistent with, although not exclusively explained by, ecological competition changing the adaptive landscape of body size evolution to induce character displacement. These results show that innovations do not necessarily instigate further differentiation among species, as is so often assumed, and suggest that competition can be a powerful force generating morphological divergence among coexisting species, even in the face of strong stabilizing selection., This study was partly supported by an Endeavour Research Fellowship to J.G.P. awarded by the Department of Education and Training, Australian Government (ID: 5471_ 2016).

Aim To understand the patterns of genetic, geographical and morphological diversification in the Arabian gecko subspecies Pristurus rupestris rupestris, and to discuss the various processes that generate and shape diversity in arid mountain ranges. Location Hajar Mountains, south-eastern Arabian Peninsula. Methods We sampled 466 individuals across the entire range of P. r. rupestris in the Hajar Mountains and sequenced one mitochondrial gene for all the samples, and one extra mitochondrial and four nuclear genes for a subset of 75 specimens. A total of 155 individuals were characterized morphologically. Multilocus coalescent-based methods were used for species delimitation, for phylogenetic inference of species trees and to investigate the diversification dynamics. Multivariate statistics were used to analyse morphological data. Results Our molecular methods revealed 14 candidate species hidden within the subspecies P. r. rupestris. Diversification started c. 15 Ma, but most of the lineages have evolved in the last 7 Ma of the mountains’ orogeny, producing a mosaic of allopatric taxa. Secondary interchange has occurred between high and low elevation lineages, and along the mountain chain between unrelated lineages formerly separated by mountain blocks. Morphological variation between the 14 candidate species is slight, but a high elevation robust morphotype is identified. Main conclusions Our results support the notion that mountain ranges constitute important centres of diversification in arid regions, and also act as ‘species pumps’ into surrounding areas. We also demonstrate that arid mountains may constitute important reservoirs of cryptic diversity, even in common and widespread species. This, and previous evidence, suggest that the Hajar Mountains are a biological hotspot and an important reservoir of diversity. We therefore suggest that these mountains should be a priority focal point for conservation in Arabia.

The taxonomy of Procellariiformes, particularly petrels and shearwaters, is still unresolved. The Manx shearwater Puffinus puffinus is one of the best studied seabirds worldwide. Most of the information known on this seabird is focused on the northern core populations where the species is abundant. However, the species shows a high number of peripheral populations, which are extremely small and difficult to study in comparison to central populations. Using an integrative approach, we provided significant evidence of phenological, morphological, acoustic, plumage colour, and genetic differentiation of the Canarian Manx shearwaters (the most southern population) from the northern breeding colonies, which is compatible with a long period of isolation. Birds from the Canary Islands breed around 2‐3 months earlier, are smaller and lighter, and show darker underwing plumage than those from northern populations. In addition, Canarian call features are different from the northern populations. Finally, genetic analyses of the mitochondrial control region indicate an incipient genetic differentiation of Canarian Manx shearwaters from the other breeding populations. The Canarian population holds a small number of breeding colonies and it is declining, so accurate taxonomic recognition critically affects conservation efforts. For all the aforementioned reasons, we propose to rank the Canarian breeding population as a new taxon by presenting the formal description of a new subspecies Puffinus puffinus canariensis ssp. nova.

The Socotra Archipelago in the Arabian Sea is considered one of the most geo-politically isolated landforms on earth and a center of endemism. The archipelago is located at the western edge of the Indian Ocean and comprises four islands: Socotra, Darsa, Samha, and Abd al Kuri. Here we provide an integrative study on Haemodracon geckos, the sole genus of geckos strictly endemic to the archipelago. The sympatric distribution of Haemodracon riebeckii and H. trachyrhinus on Socotra Island provides a unique opportunity to explore evolutionary relationships and speciation patterns, examining the interplay between possible sympatric and allopatric scenarios. We used molecular data for phylogenetic inference, species delimitation analyses, and to infer the diversification timeframe. Multivariate statistics were used to analyze morphological data. Ecological comparisons were explored for macro-niches using species distribution models and observations were used for micro-habitat use. Haemodracon species exhibit great levels of intraspecific genetic diversity. Our calibration estimates revealed that Haemodracon diverged from its closest relative, the mainland genus Asaccus, in the Eocene, before the detachment of the archipelago. The two Haemodracon species diversified in situ on Socotra Island during the Middle Miocene, after the archipelago's isolation, into the two reciprocally monophyletic recognized species. Their divergence is associated mostly with remarkable body size differences and micro-habitat segregation, with low levels of climatic and body shape divergences within their sympatric distributions. These results display how ecological, sympatric speciation, and allopatric speciation followed by secondary contact, may both have varying roles at different evolutionary phases., This work was funded by grants CGL2012-36970 and CGL2015-70390-P from the Ministerio de Economía y Competitividad, Spain (cofunded by FEDER) and grant 2014-SGR-1532 from the Secretaria d’Universitats i Recerca del Departament d’Economia i Coneixement de la Generalitat de Catalunya. MSR is funded by a FPI grant from the Ministerio de Economía y Competitividad, Spain (BES-2013-064248). JGP is supported by a “Juan de la Cierva Fellowship” (reference FJCI-2014-20380) from the Ministerio de Economía y Competitividad (Spanish Government). RV was supported by grant (SFRH/BPD/79913/2011) funded by FCT/MEC and POPH/QREN/FSE programs.

In this study we used the complete fauna of geckos of the Socotra Archipelago to test whether the three gecko genera co-occurring in the islands (Pristurus, Hemidactylus and Haemodracon) produced similar outcomes of morphological and climatic diversification. To test this, we produced a time-calibrated tree of 346 geckos including all 16 endemic species of the archipelago and 26 potential close-relatives in the continent. Our dating estimates revealed that most of the diversity of geckos in the archipelago was the consequence of in situ diversification. However not all genera shared similar patterns of diversification. While in Hemidactylus and Haemodracon this involved great differences in body size and low levels of climatic diversification (mostly involving sympatric distributions), an opposite pattern appeared in Pristurus in which most of the diversification involved shifts in climatic envelopes (mostly involving allopatric and parapatric distributions) but almost no size differentiation. Consistently with this, Pristurus was the only genus in which rates of size diversification in islands were substantially lower than in the continent. This illustrates how different groups can greatly differ in their patterns of intra-island diversification and highlights the importance of taxon-dependent factors at determining different patterns of diversification in the same insular context., This work was funded by CGL2012-36970 from the Ministerio de Economía y Competitividad, Spain (co-funded by FEDER).

Mosses are amongst the oldest and simplest plants, they can be found almost everywhere in the world, and they condition the structure and function of many ecosystems. Their sensitivity to environmental changes makes them very interesting subjects of study in ecology, and understanding them can provide insights into the evolutionary history of plants. However, the study of moss traits and their relationship with their environment is far behind that of vascular plants. We sampled 303 assemblages of aquatic and semi‐aquatic (hygrophytic) mosses growing in semi‐natural springs distributed around the northeast of the Iberian Peninsula to study how moss traits vary depending on their evolutionary history, climate and water chemistry. To do so, we analysed 30 moss species and 17 traits using phylogenetic comparative methods and an extended RLQ analysis, accounting for spatial and phylogenetic information. We hypothesized that there is a sclerophylly continuum in mosses living across a gradient of high and low water conductivity springs that may mimic sclerophylly in vascular plants that live in stressful environments. Results indicated that life‐forms and, especially, morphological traits were well preserved phylogenetically and responsive to water chemistry and climate. That combined with spatial autocorrelation in environmental variables resulted in a clustered distribution of phylogenetically closely related mosses in space. Mosses living in springs with a warm and dry climate that discharge hard water mainly presented species with needle‐like leaves, were denser, and had lower water absorption capacity. The opposite was found in cold, humid and soft water springs. Synthesis. Our results suggest that climate and water chemistry are main determinants of traits of hygrophytic mosses and of species distributions. We found evidence of a potential sclerophylly continuum in moss traits, which we hypothesize may be mainly related to physical and physiological constraints produced by water chemistry. Our findings describe moss sclerophylly in a gradient of water conductivity similar to that found in vascular plants with water availability and temperature. Further experimental studies will be required to confirm the observations found in this study. A free Plain Language Summary can be found within the Supporting Information of this article.

WOS: 000484779500005, PubMed ID: 31501432, Climatic conditions changing over time and space shape the evolution of organisms at multiple levels, including temperate lizards in the family Lacertidae. Here we reconstruct a dated phylogenetic tree of 262 lacertid species based on a supermatrix relying on novel phylogenomic datasets and fossil calibrations. Diversification of lacertids was accompanied by an increasing disparity among occupied bioclimatic niches, especially in the last 10 Ma, during a period of progressive global cooling. Temperate species also underwent a genomewide slowdown in molecular substitution rates compared to tropical and desert-adapted lacertids. Evaporative water loss and preferred temperature are correlated with bioclimatic parameters, indicating physiological adaptations to climate. Tropical, but also some populations of cool-adapted species experience maximum temperatures close to their preferred temperatures. We hypothesize these species-specific physiological preferences may constitute a handicap to prevail under rapid global warming, and contribute to explaining local lizard extinctions in cool and humid climates., Deutsche Forschungsgemeinschaft (DFG)German Research Foundation (DFG) [VE 247/11-1/MU 1760/9-1, VE 247/16-1-HO3492/6-1]; Juan de la Cierva fellowships from the Spanish 'Ministerio de Economia y Competitividad' [FJCI-2014-20380, IJCI-2016-29566]; 'Rita Levi Montalcini' program for recruitment of young researchers at the University of L'Aquila; US-National Science Foundation Emerging Frontiers program [EF-1241848]; Hassan II Academy of Sciences and Technologies (ICGVSA Project); Slovenian Research Agency Research Program [P1-0255]; Uppsala Multidisciplinary Center for Advanced Computational Science (UPPMAX) [SNIC 2017/7-275]; [NORTE-01-0145-FEDER-000007]; [ON173025 MESTD RS], We are grateful to numerous students, field assistants and technicians who supported field and laboratory work, and to G. Jones for useful comments on the manuscript. This study was supported by the Deutsche Forschungsgemeinschaft (DFG) to M.V. and J.M. (VE 247/11-1/MU 1760/9-1), and to L.R. in the framework of the "TaxonOmics" priority program (VE 247/16-1-HO3492/6-1). J.G.-P. and I. I. were supported by Juan de la Cierva fellowships from the Spanish 'Ministerio de Economia y Competitividad' (FJCI-2014-20380 and IJCI-2016-29566), D.S. by the 'Rita Levi Montalcini' program for recruitment of young researchers at the University of L'Aquila, M.A.C. by project NORTE-01-0145-FEDER-000007, B.S. and D.B.M. by the US-National Science Foundation Emerging Frontiers program (EF-1241848), J.C.I. by project ON173025 MESTD RS, T.S., A.F., and A.S. by the Hassan II Academy of Sciences and Technologies (ICGVSA Project), A.Z. by the Slovenian Research Agency Research Program P1-0255. The computations were in part performed on the Altamira supercomputer at the Institute of Physics of Cantabria (IFCA-CSIC), Spain; the Uppsala Multidisciplinary Center for Advanced Computational Science (UPPMAX) under Project SNIC 2017/7-275; as well as the Zentraler Informations-und Datenverarbeitungsservice of the Tierarztliche Hochschule Hannover (IDS-TiHo). Further, we acknowledge the Viper High Performance Computing facility of the University of Hull and its support team, especially Ahmed Elnawasany, for facilitating computational analyses.

Flying squirrels are the only group of gliding mammals with a remarkable diversity and wide geographical range. However, their evolutionary story is not well known. Thus far, identification of extinct flying squirrels has been exclusively based on dental features, which, contrary to certain postcranial characters, are not unique to them. Therefore, fossils attributed to this clade may indeed belong to other squirrel groups. Here we report the oldest fossil skeleton of a flying squirrel (11.6 Ma) that displays the gliding-related diagnostic features shared by extant forms and allows for a recalibration of the divergence time between tree and flying squirrels. Our phylogenetic analyses combining morphological and molecular data generally support older dates than previous molecular estimates (~23 Ma), being congruent with the inclusion of some of the earliest fossils (~36 Ma) into this clade. They also show that flying squirrels experienced little morphological change for almost 12 million years., eLife digest Mammals can walk, hop, swim and fly; a few, like marsupial sugar gliders or colugos, can even glide. With 52 species scattered across the Northern hemisphere, flying squirrels are by far the most successful group that adopted this way of going airborne. To drift from tree to tree, these small animals pack their own ‘parachute’: a membrane draping between their lower limbs and the long cartilage rods that extend from their wrists. Tiny specialized wrist bones, which are unique to flying squirrels, help to support the cartilaginous extensions. The origin of flying squirrels is a point of contention: while most genetic studies point towards the group splitting from tree squirrels about 23 million years ago, the oldest remains – mostly cheek teeth – suggest the animals were already soaring through forests 36 million years ago. However, recent studies show that the dental features used to distinguish between gliding and non-gliding squirrels may actually be shared by the two groups. In 2002, the digging of a dump site in Barcelona unearthed a peculiar skeleton: first a tail and two thigh bones, big enough that the researchers thought it could be the fossil of a small primate. In fact, and much to the disappointment of paleoprimatologists, further excavating revealed that it was a rodent. As the specimen – nearly an entire skeleton – was being prepared, paleontologists insisted that all the ‘dirt’ attached to the bones had to be carefully screen-washed. From the mud emerged the minuscule specialized wrist bones: the primate-turned-rodent was in fact Miopetaurista neogrivensis, an extinct flying squirrel. Here, Casanovas-Vilar et al. describe the 11.6 million years old fossil, the oldest ever found. The wrist bones reveal that the animal belongs to the group of flying squirrels that have large sizes. Evolutionary analyses that combined molecular and paleontological data demonstrated that flying squirrels evolved from tree squirrels as far back as 31 to 25 million years ago, and possibly even earlier. In addition, the results show that Miopetaurista is closely related to Petaurista, a modern group of giant flying squirrels. In fact, their skeletons are so similar that the large species that currently inhabit the tropical and subtropical forests of Asia could be considered living fossils. Molecular and paleontological data are often at odds, but this fossil shows that they can be reconciled and combined to retrace history. Discovering older fossils, or even transitional forms, could help to retrace how flying squirrels took a leap from the rest of their evolutionary tree.

Altres ajuts: MAGRAMA/OAPN-022/2008 Metabolomes, as chemical phenotypes of organisms, are likely not only shaped by the environment but also by common ancestry. If this is the case, we expect that closely related species of pines will tend to reach similar metabolomic solutions to the same environmental stressors. We examined the metabolomes of two sympatric subspecies of Pinus sylvestris in Sierra Nevada (southern Iberian Peninsula), in summer and winter and exposed to folivory by the pine processionary moth. The overall metabolomes differed between the subspecies but both tended to respond more similarly to folivory. The metabolomes of the subspecies were more dissimilar in summer than in winter, and iberica trees had higher concentrations of metabolites directly related to drought stress. Our results are consistent with the notion that certain plant metabolic responses associated with folivory have been phylogenetically conserved. The larger divergence between subspecies metabolomes in summer is likely due to the warmer and drier conditions that the northern iberica subspecies experience in Sierra Nevada. Our results provide crucial insights into how iberica populations would respond to the predicted conditions of climate change under an increased defoliation in the Mediterranean Basin.

Island colonization is often assumed to trigger extreme levels of phenotypic diversification. Yet, empirical evidence suggests that it does not always so. In this study we test this hypothesis using a completely sampled mainland-island system, the arid clade of Hemidactylus, a group of geckos mainly distributed across Africa, Arabia and the Socotra Archipelago. To such purpose, we generated a new molecular phylogeny of the group on which we mapped body size and head proportions. We then explored whether island and continental taxa shared the same morphospace and differed in their disparities and tempos of evolution. Insular species produced the most extreme sizes of the radiation, involving accelerated rates of evolution and higher disparities compared with most (but not all) of the continental groups. In contrast, head proportions exhibited constant evolutionary rates across the radiation and similar disparities in islands compared with the continent. These results, although generally consistent with the notion that islands promote high morphological disparity, reveal at the same time a complex scenario in which different traits may experience different evolutionary patterns in the same mainland-island system and continental groups do not always present low levels of morphological diversification compared to insular groups., This work was funded by CGL2012-36970 from the Ministerio de Economía y Competitividad, Spain (co-funded by FEDER). The JGP was also supported by the JAE-predoc grant of the CSIC and JS was supported by the Ministry of Culture of the Czech Republic under grant DKRVO 2016/15, National Museum, 00023272.

All data used in this study have been archived in the Dryad Digital Repository: http://dx.doi.org/10.5061/dryad.nf7697j6, Complex social communication is expected to evolve whenever animals engage in many and varied social interactions; that is, sociality should promote communicative complexity. Yet, informal comparisons among phylogenetically independent taxonomic groups seem to cast doubt on the putative role of social factors in the evolution of complex communication. Here, we provide a formal test of the sociality hypothesis alongside alternative explanations for the evolution of communicative complexity. We compiled data documenting variations in signal complexity among closely related species for several case study groups-ants, frogs, lizards and birds-and used new phylogenetic methods to investigate the factors underlying communication evolution. Social factors were only implicated in the evolution of complex visual signals in lizards. Ecology, and to some degree allometry, were most likely explanations for complexity in the vocal signals of frogs (ecology) and birds (ecology and allometry). There was some evidence for adaptive evolution in the pheromone complexity of ants, although no compelling selection pressure was identified. For most taxa, phylogenetic null models were consistently ranked above adaptive models and, for some taxa, signal complexity seems to have accumulated in species via incremental or random changes over long periods of evolutionary time. Becoming social presumably leads to the origin of social communication in animals, but its subsequent influence on the trajectory of signal evolution has been neither clear-cut nor general among taxonomic groups. © 2012 The Royal Society., J.G.P. was supported by a pre-doctoral research fellowship (JAE) from the CSIC.

Altres ajuts: MAGRAMA/OAPN-022/2008 Plants respond locally and systemically to herbivore attack. Most of the research conducted on plant-herbivore relationships at elemental and molecular levels have focused on the elemental composition or/and certain molecular compounds or specific families of defensive metabolites showing that herbivores tend to select plant individuals or species with higher nutrient concentrations and to avoid those with higher levels of defensive compounds. We performed stoichiometric and metabolomics, local and systemic, analyses in two subspecies of Pinus sylvestris under the attack by the caterpillars of the pine processionary moth, an important pest in the Mediterranean Basin. Both pine subspecies responded locally to folivory mainly by increasing the relative concentrations of terpenes and some phenolics. Systemic responses differed between subspecies and most of the metabolites presented intermediate concentrations between those of the affected parts and unattacked trees. Our results support the hypothesis that foliar nutrient concentrations are not a key factor of an alleged plant selection by adult female processionary moths for oviposition since folivory was not associated with any of the elements analyzed. Phenolic compounds did not generally increase in the attacked trees questioning thus their commonly proposed induction by folivory attack and their anti-feeding properties. Herbivory attack produced a general systemic shift in pines, including both primary and secondary metabolisms, that was less intense and chemically different from the local responses. Local pine responses were similar between subspecies while systemic responses were more distant between them.

The existence of elaborate ornamental structures in males is often assumed to reflect the outcome of female mate choice for showy males. However, female mate choice appears weak in many iguanian lizards, but males still exhibit an array of ornament-like structures around the throat. We performed a phylogenetic comparative study to assess whether these structures have originated in response to male–male competition or the need for improved signal efficiency in visually difficult environments. We found little evidence for the influence of male–male competition. Instead, forest species were more likely to exhibit colourful throat appendages than species living in open habitats, suggesting selection for signal efficiency. On at least three independent occasions, throat ornamentation has become further elaborated into a large, conspicuously coloured moving dewlap. Although the function of the dewlap is convergent, the underlying hyoid apparatus has evolved very differently, revealing the same adaptive outcome has been achieved through multiple evolutionary trajectories. More generally, our findings highlight that extravagant, ornament-like morphology can evolve in males without the direct influence of female mate choice and that failure to consider alternative hypotheses for the evolution of these structures can obscure the true origins of signal diversity among closely related taxa., This work was partly supported by funds from a UNSW SFRGP grant to M. Hagman.

Two viperid snakes, Vipera natiensis sp. n. (belonging to the Vipera aspis group) and Vipera sp., have been found in the Pliocene of Minorca. On the other hand, an indeterminate viperid extends back the fossil record of the family in the Balearic Islands, to the Middle Miocene. These fossils represent the first report of viperids from the Balearic Islands. Vertebra of Vipera natiensis sp. n. are characterised by a concave anterior border of zygosphene, dorsoventrally flattened, short (as wide as long) and blunt prezygapophyseal processes. Centrum convex in transverse section, subcentral margins poorly marked and presence of two well defined subcotylar tubercles. Hypapophysis located posteriorly on the centrum and not prolonged anteriorly, or only slightly prolonged as a keel. The dorsal border of the neural spine is slightly thickened; the posterior border of the neural spine is markedly inclined posteriorly; as a result, the spine forms a posterior point, which is more or less developed, depending on the position of the vertebra along the vertebral column. The presence of Viperidae in the Neogene of the Balearic Islands involves the existence of a terrestrial connection between this archipelago and the continent during the Early Miocene and/or Middle Miocene and perhaps the presence of a later connection during the Late Miocene.

The amphibian fauna of the Kingdom of Morocco was traditionally regarded as poor and closely related to its European counterpart. However, an increase in research during the last decades revealed a considerable degree of endemism amongst Moroccan amphibians, as well as phenotypic and genotypic inter- and intraspecific divergence. Despite this increase in knowledge, a comprehensible overview is lacking while several systematic issues have remained unresolved. We herein present a contemporary overview of the distribution, taxonomy and biogeography of Moroccan amphibians. Fourteen fieldtrips were made by the authors and colleagues between 2000 and 2012, which produced a total of 292 new distribution records. Furthermore, based on the results of the present work, we (i) review the systematics of the genus Salamandra in Morocco, including the description of a new subspecies from the Rif- and Middle Atlas Mountains, Salamandra algira splendens ssp. nov.; (ii) present data on intraspecific morphological variability of Pelobates varaldii and Pleurodeles waltl in Morocco; (iii) attempt to resolve the phylogenetic position of Bufo brongersmai and erect a new genus for this species, Barbarophryne gen. nov.; (iv) summarize and assess the availability of tadpole-specific characteristics and bioacoustical data, and (v) summarize natural history data. Copyright © 2013 Magnolia Press.

A molecular phylogeny of the sphaerodactylid geckos of the genus Pristurus is inferred based on an alignment of 1845 base pairs (bp) of concatenated mitochondrial (12S) and nuclear (acm4, cmos, rag1 and rag2) genes for 80 individuals, representing 18 of the 23-26 species, and the three subspecies of P. rupestris. The results indicate that P. rupestris is polyphyletic and includes two highly divergent clades: the eastern clade, found in coastal Iran and throughout the Hajar Mountain range in northern Oman and eastern UAE; and the western clade, distributed from central coastal Oman, through Yemen, Saudi Arabia and north to southern Jordan. Inferred haplotype networks for the four nuclear genes show that the eastern and western clades of "P. rupestris" are highly differentiated and do not share any alleles. Moreover, although the two clades are differentiated by a morphological multivariate analysis, no one character or set of characters was found to be diagnostic. Based on the molecular analysis of specimens from the type locality of P. rupestris rupestris, the name P. rupestris is applied to the eastern clade. The name that should apply to the western clade cannot be clarified until morphological and genetic data for "P. rupestris" is available from the vicinity of Bosaso, Somalia, and therefore we refer to it as Pristurus sp. 1. The phylogenetic tree of Pristurus supports the hypothesis that P. celerrimus is sister to all the other species in the analyses and that the Socotra Archipelago was independently colonized a minimum of two times.

© 2014 Elsevier Inc. The rapid development of innovative molecular tools for characterizing biodiversity is leading to an extensive and sometimes unexpected renovation of taxonomic classifications. Particularly, for species having allopatric or parapatric distributions or resulting from recent speciation processes, the absence of clear phenotypic differentiation may hinder the recognition of closely related taxa, while intraspecific polymorphism may be confused with the presence of more than one single species. In the present work, we apply different phylogenetic methods in order to infer relationships within the genus Anatololacerta, and to assess the taxonomy of this morphologically diversified group of lizards endemic to western and southern Anatolia and some neighboring Aegean islands. According to morphology, three species have been recognized (Anatololacerta anatolica, A. oertzeni and A. danfordi) as well as several subspecies, but small variation at immunological markers led some authors to join all the populations into one single taxon, A. danfordi. By selecting both mitochondrial and nuclear informative markers, we tested the effectiveness of classical >gene tree> (i.e. Bayesian Inference) vs. innovative (i.e. coalescent-based) >species tree> methods in resolving the Anatololacerta taxonomic enigma, as a case in point for similar studies on species complexes resulting from non-obvious and cryptic diversification patterns. According to our results, the gene tree method failed in resolving phylogenetic relationships among clades, whereas the multi-locus species tree approach, coupled with species delimitation methods, allowed the identification of four well distinct species. These species probably diversified in different allopatric refugia located in southern and western Anatolia, where isolated populations may have persisted during Pleistocene glacial cycles., SC is supported by grant CGL2012-36970 from the Ministerio de Economía y Competitividad, Spain (co-funded by FEDER).

All data compiled by our study have been archived in the Dryad Digital Repository http://dx.doi.org/10.5061/dryad.56vf1, The acquisition of key innovations and the invasion of new areas constitute two major processes that facilitate ecological opportunity and subsequent evolutionary diversification. Using a major lizard radiation as a model, the Australasian diplodactyloid geckos, we explored the effects of two key innovations (adhesive toepads and a snake-like phenotype) and the invasion of new environments (island colonization) in promoting the evolution of phenotypic and species diversity. We found no evidence that toepads had significantly increased evolutionary diversification, which challenges the common assumption that the evolution of toepads has been responsible for the extensive radiation of geckos. In contrast, a snakelike phenotype was associated with increased rates of body size evolution and, to a lesser extent, species diversification. However, the clearest impact on evolutionary diversification has been the colonization of New Zealand and New Caledonia, which were associated with increased rates of both body size evolution and species diversification. This highlights that colonizing new environments can drive adaptive diversification in conjunction or independently of the evolution of a key innovation. Studies wishing to confirm the putative link between a key innovation and subsequent evolutionary diversification must therefore show that it has been the acquisition of an innovation specifically, not the colonization of new areas more generally, that has prompted diversification. © 2013 European Society For Evolutionary Biology., Our study was only possible because of the morphological and molecular data collected by many researchers over many years. JGP is recipient of a predoctoral research fellowship (JAE) from the CSIC.

In most pan-Eurasiatic species complexes, two phenomena have been traditionally considered key processes of their cladogenesis and biogeography. First, it is hypothesized that the origin and development of the Central Asian Deserts generated a biogeographic barrier that fragmented past continuous distributions in Eastern and Western domains. Second, Pleistocene glaciations have been proposed as the main process driving the regional diversification within each of these domains. The European common toad and its closest relatives provide an interesting opportunity to examine the relative contributions of these paleogeographic and paleoclimatic events to the phylogeny and biogeography of a widespread Eurasiatic group. We investigate this issue by applying a multiproxy approach combining information from molecular phylogenies, a multiple correspondence analysis of allozyme data and species distribution models. Our study includes 304 specimens from 164 populations, covering most of the distributional range of the Bufo bufo species complex in the Western Palearctic. The phylogenies (ML and Bayesian analyses) were based on a total of 1988 bp of mitochondrial DNA encompassing three genes (tRNAval, 16S and ND1). A dataset with 173 species of the family Bufonidae was assembled to estimate the separation of the two pan-Eurasiatic species complexes of Bufo and to date the main biogeographic events within the Bufo bufo species complex. The allozyme study included sixteen protein systems, corresponding to 21 presumptive loci. Finally, the distribution models were based on maximum entropy. Our distribution models show that Eastern and Western species complexes are greatly isolated by the Central Asian Deserts, and our dating estimates place this divergence during the Middle Miocene, a moment in which different sources of evidence document a major upturn of the aridification rate of Central Asia. This climate-driven process likely separated the Eastern and Western species. At the level of the Western Palearctic, our dating estimates place most of the deepest phylogenetic structure before the Pleistocene, indicating that Pleistocene glaciations did not have a major role in splitting the major lineages. At a shallow level, the glacial dynamics contributed unevenly to the genetic structuring of populations, with a strong influence in the European–Caucasian populations, and a more relaxed effect in the Iberian populations., The first author is recipient of a pre-doctoral research fellowship (JAE) from the CSIC. The work was partially funded by Grants CGL2008-00827 and CGL2009-11663 from the Ministerio de Educación y Ciencia, Spain to SC. S.N. Litvinchuk was partly supported by Grants NSh-3299.2010.4 and MCB-N22n.

A molecular phylogeny of the sphaerodactylid geckos of the genus Pristurus is inferred based on an alignment of 1845 base pairs (bp) of concatenated mitochondrial (12S) and nuclear (acm4, cmos, rag1 and rag2) genes for 80 individuals, representing 18 of the 23–26 species, and the three subspecies of P. rupestris. The results indicate that P. rupestris is polyphyletic and includes two highly divergent clades: the eastern clade, found in coastal Iran and throughout the Hajar Mountain range in northern Oman and eastern UAE; and the western clade, distributed from central coastal Oman, through Yemen, Saudi Arabia and north to southern Jordan. Inferred haplotype networks for the four nuclear genes show that the eastern and western clades of “P. rupestris” are highly differentiated and do not share any alleles. Moreover, although the two clades are differentiated by a morphological multivariate analysis, no one character or set of characters was found to be diagnostic. Based on the molecular analysis of specimens from the type locality of P. rupestris rupestris, the name P. rupestris is applied to the eastern clade. The name that should apply to the western clade cannot be clarified until morphological and genetic data for “P. rupestris” is available from the vicinity of Bosaso, Somalia, and therefore we refer to it as Pristurus sp. 1. The phylogenetic tree of Pristurus supports the hypothesis that P. celerrimus is sister to all the other species in the analyses and that the Socotra Archipelago was independently colonized a minimum of two times.

All authors: Salvador Carranza , Meritxell Xipell, Pedro Tarroso, Andrew Gardner, Edwin Nicholas Arnold, Michael D. Robinson, Marc Simó-Riudalbas, Raquel Vasconcelos, Philip de Pous, Fèlix Amat, Jiří Šmíd, Roberto Sindaco, Margarita Metallinou †, Johannes Els, Juan Manuel Pleguezuelos, Luis Machado, David Donaire, Gabriel Martínez, Joan Garcia-Porta, Tomáš Mazuch, Thomas Wilms, Jürgen Gebhart, Javier Aznar, Javier Gallego, Bernd-Michael Zwanzig, Daniel Fernández-Guiberteau, Theodore Papenfuss, Saleh Al Saadi, Ali Alghafri, Sultan Khalifa, Hamed Al Farqani, Salim Bait Bilal, Iman Sulaiman Alazri, Aziza Saud Al Adhoobi, Zeyana Salim Al Omairi, Mohammed Al Shariani, Ali Al Kiyumi, Thuraya Al Sariri, Ahmed Said Al Shukaili, Suleiman Nasser Al Akhzami., In the present work, we use an exceptional database including 5,359 records of 101 species of Oman’s terrestrial reptiles together with spatial tools to infer the spatial patterns of species richness and endemicity, to infer the habitat preference of each species and to better define conservation priorities, with especial focus on the effectiveness of the protected areas in preserving this unique arid fauna. Our results indicate that the sampling effort is not only remarkable from a taxonomic point of view, with multiple observations for most species, but also for the spatial coverage achieved. The observations are distributed almost continuously across the two-dimensional climatic space of Oman defined by the mean annual temperature and the total annual precipitation and across the Principal Component Analysis (PCA) of the multivariate climatic space and are well represented within 17 out of the 20 climatic clusters grouping 10% of the explained climatic variance defined by PC1 and PC2. Species richness is highest in the Hajar and Dhofar Mountains, two of the most biodiverse areas of the Arabian Peninsula, and endemic species richness is greatest in the Jebel Akhdar, the highest part of the Hajar Mountains. Oman’s 22 protected areas cover only 3.91% of the country, including within their limits 63.37% of terrestrial reptiles and 50% of all endemics. Our analyses show that large areas of the climatic space of Oman lie outside protected areas and that seven of the 20 climatic clusters are not protected at all. The results of the gap analysis indicate that most of the species are below the conservation target of 17% or even the less restrictive 12% of their total area within a protected area in order to be considered adequately protected. Therefore, an evaluation of the coverage of the current network of protected areas and the identification of priority protected areas for reptiles using reserve design algorithms are urgently needed. Our study also shows that more than half of the species are still pending of a definitive evaluation by the International Union for Conservation of Nature (IUCN)., This work was funded by grants CGL2012-36970, CGL2015-70390-P from the Ministerio de Economía y Competitividad, Spain (cofunded by FEDER) to SC, the project Field study for the conservation of reptiles in Oman, Ministry of Environment and Climate Affairs, Oman (Ref: 22412027) to SC and grant 2014-SGR-1532 from the Secretaria d'Universitats i Recerca del Departament d'Economia i Coneixement de la Generalitat de Catalunya to SC. MSR is funded by a FPI grant from the Ministerio de Economía y Competitividad, Spain (BES-2013-064248); RV, PT and LM were funded by Fundação para a Ciência e Tecnologia (FCT) through post-doc grants (SFRH/BPD/79913/2011) to RV, (SFRH/BPD/93473/2013) to PT and PhD grant (SFRH/BD/89820/2012) to LM, financed by Programa Operacional Potencial Humano (POPH) – Quadro de Referência Estrategico Nacional (QREN) from the European Social Fund and Portuguese Ministerio da Educação e Ciência.

Islands have long been recognized as key contributors to biodiversity because they facilitate geographic isolation and ecological divergence from mainland ancestors. However, island colonization has traditionally been considered an evolutionary dead-end process, and its consequences for continental biodiversity remain understudied. Here, we use the evolutionary radiation of Columbiformes (i.e. pigeons and doves) to examine if ecological niche shifts on islands shaped biological diversification and community composition on continents. We show that the colonization of islands by continental, terrestrial-foraging lineages led to the exploitation of a new ecological niche (i.e. arboreal foraging). This transition towards arboreal foraging was associated with evolutionary adaptation towards a new morphological optimum. In addition, arboreal-foraging lineages of islands experienced an increase in speciation rates, which was associated with successful range expansions to other islands as well as back colonization of continents. Our results provide empirical evidence that diversification on continents can only be fully understood when studying the diversification processes that took place on islands, challenging the view of islands as mere sinks of evolutionary diversity.