The cumulative impacts of anthropogenic stressors vary markedly along environmental gradients

Understanding the cumulative effects of multiple stressors on biodiversity is key to managing their impacts. Stressor interactions are often studied using an additive/antagonistic/synergistic typology, aimed at identifying situations where individual stressor effects are reduced or amplified when they act in combination. Here, we analysed variation in the family richness of stream macroinvertebrates in the groups Ephemeroptera, Plecoptera and Trichoptera (EPT) at 4658 sites spanning a 32° latitudinal range in eastern Australia in relation to two largely human-induced stressors, salinity and turbidity, and two environmental gradients, temperature and slope. The cumulative and interactive effect of salinity and turbidity on EPT family richness varied across the landscape and by habitat (edge or riffle) such that we observed additive, antagonistic and synergistic outcomes depending on the environmental context. Our findings highlight the importance of understanding the consistency of multiple stressor impacts, which will involve higher-order interactions between multiple stressors and environmental factors.     

Interactive and cumulative effects of multiple human stressors in marine systems

Humans impact natural systems in a multitude of ways, yet the cumulative effect of multiple stressors on ecological communities remains largely unknown. Here we synthesized 171 studies that manipulated two or more stressors in marine and coastal systems and found that cumulative effects in individual studies were additive (26%), synergistic (36%), and antagonistic (38%). The overall interaction effect across all studies was synergistic, but interaction type varied by response level (community: antagonistic, population: synergistic), trophic level (autotrophs: antagonistic, heterotrophs: synergistic), and specific stressor pair (seven pairs additive, three pairs each synergistic and antagonistic). Addition of a third stressor changed interaction effects significantly in two‐thirds of all cases and doubled the number of synergistic interactions. Given that most studies were performed in laboratories where stressor effects can be carefully isolated, these three‐stressor results suggest that synergies may be quite common in nature where more than two stressors almost always coexist. While significant gaps exist in multiple stressor research, our results suggest an immediate need to account for stressor interactions in ecological studies and conservation planning.     

Managing for Interactions between Local and Global Stressors of Ecosystems

Global stressors, including climate change, are a major threat to ecosystems, but they cannot be halted by local actions. Ecosystem management is thus attempting to compensate for the impacts of global stressors by reducing local stressors, such as overfishing. This approach assumes that stressors interact additively or synergistically, whereby the combined effect of two stressors is at least the sum of their isolated effects. It is not clear, however, how management should proceed for antagonistic interactions among stressors, where multiple stressors do not have an additive or greater impact. Research to date has focussed on identifying synergisms among stressors, but antagonisms may be just as common. We examined the effectiveness of management when faced with different types of interactions in two systems – seagrass and fish communities – where the global stressor was climate change but the local stressors were different. When there were synergisms, mitigating local stressors delivered greater gains, whereas when there were antagonisms, management of local stressors was ineffective or even degraded ecosystems. These results suggest that reducing a local stressor can compensate for climate change impacts if there is a synergistic interaction. Conversely, if there is an antagonistic interaction, management of local stressors will have the greatest benefits in areas of refuge from climate change. A balanced research agenda, investigating both antagonistic and synergistic interaction types, is needed to inform management priorities.     

Long-term adaptation of epistatic genetic networks

Gene networks are likely to govern most traits in nature. Mutations at these genes often show functional epistatic interactions that lead to complex genetic architectures and variable fitness effects in different genetic backgrounds. Understanding how epistatic genetic systems evolve in nature remains one of the great challenges in evolutionary biology. Here we combine an analytical framework with individual-based simulations to generate novel predictions about long-term adaptation of epistatic networks. We find that relative to traits governed by independently evolving genes, adaptation with epistatic gene networks is often characterized by longer waiting times to selective sweeps, lower standing genetic variation, and larger fitness effects of adaptive mutations. This may cause epistatic networks to either adapt more slowly or more quickly relative to a nonepistatic system. Interestingly, epistatic networks may adapt faster even when epistatic effects of mutations are on average deleterious. Further, we study the evolution of epistatic properties of adaptive mutations in gene networks. Our results show that adaptive mutations with small fitness effects typically evolve positive synergistic interactions, whereas adaptive mutations with large fitness effects evolve positive synergistic and negative antagonistic interactions at approximately equal frequencies. These results provide testable predictions for adaptation of traits governed by epistatic networks and the evolution of epistasis within networks.     

Net effects of multiple stressors in freshwater ecosystems: a meta‐analysis

The accelerating rate of global change has focused attention on the cumulative impacts of novel and extreme environmental changes (i.e. stressors), especially in marine ecosystems. As integrators of local catchment and regional processes, freshwater ecosystems are also ranked highly sensitive to the net effects of multiple stressors, yet there has not been a large‐scale quantitative synthesis. We analysed data from 88 papers including 286 responses of freshwater ecosystems to paired stressors and discovered that overall, their cumulative mean effect size was less than the sum of their single effects (i.e. an antagonistic interaction). Net effects of dual stressors on diversity and functional performance response metrics were additive and antagonistic, respectively. Across individual studies, a simple vote‐counting method revealed that the net effects of stressor pairs were frequently more antagonistic (41%) than synergistic (28%), additive (16%) or reversed (15%). Here, we define a reversal as occurring when the net impact of two stressors is in the opposite direction (negative or positive) from that of the sum of their single effects. While warming paired with nutrification resulted in additive net effects, the overall mean net effect of warming combined with a second stressor was antagonistic. Most importantly, the mean net effects across all stressor pairs and response metrics were consistently antagonistic or additive, contrasting the greater prevalence of reported synergies in marine systems. Here, a possible explanation for more antagonistic responses by freshwater biota to stressors is that the inherent greater environmental variability of smaller aquatic ecosystems fosters greater potential for acclimation and co‐adaptation to multiple stressors.     

Integrating animal behaviour into research on multiple environmental stressors: a conceptual framework

While a large body of research has focused on the physiological effects of multiple environmental stressors, how behavioural and life-history plasticity mediate multiple-stressor effects remains underexplored. Behavioural plasticity can not only drive organism-level responses to stressors directly but can also mediate physiological responses. Here, we provide a conceptual framework incorporating four fundamental trade-offs that explicitly link animal behaviour to life-history-based pathways for energy allocation, shaping the impact of multiple stressors on fitness. We first address how small-scale behavioural changes can either mediate or drive conflicts between the effects of multiple stressors and alternative physiological responses. We then discuss how animal behaviour gives rise to three additional understudied and interrelated trade-offs: balancing the benefits and risks of obtaining the energy needed to cope with stressors, allocation of energy between life-history traits and stressor responses, and larger-scale escape from stressors in space or time via large-scale movement or dormancy. Finally, we outline how these trade-offs interactively affect fitness and qualitative ecological outcomes resulting from multiple stressors. Our framework suggests that explicitly considering animal behaviour should enrich our mechanistic understanding of stressor effects, help explain extensive context dependence observed in these effects, and highlight promising avenues for future empirical and theoretical research.     

SEXUAL CONFLICT AND SEXUAL SELECTION: MEASURING ANTAGONISTIC COEVOLUTION

Abstract Arnqvist (2004) raises some concerns with several of the points made by Pizzari and Snook (2003) on the study of sexually antagonistic coevolution (SAC) generated by sexual conflict, arguing that: (1) sexual conflict cannot be expressed in terms of average male and female fitness; (2) our criticism of current experimental approaches, particularly interpopulation crosses, is unjustified; and (3) the alternative experimental approach we proposed is problematic. Here we discuss and respond to these criticisms by: (1) clarifying that we can distinguish between SAC and mutualistic sexual coevolution by measuring changes in the average fitness of the reproducing subsamples of males and females of a population across generations, (2) maintaining that testing SAC using interpopulation crosses is undermined by the lack of a priori knowledge of what traits mediate SAC across isolated populations, and (3) reinforcing the advantages of our experimental approach to distinguish between sexually mutualistic and antagonistic selection.     

Antagonistic pleiotropy and mutation accumulation contribute to age‐related decline in stress response

As organisms age, the effectiveness of natural selection weakens, leading to age‐related decline in fitness‐related traits. The evolution of age‐related changes associated with senescence is likely influenced by mutation accumulation (MA) and antagonistic pleiotropy (AP). MA predicts that age‐related decline in fitness components is driven by age‐specific sets of alleles, nonnegative genetic correlations within trait across age, and an increase in the coefficient of genetic variance. AP predicts that age‐related decline in a trait is driven by alleles with positive effects on fitness in young individuals and negative effects in old individuals, and is expected to lead to negative genetic correlations within traits across age. We build on these predictions using an association mapping approach to investigate the change in additive effects of SNPs across age and among traits for multiple stress‐response fitness‐related traits, including cold stress with and without acclimation and starvation resistance. We found support for both MA and AP theories of aging in the age‐related decline in stress tolerance. Our study demonstrates that the evolution of age‐related decline in stress tolerance is driven by a combination of alleles that have age‐specific additive effects, consistent with MA, as well as nonindependent and antagonistic genetic architectures characteristic of AP.     

Manifestation of radiation effects in cold environment: data review and modeling

The peculiarities of radiation response in animals at low environmental temperatures are analyzed in the context of radiation safety of the Arctic/Northern wildlife. The paper includes a data review on radiation effects in cold environments based on international and Russian publications since 1948, which forms a supplement to the EPIC and FREDERICA data collections. In homoiothermic and heterothermic animals, imbalances in thermoregulation caused by ionizing radiation are discussed, which increase energy loss of animals, and decrease their fitness to the Arctic/Northern climate. In poikilothermic animals, both radiation damage and recovery are temperature dependant, their rates being slow in the cold environment. At low temperatures, radiation damage of biological tissues is conserved in hidden form; when the temperature of poikilothermic animal rises to a normal level, radiation injury is developed rapidly similar to acute dose response. Additionally, a mathematical model is described, demonstrating the combined effects of chronic radiation exposures and seasonal temperature variations on a fish population. Computer simulations show that at the same level of irradiation, the overall radiation damage to Arctic/Northern poikilothermic fish is higher than that to the fish from warm climate. Considering the peculiarities of radiation effects in the cold climate, the Arctic/Northern fauna might be expected to be more vulnerable to chronic radiation stress compared to temperate fauna. In the case of acute radiation exposure during winter periods, hibernation of heterothermic and cooling of poikilothermic animals may provide temporary protection from acute radiation effects.     

A veterinary perspective on One Health in the Arctic

Exposure to long-range transported industrial chemicals, climate change and diseases is posing a risk to the overall health and populations of Arctic wildlife. Since local communities are relying on the same marine food web as marine mammals in the Arctic, it requires a One Health approach to understand the holistic ecosystem health including that of humans. Here we collect and identify gaps in the current knowledge of health in the Arctic and present the veterinary perspective of One Health and ecosystem dynamics. The review shows that exposure to persistent organic pollutants (POPs) is having multiple organ-system effects across taxa, including impacts on neuroendocrine disruption, immune suppression and decreased bone density among others. Furthermore, the warming Arctic climate is suspected to influence abiotic and biotic long-range transport and exposure pathways of contaminants to the Arctic resulting in increases in POP exposure of both wildlife and human populations. Exposure to vector-borne diseases and zoonoses may increase as well through range expansion and introduction of invasive species. It will be important in the future to investigate the effects of these multiple stressors on wildlife and local people to better predict the individual-level health risks. It is within this framework that One Health approaches offer promising opportunities to survey and pinpoint environmental changes that have effects on wildlife and human health.     

Multiple stressor effects on coral reef ecosystems

Global climate change has profound implications on species distributions and ecosystem functioning. In the coastal zone, ecological responses may be driven by various biogeochemical and physical environmental factors. Synergistic interactions can occur when the combined effects of stressors exceed their individual effects. The Red Sea, characterized by strong gradients in temperature, salinity, and nutrients along the latitudinal axis provides a unique opportunity to study ecological responses over a range of these environmental variables. Using multiple linear regression models integrating in situ, satellite and oceanographic data, we investigated the response of coral reef taxa to local stressors and recent climate variability. Taxa and functional groups responded to a combination of climate (temperature, salinity, air‐sea heat fluxes, irradiance, wind speed), fishing pressure and biogeochemical (chlorophyll a and nutrients ‐ phosphate, nitrate, nitrite) factors. The regression model for each species showed interactive effects of climate, fishing pressure and nutrient variables. The nature of the effects (antagonistic or synergistic) was dependent on the species and stressor pair. Variables consistently associated with the highest number of synergistic interactions included heat flux terms, temperature, and wind speed followed by fishing pressure. Hard corals and coralline algae abundance were sensitive to changing environmental conditions where synergistic interactions decreased their percentage cover. These synergistic interactions suggest that the negative effects of fishing pressure and eutrophication may exacerbate the impact of climate change on corals. A high number of interactions were also recorded for algae, however for this group, synergistic interactions increased algal abundance. This study is unique in applying regression analysis to multiple environmental variables simultaneously to understand stressor interactions in the field. The observed responses have important implications for understanding climate change impacts on marine ecosystems and whether managing local stressors, such as nutrient enrichment and fishing activities, may help mitigate global drivers of change.     

Adaptive thermoregulation in endotherms may alter responses to climate change

Climate change is one of the major issues facing natural populations and thus a focus of recent research has been to predict the responses of organisms to these changes. Models are becoming more complex and now commonly include physiological traits of the organisms of interest. However, endothermic species have received less attention than have ectotherms in these mechanistic models. Further, it is not clear whether responses of endotherms to climate change are modified by variation in thermoregulatory characteristics associated with phenotypic plasticity and/or adaptation to past selective pressures. Here, we review the empirical data on thermal adaptation and acclimatization in endotherms and discuss how those factors may be important in models of responses to climate change. We begin with a discussion of why thermoregulation and thermal sensitivity at high body temperatures should be co-adapted. Importantly, we show that there is, in fact, considerable variation in the ability of endotherms to tolerate high body temperatures and/or high environmental temperatures, but a better understanding of this variation will likely be critical for predicting responses to future climatic scenarios. Next, we discuss why variation in thermoregulatory characteristics should be considered when modeling the effects of climate change on heterothermic endotherms. Finally, we review some biophysical and biochemical factors that will limit adaptation and acclimation in endotherms. We consider both long-term, directional climate change and short-term (but increasingly common) anomalies in climate such as extreme heat waves and we suggest areas of important future research relating to both our basic understanding of endothermic thermoregulation and the responses of endotherms to climate change.     

Climate change shifts natural selection and the adaptive potential of the perennial forb Boechera stricta in the Rocky Mountains

Heritable genetic variation is necessary for populations to evolve in response to anthropogenic climate change. However, antagonistic genetic correlations among traits may constrain the rate of adaptation, even if substantial genetic variation exists. We examine potential genetic responses to selection by comparing multivariate genetic variance–covariances of traits and fitness (multivariate Robertson–Price identities) across different environments in a reciprocal transplant experiment of the forb Boechera stricta in the Rocky Mountains. By transplanting populations into four common gardens arrayed along an elevational gradient, and exposing populations to control and snow removal treatments, we simulated future and current climates and snowmelt regimes. Genetic variation in flowering and germination phenology declined in plants moved downslope to warmer, drier sites, suggesting that these traits may have a limited ability to evolve under future climates. Simulated climate change via snow removal altered the strength of selection on flowering traits, but we found little evidence that genetic correlations among traits are likely to affect the rate of adaptation to climate change. Overall, our results suggest that climate change may alter the evolutionary potential of B. stricta, but reduced expression of genetic variation may be a larger impediment to adaptation than constraints imposed by antagonistic genetic correlations.     

Evolutionary potential of Chamaecrista fasciculata in relation to climate change. II. Genetic architecture of three populations reciprocally planted along an environmental gradient in the great plains

Climate change will test the evolutionary potential of populations. Information regarding the genetic architecture within and among populations is essential for prediction of evolutionary outcomes. However, little is known about the distribution of genetic variation for relevant traits in natural populations or alteration of genetic architecture in a changing environment. In this study, pedigreed families from three populations of the annual prairie legume Chamaecrista fasciculata were reciprocally transplanted in three environments across a broad latitudinal range in the Great Plains. The underlying premise of this work is that northern populations will in the future experience climates similar to current-day climates further south. Estimates of narrow-sense heritability ranged from 0.053 to 0.481, suggesting the potential for evolutionary change is possible for most traits. In general, the northern population harbored less genetic variation and had lower heritability for traits than the southern population. This population also experienced large reductions in fitness, as measured by estimated lifetime fecundity, when raised in either the intermediate or the southern climate, whereas the difference between the intermediate and southern population was less extreme. For fecundity, the pattern of cross-environment additive genetic correlations was antagonistic to evolutionary change in four of six cases when native and nonnative sites were compared. Six additional antagonistic positive correlations were found for the rate of phenological development and leaf thickness. Overall, the data suggest that if climate changes as predicted, the northern population will face a severe evolutionary challenge in the future because of low heritabilities, cross-environment genetic correlations antagonistic to selection, and demographic instability due to lower seed production in a hotter and drier climate.     

Fluctuating environments impact thermal tolerance in an invasive insect species Bactrocera dorsalis (Diptera: Tephritidae)

The incidence and severity of environmental stressors associated with global climate change are increasing and insects frequently face variability in temperature and moisture regimes at variable spatio-temporal scales. Coincidental with this, is increased thermal and hydric stress on insects as warming increases vapour pressure deficit (VPD), the drying power of the air. While the effects of mean temperatures on fitness are widely documented, fluctuations in both temperature and relative humidity (RH) are largely unexplored. Here, we investigated the effects of dynamic temperature and RH fluctuations (around the mean [28°C; 65% RH]) on low and high thermal tolerance of laboratory-reared adult invasive Bactrocera dorsalis (Hendel) (Diptera: Tephritidae), measured as critical thermal minima (CT_min), critical thermal maxima (CT_max), chill coma recovery time (CCRT) and heat knockdown time (HKDT). Our results show that increased environmental amplitude significantly influenced low and high temperature responses and varied across traits tested. The highest amplitude (δ12°C; 28% RH) compromised CT_min, CCRT and HKDT traits while enhancing CT_max. Similarly, acclimation to δ3°C; 7% RH compromised both low (CT_min and CCRT) and high (CT_max and HKDT) fitness traits. Variations in fitness reported here indicate significant roles of combined thermal and moisture fluctuations on B. dorsalis fitness suggesting caveats that are worthy considering when predicting species responses to climate change. These results are significant for B. dorsalis population phenology, management, quantifying vulnerability to climate variability and may help modelling future biogeographical patterns.     

Synergistic effects of climate change and local stressors: CO2 and nutrient-driven change in subtidal rocky habitats

Climate-driven change represents the cumulative effect of global through local-scale conditions, and understanding their manifestation at local scales can empower local management. Change in the dominance of habitats is often the product of local nutrient pollution that occurs at relatively local scales (i.e. catchment scale), a critical scale of management at which global impacts will manifest. We tested whether forecasted global-scale change [elevated carbon dioxide (CO₂) and subsequent ocean acidification] and local stressors (elevated nutrients) can combine to accelerate the expansion of filamentous turfs at the expense of calcifying algae (kelp understorey). Our results not only support this model of future change, but also highlight the synergistic effects of future CO₂ and nutrient concentrations on the abundance of turfs. These results suggest that global and local stressors need to be assessed in meaningful combinations so that the anticipated effects of climate change do not create the false impression that, however complex, climate change will produce smaller effects than reality. These findings empower local managers because they show that policies of reducing local stressors (e.g. nutrient pollution) can reduce the effects of global stressors not under their governance (e.g. ocean acidification). The connection between research and government policy provides an example whereby knowledge (and decision making) across local through global scales provides solutions to some of the most vexing challenges for attaining social goals of sustainability, biological conservation and economic development.     

Meta-analysis on the responses of traits of different taxonomic groups to global and local stressors

Climate change and pollution are considered as major drivers of biodiversity loss. Climate change is a global multi-stressor, whereas pollution predominantly acts on the local scale. Organisms traits provide mechanistic links between biotic responses and stressors. We reviewed and analyzed the literature on the responses of vertebrates, invertebrates, microorganisms and plants traits to climate change (437 studies) and pollution (121 studies), to assess whether there was uniformity (i.e. convergence) in the responses of traits to the multi-stressors. For climate change, the traits related to tolerance responded uniformly across taxonomic groups, indicating trait convergence. For pollution, the low number of studies hampered a comparison across taxonomic groups. However, aquatic invertebrates that are tolerant, or exhibit high dispersal or reproduction capacities increased in response to pollution, whereas body mass and size increased in phytoplankton and fish, respectively. We provide a set of traits that have the potential to predict ecosystem-wide effects of climate change and pollution.     

Do physical plant litter traits explain non‐additivity in litter mixtures? A test of the improved microenvironmental conditions theory

The decomposition rates of plant litter mixtures often deviate from the averaged rates of monocultures of their component litter species. The mechanisms behind these non‐additive effects in decomposition of litter mixtures are lively debated. One plausible explanation for non‐additive effects is given by the improved microenvironmental condition (IMC) theory. According to this theory, plant litter species, whose physical characteristics improve the microclimatic conditions for decomposers, will promote the decomposition of their co‐occurring litter species. We tested the IMC theory in relation to leaf litter and soil moisture in two contrasting moisture conditions in a dry subarctic mountain birch forest with vascular plant leaf litters of poor and high quality. The non‐additive effects in mass loss of litter mixtures increased when moisture conditions in litter and soil became more favourable for plant litter decomposition. The sign of this increase (antagonistic or synergistic) in non‐additive effects was more predictable for litter mixtures of poor litter quality. Although the specific mechanisms underlying the IMC theory depended on the litter quality of the litter mixtures, a standardized water holding capacity (WHC) was the litter trait most closely related to the non‐additive effects in mixtures of both poor and high quality litter types. Furthermore, we found that higher dissimilarity in WHC traits between the component litter species in a mixture increased synergistic effects in litter mixtures under limiting moisture conditions. However, under improved moisture conditions, increased antagonistic effects were observed. Thus, we found clear support for the IMC theory and showed that climatic conditions and leaf litter physical traits determine whether the non‐additive effects in litter mixtures are antagonistic or synergistic. Our study emphasizes the need to include litter physical traits into predictive models of mixing effects on plant litter decomposition and in general suggests climate specificity into these models.     

Individual and combined effects of fine sediment and the herbicide glyphosate on benthic macroinvertebrates and stream ecosystem function

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Food and temperature stressors have opposing effects in determining flexible migration decisions in brown trout (Salmo trutta)

With rapid global change, organisms in natural systems are exposed to a multitude of stressors that likely co‐occur, with uncertain impacts. We explored individual and cumulative effects of co‐occurring environmental stressors on the striking, yet poorly understood, phenomenon of facultative migration. We reared offspring of a brown trout population that naturally demonstrates facultative anadromy (sea migration), under different environmental stressor treatments and measured life history responses in terms of migratory tactics and freshwater maturation rates. Juvenile fish were exposed to reduced food availability, temperatures elevated to 1.8°C above natural conditions or both treatments in combination over 18 months of experimental tank rearing. When considered in isolation, reduced food had negative effects on the size, mass and condition of fish across the experiment. We detected variable effects of warm temperatures (negative effects on size and mass, but positive effect on lipids). When combined with food restriction, temperature effects on these traits were less pronounced, implying antagonistic stressor effects on morphological traits. Stressors combined additively, but had opposing effects on life history tactics: migration increased and maturation rates decreased under low food conditions, whereas the opposite occurred in the warm temperature treatment. Not all fish had expressed maturation or migration tactics by the end of the study, and the frequency of these ‘unassigned’ fish was higher in food deprivation treatments, but lower in warm treatments. Fish showing migration tactics were smaller and in poorer condition than fish showing maturation tactics, but were similar in size to unassigned fish. We further detected effects of food restriction on hypo‐osmoregulatory function of migrants that may influence the fitness benefits of the migratory tactic at sea. We also highlight that responses to multiple stressors may vary depending on the response considered. Collectively, our results indicate contrasting effects of environmental stressors on life history trajectories in a facultatively migratory species.