A species' response to spatial climatic variation does not predict its response to climate change.

The dominant paradigm for assessing ecological responses to climate change assumes that future states of individuals and populations can be predicted by current, species-wide performance variation across spatial climatic gradients. However, if the fates of ecological systems are better predicted by past responses to in situ climatic variation through time, this current analytical paradigm may be severely misleading. Empirically testing whether spatial or temporal climate responses better predict how species respond to climate change has been elusive, largely due to restrictive data requirements. Here, we leverage a newly collected network of ponderosa pine tree-ring time series to test whether statistically inferred responses to spatial versus temporal climatic variation better predict how trees have responded to recent climate change. When compared to observed tree growth responses to climate change since 1980, predictions derived from spatial climatic variation were wrong in both magnitude and direction. This was not the case for predictions derived from climatic variation through time, which were able to replicate observed responses well. Future climate scenarios through the end of the 21st century exacerbated these disparities. These results suggest that the currently dominant paradigm of forecasting the ecological impacts of climate change based on spatial climatic variation may be severely misleading over decadal to centennial timescales.

Drivers of future alien species impacts: An expert-based assessment.

Understanding the likely future impacts of biological invasions is crucial yet highly challenging given the multiple relevant environmental, socio-economic and societal contexts and drivers. In the absence of quantitative models, methods based on expert knowledge are the best option for assessing future invasion trajectories. Here, we present an expert assessment of the drivers of potential alien species impacts under contrasting scenarios and socioecological contexts through the mid-21st century. Based on responses from 36 experts in biological invasions, moderate (20%-30%) increases in invasions, compared to the current conditions, are expected to cause major impacts on biodiversity in most socioecological contexts. Three main drivers of biological invasions-transport, climate change and socio-economic change-were predicted to significantly affect future impacts of alien species on biodiversity even under a best-case scenario. Other drivers (e.g. human demography and migration in tropical and subtropical regions) were also of high importance in specific global contexts (e.g. for individual taxonomic groups or biomes). We show that some best-case scenarios can substantially reduce potential future impacts of biological invasions. However, rapid and comprehensive actions are necessary to use this potential and achieve the goals of the Post-2020 Framework of the Convention on Biological Diversity.

Experimental evidence of climate change extinction risk in Neotropical montane epiphytes.

Climate change is conjectured to endanger tropical species, particularly in biodiverse montane regions, but accurate estimates of extinction risk are limited by a lack of empirical data demonstrating tropical species' sensitivity to climate. To fill this gap, studies could match high-quality distribution data with multi-year transplant experiments. Here, we conduct field surveys of epiphyte distributions on three mountains in Central America and perform reciprocal transplant experiments on one mountain across sites that varied in elevation, temperature and aridity. We find that most species are unable to survive outside of their narrow elevational distributions. Additionally, our findings suggest starkly different outcomes from temperature conditions expected by 2100 under different climate change scenarios. Under temperatures associated with low-emission scenarios, most tropical montane epiphyte species will survive, but under emission scenarios that are moderately high, 5-36% of our study species may go extinct and 10-55% of populations may be lost. Using a test of tropical species' climate tolerances from a large field experiment, paired with detailed species distribution data across multiple mountains, our work strengthens earlier conjecture about risks of wide-spread extinctions from climate change in tropical montane ecosystems.

Coexistence between native and exotic species is facilitated by asymmetries in competitive ability and susceptibility to herbivores.

Differences between native and exotic species in competitive ability and susceptibility to herbivores are hypothesized to facilitate coexistence. However, little fieldwork has been conducted to determine whether these differences are present in invaded communities. Here, we experimentally examined whether asymmetries exist between native and exotic plants in a community invaded for over 200 years and whether removing competitors or herbivores influences coexistence. We found that natives and exotics exhibit pronounced asymmetries, as exotics are competitively superior to natives, but are more significantly impacted by herbivores. We also found that herbivore removal mediated the outcome of competitive interactions and altered patterns of dominance across our field sites. Collectively, these findings suggest that asymmetric biotic interactions between native and exotic plants can help to facilitate coexistence in invaded communities.

Multidimensional evaluation of managed relocation.

Managed relocation (MR) has rapidly emerged as a potential intervention strategy in the toolbox of biodiversity management under climate change. Previous authors have suggested that MR (also referred to as assisted colonization, assisted migration, or assisted translocation) could be a last-alternative option after interrogating a linear decision tree. We argue that numerous interacting and value-laden considerations demand a more inclusive strategy for evaluating MR. The pace of modern climate change demands decision making with imperfect information, and tools that elucidate this uncertainty and integrate scientific information and social values are urgently needed. We present a heuristic tool that incorporates both ecological and social criteria in a multidimensional decision-making framework. For visualization purposes, we collapse these criteria into 4 classes that can be depicted in graphical 2-D space. This framework offers a pragmatic approach for summarizing key dimensions of MR: capturing uncertainty in the evaluation criteria, creating transparency in the evaluation process, and recognizing the inherent tradeoffs that different stakeholders bring to evaluation of MR and its alternatives.

Unifying climate change biology across realms and taxa.

A major challenge in modern biology is to understand extinction risk from climate change across all realms. Recent research has revealed that physiological tolerance, behavioral thermoregulation, and small elevation shifts are dominant coping strategies on land, whereas large-scale latitudinal shifts are more important in the ocean. Freshwater taxa may face the highest global extinction risks. Nevertheless, some species in each realm face similar risks because of shared adaptive, dispersal, or physiological tolerances and abilities. Taking a cross-realm perspective offers unique research opportunities because confounding physical factors in one realm are often disaggregated in another realm. Cross-realm, across taxa, and other forms of climate change biology synthesis are needed to advance our understanding of emergent patterns of risk across all life.

Valuing the contributions of non-native species to people and nature.

While decision-making can benefit from considering positive and negative outcomes of change, over the past half-century, research on non-native species has focused predominately on their negative impacts. Here we provide a framework for considering the positive consequences of non-native species relative to relational, instrumental, and intrinsic values. We demonstrate that their beneficial outcomes are common and profoundly important for human well-being. Identified benefits include social cohesion, cultural identity, mental health, food and fuel production, regulation of clean waters, and attenuation of climate change. We argue that long-standing biases against non-native species within the literature have clouded the scientific process and hampered policy advances and sound public understanding. Future research should consider both costs and benefits of non-native species.

Analysis of climate paths reveals potential limitations on species range shifts.

Forecasts of species endangerment under climate change usually ignore the processes by which species ranges shift. By analysing the 'climate paths' that range shifts might follow, and two key range-shift processes--dispersal and population persistence--we show that short-term climatic and population characteristics have dramatic effects on range-shift forecasts. By employing this approach with 15 amphibian species in the western USA, we make unexpected predictions. First, inter-decadal variability in climate change can prevent range shifts by causing gaps in climate paths, even in the absence of geographic barriers. Second, the hitherto unappreciated trait of persistence during unfavourable climatic conditions is critical to species range shifts. Third, climatic fluctuations and low persistence could lead to endangerment even if the future potential range size is large. These considerations may render habitat corridors ineffectual for some species, and conservationists may need to consider managed relocation and augmentation of in situ populations.

The potential conservation value of non-native species.

Non-native species can cause the loss of biological diversity (i.e., genetic, species, and ecosystem diversity) and threaten the well-being of humans when they become invasive. In some cases, however, they can also provide conservation benefits. We examined the ways in which non-native species currently contribute to conservation objectives. These include, for example, providing habitat or food resources to rare species, serving as functional substitutes for extinct taxa, and providing desirable ecosystem functions. We speculate that non-native species might contribute to achieving conservation goals in the future because they may be more likely than native species to persist and provide ecosystem services in areas where climate and land use are changing rapidly and because they may evolve into new and endemic taxa. The management of non-native species and their potential integration into conservation plans depends on how conservation goals are set in the future. A fraction of non-native species will continue to cause biological and economic damage, and substantial uncertainty surrounds the potential future effects of all non-native species. Nevertheless, we predict the proportion of non-native species that are viewed as benign or even desirable will slowly increase over time as their potential contributions to society and to achieving conservation objectives become well recognized and realized.

Colloquium paper: species invasions and extinction: the future of native biodiversity on islands.

Predation by exotic species has caused the extinction of many native animal species on islands, whereas competition from exotic plants has caused few native plant extinctions. Exotic plant addition to islands is highly nonrandom, with an almost perfect 1 to 1 match between the number of naturalized and native plant species on oceanic islands. Here, we evaluate several alternative implications of these findings. Does the consistency of increase in plant richness across islands imply that a saturation point in species richness has been reached? If not, should we expect total plant richness to continue to increase as new species are added? Finally, is the rarity of native plant extinctions to date a misleading measure of the impact of past invasions, one that hides an extinction debt that will be paid in the future? By analyzing historical records, we show that the number of naturalized plant species has increased linearly over time on many individual islands. Further, the mean ratio of naturalized to native plant species across islands has changed steadily for nearly two centuries. These patterns suggest that many more species will become naturalized on islands in the future. We also discuss how dynamics of invasion bear upon alternative saturation scenarios and the implications these scenarios have for the future retention or extinction of native plant species. Finally, we identify invasion-motivated research gaps (propagule pressure, time-lags to extinction, abundance shifts, and loss of area) that can aid in forecasting extinction and in developing a more comprehensive theory of species extinctions.

Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography.

A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.

Globalization of human infectious disease.

Globalization has facilitated the spread of numerous infectious agents to all corners of the planet. Analysis of the Global Infectious Disease and Epidemiology Network (GIDEON) database quantitatively illustrates that the globalization of human infectious agents depends significantly on the range of hosts used. Infectious agents specific to humans are broadly and uniformly distributed, whereas zoonotic infectious agents are far more localized in their geographical distribution. Moreover, these patterns vary depending on transmission mode and infectious agent taxonomy. This dichotomy is unlikely to persist if certain aspects of globalization (for example, exotic species introductions) continue unabated. This raises a serious concern for public health and leaves nations with the task of determining the infectious agents that have the greatest potential to establish within their borders. At the advent of a century characterized by an apparent increase in emerging infectious diseases, these results have critical implications for public-health policy and future research pathways of infectious disease ecology.

Plant Biodiversity Change Across Scales During the Anthropocene.

Plant communities have undergone dramatic changes in recent centuries, although not all such changes fit with the dominant biodiversity-crisis narrative used to describe them. At the global scale, future declines in plant species diversity are highly likely given habitat conversion in the tropics, although few extinctions have been documented for the Anthropocene to date (<0.1%). Nonnative species introductions have greatly increased plant species richness in many regions of the world at the same time that they have led to the creation of new hybrid polyploid species by bringing previously isolated congeners into close contact. At the local scale, conversion of primary vegetation to agriculture has decreased plant diversity, whereas other drivers of change-e.g., climate warming, habitat fragmentation, and nitrogen deposition-have highly context-dependent effects, resulting in a distribution of temporal trends with a mean close to zero. These results prompt a reassessment of how conservation goals are defined and justified.

Species invasions exceed extinctions on islands worldwide: a comparative study of plants and birds.

Species richness is decreasing at a global scale. At subglobal scales, that is, within any defined area less extensive than the globe, species richness will increase when the number of nonnative species becoming naturalized is greater than the number of native species becoming extinct. Determining whether this has occurred is usually difficult because detailed records of species extinctions and naturalizations are rare; these records often exist, however, for oceanic islands. Here we show that species richness on oceanic islands has remained relatively unchanged for land birds, with the number of naturalizations being roughly equal to the number of extinctions, and has increased dramatically for vascular plants, with the number of naturalizations greatly exceeding the number of extinctions. In fact, for plants, the net number of species on islands has approximately doubled. We show further that these patterns are robust to differences in the history of human occupation of these islands and to the possibility of undocumented species extinctions. These results suggest that species richness may be increasing at subglobal scales for many groups and that future research should address what consequences this may have on ecological processes.

Niche syndromes, species extinction risks, and management under climate change.

The current distributions of species are often assumed to correspond with the total set of environmental conditions under which species can persist. When this assumption is incorrect, extinction risk estimated from species distribution models can be misleading. The degree to which species can tolerate or even thrive under conditions found beyond their current distributions alters extinction risks, time lags in realizing those risks, and the usefulness of alternative management strategies. To inform these issues, we propose a conceptual framework within which empirical data could be used to generate hypotheses regarding the realized, fundamental, and 'tolerance' niche of species. Although these niche components have rarely been characterized over geographic scales, we suggest that this could be done for many plant species by comparing native, naturalized, and horticultural distributions.

Balancing biodiversity in a changing environment: extinction debt, immigration credit and species turnover.

Here, we outline a conceptual framework for biodiversity dynamics following environmental change. The model incorporates lags in extinction and immigration, which lead to extinction debt and immigration credit, respectively. Collectively, these concepts enable a balanced consideration of changes in biodiversity following climate change, habitat fragmentation and other forcing events. They also reveal transient phenomena, such as biodiversity surpluses and deficits, which have important ramifications for biological conservation and the preservation of ecosystem services. Predicting such transient dynamics poses a serious conservation challenge in a time of rapid environmental change.

Local scale effects of disease on biodiversity.

To date, ecologists and conservation biologists have focused much of their attention on the population and ecosystem effects of disease at regional scales and the role that diseases play in global species extinction. Far less research has been dedicated to identifying the effects that diseases can have on local scale species assemblages. We examined the role of infectious disease in structuring local biodiversity. Our intention was to illustrate how variable outcomes can occur by focusing on three case studies: the influence of chestnut blight on forest communities dominated by chestnut trees, the influence of red-spot disease on urchin barrens and kelp forests, and the influence of sylvatic plague on grassland communities inhabited by prairie dogs. Our findings reveal that at local scales infectious disease seems to play an important, though unpredictable, role in structuring species diversity. Through our case studies, we have shown that diseases can cause drastic population declines or local extirpations in keystone species, ecosystem engineers, and otherwise abundant species. These changes in local diversity may be very important, particularly when considered alongside potentially corresponding changes in community structure and function, and we believe that future efforts to understand the importance of disease to species diversity should have an increased focus on these local scales.

Effects of exotic species on evolutionary diversification.

Exotic species invasions create almost ideal conditions for promoting evolutionary diversification: establishment of allopatric populations in new environmental conditions; altered ecological opportunities for native species; and new opportunities for hybridization between previously allopatric taxa. Here, we review recent studies of the evolutionary consequences of species invasions, revealing abundant and widespread examples of exotic species promoting evolutionary diversification via increased genetic differentiation among populations of both exotic and native species and the creation of new hybrid lineages. Our review indicates that, although the well-documented reductions to biodiversity caused by exotic species might outweigh the increases resulting from diversification, a complete understanding of the net effects of exotic species on biodiversity in the long term will require consideration of both.