Camilla Fløjgaard

Upper thermal limits of Drosophila are linked to species distributions and strongly constrained phylogenetically

Upper thermal limits vary less than lower limits among related species of terrestrial ectotherms. This pattern may reflect weak or uniform selection on upper limits, or alternatively tight evolutionary constraints. We investigated this issue in 94 Drosophila species from diverse climates and reared in a common environment to control for plastic effects that may confound species comparisons. We found substantial variation in upper thermal limits among species, negatively correlated with annual precipitation at the central point of their distribution and also with the interaction between precipitation and maximum temperature, showing that heat resistance is an important determinant of Drosophila species distributions. Species from hot and relatively dry regions had higher resistance, whereas resistance was uncorrelated with temperature in wetter regions. Using a suite of analyses we showed that phylogenetic signal in heat resistance reflects phylogenetic inertia rather than common selection pressures. Current species distributions are therefore more likely to reflect environmental sorting of lineages rather than local adaptation. Similar to previous studies, thermal safety margins were small at low latitudes, with safety margins smallest for species occupying both humid and dry tropical environments. Thus, species from a range of environments are likely to be at risk owing to climate change. Together these findings suggest that this group of insects is unlikely to buffer global change effects through marked evolutionary changes, highlighting the importance of facilitating range shifts for maintaining biodiversity.

PHYLOGENETIC CONSTRAINTS IN KEY FUNCTIONAL TRAITS BEHIND SPECIES’ CLIMATE NICHES: PATTERNS OF DESICCATION AND COLD RESISTANCE ACROSS 95 DROSOPHILA SPECIES

Species distributions are often constrained by climatic tolerances that are ultimately determined by evolutionary history and/or adaptive capacity, but these factors have rarely been partitioned. Here, we experimentally determined two key climatic niche traits (desiccation and cold resistance) for 92–95 Drosophila species and assessed their importance for geographic distributions, while controlling for acclimation, phylogeny, and spatial autocorrelation. Employing an array of phylogenetic analyses, we documented moderate‐to‐strong phylogenetic signal in both desiccation and cold resistance. Desiccation and cold resistance were clearly linked to species distributions because significant associations between traits and climatic variables persisted even after controlling for phylogeny. We used different methods to untangle whether phylogenetic signal reflected phylogenetically related species adapted to similar environments or alternatively phylogenetic inertia. For desiccation resistance, weak phylogenetic inertia was detected; ancestral trait reconstruction, however, revealed a deep divergence that could be traced back to the genus level. Despite drosophilids’ high evolutionary potential related to short generation times and high population sizes, cold resistance was found to have a moderate‐to‐high level of phylogenetic inertia, suggesting that evolutionary responses are likely to be slow. Together these findings suggest species distributions are governed by evolutionarily conservative climate responses, with limited scope for rapid adaptive responses to future climate change.

Climate, history and neutrality as drivers of mammal beta diversity in Europe: insights from multiscale deconstruction

1. Environmental sorting, historical factors and neutral dynamics may all drive beta diversity (change in species composition across space), but their relative importance remains unresolved. In the case of European mammals, key potential drivers of large-scale beta diversity include current climate, neutral dynamics and two historical factors: Pleistocene glaciations and peninsular dynamics (immigration from extra-regional eastern faunal source areas and inter-linked relictual survival and evolutionary differentiation in isolated areas). 2. We assessed the relative importance of these drivers using a novel analytical framework to deconstruct beta diversity of non-volant mammals in Europe (138 species) into its turnover (change in species composition because of species replacements) and nestedness components (change in species composition because of species richness differences) at continental and regional (250,000 km(2) ) scales. 3. We found continental-scale mammal beta diversity to be mainly caused by spatial turnover (99·9%), with only a small contribution (0·1%) from nestedness. 4. Current climate emerged as an important driver of beta diversity, given the strong continental-scale turnover, particularly in north-south direction, i.e., in line with the latitudinal climate gradient, and, more directly, the strong correlation of climate with spatial turnover at both continental and regional scales. 5. However, there was also evidence for the importance of non-climatic drivers. Notably, the compositional variation purely accounted for by space was greater than that purely accounted for by environment for both the turnover and the nestedness component of beta diversity. Furthermore, the strong longitudinal turnover within Southern Europe is in accordance with the regions long-term climatic stability having allowed multiple refugia and local evolutionary diversification. As expected from peninsular dynamics, there was increasing dissimilarity with geographic distance in an east-west direction because of nestedness, but only in Central and Northern Europe. 6. In conclusion, European mammal beta diversity mainly reflects spatial turnover and only to a limited extent nestedness and is driven by current climate in combination with historical - and perhaps, neutral - dynamics. These findings suggest that a key challenge for climate-change predictive studies will be taking the influence of non-climatic factors into account.

Mammal predator and prey species richness are strongly linked at macroscales

Predator-prey interactions play an important role for species composition and community dynamics at local scales, but their importance in shaping large-scale gradients of species richness remains unexplored. Here, we use global range maps, structural equation models (SEM), and comprehensive databases of dietary preferences and body masses of all terrestrial, non-volant mammals worldwide, to test whether (1) prey bottom-up or predator top-down relationships are important drivers of broad-scale species richness gradients once the environment and human influence have been accounted for, (2) predator-prey richness associations vary among biogeographic regions, and (3) body size influences large-scale covariation between predators and prey. SEMs including only productivity, climate, and human factors explained a high proportion of variance in prey richness (R2=0.56) but considerably less in predator richness (R2=0.13). Adding predator-to-prey or prey-to-predator paths strongly increased the explained variance in both cases (prey R2=0.79, predator R2=0.57), suggesting that predator-prey interactions play an important role in driving global diversity gradients. Prey bottom-up effects prevailed over productivity, climate, and human influence to explain predator richness, whereas productivity and climate were more important than predator top-down effects for explaining prey richness, although predator top-down effects were still significant. Global predator-prey associations were not reproduced in all regions, indicating that distinct paleoclimate and evolutionary histories (Africa and Australia) may alter species interactions across trophic levels. Stronger cross-trophic-level associations were recorded within categories of similar body size (e.g., large prey to large predators) than between them (e.g., large prey to small predators), suggesting that mass-related energetic and physiological constraints influence broad-scale richness links, especially for large-bodied mammals. Overall, our results support the idea that trophic interactions can be important drivers of large-scale species richness gradients in combination with environmental effects.

Establishing macroecological trait datasets: digitalization, extrapolation, and validation of diet preferences in terrestrial mammals worldwide

Ecological trait data are essential for understanding the broad-scale distribution of biodiversity and its response to global change. For animals, diet represents a fundamental aspect of species’ evolutionary adaptations, ecological and functional roles, and trophic interactions. However, the importance of diet for macroevolutionary and macroecological dynamics remains little explored, partly because of the lack of comprehensive trait datasets. We compiled and evaluated a comprehensive global dataset of diet preferences of mammals (“MammalDIET”). Diet information was digitized from two global and cladewide data sources and errors of data entry by multiple data recorders were assessed. We then developed a hierarchical extrapolation procedure to fill-in diet information for species with missing information. Missing data were extrapolated with information from other taxonomic levels (genus, other species within the same genus, or family) and this extrapolation was subsequently validated both internally (with a jack-knife approach applied to the compiled species-level diet data) and externally (using independent species-level diet information from a comprehensive continentwide data source). Finally, we grouped mammal species into trophic levels and dietary guilds, and their species richness as well as their proportion of total richness were mapped at a global scale for those diet categories with good validation results. The success rate of correctly digitizing data was 94%, indicating that the consistency in data entry among multiple recorders was high. Data sources provided species-level diet information for a total of 2033 species (38% of all 5364 terrestrial mammal species, based on the IUCN taxonomy). For the remaining 3331 species, diet information was mostly extrapolated from genus-level diet information (48% of all terrestrial mammal species), and only rarely from other species within the same genus (6%) or from family level (8%). Internal and external validation showed that: (1) extrapolations were most reliable for primary food items; (2) several diet categories (“Animal”, “Mammal”, “Invertebrate”, “Plant”, “Seed”, “Fruit”, and “Leaf”) had high proportions of correctly predicted diet ranks; and (3) the potential of correctly extrapolating specific diet categories varied both within and among clades. Global maps of species richness and proportion showed congruence among trophic levels, but also substantial discrepancies between dietary guilds. MammalDIET provides a comprehensive, unique and freely available dataset on diet preferences for all terrestrial mammals worldwide. It enables broad-scale analyses for specific trophic levels and dietary guilds, and a first assessment of trait conservatism in mammalian diet preferences at a global scale. The digitalization, extrapolation and validation procedures could be transferable to other trait data and taxa.

Potential 21st century changes to the mammal fauna of Denmark ? implications of climate change, land-use, and invasive species

The moderate temperature increase of 0.74 °C in the 20th century has caused latitudinal and altitudinal range shifts in many species including mammals. Therefore, given the more dramatic temperature increase predicted for the 21st century, we can therefore expect even stronger range shifts as well. However, European mammals are already faced with other anthropogenic pressures, notably habitat loss, pollution, overexploitation, and invasive species, and will have to face the combined challenge posed by climate change in a landscape highly influenced by human activities. As an example of the possible consequences of land use, invasive species, and climate change for the regional-scale mammal species composition, we here focus on the potential 21st century changes to the mammal fauna of Denmark. Supported by species distribution modelling, we present a discussion of the possible changes to the Danish mammal fauna: Which species are likely to become locally extinct? Which new species are most likely to immigrate? And, what is the potential threat from invasive species? We find that future climate change is likely to cause a general enrichment of the Danish mammal fauna by the potential immigration of seventeen new species. Only the northern birch mouse (Sicista betulina) is at risk of extinction from climate change predicted. The European native mammals are not anticipated to contribute to the invasive-species problem as they coexist with most Danish species in other parts of Europe. However, non-European invasive species are also likely to enter the Danish fauna and may negatively impact the native species.

Big moving day for biodiversity? A macroecological assessment of the scope for assisted colonization as a conservation strategy under global warming

Future climate change constitutes a major threat to Earths biodiversity. If anthropogenic greenhouse gas emissions continue unabated, 21st century climate change is likely to exceed the natural adaptive capacity of many natural ecosystems and a large proportion of species may risk extinction. A recurrent finding is that the degree of negative impact depends strongly on the dispersal potential of the species. However, there is a growing realization that many, if not most species would be unlikely to disperse as fast and far as required. As a consequence, it has been proposed that species at risk should be actively translocated into unoccupied, but environmentally suitable areas that are likely to stay suitable over the next 100 or more years (assisted colonization or assisted migration). This solution is controversial, though, reflecting negative experiences with introduced exotics and probably also the traditional emphasis in conservation management on preserving a certain local, often historical situation with a static species composition, and a tendency among ecologists to think of biological communities as generally saturated with species. Using the European flora as a case study, we here estimate the main environmental controls of plant species richness, assess how the maximum observed species richness depends on these environmental controls, and based here on estimate how many species could at least be added to an area before further species additions would perhaps inevitably lead to corresponding losses locally. Our results suggest that there is substantial room for additional plant species across most areas of Europe, indicating that there is considerable scope for implementing assisted colonization as a proactive conservation strategy under global warming without necessarily implicating negative effects on the native flora in the areas targeted for establishment of translocated populations. Notably, our results suggest that 50% of the cells in Northern Europe, the likely target area for many translocations, could harbor at least 1/3 as many additional species as they have native species. However, we also emphasize that other, more traditional conservation strategies should also be strengthened, notably providing more space for nature and reducing nitrogen deposition to increase population resilience and facilitate unassisted colonization. Furthermore, any implementation of assisted colonization should be done cautiously, with a careful analysis on a species-by-species case.

Are ungulates in forests concerns or key species for conservation and biodiversity? Reply to Boulanger et al. (DOI: 10.1111/gcb.13899)

Increasing species richness of light demanding species in forests may not be a conservation concern if we accept a macroecological and evolutionary baseline for biodiversity. Most of the current biodiversity in Europe has evolved in the Pleistocene or earlier, and in ecosystems markedly influenced by dynamic natural processes, including grazing. Many threatened species are associated with high-light forest environments such as forest glades and edges, as these have strongly declined at least partially due to the decline of large herbivores in European forests. Hence, moderate grazing in forests should be an ecological baseline and conservation target rather than a concern.

Using dark diversity and plant characteristics to guide conservation and restoration

Summary 1.Dark diversity is a promising concept for prioritising management efforts as it focuses on species that are present in the regional pool, but locally absent even though environmental requirements are met. Currently, we lack knowledge of what characterises species belonging to the dark diversity more often than others, although this is important knowledge for restoration and conservation actions. 2.We applied the concept to a massive national (Danish) plant diversity database, containing 236,923 records from 15,160 surveys involving 564 species. This enabled the first geographically comprehensive (43,000 km2) assessment of dark diversity, at a spatial resolution relevant for conservation and restoration planning (78 m2) across multiple terrestrial habitats, thereby maximising the practical applications of this concept. The probability for a given plant species to belong to the dark diversity was computed and logistically regressed against variables representing its ecological preferences (e.g. nutrient availability), strategies (competitor, stress tolerant, ruderal), mycorrhizal relationships, establishment capacities (seed mass) and dispersal abilities. 3.Forty-six percent of the species had a high probability (>95%) of being part of dark diversity, whereas for 7% of the species this probability was less than 60%. 4.Typical dark diversity plants tended to depend on mycorrhiza, were mostly adapted to low light and low nutrient levels, had poor dispersal abilities and were ruderals and stress intolerant. 5.Synthesis and applications. Characterising species that are more often absent from suitable sites than others (dark diversity species) has important implications for the planning and management of natural ecosystems. From our study, practitioners gain insight into the factors triggering the absence of individual plant species in a seemingly suitable habitat. We highlight the need to carefully consider mycorrhizal inoculations with a suitable assemblage of fungi to promote the establishment success of dark diversity plants. Additionally, time lags in plant species dispersal and establishment as well as spatial connectivity in fragmented habitats are central to consider in nature management although assisted migration might also aid poor dispersers. Finally, nutrient-poor localities are probably important “islets” allowing nitrophobic dark diversity plant species to thrive within agricultural landscapes that are generally nutrient rich. This article is protected by copyright. All rights reserved.

Dark diversity illuminates the dim side of conservation and restoration

1 Dark diversity is a promising concept for prioritizing management efforts as it focuses on missing species, i.e., species present in the regional pool, but locally absent despite suitable environmental conditions. 2 We applied the concept to a massive national plant diversity database (236,923 records from 15,160 surveys involving 564 species) to provide the first geographically comprehensive assessment of dark diversity across a large area (43,000 km2), at a spatial scale (~75 m2) relevant for conservation and restoration planning and across multiple terrestrial habitats, thus maximising its practical application potential. The likelihood for a given plant species to belong to the dark diversity pool was computed and logistically regressed against its ecological preferences (nutrient availability, pH etc.), strategies (competitor, stress tolerance, ruderal), mycorrhizal dependence and infection percentage, seed mass and maximum dispersal distance. 3 Forty-six percent of the species were absent in >95 % of the suitable sites, whereas 7 % of the species were absent in less than 60 % of sites that were deemed suitable. 4 Species that were more likely to belong to the dark diversity tended to depend on mycorrhiza, were mostly adapted to low light and nutrient levels, had poor dispersal abilities, were ruderals and had a low stress tolerance. Synthesis and applications Our findings have important implications for the planning and management of natural ecosystems requiring detailed knowledge of what triggers the presence/absence of individual plant species in a seemingly suitable habitat. We conclude that practitioners may need to carefully consider mycorrhizal inoculations with a suitable assemblage of fungi for certain plant species to become established. Also assisted migration might be necessary to help poor dispersers although spatial and temporal processes are also important to have in mind. Finally, it is important to vary nutrient loads making room for plant species to colonise both nutrient-poor and nutrient-rich localities.