Impacts of global change on species distributions: obstacles and solutions to integrate climate and land use

Aim The impact of multiple stressors on biodiversity is one of the most pressing questions in ecology and biodiversity conservation. Here we critically assess how often and efficiently two main drivers of global change have been simultaneously integrated into research, with the aim of providing practical solutions for better integration in the future. We focus on the integration of climate change (CC) and land‐use change (LUC) when studying changes in species distributions. Location Global. Methods We analysed the peer‐reviewed literature on the effects of CC and LUC on observed changes in species distributions, i.e. including species range and abundance, between 2000 and 2014. Results Studies integrating CC and LUC remain extremely scarce, which hampers our ability to develop appropriate conservation strategies. The lack of CC–LUC integration is likely to be a result of insufficient recognition of the co‐occurrence of CC and LUC at all scales, covariation and interactions between CC and LUC, as well as correlations between species thermal and habitat requirements. Practical guidelines for the study of these interactive effects include considering multiple drivers and processes when designing studies, using available long‐term datasets on multiple drivers, revisiting single‐driver studies with additional drivers or conducting comparative studies and meta‐analyses. Combining various methodological approaches, including time lags and adaptation processes, represent further avenues to improve global change science. Main conclusions Despite repeated claims for a better integration of multiple drivers, the effects of CC and LUC on species distributions and abundances have been mostly studied in isolation, which calls for a shift of standards towards more integrative global change science. The guidelines proposed here will encourage study designs that account for multiple drivers and improve our understanding of synergies or antagonisms among drivers.     

土壤线虫对气候变化的响应研究进展

全球变化对陆地生态系统功能具有重要而深远的影响。陆地生态系统地下部分具有重要的生态功能,其组成及结构对气候变化的响应将进一步减缓或加剧全球化进程。土壤线虫在各类生态系统中分布十分广泛,是地下食物网的重要组分,在维持土壤生物多样性及营养物质循环过程中发挥重要作用,其组成及结构对不同气候变化驱动因子的响应机制与模式不尽相同。增温及降水格局变化主要是通过改变线虫生境而直接影响其种群密度与结构,两者通常表现为正效应且作用效果随处理时间的延长而增强。CO2与大气氮沉降主要是通过影响地上植被,凋落物质量,土壤理化性质等间接过程影响土壤线虫。同时,不同的全球变化因子之间存在着复杂的交互作用,深入理解这些因子之间交互作用对线虫群落的影响模式与机制对于探讨未来气候变化情景下生态统生物多样性及养分循环过程具有重要的理论指导意义。     

The devil is in the detail: Nonadditive and context‐dependent plant population responses to increasing temperature and precipitation

In climate change ecology, simplistic research approaches may yield unrealistically simplistic answers to often more complicated problems. In particular, the complexity of vegetation responses to global climate change begs a better understanding of the impacts of concomitant changes in several climatic drivers, how these impacts vary across different climatic contexts, and of the demographic processes underlying population changes. Using a replicated, factorial, whole‐community transplant experiment, we investigated regional variation in demographic responses of plant populations to increased temperature and/or precipitation. Across four perennial forb species and 12 sites, we found strong responses to both temperature and precipitation change. Changes in population growth rates were mainly due to changes in survival and clonality. In three of the four study species, the combined increase in temperature and precipitation reflected nonadditive, antagonistic interactions of the single climatic changes for population growth rate and survival, while the interactions were additive and synergistic for clonality. This disparity affects the persistence of genotypes, but also suggests that the mechanisms behind the responses of the vital rates differ. In addition, survival effects varied systematically with climatic context, with wetter and warmer + wetter transplants showing less positive or more negative responses at warmer sites. The detailed demographic approach yields important mechanistic insights into how concomitant changes in temperature and precipitation affect plants, which makes our results generalizable beyond the four study species. Our comprehensive study design illustrates the power of replicated field experiments in disentangling the complex relationships and patterns that govern climate change impacts across real‐world species and landscapes.     

Flashiness and Flooding of Two Lakes in the Upper Midwest During a Century of Urbanization and Climate Change

Globally, ecosystem services are threatened by increasing urbanization and more variable precipitation patterns driven by climate change. However, how these drivers interact over long-time scales to affect underlying processes remains poorly understood, hindering our ability to predict their long-term consequences. Here, we use long-term data spanning nearly a century to investigate changes in hydrologic attributes for two lakes in the Upper Midwest with urbanizing watersheds. We quantified flashiness—the variability of runoff rate, volume, or stage-level of waterways—to investigate the concurrent impacts of urbanization and climate change on flashiness and flooding potential. Our results indicate that flashiness generally increased for both lakes over the period of 1916–2013, although this overall trend consists of sub-periods of increase and decrease. Increasing impervious surface area has been the stronger driver of flashiness historically; however, our results suggest that the impact of urbanization may reach a threshold, such that saturation effects would cause large magnitude precipitation events to become a relatively stronger driver of flashiness. Increasing flashiness indicates an increase in flooding potential, documented by increases in the 10- and 100-year flood threshold levels as large as 30 cm. Since flashiness is strongly related to the provisioning of multiple ecosystem services, the methodology and results presented here provide a unique approach to gain insight into how non-linear interactions between global change drivers, at multiple time scales, impact the simultaneous provision of multiple services.     

Experimental strategies to assess the biological ramifications of multiple drivers of global ocean change—A review

Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy‐making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process‐oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science‐based policy formulation.     

Soil microbial responses to warming and increased precipitation and their implications for ecosystem C cycling

A better understanding of soil microbial ecology is critical to gaining an understanding of terrestrial carbon (C) cycle–climate change feedbacks. However, current knowledge limits our ability to predict microbial community dynamics in the face of multiple global change drivers and their implications for respiratory loss of soil carbon. Whether microorganisms will acclimate to climate warming and ameliorate predicted respiratory C losses is still debated. It also remains unclear how precipitation, another important climate change driver, will interact with warming to affect microorganisms and their regulation of respiratory C loss. We explore the dynamics of microorganisms and their contributions to respiratory C loss using a 4-year (2006–2009) field experiment in a semi-arid grassland with increased temperature and precipitation in a full factorial design. We found no response of mass-specific (per unit microbial biomass C) heterotrophic respiration to warming, suggesting that respiratory C loss is directly from microbial growth rather than total physiological respiratory responses to warming. Increased precipitation did stimulate both microbial biomass and mass-specific respiration, both of which make large contributions to respiratory loss of soil carbon. Taken together, these results suggest that, in semi-arid grasslands, soil moisture and related substrate availability may inhibit physiological respiratory responses to warming (where soil moisture was significantly lower), while they are not inhibited under elevated precipitation. Although we found no total physiological response to warming, warming increased bacterial C utilization (measured by BIOLOG EcoPlates) and increased bacterial oxidation of carbohydrates and phenols. Non-metric multidimensional scaling analysis as well as ANOVA testing showed that warming or increased precipitation did not change microbial community structure, which could suggest that microbial communities in semi-arid grasslands are already adapted to fluctuating climatic conditions. In summary, our results support the idea that microbial responses to climate change are multifaceted and, even with no large shifts in community structure, microbial mediation of soil carbon loss could still occur under future climate scenarios.     

Interaction matters: Bottom-up driver interdependencies alter the projected response of phytoplankton communities to climate change

Phytoplankton growth is controlled by multiple environmental drivers, which are all modified by climate change. While numerous experimental studies identify interactive effects between drivers, large-scale ocean biogeochemistry models mostly account for growth responses to each driver separately and leave the results of these experimental multiple-driver studies largely unused. Here, we amend phytoplankton growth functions in a biogeochemical model by dual-driver interactions (CO₂ and temperature, CO₂ and light), based on data of a published meta-analysis on multiple-driver laboratory experiments. The effect of this parametrization on phytoplankton biomass and community composition is tested using present-day and future high-emission (SSP5-8.5) climate forcing. While the projected decrease in future total global phytoplankton biomass in simulations with driver interactions is similar to that in control simulations without driver interactions (5%–6%), interactive driver effects are group-specific. Globally, diatom biomass decreases more with interactive effects compared with the control simulation (−8.1% with interactions vs. no change without interactions). Small-phytoplankton biomass, by contrast, decreases less with on-going climate change when the model accounts for driver interactions (−5.0% vs. −9.0%). The response of global coccolithophore biomass to future climate conditions is even reversed when interactions are considered (+33.2% instead of −10.8%). Regionally, the largest difference in the future phytoplankton community composition between the simulations with and without driver interactions is detected in the Southern Ocean, where diatom biomass decreases (−7.5%) instead of increases (+14.5%), raising the share of small phytoplankton and coccolithophores of total phytoplankton biomass. Hence, interactive effects impact the phytoplankton community structure and related biogeochemical fluxes in a future ocean. Our approach is a first step to integrate the mechanistic understanding of interacting driver effects on phytoplankton growth gained by numerous laboratory experiments into a global ocean biogeochemistry model, aiming toward more realistic future projections of phytoplankton biomass and community composition.     

Global change and species interactions in terrestrial ecosystems

The main drivers of global environmental change (CO₂ enrichment, nitrogen deposition, climate, biotic invasions and land use) cause extinctions and alter species distributions, and recent evidence shows that they exert pervasive impacts on various antagonistic and mutualistic interactions among species. In this review, we synthesize data from 688 published studies to show that these drivers often alter competitive interactions among plants and animals, exert multitrophic effects on the decomposer food web, increase intensity of pathogen infection, weaken mutualisms involving plants, and enhance herbivory while having variable effects on predation. A recurrent finding is that there is substantial variability among studies in both the magnitude and direction of effects of any given GEC driver on any given type of biotic interaction. Further, we show that higher order effects among multiple drivers acting simultaneously create challenges in predicting future responses to global environmental change, and that extrapolating these complex impacts across entire networks of species interactions yields unanticipated effects on ecosystems. Finally, we conclude that in order to reliably predict the effects of GEC on community and ecosystem processes, the greatest single challenge will be to determine how biotic and abiotic context alters the direction and magnitude of GEC effects on biotic interactions.     

Direct and indirect interactions with vegetation shape crocodylian ecology at multiple scales

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Tree mortality in the eastern and central United States: patterns and drivers

Substantial uncertainty surrounds how forest ecosystems will respond to the simultaneous impacts of multiple global change drivers. Long‐term forest dynamics are sensitive to changes in tree mortality rates; however, we lack an understanding of the relative importance of the factors that affect tree mortality across different spatial and temporal scales. We used the US Forest Service Forest Inventory and Analysis database to evaluate the drivers of tree mortality for eastern temperate forest at the individual‐level across spatial scales from tree to landscape to region. We investigated 13 covariates in four categories: climate, air pollutants, topography, and stand characteristics. Overall, we found that tree mortality was most sensitive to stand characteristics and air pollutants. Different functional groups also varied considerably in their sensitivity to environmental drivers. This research highlights the importance of considering the interactions among multiple global change agents in shaping forest ecosystems.     

Separating direct and indirect effects of global change: a population dynamic modeling approach using readily available field data

Two sources of complexity make predicting plant community response to global change particularly challenging. First, realistic global change scenarios involve multiple drivers of environmental change that can interact with one another to produce non‐additive effects. Second, in addition to these direct effects, global change drivers can indirectly affect plants by modifying species interactions. In order to tackle both of these challenges, we propose a novel population modeling approach, requiring only measurements of abundance and climate over time. To demonstrate the applicability of this approach, we model population dynamics of eight abundant plant species in a multifactorial global change experiment in alpine tundra where we manipulated nitrogen, precipitation, and temperature over 7 years. We test whether indirect and interactive effects are important to population dynamics and whether explicitly incorporating species interactions can change predictions when models are forecast under future climate change scenarios. For three of the eight species, population dynamics were best explained by direct effect models, for one species neither direct nor indirect effects were important, and for the other four species indirect effects mattered. Overall, global change had negative effects on species population growth, although species responded to different global change drivers, and single‐factor effects were slightly more common than interactive direct effects. When the fitted population dynamic models were extrapolated under changing climatic conditions to the end of the century, forecasts of community dynamics and diversity loss were largely similar using direct effect models that do not explicitly incorporate species interactions or best‐fit models; however, inclusion of species interactions was important in refining the predictions for two of the species. The modeling approach proposed here is a powerful way of analyzing readily available datasets which should be added to our toolbox to tease apart complex drivers of global change.     

Climate warming interacts with other global change drivers to influence plant phenology: A meta-analysis of experimental studies

Shifts in plant phenology influence ecosystem structures and functions, yet how multiple global change drivers interact to affect phenology remains elusive. We conducted a meta-analysis of 242 published articles to assess interactions between warming (W) and other global change drivers including nitrogen addition (N), increased precipitation (IP), decreased precipitation (DP) and elevated CO₂ (eCO₂) on multiple phenophases in experimental studies. We show that leaf out and first flowering were most strongly affected by warming, while warming and decreased precipitation were the most pronounced drivers for leaf colouring. Moreover, interactions between warming and other global change drivers were common and both synergistic and antagonistic interactions were observed: interactions W + IP and W + eCO₂ were frequently synergistic, whereas interactions W + N and W + DP were mostly antagonistic. These findings demonstrate that global change drivers often affect plant phenology interactively. Incorporating the multitude of interactions into models is crucial for accurately predicting plant responses to global changes.     

A single climate driver has direct and indirect effects on insect population dynamics

Weather drives population dynamics directly, through effects on vital rates, or indirectly, through effects on the population's competitors, predators or prey and thence on vital rates. Indirect effects may include non-additive interactions with density dependence. Detection of climate drivers is critical to predicting climate change effects, but identification of potential drivers may depend on knowing the underlying mechanisms. For the butterfly Speyeria mormonia, one climate driver, snow melt date, has multiple effects on population growth. Snow melt date in year t has density-dependent indirect effects. Through frost effects, early snow melt decreases floral resources, thence per-capita nectar availability, which determines fecundity in the lab. Snow melt date in year t?+?1 has density-independent direct effects. These effects explain 84% of the variation in population growth rate. One climate parameter thus has multiple effects on the dynamics of a species with non-overlapping generations, with one effect not detectable without understanding the underlying mechanism.     

Warming and top predator loss drive ecosystem multifunctionality

Global change affects ecosystem functioning both directly by modifications in physicochemical processes, and indirectly, via changes in biotic metabolism and interactions. Unclear, however, is how multiple anthropogenic drivers affect different components of community structure and the performance of multiple ecosystem functions (ecosystem multifunctionality). We manipulated small natural freshwater ecosystems to investigate how warming and top predator loss affect seven ecosystem functions representing two major dimensions of ecosystem functioning, productivity and metabolism. We investigated their direct and indirect effects on community diversity and standing stock of multitrophic macro and microorganisms. Warming directly increased multifunctional ecosystem productivity and metabolism. In contrast, top predator loss indirectly affected multifunctional ecosystem productivity via changes in the diversity of detritivorous macroinvertebrates, but did not affect ecosystem metabolism. In addition to demonstrating how multiple anthropogenic drivers have different impacts, via different pathways, on ecosystem multifunctionality components, our work should further spur advances in predicting responses of ecosystems to multiple simultaneous environmental changes.     

Global environmental change effects on ecosystems: the importance of land‐use legacies

One of the major challenges in ecology is to predict how multiple global environmental changes will affect future ecosystem patterns (e.g. plant community composition) and processes (e.g. nutrient cycling). Here, we highlight arguments for the necessary inclusion of land‐use legacies in this endeavour. Alterations in resources and conditions engendered by previous land use, together with influences on plant community processes such as dispersal, selection, drift and speciation, have steered communities and ecosystem functions onto trajectories of change. These trajectories may be modulated by contemporary environmental changes such as climate warming and nitrogen deposition. We performed a literature review which suggests that these potential interactions have rarely been investigated. This crucial oversight is potentially due to an assumption that knowledge of the contemporary state allows accurate projection into the future. Lessons from other complex dynamic systems, and the recent recognition of the importance of previous conditions in explaining contemporary and future ecosystem properties, demand the testing of this assumption. Vegetation resurvey databases across gradients of land use and environmental change, complemented by rigorous experiments, offer a means to test for interactions between land‐use legacies and multiple environmental changes. Implementing these tests in the context of a trait‐based framework will allow biologists to synthesize compositional and functional ecosystem responses. This will further our understanding of the importance of land‐use legacies in determining future ecosystem properties, and soundly inform conservation and restoration management actions.     

Tropical grassy biomes: linking ecology,human use and conservation

Tropical grassy biomes (TGBs) are changing rapidly the world over through a coalescence of high rates of land-use change, global change and altered disturbance regimes that maintain the ecosystem structure and function of these biomes. Our theme issue brings together the latest research examining the characterization, complex ecology, drivers of change, and human use and ecosystem services of TGBs. Recent advances in ecology and evolution have facilitated a new perspective on these biomes. However, there continues to be controversies over their classification and state dynamics that demonstrate critical data and knowledge gaps in our quantitative understanding of these geographically dispersed regions. We highlight an urgent need to improve ecological understanding in order to effectively predict the sensitivity and resilience of TGBs under future scenarios of global change. With human reliance on TGBs increasing and their propensity for change, ecological and evolutionary understanding of these biomes is central to the dual goals of sustaining their ecological integrity and the diverse services these landscapes provide to millions of people.This article is part of the themed issue ‘Tropical grassy biomes: linking ecology, human use and conservation’.     

Global change biology: A primer

Because of human action, the Earth has entered an era where profound changes in the global environment are creating novel conditions that will be discernable far into the future. One consequence may be a large reduction of the Earth's biodiversity, potentially representing a sixth mass extinction. With effective stewardship, the global change drivers that threaten the Earth's biota could be alleviated, but this requires clear understanding of the drivers, their interactions, and how they impact ecological communities. This review identifies 10 anthropogenic global change drivers and discusses how six of the drivers (atmospheric CO₂ enrichment, climate change, land transformation, species exploitation, exotic species invasions, eutrophication) impact Earth's biodiversity. Driver impacts on a particular species could be positive or negative. In either case, they initiate secondary responses that cascade along ecological lines of connection and in doing so magnify the initial impact. The unique nature of the threat to the Earth's biodiversity is not simply due to the magnitude of each driver, but due to the speed of change, the novelty of the drivers, and their interactions. Emphasizing one driver, notably climate change, is problematic because the other global change drivers also degrade biodiversity and together threaten the stability of the biosphere. As the main academic journal addressing global change effects on living systems, GCB is well positioned to provide leadership in solving the global change challenge. If humanity cannot meet the challenge, then GCB is positioned to serve as a leading chronicle of the sixth mass extinction to occur on planet Earth.     

Individual to community-level faunal responses to environmental change from a marine fossil record of Early Miocene global warming

Modern climate change has a strong potential to shift earth systems and biological communities into novel states that have no present-day analog, leaving ecologists with no observational basis to predict the likely biotic effects. Fossil records contain long time-series of past environmental changes outside the range of modern observation, which are vital for predicting future ecological responses, and are capable of (a) providing detailed information on rates of ecological change, (b) illuminating the environmental drivers of those changes, and (c) recording the effects of environmental change on individual physiological rates. Outcrops of Early Miocene Newport Member of the Astoria Formation (Oregon) provide one such time series. This record of benthic foraminiferal and molluscan community change from continental shelf depths spans a past interval environmental change (≈ 20.3-16.7 mya) during which the region warmed 2.1-4.5°C, surface productivity and benthic organic carbon flux increased, and benthic oxygenation decreased, perhaps driven by intensified upwelling as on the modern Oregon coast. The Newport Member record shows that (a) ecological responses to natural environmental change can be abrupt, (b) productivity can be the primary driver of faunal change during global warming, (c) molluscs had a threshold response to productivity change while foraminifera changed gradually, and (d) changes in bivalve body size and growth rates parallel changes in taxonomic composition at the community level, indicating that, either directly or indirectly through some other biological parameter, the physiological tolerances of species do influence community change. Ecological studies in modern and fossil records that consider multiple ecological levels, environmental parameters, and taxonomic groups can provide critical information for predicting future ecological change and evaluating species vulnerability.     

Towards an eco‐phylogenetic framework for infectious disease ecology

Identifying patterns and drivers of infectious disease dynamics across multiple scales is a fundamental challenge for modern science. There is growing awareness that it is necessary to incorporate multi‐host and/or multi‐parasite interactions to understand and predict current and future disease threats better, and new tools are needed to help address this task. Eco‐phylogenetics (phylogenetic community ecology) provides one avenue for exploring multi‐host multi‐parasite systems, yet the incorporation of eco‐phylogenetic concepts and methods into studies of host pathogen dynamics has lagged behind. Eco‐phylogenetics is a transformative approach that uses evolutionary history to infer present‐day dynamics. Here, we present an eco‐phylogenetic framework to reveal insights into parasite communities and infectious disease dynamics across spatial and temporal scales. We illustrate how eco‐phylogenetic methods can help untangle the mechanisms of host–parasite dynamics from individual (e.g. co‐infection) to landscape scales (e.g. parasite/host community structure). An improved ecological understanding of multi‐host and multi‐pathogen dynamics across scales will increase our ability to predict disease threats.