Poised to prosper? A cross‐system comparison of climate change effects on native and non‐native species performance

Climate change and biological invasions are primary threats to global biodiversity that may interact in the future. To date, the hypothesis that climate change will favour non‐native species has been examined exclusively through local comparisons of single or few species. Here, we take a meta‐analytical approach to broadly evaluate whether non‐native species are poised to respond more positively than native species to future climatic conditions. We compiled a database of studies in aquatic and terrestrial ecosystems that reported performance measures of non‐native (157 species) and co‐occurring native species (204 species) under different temperature, CO₂ and precipitation conditions. Our analyses revealed that in terrestrial (primarily plant) systems, native and non‐native species responded similarly to environmental changes. By contrast, in aquatic (primarily animal) systems, increases in temperature and CO₂ largely inhibited native species. There was a general trend towards stronger responses among non‐native species, including enhanced positive responses to more favourable conditions and stronger negative responses to less favourable conditions. As climate change proceeds, aquatic systems may be particularly vulnerable to invasion. Across systems, there could be a higher risk of invasion at sites becoming more climatically hospitable, whereas sites shifting towards harsher conditions may become more resistant to invasions.     

Will climate change promote future invasions?

Biological invasion is increasingly recognized as one of the greatest threats to biodiversity. Using ensemble forecasts from species distribution models to project future suitable areas of the 100 of the world's worst invasive species defined by the International Union for the Conservation of Nature, we show that both climate and land use changes will likely cause drastic species range shifts. Looking at potential spatial aggregation of invasive species, we identify three future hotspots of invasion in Europe, northeastern North America, and Oceania. We also emphasize that some regions could lose a significant number of invasive alien species, creating opportunities for ecosystem restoration. From the list of 100, scenarios of potential range distributions show a consistent shrinking for invasive amphibians and birds, while for aquatic and terrestrial invertebrates distributions are projected to substantially increase in most cases. Given the harmful impacts these invasive species currently have on ecosystems, these species will likely dramatically influence the future of biodiversity.     

Will all species with temperature‐dependent sex determination respond the same way to climate change? A reply to Kallimanis (2010)

Recently, Kallimanis (2010) published a paper proposing a mechanism by which temperature‐dependent sex determination (TSD) may play a key role at facilitating species with this strategy to track their climatic niches across space under climate change. Kallimanis hypothesized that TSD species currently inhabiting stable climatic conditions show reduced population growth rates at the edges of their distributional ranges; under warming conditions, these populations will experience faster growth rates and thus are able to colonize new suitable sites. These ideas are based on the assumption that populations of TSD species have balanced sex ratios at the core of their geographic ranges and biased proportions at the edges. However, Kallimanis’ model overlooks complex processes that may produce a more broadly and less predictable aftermath of climate change on TSD species, so we discuss some of his postulates and underlying assumptions. Kallimanis’ model is based only on one of three known TSD strategies in reptiles, thus it lacks generality; and it does not consider the phenological, behavioral, and physiological strategies that TSD species exhibit across their geographic ranges to buffer the potential impacts of climatic variation over the whole reproductive process. We conclude that simple models such as the one proposed by Kallimanis are not broadly applicable; hence, forecasts of TSD species’ responses to climate change will need to be more specific to groups with similar ecologies and modes of TSD.     

Would climate change drive species out of reserves? An assessment of existing reserve‐selection methods

Concern for climate change has not yet been integrated in protocols for reserve selection. However if climate changes as projected, there is a possibility that current reserve‐selection methods might provide solutions that are inadequate to ensure species' long‐term persistence within reserves. We assessed, for the first time, the ability of existing reserve‐selection methods to secure species in a climate‐change context. Six methods using a different combination of criteria (representation, suitability and reserve clustering) are compared. The assessment is carried out using European distributions of 1200 plant species and considering two extreme scenarios of response to climate change: no dispersal and universal dispersal. With our data, 6–11% of species modelled would be potentially lost from selected reserves in a 50‐year period. Measured uncertainties varied in 6% being 1–3% attributed to dispersal assumptions and 2–5% to the choice of reserve‐selection method. Suitability approaches to reserve selection performed best, while reserve clustering performed poorly. We also found that 5% of species modelled would lose their entire climatic envelope in the studied area; 2% of the species modelled would have nonoverlapping distributions; 93% of the species modelled would maintain varying levels of overlapping distributions. We conclude there are opportunities to minimize species' extinctions within reserves but new approaches are needed to account for impacts of climate change on species; especially for those projected to have temporally nonoverlapping distributions.     

Will climate change favor exotic grasses over native ecosystem engineer species in the Amazon Basin?

Several anthropic disturbances, including deforestation, fires, the building of roads and dams, have intensified in Amazon in last decades. These disturbances contribute to an increase in the occurrence and intensity of extreme events, such as more frequent floods and more severe droughts, due to climate change. Along the Amazonian rivers, aquatic herbaceous plants, mainly of the Poaceae family, are very abundant and produce up to three times more biomass than the adjacent flooded forests, and some are considered ecosystem engineers given their structuring role in these environments. Invasive grasses have spread through the Neotropics and are gradually entering the Amazon via the Arc of Deforestation. These invasive species often attain high coverage, suppress other species, and become dominant in both disturbed and pristine habitats. The aim of this study was to establish the current and future distribution patterns of two native ecosystem engineer species (Echinochloa polystachya and Paspalum fasciculatum) and two invasive species (Urochloa brizantha and Urochloa decumbens) in the Amazon Basin. To predict the future climate, we used three scenarios, namely SSP1–2.6, SSP3–7.0 and SSP5–8.5 for the years 2040, 2080 and 2100, to project climatically suitable areas. The current climatically suitable range for the native ecosystem engineer species was estimated at 33–35% of the Amazon Basin, while the invasive ones have a range of 53–84% in potential climatically suitable areas. A decrease in the areas of suitability of the two ecosystem engineer species, E. polystachya and P. fasciculatum, was observed in all scenarios and years, while only the invasive U. brizantha showed an increase in suitable areas in all years. These results raise concerns about the invasion of grasses with high aggressive potential that could result in the exclusion of native ecosystem engineer species and their ecological roles.     

Will variation among genetic individuals influence species responses to global climate change?

Increased anthropogenic CO₂ emissions in the last two centuries have lead to rising sea surface temperature and falling ocean pH, and it is predicted that current global trends will worsen over the next few decades. There is limited understanding of how genetic variation among individuals will influence the responses of populations and species to these changes. A microcosm system was set up to study the effects of predicted temperature and CO₂ levels on the bryozoan Celleporella hyalina. In this marine species, colonies grow by the addition of male, female and feeding modular individuals (zooids) and can be physically subdivided to produce a clone of genetically identical colonies. We studied colony growth rate (the addition of zooids), reproductive investment (the ratio of sexual to feeding zooids) and sex ratio (male to female zooids) in four genetically distinct clonal lines. There was a significant effect of clone on growth rate, reproductive investment and sex ratio, with clones showing contrasting responses to the various temperature and pH combinations. Overall, decreasing pH and increasing temperature caused reduction of growth, and eventual cessation of growth was often observed at the highest temperature, especially during the latter half of the 15‐day trials. Reproductive investment increased with increasing temperature and decreasing pH, varying more widely with temperature at the lowest pH. The increased production of males, a general stress response of the bryozoan, was seen upon exposure to reduced pH, but was not expressed at the highest temperature tested, presumably due to the frequent cessation of growth. Further to the significant effect of pH on the measured whole‐colony parameters, observation by scanning electron microscopy revealed surface pitting of the calcified exoskeleton in colonies that were exposed to increased acidity. Studying ecologically relevant processes of growth and reproduction, we demonstrate the existence of relevant levels of variation among genetic individuals which may enable future adaptation via non‐mutational natural selection to falling pH and rising temperature.     

The role of nitrogen in climate change and the impacts of nitrogen–climate interactions in the United States: foreword to thematic issue

Producing food, transportation, and energy for seven billion people has led to large and widespread increases in the use of synthetic nitrogen (N) fertilizers and fossil fuel combustion, resulting in a leakage of N into the environment as various forms of air and water pollution. The global N cycle is more severely altered by human activity than the global carbon (C) cycle, and reactive N dynamics affect all aspects of climate change considerations, including mitigation, adaptation, and impacts. In this special issue of Biogeochemistry, we present a review of the climate–nitrogen interactions based on a technical report for the United States National Climate Assessment presented as individual papers for terrestrial and aquatic ecosystems, agriculture and human health within the US. We provide a brief overview of each of the paper’s main points and conclusions is presented in this foreword summary.     

How do climate change experiments alter plot‐scale climate?

To understand and forecast biological responses to climate change, scientists frequently use field experiments that alter temperature and precipitation. Climate manipulations can manifest in complex ways, however, challenging interpretations of biological responses. We reviewed publications to compile a database of daily plot‐scale climate data from 15 active‐warming experiments. We find that the common practices of analysing treatments as mean or categorical changes (e.g. warmed vs. unwarmed) masks important variation in treatment effects over space and time. Our synthesis showed that measured mean warming, in plots with the same target warming within a study, differed by up to 1.6 C (63% of target), on average, across six studies with blocked designs. Variation was high across sites and designs: for example, plots differed by 1.1 C (47% of target) on average, for infrared studies with feedback control (n = 3) vs. by 2.2 C (80% of target) on average for infrared with constant wattage designs (n = 2). Warming treatments produce non‐temperature effects as well, such as soil drying. The combination of these direct and indirect effects is complex and can have important biological consequences. With a case study of plant phenology across five experiments in our database, we show how accounting for drier soils with warming tripled the estimated sensitivity of budburst to temperature. We provide recommendations for future analyses, experimental design, and data sharing to improve our mechanistic understanding from climate change experiments, and thus their utility to accurately forecast species’ responses.     

Molecular and quantitative trait variation across the native range of the invasive species Hypericum canariense: evidence for ancient patterns of colonization via pre‐adaptation?

To understand the success of invasive species, it is important to know whether colonization events are facilitated by adaptive evolution or are limited to sites where a species is pre-adapted to thrive. Studies of the ancient colonization patterns of an invader in its native range provide an opportunity to examine its natural history of adaptation and colonization. This study uses molecular (internal transcribed spacer sequence and amplified fragment length polymorphism) and common garden approaches to assess the ancient patterns of establishment and quantitative trait evolution in the invasive shrub Hypericum canariense. This species has an unusually small and discrete native range in the Canary Islands. Our data reveal two genetic varieties with divergent life histories and different colonization patterns across the islands. Although molecular divergence within each variety is large (pairwise FST from 0.18 to 0.32 between islands) and nearly as great as divergence between them, life-history traits show striking uniformity within varieties. The discrepancy between molecular and life-history trait divergence points to the action of stabilizing selection within varieties and the influence of pre-adaptation on patterns of colonization. The colonization history of H. canariense reflects how the relationship between selective environments in founding and source populations can dictate establishment by particular lineages and their subsequent evolutionary stasis or change.     

Has past climate change affected cold‐specialized species differentially through space and time?

Quaternary climate change has been strongly linked to distributional shifts and recent species diversification. Montane species, in particular, have experienced enhanced isolation and rapid genetic divergence during glacial fluctuations, and these processes have resulted in a disproportionate number of neo‐endemic species forming in high‐elevation habitats. In temperate montane environments, a general model of alpine population history is well supported, where cold‐specialized species track favourable climate conditions downslope during glacial episodes and upslope during warmer interglacial periods, which leads to a climate‐driven population or species diversification pump. However, it remains unclear how geography mediates distributional changes and whether certain episodes of glacial history have differentially impacted rates of diversification. We address these questions by examining phylogenomic data in a North American clade of flightless, cold‐specialized insects, the ice crawlers (Insecta: Grylloblattodea: Grylloblattidae: Grylloblatta). These low‐vagility organisms have the potential to reveal highly localized refugia and patterns of spatial recolonization, as well as a longer history of in situ diversification. Using continuous phylogeographic analysis of species groups, we show that all species tend to retreat to nearby low‐elevation habitats across western North America during episodes of glaciation, but species at high latitude exhibit larger distributional shifts. Lineage diversification was examined over the course of the Neogene and Quaternary periods, with statistical analysis supporting a direct association between climate variation and diversification rate. Major increases in lineage diversification appear to be correlated with warm and dry periods, rather than with extreme glacial events. Finally, we identify substantial cryptic diversity among ice crawlers, leading to high endemism across their range. This diversity provides new insights into highly localized glacial refugia for cold‐specialized species across western North America.     

Going with the flow? Threatened species management and legislation in the face of climate change

Summary   The potential impacts of climate change on threatened species, populations and communities are considered. It is suggested that minor changes to legislation will be required to address the consequences of movement of threatened species but that threatened species legislation will remain relevant as an important tool for prioritizing conservation actions. The importance of taking proactive steps now to permit future movement of species and communities across fragmented landscapes is emphasized.     

Will climate change reduce the effects of a pesticide on amphibians?: partitioning effects on exposure and susceptibility to contaminants

Several studies suggest that global climate change could increase the toxicity of contaminants, but none of these studies explicitly integrate the effects of climate change on both susceptibility and duration of exposure to pollution. For many amphibian and aquatic insect species, exposure to contaminants is probably greatest during their fully aquatic embryonic and larval stages because these stages cannot readily escape water bodies where many contaminants accumulate and concentrate. Hence, by accelerating embryonic and larval development, global warming might reduce the duration of contaminant exposure for these taxa. To test this hypothesis, we isolated the effects of a temperature gradient (13–25 °C) on susceptibility (toxicity at a controlled exposure duration) and exposure of the streamside salamander, Ambystoma barbouri, to the herbicide atrazine (0, 4, 40, and 400 μg L^?1) by quantifying growth, survival, hatching, and metamorphosis under an atrazine exposure duration that was either constant or that depended on time to metamorphosis (and thus temperature). Increasing atrazine concentrations reduced growth, delayed hatching and metamorphosis, and decreased embryonic and larval survival. Increasing temperatures enhanced growth, accelerated development, and reduced survival for embryos but not larvae. With the exception of growth, increasing temperatures generally did not enhance the toxicity of atrazine, but they did generally ameliorate the adverse effects of atrazine by accelerating development and reducing the duration of atrazine exposure. The actual effects of climate change on contaminants remains difficult to predict because temperature changes can affect chemical use, uptake, excretion, biotransformation, fate, transport, and bioavailability. However, this work highlights the importance of explicitly considering how climate change will affect both exposure and toxicity to contaminants to accurately assess risk.     

Will changes in phenology track climate change? A study of growth initiation timing in coast Douglas‐fir

Under climate change, the reduction of frost risk, onset of warm temperatures and depletion of soil moisture are all likely to occur earlier in the year in many temperate regions. The resilience of tree species will depend on their ability to track these changes in climate with shifts in phenology that lead to earlier growth initiation in the spring. Exposure to warm temperatures (‘forcing’) typically triggers growth initiation, but many trees also require exposure to cool temperatures (‘chilling’) while dormant to readily initiate growth in the spring. If warming increases forcing and decreases chilling, climate change could maintain, advance or delay growth initiation phenology relative to the onset of favorable conditions. We modeled the timing of height‐ and diameter‐growth initiation in coast Douglas‐fir (an ecologically and economically vital tree in western North America) to determine whether changes in phenology are likely to track changes in climate using data from field‐based and controlled‐environment studies, which included conditions warmer than those currently experienced in the tree's range. For high latitude and elevation portions of the tree's range, our models predicted that warming will lead to earlier growth initiation and allow trees to track changes in the onset of the warm but still moist conditions that favor growth, generally without substantially greater exposure to frost. In contrast, toward lower latitude and elevation range limits, the models predicted that warming will lead to delayed growth initiation relative to changes in climate due to reduced chilling, with trees failing to capture favorable conditions in the earlier parts of the spring. This maladaptive response to climate change was more prevalent for diameter‐growth initiation than height‐growth initiation. The decoupling of growth initiation with the onset of favorable climatic conditions could reduce the resilience of coast Douglas‐fir to climate change at the warm edges of its distribution.     

How does climate warming affect plant‐pollinator interactions?

Climate warming affects the phenology, local abundance and large-scale distribution of plants and pollinators. Despite this, there is still limited knowledge of how elevated temperatures affect plant-pollinator mutualisms and how changed availability of mutualistic partners influences the persistence of interacting species. Here we review the evidence of climate warming effects on plants and pollinators and discuss how their interactions may be affected by increased temperatures. The onset of flowering in plants and first appearance dates of pollinators in several cases appear to advance linearly in response to recent temperature increases. Phenological responses to climate warming may therefore occur at parallel magnitudes in plants and pollinators, although considerable variation in responses across species should be expected. Despite the overall similarities in responses, a few studies have shown that climate warming may generate temporal mismatches among the mutualistic partners. Mismatches in pollination interactions are still rarely explored and their demographic consequences are largely unknown. Studies on multi-species plant-pollinator assemblages indicate that the overall structure of pollination networks probably are robust against perturbations caused by climate warming. We suggest potential ways of studying warming-caused mismatches and their consequences for plant-pollinator interactions, and highlight the strengths and limitations of such approaches.