Animal behaviour shapes the ecological effects of ocean acidification and warming: moving from individual to community‐level responses

Biological communities are shaped by complex interactions between organisms and their environment as well as interactions with other species. Humans are rapidly changing the marine environment through increasing greenhouse gas emissions, resulting in ocean warming and acidification. The first response by animals to environmental change is predominantly through modification of their behaviour, which in turn affects species interactions and ecological processes. Yet, many climate change studies ignore animal behaviour. Furthermore, our current knowledge of how global change alters animal behaviour is mostly restricted to single species, life phases and stressors, leading to an incomplete view of how coinciding climate stressors can affect the ecological interactions that structure biological communities. Here, we first review studies on the effects of warming and acidification on the behaviour of marine animals. We demonstrate how pervasive the effects of global change are on a wide range of critical behaviours that determine the persistence of species and their success in ecological communities. We then evaluate several approaches to studying the ecological effects of warming and acidification, and identify knowledge gaps that need to be filled, to better understand how global change will affect marine populations and communities through altered animal behaviours. Our review provides a synthesis of the far‐reaching consequences that behavioural changes could have for marine ecosystems in a rapidly changing environment. Without considering the pervasive effects of climate change on animal behaviour we will limit our ability to forecast the impacts of ocean change and provide insights that can aid management strategies.     

Future warming and acidification effects on anti‐fouling and anti‐herbivory traits of the brown alga Fucus vesiculosus (Phaeophyceae)

Human‐induced ocean warming and acidification have received increasing attention over the past decade and are considered to have substantial consequences for a broad range of marine species and their interactions. Understanding how these interactions shift in response to climate change is particularly important with regard to foundation species, such as the brown alga Fucus vesiculosus. This macroalga represents the dominant habitat former on coastal rocky substrata of the Baltic Sea, fulfilling functions essential for the entire benthic community. Its ability to withstand extensive fouling and herbivory regulates the associated community and ecosystem dynamics. This study tested the interactive effects of future warming, acidification, and seasonality on the interactions of a marine macroalga with potential foulers and consumers. F. vesiculosus rockweeds were exposed to different combinations of conditions predicted regionally for the year 2100 (+?5°C, +?700 μatm CO₂) using multifactorial long‐term experiments in novel outdoor benthic mesocosms (“Benthocosms”) over 9–12‐week periods in four seasons. Possible shifts in the macroalgal susceptibility to fouling and consumption were tested using consecutive bioassays. Algal susceptibility to fouling and grazing varied substantially among seasons and between treatments. In all seasons, warming predominantly affected anti‐fouling and anti‐herbivory interactions while acidification had a subtle nonsignificant influence. Interestingly, anti‐microfouling activity was highest during winter under warming, while anti‐macrofouling and anti‐herbivory activities were highest in the summer under warming. These contrasting findings indicate that seasonal changes in anti‐fouling and anti‐herbivory traits may interact with ocean warming in altering F. vesiculosus community composition in the future.     

Ocean acidification ameliorates harmful effects of warming in primary consumer

Climate change‐induced warming and ocean acidification are considered two imminent threats to marine biodiversity and current ecosystem structures. Here, we have for the first time examined an animal's response to a complete life cycle of exposure to co‐occurring warming (+3°C) and ocean acidification (+1,600 μatm CO₂), using the key subarctic planktonic copepod, Calanus finmarchicus, as a model species. The animals were generally negatively affected by warming, which significantly reduced the females’ energy status and reproductive parameters (respectively, 95% and 69%–87% vs. control). Unexpectedly, simultaneous acidification partially offset the negative effect of warming in an antagonistic manner, significantly improving reproductive parameters and hatching success (233%–340% improvement vs. single warming exposure). The results provide proof of concept that ocean acidification may partially offset negative effects caused by warming in some species. Possible explanations and ecological implications for the observed antagonistic effect are discussed.     

Temperature,but not pH,compromises sea urchin fertilization and early development under near-future climate change scenarios

Global warming is causing ocean warming and acidification. The distribution of Heliocidaris erythrogramma coincides with the eastern Australia climate change hot spot, where disproportionate warming makes marine biota particularly vulnerable to climate change. In keeping with near-future climate change scenarios, we determined the interactive effects of warming and acidification on fertilization and development of this echinoid. Experimental treatments (20–26°C, pH 7.6–8.2) were tested in all combinations for the ‘business-as-usual’ scenario, with 20°C/pH 8.2 being ambient. Percentage of fertilization was high (>89%) across all treatments. There was no difference in percentage of normal development in any pH treatment. In elevated temperature conditions, +4°C reduced cleavage by 40 per cent and +6°C by a further 20 per cent. Normal gastrulation fell below 4 per cent at +6°C. At 26°C, development was impaired. As the first study of interactive effects of temperature and pH on sea urchin development, we confirm the thermotolerance and pH resilience of fertilization and embryogenesis within predicted climate change scenarios, with negative effects at upper limits of ocean warming. Our findings place single stressor studies in context and emphasize the need for experiments that address ocean warming and acidification concurrently. Although ocean acidification research has focused on impaired calcification, embryos may not reach the skeletogenic stage in a warm ocean.     

Combined effects of two ocean change stressors, warming and acidification, on fertilization and early development of the Antarctic echinoid Sterechinus neumayeri

The effects of concurrent ocean warming and acidification on Antarctic marine benthos warrant investigation as little is known about potential synergies between these climate change stressors. We examined the interactive effects of warming and acidification on fertilization and embryonic development of the ecologically important sea urchin Sterechinus neumayeri reared from fertilization in elevated temperature (+1.5°C and 3°C) and decreased pH (?0.3 and ?0.5 pH units) treatments. Fertilization using gametes from multiple males and females, to represent populations of spawners, was resilient to acidification at ambient temperature (0°C). At elevated temperatures, there was a negative interactive effect of temperature and pH on percentage of fertilization (11% reduction at 3°C). For cleavage stage embryos, there was a significant, but small reduction (6%) in the percentage of normal embryos at pH 7.5. For blastulae, a 10–11% decrease in normal development occurred in the +3°C treatments across all pH levels. Our results highlight the importance of considering the impacts of both temperature and pH in assessing the life history response of S. neumayeri in a changing polar ocean. While fertilization and development to the blastula stage were robust to levels of temperature and pH change predicted over coming decades, deleterious interactive effects were evident between these stressors at levels projected to occur by 2100 and beyond.     

The future of the northeast Atlantic benthic flora in a high CO2 world

Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO₂ emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf‐forming seaweeds.     

Complex and interactive effects of ocean acidification and warming on the life span of a marine trematode parasite

Human activities have caused an increase in atmospheric CO₂ over the last 250 years, leading to unprecedented rates of change in seawater pH and temperature. These global scale processes are now commonly referred to as ocean acidification and warming, and have the potential to substantially alter the physiological performance of many marine organisms. It is vital that the effects of ocean acidification and warming on marine organisms are explored so that we can predict how marine communities may change in future. In particular, the effect of ocean acidification and warming on host-parasite dynamics is poorly understood, despite the ecological importance of these relationships. Here, we explore the response of one himasthlid trematode, Himasthla sp., an abundant and broadly distributed species of marine parasite, to combinations of elevated temperature and pCO₂ that represent physiological extremes, pre-industrial conditions, and end of century predictions. Specifically, we quantified the life span of the free-living cercarial stage under elevated temperature and pCO₂, focussing our research on functional life span (the time cercariae spend actively swimming) and absolute life span (the period before death). We found that the effects of temperature and pCO₂ were complex and interactive. Overall, increased temperature negatively affected functional and absolute life span, e.g. across all pCO₂ treatments the average time to 50% cessation of active swimming was approximately 8 h at 5 °C, 6 h at 15 °C, 4 h at 25 °C, and 2 h at 40 °C. The effect of pCO₂, which significantly affected absolute life span, was highly variable across temperature treatments. These results strongly suggest that ocean acidification and warming may alter the transmission success of trematode cercariae, and potentially reduce the input of cercariae to marine zooplankton. Either outcome could substantially alter the community structure of coastal marine systems.     

Future ocean climate homogenizes communities across habitats through diversity loss and rise of generalist species

Predictions of the effects of global change on ecological communities are largely based on single habitats. Yet in nature, habitats are interconnected through the exchange of energy and organisms, and the responses of local communities may not extend to emerging community networks (i.e., metacommunities). Using large mesocosms and meiofauna communities as a model system, we investigated the interactive effects of ocean warming and acidification on the structure of marine metacommunities from three shallow‐water habitats: sandy soft‐bottoms, marine vegetation, and rocky reef substrates. Primary producers and detritus—key food sources for meiofauna—increased in biomass under the combined effect of temperature and acidification. The enhanced bottom‐up forcing boosted nematode densities but impoverished the functional and trophic diversity of nematode metacommunities. The combined climate stressors further homogenized meiofauna communities across habitats. Under present‐day conditions metacommunities were structured by habitat type, but under future conditions they showed an unstructured random pattern with fast‐growing generalist species dominating the communities of all habitats. Homogenization was likely driven by local species extinctions, reducing interspecific competition that otherwise could have prevented single species from dominating multiple niches. Our findings reveal that climate change may simplify metacommunity structure and prompt biodiversity loss, which may affect the biological organization and resilience of marine communities.     

Noncalcifying larvae in a changing ocean: warming,not acidification/hypercapnia,is the dominant stressor on development of the sea star Meridiastra calcar

Climate change driven ocean warming and acidification is potentially detrimental to the sensitive planktonic life stages of benthic marine invertebrates. Research has focused on the effects of acidification on calcifying larvae with a paucity of data on species with alternate developmental strategies and on the interactive effects of warming and acidification. To determine the impact of climate change on a conspicuous component of the intertidal fauna of southeast Australia, the development of the noncalcifying lecithotrophic larvae of the sea star Meridiastra calcar was investigated in the setting of predicted ocean warming (+2 to 4 ^°C) and acidification (?0.4 to 0.6 pH units) for 2100 and beyond in all combinations of stressors. Temperature and pH were monitored in the habitat of M. calcar to place experiments in context with current environmental conditions. There was no effect of temperature or pH on cleavage stage embryos but later development (gastrula‐larvae) was negatively effected by a +2 to 4 ^°C warming and there was a negative effect of ?0.6 pH units on embryos reaching the hatched gastrula stage. Mortality and abnormal development in larvae increased significantly even with +2 ^°C warming and larval growth was impaired at +4 ^°C. For the range of temperature and pH conditions tested, there were no interactive effects of stressors across all stages monitored. For M. calcar, warming not acidification was the dominant stressor. A regression model incorporating data from this study and projected increasing SST for the region suggests an increase in larval mortality to 70% for M. calcar by 2100 in the absence of acclimation and adaptation. The broad distribution of this species in eastern Australia encompassing subtropical to cold temperate thermal regimes provides the possibility that local M. calcar populations may be sustained in a warming world through poleward migration of thermotolerant propagules, facilitated by the strong southward flow of the East Australian Current.     

Mechanical robustness of the calcareous tubeworm Hydroides elegans: warming mitigates the adverse effects of ocean acidification

Development of antifouling strategies requires knowledge of how fouling organisms would respond to climate change associated environmental stressors. Here, a calcareous tube built by the tubeworm, Hydroides elegans, was used as an example to evaluate the individual and interactive effects of ocean acidification (OA), warming and reduced salinity on the mechanical properties of a tube. Tubeworms produce a mechanically weaker tube with less resistance to simulated predator attack under OA (pH 7.8). Warming (29°C) increased tube volume, tube mineral density and the tube’s resistance to a simulated predatory attack. A weakening effect by OA did not make the removal of tubeworms easier except for the earliest stage, in which warming had the least effect. Reduced salinity (27 psu) did not affect tubes. This study showed that both mechanical analysis and computational modeling can be integrated with biofouling research to provide insights into how fouling communities might develop in future ocean conditions.     

Temperature effects on oxygen thresholds for hypoxia in marine benthic organisms

The effect of warming on the oxygen requirements and the survival of benthic organisms under hypoxia was tested using a meta‐analysis of published results of experiments evaluating the effects of temperature on the median lethal time and median lethal concentration of benthic macrofauna under hypoxia. The meta‐analysis confirmed that survival times under hypoxia were reduced by on average 74% and that median lethal concentration increased by on average 16% when marine benthic organisms were exposed to warmer temperatures. Warming reduced survival times of marine benthic macrofauna under hypoxia by a median of 3.95±1.67 h °C^?1 and increased the oxygen thresholds for hypoxia‐driven mortality by a median of 1.02±0.15% saturation °C^?1 or 0.07±0.01 mg O₂ L^?1 °C^?1. The corresponding Q₁₀ values averaged 3.01±0.29 for the median survival time and 2.09±0.20 for the median lethal oxygen concentration. Use of these Q₁₀ values predicts that the 4 °C warming expected during the 21st century will lead to survival times 35.6% lower under hypoxia and that the threshold oxygen concentrations for high mortality to occur will increase by, on average, 25.5% if bottom water temperature increased by 4 °C. Hence, ocean warming is expected to increase the vulnerability of benthic macrofauna to reduced oxygen concentrations and expand the area of coastal ecosystems affected by hypoxia.     

Adjustments in fatty acid composition is a mechanism that can explain resilience to marine heatwaves and future ocean conditions in the habitat‐forming seaweed Phyllospora comosa (Labillardière) C.Agardh

Marine heatwaves are extreme events that can have profound and lasting impacts on marine species. Field observations have shown seaweeds to be highly susceptible to marine heatwaves, but the physiological drivers of this susceptibility are poorly understood. Furthermore, the effects of marine heatwaves in conjunction with ocean warming and acidification are yet to be investigated. To address this knowledge gap, we conducted a laboratory culture experiment in which we tested the growth and physiological responses of Phyllospora comosa juveniles from the southern extent of its range (43–31°S) to marine heatwaves, ocean warming and acidification. We used a ‘collapsed factorial design’ in which marine heatwaves were superimposed on current (today's pH and temperature) and future (pH and temperature projected by 2100) ocean conditions. Responses were tested both during the heatwaves, and after a 7‐day recovery period. Heatwaves reduced net photosynthetic rates in both current and future conditions, while respiration rates were elevated under heatwaves in the current conditions only. Following the recovery period, there was little evidence of heatwaves having lasting negative effects on growth, photosynthesis or respiration. Exposure to heatwaves, future ocean conditions or both caused an increase in the degree of saturation of fatty acids. This adjustment may have counteracted negative effects of elevated temperatures by decreasing membrane fluidity, which increases at higher temperatures. Furthermore, P. comosa appeared to down‐regulate the energetically expensive carbon dioxide concentrating mechanism in the future conditions with a reduction in δ¹³C values detected in these treatments. Any saved energy arising from this down‐regulation was not invested in growth and was likely invested in the adjustment of fatty acid composition. This adjustment is a mechanism by which P. comosa and other seaweeds may tolerate the negative effects of ocean warming and marine heatwaves through benefits arising from ocean acidification.     

Unshelled abalone and corrupted urchins: development of marine calcifiers in a changing ocean

The most fragile skeletons produced by benthic marine calcifiers are those that larvae and juveniles make to support their bodies. Ocean warming, acidification, decreased carbonate saturation and their interactive effects are likely to impair skeletogenesis. Failure to produce skeleton in a changing ocean has negative implications for a diversity of marine species. We examined the interactive effects of warming and acidification on an abalone (Haliotis coccoradiata) and a sea urchin (Heliocidaris erythrogramma) reared from fertilization in temperature and pH/pCO(2) treatments in a climatically and regionally relevant setting. Exposure of ectodermal (abalone) and mesodermal (echinoid) calcifying systems to warming (+2°C to 4°C) and acidification (pH 7.6-7.8) resulted in unshelled larvae and abnormal juveniles. Haliotis development was most sensitive with no interaction between stressors. For Heliocidaris, the percentage of normal juveniles decreased in response to both stressors, although a +2°C warming diminished the negative effect of low pH. The number of spines produced decreased with increasing acidification/pCO(2), and the interactive effect between stressors indicated that a +2°C warming reduced the negative effects of low pH. At +4°C, the developmental thermal tolerance was breached. Our results show that projected near-future climate change will have deleterious effects on development with differences in vulnerability in the two species.     

Projected marine climate change: effects on copepod oxidative status and reproduction

Zooplankton are an important link between primary producers and fish. Therefore, it is crucial to address their responses when predicting effects of climate change on pelagic ecosystems. For realistic community‐level predictions, several biotic and abiotic climate‐related variables should be examined in combination. We studied the combined effects of ocean acidification and global warming predicted for year 2100 with toxic cyanobacteria on the calanoid copepod, Acartia bifilosa. Acidification together with higher temperature reduced copepod antioxidant capacity. Higher temperature also decreased egg viability, nauplii development, and oxidative status. Exposure to cyanobacteria and its toxin had a negative effect on egg production but, a positive effect on oxidative status and egg viability, giving no net effects on viable egg production. Additionally, nauplii development was enhanced by the presence of cyanobacteria, which partially alleviated the otherwise negative effects of increased temperature and decreased pH on the copepod recruitment. The interactive effects of temperature, acidification, and cyanobacteria on copepods highlight the importance of testing combined effects of climate‐related factors when predicting biological responses.     

Individual and interactive effects of ocean acidification,global warming,and UV radiation on phytoplankton

Rising carbon dioxide (CO₂) concentrations in the atmosphere result in increasing global temperatures and ocean warming (OW). Concomitantly, dissolution of anthropogenic CO₂ declines seawater pH, resulting in ocean acidification (OA) and altering marine chemical environments. The marine biological carbon pump driven by marine photosynthesis plays an important role for oceanic carbon sinks. Therefore, how ocean climate changes affect the amount of carbon fixation by primary producers is closely related to future ocean carbon uptake. OA may upregulate metabolic pathways in phytoplankton, such as upregulating ß-oxidation and the tricarboxylic acid cycle, resulting in increased accumulation of toxic phenolic compounds. Ocean warming decreases global phytoplankton productivity; however, regionally, it may stimulate primary productivity and change phytoplankton community composition, due to different physical and chemical environmental requirements of species. It is still controversial how OA and OW interactively affect marine carbon fixation by photosynthetic organisms. OA impairs the process of calcification in calcifying phytoplankton and aggravate ultraviolet (UV)-induced harms to the cells. Increasing temperatures enhance the activity of cellular repair mechanisms, which mitigates UV-induced damage. The effects of OA, warming, enhanced exposure to UV-B as well as the interactions of these environmental stress factors on phytoplankton productivity and community composition, are discussed in this review.     

Circumpolar dynamics of a marine top‐predator track ocean warming rates

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long‐lived, wide‐ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black‐legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea‐surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large‐scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom‐up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large‐scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.     

Meta‐analysis reveals complex marine biological responses to the interactive effects of ocean acidification and warming

Ocean acidification and warming are considered two of the greatest threats to marine biodiversity, yet the combined effect of these stressors on marine organisms remains largely unclear. Using a meta‐analytical approach, we assessed the biological responses of marine organisms to the effects of ocean acidification and warming in isolation and combination. As expected biological responses varied across taxonomic groups, life‐history stages, and trophic levels, but importantly, combining stressors generally exhibited a stronger biological (either positive or negative) effect. Using a subset of orthogonal studies, we show that four of five of the biological responses measured (calcification, photosynthesis, reproduction, and survival, but not growth) interacted synergistically when warming and acidification were combined. The observed synergisms between interacting stressors suggest that care must be made in making inferences from single‐stressor studies. Our findings clearly have implications for the development of adaptive management strategies particularly given that the frequency of stressors interacting in marine systems will be likely to intensify in the future. There is now an urgent need to move toward more robust, holistic, and ecologically realistic climate change experiments that incorporate interactions. Without them accurate predictions about the likely deleterious impacts to marine biodiversity and ecosystem functioning over the next century will not be possible.     

Antagonistic effects of ocean acidification and warming on hunting sharks

Ocean warming and acidification alter the physiological performance and behaviour of many small‐bodied fishes, yet the potential interactive effects of these stressors on larger predators remains poorly understood. In particular, the combined effects of elevated temperature on metabolism and of elevated CO₂ on the behaviour of large predators may not only affect their foraging behaviour, but also the communities in which their prey live. We used a factorial design to assess how projected warming and acidification create synergies or antagonisms between physiological and behavioural processes, such as swimming activity and feeding behaviour through odour tracking and vision. Temperature increased swimming activity during feeding, independent of CO₂. Although temperature also increased motivational drive to locate and accept prey, elevated CO₂ negated chemical and visual behavioural responses that enable effective hunting. Fundamental to these effects was the negligible effect of high CO₂ in isolation, but its power to negate the positive effects of temperature when brought in conjunction. The reduced potential to locate prey due to the interactive effects of ocean acidification and warming, in combination with increases in energetic demand, suggests that energetic tradeoffs will be needed for sharks to sustain themselves at an individual and population level in a future ocean.     

Effects of ocean acidification on Antarctic marine organisms: A meta‐analysis

Southern Ocean waters are among the most vulnerable to ocean acidification. The projected increase in the CO₂ level will cause changes in carbonate chemistry that are likely to be damaging to organisms inhabiting these waters. A meta‐analysis was undertaken to examine the vulnerability of Antarctic marine biota occupying waters south of 60°S to ocean acidification. This meta‐analysis showed that ocean acidification negatively affects autotrophic organisms, mainly phytoplankton, at CO₂ levels above 1,000 μatm and invertebrates above 1,500 μatm, but positively affects bacterial abundance. The sensitivity of phytoplankton to ocean acidification was influenced by the experimental procedure used. Natural, mixed communities were more sensitive than single species in culture and showed a decline in chlorophyll a concentration, productivity, and photosynthetic health, as well as a shift in community composition at CO₂ levels above 1,000 μatm. Invertebrates showed reduced fertilization rates and increased occurrence of larval abnormalities, as well as decreased calcification rates and increased shell dissolution with any increase in CO₂ level above 1,500 μatm. Assessment of the vulnerability of fish and macroalgae to ocean acidification was limited by the number of studies available. Overall, this analysis indicates that many marine organisms in the Southern Ocean are likely to be susceptible to ocean acidification and thereby likely to change their contribution to ecosystem services in the future. Further studies are required to address the poor spatial coverage, lack of community or ecosystem‐level studies, and the largely unknown potential for organisms to acclimate and/or adapt to the changing conditions.     

The effects of elevated temperature and dissolved ρCO2 on a marine foundation species

Understanding how climate change and other environmental stressors will affect species is a fundamental concern of modern ecology. Indeed, numerous studies have documented how climate stressors affect species distributions and population persistence. However, relatively few studies have investigated how multiple climate stressors might affect species. In this study, we investigate the impacts of how two climate change factors affect an important foundation species. Specifically, we tested how ocean acidification from dissolution of CO₂ and increased sea surface temperatures affect multiple characteristics of juvenile eastern oysters (Crassostrea virginica). We found strong impacts of each stressor, but no interaction between the two. Simulated warming to mimic heat stressed summers reduced oyster growth, survival, and filtration rates. Additionally, we found that CO₂‐induced acidification reduced strength of oyster shells, which could potentially facilitate crab predation. As past studies have detected few impacts of these stressors on adult oysters, these results indicate that early life stages of calcareous marine organisms may be more susceptible to effects of ocean acidification and global warming. Overall, these data show that predicted changes in temperature and CO₂ can differentially influence direct effects on individual species, which could have important implications for the nature of their trophic interactions.