Responses of seagrass to anthropogenic and natural disturbances do not equally translate to its consumers

Coastal communities are under threat from many and often co‐occurring local (e.g., pollution, eutrophication) and global stressors (e.g., climate change), yet understanding the interactive and cumulative impacts of multiple stressors in ecosystem function is far from being accomplished. Ecological redundancy may be key for ecosystem resilience, but there are still many gaps in our understanding of interspecific differences within a functional group, particularly regarding response diversity, that is, whether members of a functional group respond equally or differently to anthropogenic stressors. Herbivores are critical in determining plant community structure and the transfer of energy up the food web. Human disturbances may alter the ecological role of herbivory by modifying the defense strategies of plants and thus the feeding patterns and performance of herbivores. We conducted a suite of experiments to examine the independent and interactive effects of anthropogenic (nutrient and CO₂ additions) and natural (simulated herbivory) disturbances on a seagrass and its interaction with two common generalist consumers to understand how multiple disturbances can impact both a foundation species and a key ecological function (herbivory) and to assess the potential existence of response diversity to anthropogenic and natural changes in these systems. While all three disturbances modified seagrass defense traits, there were contrasting responses of herbivores to such plant changes. Both CO₂ and nutrient additions influenced herbivore feeding behavior, yet while sea urchins preferred nutrient‐enriched seagrass tissue (regardless of other experimental treatments), isopods were deterred by these same plant tissues. In contrast, carbon enrichment deterred sea urchins and attracted isopods, while simulated herbivory only influenced isopod feeding choice. These contrasting responses of herbivores to disturbance‐induced changes in seagrass help to better understand the ecological functioning of seagrass ecosystems in the face of human disturbances and may have important implications regarding the resilience and conservation of these threatened ecosystems.     

Assessing physiological tipping point of sea urchin larvae exposed to a broad range of pH

Our ability to project the impact of global change on marine ecosystem is limited by our poor understanding on how to predict species sensitivity. For example, the impact of ocean acidification is highly species‐specific, even in closely related taxa. The aim of this study was to test the hypothesis that the tolerance range of a given species to decreased pH corresponds to their natural range of exposure. Larvae of the green sea urchin Strongylocentrotus droebachiensis were cultured from fertilization to metamorphic competence (29 days) under a wide range of pH (from pH_T = 8.0/pCO₂ ≈ 480 μatm to pH_T = 6.5/pCO₂ ≈ 20 000 μatm) covering present (from pH_T 8.7 to 7.6), projected near‐future variability (from pH_T 8.3 to 7.2) and beyond. Decreasing pH impacted all tested parameters (mortality, symmetry, growth, morphometry and respiration). Development of normal, although showing morphological plasticity, swimming larvae was possible as low as pH_T ≥ 7.0. Within that range, decreasing pH increased mortality and asymmetry and decreased body length (BL) growth rate. Larvae raised at lowered pH and with similar BL had shorter arms and a wider body. Relative to a given BL, respiration rates and stomach volume both increased with decreasing pH suggesting changes in energy budget. At the lowest pHs (pH_T ≤ 6.5), all the tested parameters were strongly negatively affected and no larva survived past 13 days post fertilization. In conclusion, sea urchin larvae appeared to be highly plastic when exposed to decreased pH until a physiological tipping point at pH_T = 7.0. However, this plasticity was associated with direct (increased mortality) and indirect (decreased growth) consequences for fitness.     

Irukandji jellyfish polyps exhibit tolerance to interacting climate change stressors

Increasing ocean temperatures and strengthening boundary currents have caused the poleward migration of many marine species. Cubozoan jellyfish known to cause Irukandji syndrome have historically been confined to tropical waters but may be expanding into subtropical regions. Here, we examine the interactive effects of warming and acidification on the population dynamics of polyps of an Irukandji jellyfish, Alatina nr mordens, and the formation of statoliths in newly metamorphosed medusae, to determine if this jellyfish could tolerate future conditions predicted for southeast Queensland (SEQ), Australia. Two experiments, examining the orthogonal factors of temperature and pH, were undertaken. Experiment 1 mimicked the current, ca. 2050 and ca. 2100 summer temperature and pH conditions predicted for SEQ using A1F1 scenarios (temperature: 25, 27, 29 °C; pH: 7.9, 7.8, 7.6) and Experiment 2 mimicked current and future winter conditions (18 and 22 °C, pH 7.9, 7.8, 7.6). All polyps in Experiment 1 survived and budded. Fewer polyps budded in the lower pH treatments; however, patterns varied slightly among temperature treatments. Statoliths at pH 7.6 were 24% narrower than those at pH 7.8 and 7.9. Most polyps survived the winter conditions mimicked by Experiment 2 but only polyps in the 22 °C, pH 7.9 treatment increased significantly. The current absence of A. nr mordens medusae in SEQ, despite the polyps' ability to tolerate the current temperature and pH conditions, suggests that ecological, rather than abiotic factors currently limit their distribution. Observations that budding was lower under low pH treatments suggest that rates of asexual reproduction will likely be much slower in the future. We consider that A. nr mordens polyps are likely to tolerate future conditions but are unlikely to thrive in the long term. However, if polyps can overcome potential ecological boundaries and acidification proceeds slowly A. nr mordens could expand polewards in the short term.     

Biomineralization plasticity and environmental heterogeneity predict geographical resilience patterns of foundation species to future change

Although geographical patterns of species' sensitivity to environmental changes are defined by interacting multiple stressors, little is known about compensatory processes shaping regional differences in organismal vulnerability. Here, we examine large‐scale spatial variations in biomineralization under heterogeneous environmental gradients of temperature, salinity and food availability across a 30° latitudinal range (3,334 km), to test whether plasticity in calcareous shell production and composition, from juveniles to large adults, mediates geographical patterns of resilience to climate change in critical foundation species, the mussels Mytilus edulis and M. trossulus. We find shell calcification decreased towards high latitude, with mussels producing thinner shells with a higher organic content in polar than temperate regions. Salinity was the best predictor of within‐region differences in mussel shell deposition, mineral and organic composition. In polar, subpolar, and Baltic low‐salinity environments, mussels produced thin shells with a thicker external organic layer (periostracum), and an increased proportion of calcite (prismatic layer, as opposed to aragonite) and organic matrix, providing potentially higher resistance against dissolution in more corrosive waters. Conversely, in temperate, higher salinity regimes, thicker, more calcified shells with a higher aragonite (nacreous layer) proportion were deposited, which suggests enhanced protection under increased predation pressure. Interacting effects of salinity and food availability on mussel shell composition predict the deposition of a thicker periostracum and organic‐enriched prismatic layer under forecasted future environmental conditions, suggesting a capacity for increased protection of high‐latitude populations from ocean acidification. These findings support biomineralization plasticity as a potentially advantageous compensatory mechanism conferring Mytilus species a protective capacity for quantitative and qualitative trade‐offs in shell deposition as a response to regional alterations of abiotic and biotic conditions in future environments. Our work illustrates that compensatory mechanisms, driving plastic responses to the spatial structure of multiple stressors, can define geographical patterns of unanticipated species resilience to global environmental change.     

Quantitative analysis of oyster larval proteome provides new insights into the effects of multiple climate change stressors

The metamorphosis of planktonic larvae of the Pacific oyster (Crassostrea gigas) underpins their complex life‐history strategy by switching on the molecular machinery required for sessile life and building calcite shells. Metamorphosis becomes a survival bottleneck, which will be pressured by different anthropogenically induced climate change‐related variables. Therefore, it is important to understand how metamorphosing larvae interact with emerging climate change stressors. To predict how larvae might be affected in a future ocean, we examined changes in the proteome of metamorphosing larvae under multiple stressors: decreased pH (pH 7.4), increased temperature (30 °C), and reduced salinity (15 psu). Quantitative protein expression profiling using iTRAQ‐LC‐MS/MS identified more than 1300 proteins. Decreased pH had a negative effect on metamorphosis by down‐regulating several proteins involved in energy production, metabolism, and protein synthesis. However, warming switched on these down‐regulated pathways at pH 7.4. Under multiple stressors, cell signaling, energy production, growth, and developmental pathways were up‐regulated, although metamorphosis was still reduced. Despite the lack of lethal effects, significant physiological responses to both individual and interacting climate change related stressors were observed at proteome level. The metamorphosing larvae of the C. gigas population in the Yellow Sea appear to have adequate phenotypic plasticity at the proteome level to survive in future coastal oceans, but with developmental and physiological costs.     

Regulation of phosphate uptake reveals cyanobacterial bloom resilience to shifting N:P ratios

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Vulnerability of global coral reef habitat suitability to ocean warming,acidification and eutrophication

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ω_arag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ω_arag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to ‘buy time’, while international agreements are found to abate global stressors.     

Viral attack exacerbates the susceptibility of a bloom‐forming alga to ocean acidification

Both ocean acidification and viral infection bring about changes in marine phytoplankton physiological activities and community composition. However, little information is available on how the relationship between phytoplankton and viruses may be affected by ocean acidification and what impacts this might have on photosynthesis‐driven marine biological CO₂ pump. Here, we show that when the harmful bloom alga Phaeocystis globosa is infected with viruses under future ocean conditions, its photosynthetic performance further decreased and cells became more susceptible to stressful light levels, showing enhanced photoinhibition and reduced carbon fixation, up‐regulation of mitochondrial respiration and decreased virus burst size. Our results indicate that ocean acidification exacerbates the impacts of viral attack on P. globosa, which implies that, while ocean acidification directly influences marine primary producers, it may also affect them indirectly by altering their relationship with viruses. Therefore, viruses as a biotic stressor need to be invoked when considering the overall impacts of climate change on marine productivity and carbon sequestration.     

The impact of CO2 emission scenarios and nutrient enrichment on a common coral reef macroalga is modified by temporal effects

Future coral reefs are expected to be subject to higher pCO₂ and temperature due to anthropogenic greenhouse gas emissions. Such global stressors are often paired with local stressors thereby potentially modifying the response of organisms. Benthic macroalgae are strong competitors to corals and are assumed to do well under future conditions. The present study aimed to assess the impact of past and future CO₂ emission scenarios as well as nutrient enrichment on the growth, productivity, pigment, and tissue nutrient content of the common tropical brown alga Chnoospora implexa. Two experiments were conducted to assess the differential impacts of the manipulated conditions in winter and spring. Chnoospora implexa's growth rate averaged over winter and spring declined with increasing pCO₂ and temperature. Furthermore, nutrient enrichment did not affect growth. Highest growth was observed under spring pre‐industrial (PI) conditions, while slightly reduced growth was observed under winter A1FI (“business‐as‐usual”) scenarios. Productivity was not a good proxy for growth, as net O₂ flux increased under A1FI conditions. Nutrient enrichment, whilst not affecting growth, led to luxury nutrient uptake that was greater in winter than in spring. The findings suggest that in contrast with previous work, C. implexa is not likely to show enhanced growth under future conditions in isolation or in conjunction with nutrient enrichment. Instead, the results suggest that greatest growth rates for this species appear to be a feature of the PI past, with A1FI winter conditions leading to potential decreases in the abundance of this species from present day levels.     

Amelioration of ocean acidification and warming effects through physiological buffering of a macroalgae

Concurrent anthropogenic global climate change and ocean acidification are expected to have a negative impact on calcifying marine organisms. While knowledge of biological responses of organisms to oceanic stress has emerged from single‐species experiments, these do not capture ecologically relevant scenarios where the potential for multi‐organism physiological interactions is assessed. Marine algae provide an interesting case study, as their photosynthetic activity elevates pH in the surrounding microenvironment, potentially buffering more acidic conditions for associated epiphytes. We present findings that indicate increased tolerance of an important epiphytic foraminifera, Marginopora vertebralis, to the effects of increased temperature (±3°C) and pCO₂ (~1,000 µatm) when associated with its common algal host, Laurencia intricata. Specimens of M. vertebralis were incubated for 15 days in flow‐through aquaria simulating current and end‐of‐century temperature and pH conditions. Physiological measures of growth (change in wet weight), calcification (measured change in total alkalinity in closed bottles), photochemical efficiency (Fv/Fm), total chlorophyll, photosynthesis (oxygen flux), and respiration were determined. When incubated in isolation, M. vertebralis exhibited reduced growth in end‐of‐century projections of ocean acidification conditions, while calcification rates were lowest in the high‐temperature, low‐pH treatment. Interestingly, association with L. intricata ameliorated these stress effects with the growth and calcification rates of M. vertebralis being similar to those observed in ambient conditions. Total chlorophyll levels in M. vertebralis decreased when in association with L. intricata, while maximum photochemical efficiency increased in ambient conditions. Net production estimates remained similar between M. vertebralis in isolation and in association with L. intricata, although both production and respiration rates of M. vertebralis were significantly higher when associated with L. intricata. These results indicate that the association with L. intricata increases the resilience of M. vertebralis to climate change stress, providing one of the first examples of physiological buffering by a marine alga that can ameliorate the negative effects of changing ocean conditions.     

Contrasting effects of rising CO2 on primary production and ecological stoichiometry at different nutrient levels

Although rising CO₂ concentrations are thought to promote the growth and alter the carbon : nutrient stoichiometry of primary producers, several studies have reported conflicting results. To reconcile these contrasting results, we tested the following hypotheses: rising CO₂ levels (1) will increase phytoplankton biomass more at high nutrient loads than at low nutrient loads, but (2) will increase their carbon : nutrient stoichiometry more at low than at high nutrient loads. We formulated a mathematical model to predict dynamic changes in phytoplankton population density, elemental stoichiometry and inorganic carbon chemistry in response to rising CO₂. The model was tested in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa. The model predictions and experimental results confirmed the hypotheses. Our findings provide a novel theoretical framework to understand and predict effects of rising CO₂ concentrations on primary producers and their nutritional quality as food for herbivores under different nutrient conditions.     

Sensitivity of sea urchin fertilization to pH varies across a natural pH mosaic

In the coastal ocean, temporal fluctuations in pH vary dramatically across biogeographic ranges. How such spatial differences in pH variability regimes might shape ocean acidification resistance in marine species remains unknown. We assessed the pH sensitivity of the sea urchin Strongylocentrotus purpuratus in the context of ocean pH variability. Using unique male–female pairs, originating from three sites with similar mean pH but different variability and frequency of low pH (pH_T ≤ 7.8) exposures, fertilization was tested across a range of pH (pH_T 7.61–8.03) and sperm concentrations. High fertilization success was maintained at low pH via a slight right shift in the fertilization function across sperm concentration. This pH effect differed by site. Urchins from the site with the narrowest pH variability regime exhibited the greatest pH sensitivity. At this site, mechanistic fertilization dynamics models support a decrease in sperm–egg interaction rate with decreasing pH. The site differences in pH sensitivity build upon recent evidence of local pH adaptation in S. purpuratus and highlight the need to incorporate environmental variability in the study of global change biology.     

Effects of multiple drivers of ocean global change on the physiology and functional gene expression of the coccolithophore Emiliania huxleyi

Ongoing ocean global change due to anthropogenic activities is causing multiple chemical and physical seawater properties to change simultaneously, which may affect the physiology of marine phytoplankton. The coccolithophore Emiliania huxleyi is a model species often employed in the study of the marine carbon cycle. The effect of ocean acidification (OA) on coccolithophore calcification has been extensively studied; however, physiological responses to multiple environmental drivers are still largely unknown. Here we examined two‐way and multiple driver effects of OA and other key environmental drivers—nitrate, phosphate, irradiance, and temperature—on the growth, photosynthetic, and calcification rates, and the elemental composition of E. huxleyi. In addition, changes in functional gene expression were examined to understand the molecular mechanisms underpinning the physiological responses. The single driver manipulation experiments suggest decreased nitrate supply being the most important driver regulating E. huxleyi physiology, by significantly reducing the growth, photosynthetic, and calcification rates. In addition, the interaction of OA and decreased nitrate supply (projected for year 2100) had more negative synergistic effects on E. huxleyi physiology than all other two‐way factorial manipulations, suggesting a linkage between the single dominant driver (nitrate) effects and interactive effects with other drivers. Simultaneous manipulation of all five environmental drivers to the conditions of the projected year 2100 had the largest negative effects on most of the physiological metrics. Furthermore, functional genes associated with inorganic carbon acquisition (RubisCO, AEL1, and δCA) and calcification (CAX3, AEL1, PATP, and NhaA2) were most downregulated by the multiple driver manipulation, revealing linkages between responses of functional gene expression and associated physiological metrics. These findings together indicate that for more holistic projections of coccolithophore responses to future ocean global change, it is necessary to understand the relative importance of environmental drivers both individually (i.e., mechanistic understanding) and interactively (i.e., cumulative effect) on coccolithophore physiology.     

Response of foundation macrophytes to near‐natural simulated marine heatwaves

Marine heatwaves have been observed worldwide and are expected to increase in both frequency and intensity due to climate change. Such events may cause ecosystem reconfigurations arising from species range contraction or redistribution, with ecological, economic and social implications. Macrophytes such as the brown seaweed Fucus vesiculosus and the seagrass Zostera marina are foundation species in many coastal ecosystems of the temperate northern hemisphere. Hence, their response to extreme events can potentially determine the fate of associated ecosystems. Macrophyte functioning is intimately linked to the maintenance of photosynthesis, growth and reproduction, and resistance against pathogens, epibionts and grazers. We investigated morphological, physiological, pathological and chemical defence responses of western Baltic Sea F. vesiculosus and Z. marina populations to simulated near‐natural marine heatwaves. Along with (a) the control, which constituted no heatwave but natural stochastic temperature variability (0HW), two treatments were applied: (b) two late‐spring heatwaves (June, July) followed by a summer heatwave (August; 3HW) and (c) a summer heatwave only (1HW). The 3HW treatment was applied to test whether preconditioning events can modulate the potential sensitivity to the summer heatwave. Despite the variety of responses measured in both species, only Z. marina growth was impaired by the accumulative heat stress imposed by the 3HW treatment. Photosynthetic rate, however, remained high after the last heatwave indicating potential for recovery. Only epibacterial abundance was significantly affected in F. vesiculosus. Hence both macrophytes, and in particular F. vesiculosus, seem to be fairly tolerant to short‐term marine heatwaves at least at the intensities applied in this experiment (up to 5°C above mean temperature over a period of 9 days). This may partly be due to the fact that F. vesiculosus grows in a highly variable environment, and may have a high phenotypic plasticity.     

Identifying species at extinction risk using global models of anthropogenic impact

The International Union for Conservation of Nature Red List of Endangered Species employs a robust, standardized approach to assess extinction threat focussed on taxa approaching an end‐point in population decline. Used alone, we argue this enforces a reactive approach to conservation. Species not assessed as threatened but which occur predominantly in areas with high levels of anthropogenic impact may require proactive conservation management to prevent loss. We matched distribution and bathymetric range data from the global Red List assessment of 632 species of marine cone snails with human impacts and projected ocean thermal stress and aragonite saturation (a proxy for ocean acidification). Our results show 67 species categorized as ‘Least Concern’ have 70% or more of their occupancy in places subject to high and very high levels of human impact with 18 highly restricted species (range <100 km²) living exclusively in such places. Using a range‐rarity scoring method we identified where clusters of endemic species are subject to all three stressors: high human impact, declining aragonite saturation levels and elevated thermal stress. Our approach reinforces Red List threatened status, highlights candidate species for reassessment, contributes important evidential data to minimize data deficiency and identifies regions and species for proactive conservation.     

Long-term response and recovery to nutrient addition of a partitioned arctic lake

1. To study the bottom‐up linkages in arctic lakes, we treated one side of a partitioned lake with inorganic nitrogen and phosphorus for a 6‐week period each summer for 6 years starting in the summer of 1985. We took a variety of weekly measurements to determine the impact of the nutrient loading on the lake and continued weekly measurements for 2–6 years after the cessation of nutrient loading to observe the recovery of the treated side. The loading rates (2.91 mmol N m^?2 day^?1 and 0.23 mmol P m^?2 day^?1) were five times the calculated loading rates for Toolik Lake, located nearby. 2. In all 6 years of nutrient addition, phytoplankton biomass and productivity were greater in the treated sector than the reference sector. In the first 4 years of nutrient addition there was no flux of phosphorus from the mineral‐rich sediments. This changed in the last 2 years of nutrient addition as phosphorus was released to the lake. 3. The response of the animal community to increased plant production was mixed. One of the four macro‐zooplankton species (Daphnia longiremis) increased in number by about twofold in the first 5 years. However, the copepod Cyclops scutifer showed no response during the treatment phase of the study. The benthic invertebrate response was also mixed. After a 2‐year lag time the snail Lymnaea elodes increased in the treated lake sector but chironomids did not. 4. Ecosystem response to fertilisation was not controlled solely by nutrient addition because phosphorus was not recycled from the sediments until the last 2 years of nutrient addition. Phytoplankton still showed the effects of nutrient addition in the recovery period and the hypolimnion of the treated sector was still anaerobic starting at 6 m in 1996.     

Linking genotype to phenotype in a changing ocean: inferring the genomic architecture of a blue mussel stress response with genome‐wide association

A key component to understanding the evolutionary response to a changing climate is linking underlying genetic variation to phenotypic variation in stress response. Here, we use a genome‐wide association approach (GWAS) to understand the genetic architecture of calcification rates under simulated climate stress. We take advantage of the genomic gradient across the blue mussel hybrid zone (Mytilus edulis and Mytilus trossulus) in the Gulf of Maine (GOM) to link genetic variation with variance in calcification rates in response to simulated climate change. Falling calcium carbonate saturation states are predicted to negatively impact many marine organisms that build calcium carbonate shells – like blue mussels. We sampled wild mussels and measured net calcification phenotypes after exposing mussels to a ‘climate change’ common garden, where we raised temperature by 3°C, decreased pH by 0.2 units and limited food supply by filtering out planktonic particles >5 μm, compared to ambient GOM conditions in the summer. This climate change exposure greatly increased phenotypic variation in net calcification rates compared to ambient conditions. We then used regression models to link the phenotypic variation with over 170 000 single nucleotide polymorphism loci (SNPs) generated by genotype by sequencing to identify genomic locations associated with calcification phenotype, and estimate heritability and architecture of the trait. We identified at least one of potentially 2–10 genomic regions responsible for 30% of the phenotypic variation in calcification rates that are potential targets of natural selection by climate change. Our simulations suggest a power of 13.7% with our study's average effective sample size of 118 individuals and rare alleles, but a power of >90% when effective sample size is 900.     

Scaling up experimental ocean acidification and warming research: from individuals to the ecosystem

Understanding long‐term, ecosystem‐level impacts of climate change is challenging because experimental research frequently focuses on short‐term, individual‐level impacts in isolation. We address this shortcoming first through an interdisciplinary ensemble of novel experimental techniques to investigate the impacts of 14‐month exposure to ocean acidification and warming (OAW) on the physiology, activity, predatory behaviour and susceptibility to predation of an important marine gastropod (Nucella lapillus). We simultaneously estimated the potential impacts of these global drivers on N. lapillus population dynamics and dispersal parameters. We then used these data to parameterize a dynamic bioclimatic envelope model, to investigate the consequences of OAW on the distribution of the species in the wider NE Atlantic region by 2100. The model accounts also for changes in the distribution of resources, suitable habitat and environment simulated by finely resolved biogeochemical models, under three IPCC global emissions scenarios. The experiments showed that temperature had the greatest impact on individual‐level responses, while acidification had a similarly important role in the mediation of predatory behaviour and susceptibility to predators. Changes in Nucella predatory behaviour appeared to serve as a strategy to mitigate individual‐level impacts of acidification, but the development of this response may be limited in the presence of predators. The model projected significant large‐scale changes in the distribution of Nucella by the year 2100 that were exacerbated by rising greenhouse gas emissions. These changes were spatially heterogeneous, as the degree of impact of OAW on the combination of responses considered by the model varied depending on local‐environmental conditions and resource availability. Such changes in macro‐scale distributions cannot be predicted by investigating individual‐level impacts in isolation, or by considering climate stressors separately. Scaling up the results of experimental climate change research requires approaches that account for long‐term, multiscale responses to multiple stressors, in an ecosystem context.