Vulnerability of terrestrial vertebrate food webs to anthropogenic threats in Europe.

Vertebrate species worldwide are currently facing significant declines in many populations. Although we have gained substantial knowledge about the direct threats that affect individual species, these threats only represent a fraction of the broader vertebrate threat profile, which is also shaped by species interactions. For example, threats faced by prey species can jeopardize the survival of their predators due to food resource scarcity. Yet, indirect threats arising from species interactions have received limited investigation thus far. In this study, we investigate the indirect consequences of anthropogenic threats on biodiversity in the context of European vertebrate food webs. We integrated data on trophic interactions among over 800 terrestrial vertebrates, along with their associated human-induced threats. We quantified and mapped the vulnerability of various components of the food web, including species, interactions, and trophic groups to six major threats: pollution, agricultural intensification, climate change, direct exploitation, urbanization, and invasive alien species and diseases. Direct exploitation and agricultural intensification were two major threats for terrestrial vertebrate food webs: affecting 34% and 31% of species, respectively, they threaten 85% and 69% of interactions in Europe. By integrating network ecology with threat impact assessments, our study contributes to a better understanding of the magnitude of anthropogenic impacts on biodiversity.

Land-use intensity influences European tetrapod food webs.

Land use intensification favours particular trophic groups which can induce architectural changes in food webs. These changes can impact ecosystem functions, services, stability and resilience. However, the imprint of land management intensity on food-web architecture has rarely been characterized across large spatial extent and various land uses. We investigated the influence of land management intensity on six facets of food-web architecture, namely apex and basal species proportions, connectance, omnivory, trophic chain lengths and compartmentalization, for 67,051 European terrestrial vertebrate communities. We also assessed the dependency of this influence of intensification on land use and climate. In addition to more commonly considered climatic factors, the architecture of food webs was notably influenced by land use and management intensity. Intensification tended to strongly lower the proportion of apex predators consistently across contexts. In general, intensification also tended to lower proportions of basal species, favoured mesopredators, decreased food webs compartmentalization whereas it increased their connectance. However, the response of food webs to intensification was different for some contexts. Intensification sharply decreased connectance in Mediterranean and Alpine settlements, and it increased basal tetrapod proportions and compartmentalization in Mediterranean forest and Atlantic croplands. Besides, intensive urbanization especially favoured longer trophic chains and lower omnivory. By favouring mesopredators in most contexts, intensification could undermine basal tetrapods, the cascading effects of which need to be assessed. Our results support the importance of protecting top predators where possible and raise questions about the long-term stability of food webs in the face of human-induced pressures.

Increase of nesting habitat suitability for green turtles in a warming Mediterranean Sea.

Climate change is reshaping global ecosystems at an unprecedented rate, with major impacts on biodiversity. Therefore, understanding how organisms can withstand change is key to identify priority conservation objectives. Marine ectotherms are being extremely impacted because their biology and phenology are directly related to temperature. Among these species, sea turtles are particularly problematic because they roam over both marine and terrestrial habitats throughout their life cycles. Focusing on green turtles (Chelonia mydas) in the Mediterranean Sea, we investigated the future potential changes of nesting grounds through time, assuming that marine turtles would shift their nesting locations. We modeled the current distribution of nesting grounds including both terrestrial and marine variables, and we projected the potential nesting distribution across the Mediterranean basin under alternative future greenhouse gas emission scenario (2000-2100). Our models show an increase in nesting probability in the western Mediterranean Sea, irrespective of the climate scenario we consider. Contrary to what is found in most global change studies, the worse the climate change scenario, the more suitable areas for green turtles will potentially increase. The most important predictors were anthropogenic variables, which negatively affect nesting probability, and sea surface temperature, positively linked to nesting probability, up to a maximum of 24-25 °C. The importance of the western Mediterranean beaches as potential nesting areas for sea turtles in the near future clearly call for a proactive conservation and management effort, focusing on monitoring actions (to document the potential range expansion) and threat detection.

Dissimilarity of vertebrate trophic interactions reveals spatial uniqueness but functional redundancy across Europe.

Identifying areas that contain species assemblages not found elsewhere in a region is central to conservation planning.1,2 Species assemblages contain networks of species interactions that underpin species dynamics,3,4 ecosystem processes, and contributions to people.5,6,7 Yet the uniqueness of interaction networks in a regional context has rarely been assessed. Here, we estimated the spatial uniqueness of 10,000 terrestrial vertebrate trophic networks across Europe (1,164 species, 50,408 potential interactions8) based on the amount of similarity between all local networks mapped at a 10 km resolution. Our results revealed more unique networks in the Arctic bioregion, but also in southern Europe and isolated islands. We then contrasted the uniqueness of trophic networks with their vulnerability to human footprint and future climate change and measured their coverage within protected areas. This analysis revealed that unique networks situated in southern Europe were particularly exposed to human footprint and that unique networks in the Arctic might be at risk from future climate change. However, considering interaction networks at the level of trophic groups, rather than species, revealed that the general structure of trophic networks was redundant across the continent, in contrast to species' interactions. We argue that proactive European conservation strategies might gain relevance by turning their eyes toward interaction networks that are both unique and vulnerable.

Mediterranean fish communities are struggling to adapt to global warming. Evidence from the western coast of Italy.

Climate change has significant impacts on marine ecosystems, resulting in disruptions in biological interactions, shifts in community composition, and changes in the physiology of fish and other marine organisms. In this study conducted in the central Mediterranean Sea, the mean temperature of the catch (MTC) was employed as an indicator to investigate the climatological factors influencing the fish community. The MTC, which utilizes species-preferred temperatures, was calculated using bottom temperature (BT) data weighted against scientific catches. The estimated MTC increasing rates were 0.01 °C year-1 for the entire community, 0.017 °C year-1 for the shelf break, and 0.004 °C year-1 for the continental slope assemblage. We found that MTC is increasing at a lower rate compared to BT, suggesting a progressive under-adaptation of the fish community that seems not fully able to keep up with the ongoing pace of warming. The study identified sea surface temperature and bottom temperature as key drivers of changes in fish community composition. Notably, the fish community composition exhibited drastic changes over the studied period, and we suggest that the MTC can be a useful index to monitor such changes within the context of the EU's climate change adaptation strategy.

A dataset on African bats' functional traits.

Trait-based approaches are becoming extremely common in ecological modeling and the availability of traits databases is increasing. However, data availability is often biased towards particular regions and taxa, with many taxa (e.g., bats) often under-represented. Here, we present the AfroBaT dataset, a compilation of trait data on 320 African bat species containing 76,914 values for 86 traits focusing on morphology, reproduction, life-history, trophic ecology, and species distributions. All data were gathered from published literature following the ecological trait-data standard procedure. Missing data for both numerical and categorical traits were imputed with a machine learning approach including species phylogeny. Trophic ecology traits showed the highest coverage in the literature (72% of the species averaged over all traits), while reproductive traits the lowest. Our data imputation improved the coverage of AfroBaT especially for reproductive traits, going from 27% to 58% of the species covered. AfroBaT has a range of potential applications in macroecology and community ecology, and the availability of open-access data on African bats will enable collaboration and data-sharing among researchers.

Conical and sabertoothed cats as an exception to craniofacial evolutionary allometry.

Among evolutionary trends shaping phenotypic diversity over macroevolutionary scales, CREA (CRaniofacial Evolutionary Allometry) describes a tendency, among closely related species, for the smaller-sized of the group to have proportionally shorter rostra and larger braincases. Here, we used a phylogenetically broad cranial dataset, 3D geometric morphometrics, and phylogenetic comparative methods to assess the validity and strength of CREA in extinct and living felids. To test for the influence of biomechanical constraints, we quantified the impact of relative canine height on cranial shape evolution. Our results provided support to CREA at the family level. Yet, whereas felines support the rule, big cats, like Pantherinae and Machairodontinae, conform weakly if not at all with CREA predictions. Our findings suggest that Machairodontinae constitute one of the first well-supported exceptions to this biological rule currently known, probably in response to the biomechanical demands and developmental changes linked with their peculiar rostral adaptations. Our results suggest that the acquisition of extreme features concerning biomechanics, evo-devo constraints, and/or ecology is likely to be associated with peculiar patterns of morphological evolution, determining potential exceptions to common biological rules, for instance, by inducing variations in common patterns of evolutionary integration due to heterochronic changes under ratchet-like evolution.

Operationalizing One Health: Environmental Solutions for Pandemic Prevention.

Human pressure on the environment is increasing the frequency, diversity, and spatial extent of disease outbreaks. Despite international recognition, the interconnection between the health of the environment, animals, and humans has been historically overlooked. Past and current initiatives have often neglected prevention under the One Health preparedness cycle, largely focusing on post-spillover stages. We argue that pandemic prevention initiatives have yet to produce actionable targets and indicators, connected to overarching goals, like it has been done for biodiversity loss and climate change. We show how the Driver-Pressure-State-Impact-Response framework, already employed by the Convention on Biological Diversity, can be repurposed to operationalize pandemic prevention. Global responses for pandemic prevention should strive for complementarity and synergies among initiatives, better articulating prevention under One Health. Without agreed-upon goals underpinning specific targets and interventions, current global efforts are unlikely to function at the speed and scale necessary to decrease the risk of disease outbreaks that might lead to pandemics. Threats to the environment are not always abatable, but decreasing the likelihood that environmental pressure leads to pandemics, and developing strategies to mitigate these impacts, are both attainable goals.

Not only climate: The importance of biotic interactions in shaping species distributions at macro scales.

Abiotic factors are usually considered key drivers of species distribution at macro scales, while biotic interactions are mostly used at local scales. A few studies have explored the role of biotic interactions at macro scales, but all considered a limited number of species and obligate interactions. We examine the role of biotic interactions in large-scale SDMs by testing two main hypotheses: (1) biotic factors in SDMs can have an important role at continental scale; (2) the inclusion of biotic factors in large-scale SDMs is important also for generalist species. We used a maximum entropy algorithm to model the distribution of 177 bat species in Africa calibrating two SDMs for each species: one considering only abiotic variables (noBIO-SDMs) and the other (BIO-SDMs) including also biotic variables (trophic resource richness). We focused the interpretation of our results on variable importance and response curves. For each species, we also compared the potential distribution measuring the percentage of change between the two models in each pixel of the study area. All models gave AUC >0.7, with values on average higher in BIO-SDMs compared to noBIO-SDMs. Trophic resources showed an importance overall higher level than all abiotic predictors in most of the species (~68%), including generalist species. Response curves were highly interpretable in all models, confirming the ecological reliability of our models. Model comparison between the two models showed a change in potential distribution for more than 80% of the species, particularly in tropical forests and shrublands. Our results highlight the importance of considering biotic interactions in SDMs at macro scales. We demonstrated that a generic biotic proxy can be important for modeling species distribution when species-specific data are not available, but we envision that a multi-scale analysis combined with a better knowledge of the species might provide a better understanding of the role of biotic interactions.

The diversity of biotic interactions complements functional and phylogenetic facets of biodiversity.

Taxonomic, functional, and phylogenetic diversities are important facets of biodiversity. Studying them together has improved our understanding of community dynamics, ecosystem functioning, and conservation values.1-3 In contrast to species, traits, and phylogenies, the diversity of biotic interactions has so far been largely ignored as a biodiversity facet in large-scale studies. This neglect represents a crucial shortfall because biotic interactions shape community dynamics, drive important aspects of ecosystem functioning,4-7 provide services to humans, and have intrinsic conservation value.8,9 Hence, the diversity of interactions can provide crucial and unique information with respect to other diversity facets. Here, we leveraged large datasets of trophic interactions, functional traits, phylogenies, and spatial distributions of >1,000 terrestrial vertebrate species across Europe at a 10-km resolution. We computed the diversity of interactions (interaction diversity [ID]) in addition to functional diversity (FD) and phylogenetic diversity (PD). After controlling for species richness, surplus and deficits of ID were neither correlated with FD nor with PD, thus representing unique and complementary information to the commonly studied facets of diversity. A three-dimensional mapping allowed for visualizing different combinations of ID-FD-PD simultaneously. Interestingly, the spatial distribution of these diversity combinations closely matched the boundaries between 10 European biogeographic regions and revealed new interaction-rich areas in the European Boreal region and interaction-poor areas in Central Europe. Our study demonstrates that the diversity of interactions adds new and ecologically relevant information to multifacetted, large-scale diversity studies with implications for understanding eco-evolutionary processes and informing conservation planning.

Addressing the Eltonian shortfall with trait-based interaction models.

We have very limited knowledge of how species interact in most communities and ecosystems despite trophic relationships being fundamental for linking biodiversity to ecosystem functioning. A promising approach to fill this gap is to predict interactions based on functional traits, but many questions remain about how well we can predict interactions for different taxa, ecosystems and amounts of input data. Here, we built a new traits-based model of trophic interactions for European vertebrates and found that even models calibrated with 0.1% of the interactions (100 out of 71 k) estimated the full European vertebrate food web reasonably well. However, predators were easier to predict than prey, especially for some clades (e.g. fowl and storks) and local food web connectance was consistently overestimated. Our results demonstrate the ability to rapidly generate food webs when empirical data are lacking-an important step towards a more complete and spatially explicit description of food webs.

Balancing conservation priorities for nature and for people in Europe.

There is an urgent need to protect key areas for biodiversity and nature's contributions to people (NCP). However, different values of nature are rarely considered together in conservation planning. Here, we explore potential priority areas in Europe for biodiversity (all terrestrial vertebrates) and a set of cultural and regulating NCP while considering demand for these NCP. We quantify the spatial overlap between these priorities and their performance in representing different values of nature. We show that different priorities rarely coincide, except in certain irreplaceable ecosystems. Notably, priorities for biodiversity better represent NCP than the reverse. Theoretically, protecting an extra 5% of land has the potential to double conservation gains for biodiversity while also maintaining some essential NCP, leading to co-benefits for both nature and people.

Testing the occurrence of convergence in the craniomandibular shape evolution of living carnivorans.

Convergence consists in the independent evolution of similar traits in distantly related species. The mammalian craniomandibular complex constitutes an ideal biological structure to investigate ecomorphological dynamics and the carnivorans, due to their phenotypic variability and ecological flexibility, offer an interesting case study to explore the occurrence of convergent evolution. Here, we applied multiple pattern-based metrics to test the occurrence of convergence in the craniomandibular shape of extant carnivorans. To this aim, we tested for convergence in many dietary groups and analyzed several cases of carnivoran convergence concerning either ecologically equivalent species or ecologically similar species of different body sizes described in the literature. Our results validate the occurrence of convergence in ecologically equivalent species in a few cases (as well as in the case of giant and red pandas), but almost never support the occurrence of convergent evolution in dietary categories of living carnivorans. Therefore, convergent evolution in this clade appears to be a rare phenomenon. This is probably the consequence of a complex interplay of one-to-many, many-to-one, and many-to-many relationships taking place between ecology, biomechanics, and morphology.

Reconstructing hotspots of genetic diversity from glacial refugia and subsequent dispersal in Italian common toads (Bufo bufo).

Genetic diversity feeds the evolutionary process and allows populations to adapt to environmental changes. However, we still lack a thorough understanding of why hotspots of genetic diversity are so 'hot'. Here, we analysed the relative contribution of bioclimatic stability and genetic admixture between divergent lineages in shaping spatial patterns of genetic diversity in the common toad Bufo bufo along the Italian peninsula. We combined population genetic, phylogeographic and species distribution modelling (SDM) approaches to map ancestral areas, glacial refugia, and secondary contact zones. We consistently identified three phylogeographic lineages, distributed in northern, central and southern Italy. These lineages expanded from their ancestral areas and established secondary contact zones, before the last interglacial. SDM identified widespread glacial refugia in peninsular Italy, sometimes located under the present-day sea-level. Generalized linear models indicated genetic admixture as the only significant predictor of the levels of population genetic diversity. Our results show that glacial refugia contributed to preserving both levels and patterns of genetic diversity across glacial-interglacial cycles, but not to their formation, and highlight a general principle emerging in Mediterranean species: higher levels of genetic diversity mark populations with substantial contributions from multiple genetic lineages, irrespective of the location of glacial refugia.

The spatial scaling of food web structure across European biogeographical regions.

The species-area relationship (SAR) is one of the most well-established scaling patterns in ecology. Its implications for understanding how communities change across spatial gradients are numerous, including the effects of habitat loss on biodiversity. However, ecological communities are not mere collections of species. They are the result of interactions between these species forming complex networks that tie them together. Should we aim to grasp the spatial scaling of biodiversity as a whole, it is fundamental to understand the changes in the structure of interaction networks with area. In spite of a few empirical and theoretical studies that address this challenge, we still do not know much about how network structure changes with area, or what are the main environmental drivers of these changes. Here, using the meta-network of potential interactions between all terrestrial vertebrates in Europe (1140 species and 67 201 feeding interactions), we analysed network-area relationships (NARs) that summarize how network properties scale with area. We do this across ten biogeographical regions, which differ in environmental characteristics. We found that the spatial scaling of network complexity strongly varied across biogeographical regions. However, once the variation in SARs was accounted for, differences in the shape of NARs vanished. On the other hand, the proportion of species across trophic levels remained remarkably constant across biogeographical regions and spatial scales, despite the great variation in species richness. Spatial variation in mean annual temperature and habitat clustering were the main environmental determinants of the shape of both SARs and NARs across Europe. Our results suggest new avenues in the exploration of the effects of environmental factors on the spatial scaling of biodiversity. We argue that NARs can provide new insights to analyse and understand ecological communities.

A Major Change in Rate of Climate Niche Envelope Evolution during Hominid History.

Homo sapiens is the only species alive able to take advantage of its cognitive abilities to inhabit almost all environments on Earth. Humans are able to culturally construct, rather than biologically inherit, their occupied climatic niche to a degree unparalleled within the animal kingdom. Precisely, when hominins acquired such an ability remains unknown, and scholars disagree on the extent to which our ancestors shared this same ability. Here, we settle this issue using fine-grained paleoclimatic data, extensive archaeological data, and phylogenetic comparative methods. Our results indicate that whereas early hominins were forced to live under physiologically suitable climatic conditions, with the emergence of H. heidelbergensis, the Homo climatic niche expanded beyond its natural limits, despite progressive harshening in global climates. This indicates that technological innovations providing effective exploitation of cold and seasonal habitats predated the emergence of Homo sapiens.

Meta-Omics Reveals Genetic Flexibility of Diatom Nitrogen Transporters in Response to Environmental Changes.

Diatoms (Bacillariophyta), one of the most abundant and diverse groups of marine phytoplankton, respond rapidly to the supply of new nutrients, often out-competing other phytoplankton. Herein, we integrated analyses of the evolution, distribution, and expression modulation of two gene families involved in diatom nitrogen uptake (DiAMT1 and DiNRT2), in order to infer the main drivers of divergence in a key functional trait of phytoplankton. Our results suggest that major steps in the evolution of the two gene families reflected key events triggering diatom radiation and diversification. Their expression is modulated in the contemporary ocean by seawater temperature, nitrate, and iron concentrations. Moreover, the differences in diversity and expression of these gene families throughout the water column hint at a possible link with bacterial activity. This study represents a proof-of-concept of how a holistic approach may shed light on the functional biology of organisms in their natural environment.

Standards for distribution models in biodiversity assessments.

Demand for models in biodiversity assessments is rising, but which models are adequate for the task? We propose a set of best-practice standards and detailed guidelines enabling scoring of studies based on species distribution models for use in biodiversity assessments. We reviewed and scored 400 modeling studies over the past 20 years using the proposed standards and guidelines. We detected low model adequacy overall, but with a marked tendency of improvement over time in model building and, to a lesser degree, in biological data and model evaluation. We argue that implementation of agreed-upon standards for models in biodiversity assessments would promote transparency and repeatability, eventually leading to higher quality of the models and the inferences used in assessments. We encourage broad community participation toward the expansion and ongoing development of the proposed standards and guidelines.

Environment and evolutionary history shape phylogenetic turnover in European tetrapods.

Phylogenetic turnover quantifies the evolutionary distance among species assemblages and is central to understanding the main drivers shaping biodiversity. It is affected both by geographic and environmental distance between sites. Therefore, analyzing phylogenetic turnover in environmental space requires removing the effect of geographic distance. Here, we apply a novel approach by deciphering phylogenetic turnover of European tetrapods in environmental space after removing geographic land distance effects. We demonstrate that phylogenetic turnover is strongly structured in environmental space, particularly in ectothermic tetrapods, and is well explained by macroecological characteristics such as niche size, species richness and relative phylogenetic diversity. In ectotherms, rather recent evolutionary processes were important in structuring phylogenetic turnover along environmental gradients. In contrast, early evolutionary processes had already shaped the current structure of phylogenetic turnover in endotherms. Our approach enables the disentangling of the idiosyncrasies of evolutionary processes such as the degree of niche conservatism and diversification rates in structuring biodiversity.