Biodiversity change under adaptive community dynamics.

Compositional change is a ubiquitous response of ecological communities to environmental drivers of global change, but is often regarded as evidence of declining "biotic integrity" relative to historical baselines. Adaptive compositional change, however, is a foundational idea in evolutionary biology, whereby changes in gene frequencies within species boost population-level fitness, allowing populations to persist as the environment changes. Here, we present an analogous idea for ecological communities based on core concepts of fitness and selection. Changes in community composition (i.e., frequencies of genetic differences among species) in response to environmental change should normally increase the average fitnessof community members. We refer to compositional changes that improve the functional match, or "fit," between organisms' traits and their environment as adaptive community dynamics. Environmental change (e.g., land-use change) commonly reduces the fit between antecedent communities and new environments. Subsequent change in community composition in response to environmental changes, however, should normally increase community-level fit, as the success of at least some constituent species increases. We argue that adaptive community dynamics are likely to improve or maintain ecosystem function (e.g., by maintaining productivity). Adaptive community responses may simultaneously produce some changes that are considered societally desirable (e.g., increased carbon storage) and others that are undesirable (e.g., declines of certain species), just as evolutionary responses within species may be deemed desirable (e.g., evolutionary rescue of an endangered species) or undesirable (e.g., enhanced virulence of an agricultural pest). When assessing possible management interventions, it is important to distinguish between drivers of environmental change (e.g., undesired climate warming) and adaptive community responses, which may generate some desirable outcomes. Efforts to facilitate, accept, or resist ecological change require separate consideration of drivers and responses, and may highlight the need to reconsider preferences for historical baseline communities over communities that are better adapted to the new conditions.

Increased dispersal explains increasing local diversity with global biodiversity declines.

The narrative of biodiversity decline in response to human impacts is overly simplistic because different aspects of biodiversity show different trajectories at different spatial scales. It is also debated whether human-caused biodiversity changes lead to subsequent, accelerating change (cascades) in ecological communities, or alternatively build increasingly robust community networks with decreasing extinction rates and reduced invasibility. Mechanistic approaches are needed that simultaneously reconcile different aspects of biodiversity change, and explore the robustness of communities to further change. We develop a trophically structured, mainland-archipelago metacommunity model of community assembly. Varying the parameters across model simulations shows that local alpha diversity (the number of species per island) and regional gamma diversity (the total number of species in the archipelago) depend on both the rate of extirpation per island and on the rate of dispersal between islands within the archipelago. In particular, local diversity increases with increased dispersal and heterogeneity between islands, but regional diversity declines because the islands become biotically similar and local one-island and few-island species are excluded (homogenisation, or reduced beta diversity). This mirrors changes observed empirically: real islands have gained species (increased local and island-scale community diversity) with increased human-assisted transfers of species, but global diversity has declined with the loss of endemic species. However, biological invasions may be self-limiting. High-dispersal, high local-diversity model communities become resistant to subsequent invasions, generating robust species-community networks unless dispersal is extremely high. A mixed-up world is likely to lose many species, but the resulting ecological communities may nonetheless be relatively robust.

Lost, gained, and regained functional and phylogenetic diversity of European mammals since 8000 years ago.

Mammals have experienced high levels of human-mediated extirpations but have also been widely introduced to new locations, and some have recovered from historic persecution. Both of these processes-losses and gains-have resulted in concern about functional losses and changes in ecological communities as new ecological states develop. The question of whether species turnover inevitably leads to declines in functional and phylogenetic diversity depends, however, on the traits and phylogenetic distinctiveness of the species that are lost, gained, or regained. Comparing ~8000 years ago with the last century, we show that extirpations and range retractions have indeed reduced the functional and phylogenetic diversity of mammals in most European regions (countries and island groups), but species recoveries and the introduction of non-native species have increased functional and phylogenetic diversity by equivalent or greater amounts in many regions. Overall, across Europe, species richness increased in 41 regions over the last 8000 years and declined in 1; phylogenetic diversity increased in 33 and declined in 12, while functional diversity results showed 20 increases and 25 decreases. The balance of losses (extirpations) and gains (introductions, range expansions) has, however, led to net increases in functional diversity on many islands, where the original diversity was low, and across most of western Europe. Historically extirpated mega- and mesofaunal species have recolonized or been reintroduced to many European regions, contributing to recent functional and phylogenetic diversity recovery. If conservation rewilding projects continue to reintroduce regionally extirpated species and domestic descendants of "extinct" species to provide replacement grazing, browsing, and predation, there is potential to generate net functional and phylogenetic diversity gains (relative to 8000 years ago) in most European regions.

A millennium of increasing diversity of ecosystems until the mid-20th century.

Land-use change is widely regarded as a simplifying and homogenising force in nature. In contrast, analysing global land-use reconstructions from the 10th to 20th centuries, we found progressive increases in the number, evenness, and diversity of ecosystems (including human-modified land-use types) present across most of the Earth's land surface. Ecosystem diversity increased more rapidly after ~1700 CE, then slowed or slightly declined (depending on the metric) following the mid-20th century acceleration of human impacts. The results also reveal increasing spatial differentiation, rather than homogenisation, in both the presence-absence and area-coverage of different ecosystem types at sub-global scales-at least, prior to the mid-20th century. Nonetheless, geographic homogenization was revealed for a subset of analyses at a global scale, reflecting the now-global presence of certain human-modified ecosystem types. Our results suggest that, while human land-use changes have caused declines in relatively undisturbed or "primary" ecosystem types, they have also driven increases in ecosystem diversity over the last millennium.

Introduced plants as novel Anthropocene habitats for insects.

Major environmental changes in the history of life on Earth have given rise to novel habitats, which gradually accumulate species. Human-induced change is no exception, yet the rules governing species accumulation in anthropogenic habitats are not fully developed. Here we propose that nonnative plants introduced to Great Britain may function as analogues of novel anthropogenic habitats for insects and mites, analysing a combination of local-scale experimental plot data and geographic-scale data contained within the Great Britain Database of Insects and their Food Plants. We find that novel plant habitats accumulate the greatest diversity of insect taxa when they are widespread and show some resemblance to plant habitats which have been present historically (based on the relatedness between native and nonnative plant species), with insect generalists colonizing from a wider range of sources. Despite reduced per-plant diversity, nonnative plants can support distinctive insect communities, sometimes including insect taxa that are otherwise rare or absent. Thus, novel plant habitats may contribute to, and potentially maintain, broader-scale (assemblage) diversity in regions that contain mixtures of long-standing and novel plant habitats.

Synergistic and antagonistic effects of land use and non-native species on community responses to climate change.

Climate change, land-use change and introductions of non-native species are key determinants of biodiversity change worldwide. However, the extent to which anthropogenic drivers of environmental change interact to affect biological communities is largely unknown, especially over longer time periods. Here, we show that plant community composition in 996 Swedish landscapes has consistently shifted to reflect the warmer and wetter climate that the region has experienced during the second half of the 20th century. Using community climatic indices, which reflect the average climatic associations of the species within each landscape at each time period, we found that species compositions in 74% of landscapes now have a higher representation of warm-associated species than they did previously, while 84% of landscapes now host more species associated with higher levels of precipitation. In addition to a warmer and wetter climate, there have also been large shifts in land use across the region, while the fraction of non-native species has increased in the majority of landscapes. Climatic warming at the landscape level appeared to favour the colonization of warm-associated species, while also potentially driving losses in cool-associated species. However, the resulting increases in community thermal means were apparently buffered by landscape simplification (reduction in habitat heterogeneity within landscapes) in the form of increased forest cover. Increases in non-native species, which generally originate from warmer climates than Sweden, were a strong driver of community-level warming. In terms of precipitation, both landscape simplification and increases in non-natives appeared to favour species associated with drier climatic conditions, to some extent counteracting the climate-driven shift towards wetter communities. Anthropogenic drivers can act both synergistically and antagonistically to determine trajectories of change in biological communities over time. Therefore, it is important to consider multiple drivers of global change when trying to understand, manage and predict biodiversity in the future.

Extinction and climate change.

Arising from F. He & S. P. Hubbell 473, 368-371 (2011). Statistical relationships between habitat area and the number of species observed (species-area relationships, SARs) are sometimes used to assess extinction risks following habitat destruction or loss of climatic suitability. He and Hubbell argue that the numbers of species confined to-rather than observed in-different areas (endemics-area relationships, EARs) should be used instead of SARs, and that SAR-based extinction estimates in the literature are too high. We suggest that He and Hubbell's SAR estimates are biased, that the empirical data they use are not appropriate to calculate extinction risks, and that their statements about extinction risks from climate change do not take into account non-SAR-based estimates or recent observations. Species have already responded to climate change in a manner consistent with high future extinction risks.

Non-native plants add to the British flora without negative consequences for native diversity.

Plants are commonly listed as invasive species, presuming that they cause harm at both global and regional scales. Approximately 40% of all species listed as invasive within Britain are plants. However, invasive plants are rarely linked to the national or global extinction of native plant species. The possible explanation is that competitive exclusion takes place slowly and that invasive plants will eventually eliminate native species (the "time-to-exclusion hypothesis"). Using the extensive British Countryside Survey Data, we find that changes to plant occurrence and cover between 1990 and 2007 at 479 British sites do not differ between native and non-native plant species. More than 80% of the plant species that are widespread enough to be sampled are native species; hence, total cover changes have been dominated by native species (total cover increases by native species are more than nine times greater than those by non-native species). This implies that factors other than plant "invasions" are the key drivers of vegetation change. We also find that the diversity of native species is increasing in locations where the diversity of non-native species is increasing, suggesting that high diversities of native and non-native plant species are compatible with one another. We reject the time-to-exclusion hypothesis as the reason why extinctions have not been observed and suggest that non-native plant species are not a threat to floral diversity in Britain. Further research is needed in island-like environments, but we question whether it is appropriate that more than three-quarters of taxa listed globally as invasive species are plants.

Climate change vulnerability for species-Assessing the assessments.

Climate change vulnerability assessments are commonly used to identify species at risk from global climate change, but the wide range of methodologies available makes it difficult for end users, such as conservation practitioners or policymakers, to decide which method to use as a basis for decision-making. In this study, we evaluate whether different assessments consistently assign species to the same risk categories and whether any of the existing methodologies perform well at identifying climate-threatened species. We compare the outputs of 12 climate change vulnerability assessment methodologies, using both real and simulated species, and validate the methods using historic data for British birds and butterflies (i.e. using historical data to assign risks and more recent data for validation). Our results show that the different vulnerability assessment methods are not consistent with one another; different risk categories are assigned for both the real and simulated sets of species. Validation of the different vulnerability assessments suggests that methods incorporating historic trend data into the assessment perform best at predicting distribution trends in subsequent time periods. This study demonstrates that climate change vulnerability assessments should not be used interchangeably due to the poor overall agreement between methods when considering the same species. The results of our validation provide more support for the use of trend-based rather than purely trait-based approaches, although further validation will be required as data become available.

Macro- and microclimatic interactions can drive variation in species' habitat associations.

Many species are more restricted in their habitat associations at the leading edges of their range margins, but some species have broadened their habitat associations in these regions during recent climate change. We examine the effects of multiple, interacting climatic variables on spatial and temporal patterns of species' habitat associations, using the speckled wood butterfly, Pararge aegeria, in Britain, as our model taxon. Our analyses reveal that this species, traditionally regarded as a woodland-dependent insect, is less restricted to woodland in regions with warmer winters and warmer and wetter summers. In addition, over the past 40 years of climate change, the species has become less restricted to woodland in locations where temperature and summer rainfall have increased most. We show that these patterns arise mechanistically because larval growth rates are slower in open (i.e. nonwoodland) habitats associated with colder microclimates in winter and greater host plant desiccation in summer. We conclude that macro- and microclimatic interactions drive variation in species' habitat associations, which for our study species resulted predominantly in a widening of habitat associations under climate change. However, species vary in their climatic and nonclimatic requirements, and so complex spatial and temporal patterns of changes in habitat associations are likely to be observed in future as the climate changes.

Refugia and connectivity sustain amphibian metapopulations afflicted by disease.

Metapopulation persistence in fragmented landscapes depends on habitat patches that can support resilient local populations and sufficient connectivity between patches. Yet epidemiological theory for metapopulations has largely overlooked the capacity of particular patches to act as refuges from disease, and has suggested that connectivity can undermine persistence. Here, we show that relatively warm and saline wetlands are environmental refuges from chytridiomycosis for an endangered Australian frog, and act jointly with connectivity to sustain frog metapopulations. We coupled models of microclimate and infection probability to map chytrid prevalence, and demonstrate a strong negative relationship between chytrid prevalence and the persistence of frog populations. Simulations confirm that frog metapopulations are likely to go extinct when they lack environmental refuges from disease and lose connectivity between patches. This study demonstrates that environmental heterogeneity can mediate host-pathogen interactions in fragmented landscapes, and provides evidence that connectivity principally supports host metapopulations afflicted by facultative pathogens.

Protected areas facilitate species' range expansions.

The benefits of protected areas (PAs) for biodiversity have been questioned in the context of climate change because PAs are static, whereas the distributions of species are dynamic. Current PAs may, however, continue to be important if they provide suitable locations for species to colonize at their leading-edge range boundaries, thereby enabling spread into new regions. Here, we present an empirical assessment of the role of PAs as targets for colonization during recent range expansions. Records from intensive surveys revealed that seven bird and butterfly species have colonized PAs 4.2 (median) times more frequently than expected from the availability of PAs in the landscapes colonized. Records of an additional 256 invertebrate species with less-intensive surveys supported these findings and showed that 98% of species are disproportionately associated with PAs in newly colonized parts of their ranges. Although colonizing species favor PAs in general, species vary greatly in their reliance on PAs, reflecting differences in the dependence of individual species on particular habitats and other conditions that are available only in PAs. These findings highlight the importance of current PAs for facilitating range expansions and show that a small subset of the landscape receives a high proportion of colonizations by range-expanding species.

Evolution on the move: specialization on widespread resources associated with rapid range expansion in response to climate change.

Generalist species and phenotypes are expected to perform best under rapid environmental change. In contrast to this view that generalists will inherit the Earth, we find that increased use of a single host plant is associated with the recent climate-driven range expansion of the UK brown argus butterfly. Field assays of female host plant preference across the UK reveal a diversity of adaptations to host plants in long-established parts of the range, whereas butterflies in recently colonized areas are more specialized, consistently preferring to lay eggs on one host plant species that is geographically widespread throughout the region of expansion, despite being locally rare. By common-garden rearing of females' offspring, we also show an increase in dispersal propensity associated with the colonization of new sites. Range expansion is therefore associated with an increase in the spatial scale of adaptation as dispersive specialists selectively spread into new regions. Major restructuring of patterns of local adaptation is likely to occur across many taxa with climate change, as lineages suited to regional colonization rather than local success emerge and expand.

Precipitation and winter temperature predict long-term range-scale abundance changes in Western North American birds.

Predicting biodiversity responses to climate change remains a difficult challenge, especially in climatically complex regions where precipitation is a limiting factor. Though statistical climatic envelope models are frequently used to project future scenarios for species distributions under climate change, these models are rarely tested using empirical data. We used long-term data on bird distributions and abundance covering five states in the western US and in the Canadian province of British Columbia to test the capacity of statistical models to predict temporal changes in bird populations over a 32-year period. Using boosted regression trees, we built presence-absence and abundance models that related the presence and abundance of 132 bird species to spatial variation in climatic conditions. Presence/absence models built using 1970-1974 data forecast the distributions of the majority of species in the later time period, 1998-2002 (mean AUC = 0.79 ± 0.01). Hindcast models performed equivalently (mean AUC = 0.82 ± 0.01). Correlations between observed and predicted abundances were also statistically significant for most species (forecast mean Spearman's ρ = 0.34 ± 0.02, hindcast = 0.39 ± 0.02). The most stringent test is to test predicted changes in geographic patterns through time. Observed changes in abundance patterns were significantly positively correlated with those predicted for 59% of species (mean Spearman's ρ = 0.28 ± 0.02, across all species). Three precipitation variables (for the wettest month, breeding season, and driest month) and minimum temperature of the coldest month were the most important predictors of bird distributions and abundances in this region, and hence of abundance changes through time. Our results suggest that models describing associations between climatic variables and abundance patterns can predict changes through time for some species, and that changes in precipitation and winter temperature appear to have already driven shifts in the geographic patterns of abundance of bird populations in western North America.

Range expansion through fragmented landscapes under a variable climate.

Ecological responses to climate change may depend on complex patterns of variability in weather and local microclimate that overlay global increases in mean temperature. Here, we show that high-resolution temporal and spatial variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. Using fine-resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat patches at the species' range margin for 27 years. Population and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual changes to population density and patch occupancy dynamics during the species' range expansion from 1982 to 2009. Our findings reveal how fine-scale, short-term environmental variability drives rates and patterns of range expansion through spatially localised, intermittent episodes of expansion and contraction. Incorporating dynamic microclimates can thus improve models of species range shifts at spatial and temporal scales relevant to conservation interventions.

Reconciling biodiversity and carbon conservation.

Climate change is leading to the development of land-based mitigation and adaptation strategies that are likely to have substantial impacts on global biodiversity. Of these, approaches to maintain carbon within existing natural ecosystems could have particularly large benefits for biodiversity. However, the geographical distributions of terrestrial carbon stocks and biodiversity differ. Using conservation planning analyses for the New World and Britain, we conclude that a carbon-only strategy would not be effective at conserving biodiversity, as have previous studies. Nonetheless, we find that a combined carbon-biodiversity strategy could simultaneously protect 90% of carbon stocks (relative to a carbon-only conservation strategy) and > 90% of the biodiversity (relative to a biodiversity-only strategy) in both regions. This combined approach encapsulates the principle of complementarity, whereby locations that contain different sets of species are prioritised, and hence disproportionately safeguard localised species that are not protected effectively by carbon-only strategies. It is efficient because localised species are concentrated into small parts of the terrestrial land surface, whereas carbon is somewhat more evenly distributed; and carbon stocks protected in one location are equivalent to those protected elsewhere. Efficient compromises can only be achieved when biodiversity and carbon are incorporated together within a spatial planning process.

Protected areas act as establishment centres for species colonizing the UK.

Protected area (PA) networks will remain valuable for conservation, as the global environment changes, if they facilitate the colonization of new regions by species that are shifting their geographical ranges. We tested the extent to which wetland bird species colonizing the UK since 1960 have exploited PAs. Colonization commenced in a PA for all six species that established permanent (greater than 10 years) breeding populations in the UK during this period. Subsequently, birds started to breed outside as well as inside PAs: the colonizing species showing declining fractions of breeding within PAs over time, a trend not seen in already-resident species. PAs were valuable as 'landing pads' for range-shifting species first arriving in a new region, and then as 'establishment centres' from which viable populations spread. Given future projections of range change across a broad range of taxonomic groups, this role for PAs can be expected to become increasingly important.