Vulnerability of the calcifying larval stage of the Antarctic sea urchin Sterechinus neumayeri to near‐future ocean acidification and warming

Stenothermal polar benthic marine invertebrates are highly sensitive to environmental perturbations but little is known about potential synergistic effects of concurrent ocean warming and acidification on development of their embryos and larvae. We examined the effects of these stressors on development to the calcifying larval stage in the Antarctic sea urchin Sterechinus neumayeri in embryos reared in present and future (2100+) ocean conditions from fertilization. Embryos were reared in 2 temperature (ambient: ?1.0 °C, + 2 °C : 1.0 °C) and 3 pH (ambient: pH 8.0, ?0.2–0.4 pH units: 7.8,7.6) levels. Principle coordinates analysis on five larval metrics showed a significant effect of temperature and pH on the pattern of growth. Within each temperature, larvae were separated by pH treatment, a pattern primarily influenced by larval arm and body length. Growth was accelerated by temperature with a 20–28% increase in postoral (PO) length at +2 °C across all pH levels. Growth was strongly depressed by reduced pH with a 8–19% decrease in PO length at pH 7.6–7.8 at both temperatures. The boost in growth caused by warming resulted in larvae that were larger than would be observed if acidification was examined in the absence of warming. However, there was no significant interaction between these stressors. The increase in left‐right asymmetry and altered body allometry indicated that decreased pH disrupted developmental patterning and acted as a teratogen (agent causing developmental malformation). Decreased developmental success with just a 2 °C warming indicates that development in S. neumayeri is particularly sensitive to increased temperature. Increased temperature also altered larval allometry. Altered body shape impairs swimming and feeding in echinoplutei. In the absence of adaptation, it appears that the larval phase may be a bottleneck for survivorship of S. neumayeri in a changing ocean in a location where poleward migration to escape inhospitable conditions is not possible.     

Signals of resilience to ocean change: high thermal tolerance of early stage Antarctic sea urchins (Sterechinus neumayeri) reared under present-day and future pCO2 and temperature

We tested the hypothesis that development of the Antarctic urchin Sterechinus neumayeri under future ocean conditions of warming and acidification would incur physiological costs, reducing the tolerance of a secondary stressor. The aim of this study is twofold: (1) quantify current austral spring temperature and pH near sea urchin habitat at Cape Evans in McMurdo Sound, Antarctica and (2) spawn S. neumayeri in the laboratory and raise early developmental stages (EDSs) under ambient (?0.7 °C; 400 µatm pCO₂) and future (+2.6 °C; 650 and 1,000 µatm pCO₂) ocean conditions and expose four EDSs (blastula, gastrula, prism, and 4-arm echinopluteus) to a one hour acute heat stress and assess survivorship. Results of field data from 2011 to 2012 show extremely stable inter-annual pH conditions ranging from 7.99 to 8.08, suggesting that future ocean acidification will drastically alter the pH-seascape for S. neumayeri. In the laboratory, S. neumayeri EDSs appear to be tolerant of temperatures and pCO₂ levels above their current habitat conditions. EDSs survived acute heat exposures >20 °C above habitat temperatures of ?1.9 °C. No pCO₂ effect was observed for EDSs reared at ?0.7 °C. When reared at +2.6 °C, small but significant pCO₂ effects were observed at the blastula and prism stage, suggesting that multiple stressors are more detrimental than single stressors. While surprisingly tolerant overall, blastulae were the most sensitive stage to ocean warming and acidification. We conclude that S. neumayeri may be unexpectedly physiologically tolerant of future ocean conditions.     

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.     

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.     

Adaptive Capacity of the Habitat Modifying Sea Urchin Centrostephanus rodgersii to Ocean Warming and Ocean Acidification: Performance of Early Embryos

Background

Predicting effects of rapid climate change on populations depends on measuring the effects of climate stressors on performance, and potential for adaptation. Adaptation to stressful climatic conditions requires heritable genetic variance for stress tolerance present in populations.

Methodology/Principal Findings

We quantified genetic variation in tolerance of early development of the ecologically important sea urchin Centrostephanus rodgersii to near-future (2100) ocean conditions projected for the southeast Australian global change hot spot. Multiple dam-sire crosses were used to quantify the interactive effects of warming (+2–4°C) and acidification (−0.3−0.5 pH units) across twenty-seven family lines. Acidification, but not temperature, decreased the percentage of cleavage stage embryos. In contrast, temperature, but not acidification decreased the percentage of gastrulation. Cleavage success in response to both stressors was strongly affected by sire identity. Sire and dam identity significantly affected gastrulation and both interacted with temperature to determine developmental success. Positive genetic correlations for gastrulation indicated that genotypes that did well at lower pH also did well in higher temperatures.

Conclusions/Significance

Significant genotype (sire) by environment interactions for both stressors at gastrulation indicated the presence of heritable variation in thermal tolerance and the ability of embryos to respond to changing environments. The significant influence of dam may be due to maternal provisioning (maternal genotype or environment) and/or offspring genotype. It appears that early development in this ecologically important sea urchin is not constrained in adapting to the multiple stressors of ocean warming and acidification. The presence of tolerant genotypes indicates the potential to adapt to concurrent warming and acidification, contributing to the resilience of C. rodgersii in a changing ocean.     

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.     

Ocean acidification has little effect on developmental thermal windows of echinoderms from Antarctica to the tropics

As the ocean warms, thermal tolerance of developmental stages may be a key driver of changes in the geographical distributions and abundance of marine invertebrates. Additional stressors such as ocean acidification may influence developmental thermal windows and are therefore important considerations for predicting distributions of species under climate change scenarios. The effects of reduced seawater pH on the thermal windows of fertilization, embryology and larval morphology were examined using five echinoderm species: two polar (Sterechinus neumayeri and Odontaster validus), two temperate (Fellaster zelandiae and Patiriella regularis) and one tropical (Arachnoides placenta). Responses were examined across 12–13 temperatures ranging from ?1.1 °C to 5.7 °C (S. neumayeri), ?0.5 °C to 10.7 °C (O. validus), 5.8 °C to 27 °C (F. zelandiae), 6.0 °C to 27.1 °C (P. regularis) and 13.9 °C to 34.8 °C (A. placenta) under present‐day and near‐future (2100+) ocean acidification conditions (‐0.3 pH units) and for three important early developmental stages 1) fertilization, 2) embryo (prehatching) and 3) larval development. Thermal windows for fertilization were broad and were not influenced by a pH decrease. Embryological development was less thermotolerant. For O. validus, P. regularis and A. placenta, low pH reduced normal development, albeit with no effect on thermal windows. Larval development in all five species was affected by both temperature and pH; however, thermal tolerance was not reduced by pH. Results of this study suggest that in terms of fertilization and development, temperature will remain as the most important factor influencing species' latitudinal distributions as the ocean continues to warm and decrease in pH, and that there is little evidence of a synergistic effect of temperature and ocean acidification on the thermal control of species ranges.     

Effects of ocean warming and acidification on survival,growth and skeletal development in the early benthic juvenile sea urchin (Heliocidaris erythrogramma)

Co‐occurring ocean warming, acidification and reduced carbonate mineral saturation have significant impacts on marine biota, especially calcifying organisms. The effects of these stressors on development and calcification in newly metamorphosed juveniles (ca. 0.5 mm test diameter) of the intertidal sea urchin Heliocidaris erythrogramma, an ecologically important species in temperate Australia, were investigated in context with present and projected future conditions. Habitat temperature and pH/pCO₂ were documented to place experiments in a biologically and ecologically relevant context. These parameters fluctuated diurnally up to 10 °C and 0.45 pH units. The juveniles were exposed to three temperature (21, 23 and 25 °C) and four pH (8.1, 7.8, 7.6 and 7.4) treatments in all combinations, representing ambient sea surface conditions (21 °C, pH 8.1; pCO₂ 397; Ω_Ca 4.7; Ω_Ar 3.1), near‐future projected change (+2–4 °C, ?0.3–0.5 pH units; pCO₂ 400–1820; Ω_Ca 5.0–1.6; Ω_Ar 3.3–1.1), and extreme conditions experienced at low tide (+4 °C, ?0.3–0.7 pH units; pCO₂ 2850–2967; Ω_Ca 1.1–1.0; Ω_Ar 0.7–0.6). The lowest pH treatment (pH 7.4) was used to assess tolerance levels. Juvenile survival and test growth were resilient to current and near‐future warming and acidification. Spine development, however, was negatively affected by near‐future increased temperature (+2–4 °C) and extreme acidification (pH 7.4), with a complex interaction between stressors. Near‐future warming was the more significant stressor. Spine tips were dissolved in the pH 7.4 treatments. Adaptation to fluctuating temperature‐pH conditions in the intertidal may convey resilience to juvenile H. erythrogramma to changing ocean conditions, however, ocean warming and acidification may shift baseline intertidal temperature and pH/pCO₂ to levels that exceed tolerance limits.     

Near-future ocean acidification enhances the feeding rate and development of the herbivorous juveniles of the crown-of-thorns starfish, <Emphasis Type="Italic">Acanthaster planci</Emphasis>

Population outbreaks of the corallivorous crown-of-thorns starfish, Acanthaster planci, are a major contributor to the decline in coral reef across the Indo-Pacific. The success of A. planci and other reef species in a changing ocean will be influenced by juvenile performance because the naturally high mortality experienced at this sensitive life history stage maybe exacerbated by ocean warming and acidification. We investigated the effects of increased temperature and acidification on growth of newly metamorphosed juvenile A. planci and their feeding rates on crustose coralline algae (CCA) during the initial herbivorous phase of their life history. The juveniles were exposed to three temperature (26, 28, 30 °C) and three pH (NIST scale: 8.1, 7.8, 7.6) levels in a flow-through cross-factorial experiment. There were positive but independent effects of warming and acidification on juvenile growth and feeding. Early juveniles were highly tolerant to moderate increases in temperature (+2 °C above ambient) with the highest growth at 30 °C. Growth and feeding rates of A. planci on CCA were highest at pH 7.6. Thus, ocean warming and acidification may enhance the success of A. planci juveniles. In contrast to its coral prey, at this vulnerable developmental stage, A. planci appears to be highly resilient to future ocean change. Success of juveniles in a future ocean may have carry-over effects into the coral-eating life stage, increasing the threat to coral reef systems.     

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.     

Impact of Ocean Warming and Ocean Acidification on Larval Development and Calcification in the Sea Urchin Tripneustes gratilla

Background

As the oceans simultaneously warm, acidify and increase in P _CO2, prospects for marine biota are of concern. Calcifying species may find it difficult to produce their skeleton because ocean acidification decreases calcium carbonate saturation and accompanying hypercapnia suppresses metabolism. However, this may be buffered by enhanced growth and metabolism due to warming.

Methodology/Principal Findings

We examined the interactive effects of near-future ocean warming and increased acidification/P _CO2 on larval development in the tropical sea urchin Tripneustes gratilla. Larvae were reared in multifactorial experiments in flow-through conditions in all combinations of three temperature and three pH/P _CO2 treatments. Experiments were placed in the setting of projected near future conditions for SE Australia, a global change hot spot. Increased acidity/P _CO2 and decreased carbonate mineral saturation significantly reduced larval growth resulting in decreased skeletal length. Increased temperature (+3°C) stimulated growth, producing significantly bigger larvae across all pH/P _CO2 treatments up to a thermal threshold (+6°C). Increased acidity (-0.3-0.5 pH units) and hypercapnia significantly reduced larval calcification. A +3°C warming diminished the negative effects of acidification and hypercapnia on larval growth.

Conclusions and Significance

This study of the effects of ocean warming and CO₂ driven acidification on development and calcification of marine invertebrate larvae reared in experimental conditions from the outset of development (fertilization) shows the positive and negative effects of these stressors. In simultaneous exposure to stressors the dwarfing effects of acidification were dominant. Reduction in size of sea urchin larvae in a high P _CO2 ocean would likely impair their performance with negative consequent effects for benthic adult populations.     

Larvae of the coral eating crown‐of‐thorns starfish,Acanthaster planci in a warmer‐high CO2 ocean

Outbreaks of crown‐of‐thorns starfish (COTS), Acanthaster planci, contribute to major declines of coral reef ecosystems throughout the Indo‐Pacific. As the oceans warm and decrease in pH due to increased anthropogenic CO₂ production_, coral reefs are also susceptible to bleaching, disease and reduced calcification. The impacts of ocean acidification and warming may be exacerbated by COTS predation, but it is not known how this major predator will fare in a changing ocean. Because larval success is a key driver of population outbreaks, we investigated the sensitivities of larval A. planci to increased temperature (2–4 °C above ambient) and acidification (0.3–0.5 pH units below ambient) in flow‐through cross‐factorial experiments (3 temperature × 3 pH/pCO₂ levels). There was no effect of increased temperature or acidification on fertilization or very early development. Larvae reared in the optimal temperature (28 °C) were the largest across all pH treatments. Development to advanced larva was negatively affected by the high temperature treatment (30 °C) and by both experimental pH levels (pH 7.6, 7.8). Thus, planktonic life stages of A. planci may be negatively impacted by near‐future global change. Increased temperature and reduced pH had an additive negative effect on reducing larval size. The 30 °C treatment exceeded larval tolerance regardless of pH. As 30 °C sea surface temperatures may become the norm in low latitude tropical regions, poleward migration of A. planci may be expected as they follow optimal isotherms. In the absence of acclimation or adaptation, declines in low latitude populations may occur. Poleward migration will be facilitated by strong western boundary currents, with possible negative flow‐on effects on high latitude coral reefs. The contrasting responses of the larvae of A. planci and those of its coral prey to ocean acidification and warming are considered in context with potential future change in tropical reef ecosystems.     

Developmental and physiological challenges of octopus (Octopus vulgaris) early life stages under ocean warming

The ability to understand and predict the effects of ocean warming (under realistic scenarios) on marine biota is of paramount importance, especially at the most vulnerable early life stages. Here we investigated the impact of predicted environmental warming (+3 °C) on the development, metabolism, heat shock response and antioxidant defense mechanisms of the early stages of the common octopus, Octopus vulgaris. As expected, warming shortened embryonic developmental time by 13 days, from 38 days at 18 °C to 25 days at 21 °C. Concomitantly, survival decreased significantly (~29.9 %). Size at hatching varied inversely with temperature, and the percentage of smaller premature paralarvae increased drastically, from 0 % at 18 °C to 17.8 % at 21 °C. The metabolic costs of the transition from an encapsulated embryo to a free planktonic form increased significantly with warming, and HSP70 concentrations and glutathione S-transferase activity levels were significantly magnified from late embryonic to paralarval stages. Yet, despite the presence of effective antioxidant defense mechanisms, ocean warming led to an augmentation of malondialdehyde levels (an indicative of enhanced ROS action), a process considered to be one of the most frequent cellular injury mechanisms. Thus, the present study provides clues about how the magnitude and rate of ocean warming will challenge the buffering capacities of octopus embryos and hatchlings’ physiology. The prediction and understanding of the biochemical and physiological responses to warmer temperatures (under realistic scenarios) is crucial for the management of highly commercial and ecologically important species, such as O. vulgaris.     

At the tipping point: Differential influences of warming and deoxygenation on the survival,emergence, and respiration of cosmopolitan clams

Although warming and low dissolved oxygen (DO) levels are co‐occurring significant climatic stressors in the ocean, the combined effects of these stressors on marine benthic animals have not been well established. Here, we tested the effects of elevated temperatures and low dissolved oxygen levels on the survival, emerging behavior from sediment, and the respiration of juvenile cosmopolitan Manila clams (Venerupis philippinarum) by exposing them to two temperatures (20 and 23.5°C) and DO levels (3.5 and 6–7 mg/L). Although within previously described tolerable ranges of temperature and DO, this 3.5°C increase in temperature combined with a 50% decrease in DO had a devastating effect on the survival of clams (85% mortality after 8 days). The mortality of clams under normoxia at 23.5°C appeared to be higher than under the low DO condition at 20°C. On the other hand, more clams emerged from sediment under the low DO condition at 20°C than under any other conditions. Oxygen consumption rates were not significantly affected by different conditions. Our results suggest temperature elevation combined with low oxygen additively increases stress on Manila clams and that warming is at least as stressful as low DO in terms of mortality. However, low DO poses another threat as it may induce emergence from sediment, and, thus increase predation risk. This is the first evidence that a combination of warming and deoxygenation stressors should reduce population survival of clams much more so than changes in a single stressor.     

Complex Responses of Intertidal Molluscan Embryos to a Warming and Acidifying Ocean in the Presence of UV Radiation

Climate change and ocean acidification will expose marine organisms to synchronous multiple stressors, with early life stages being potentially most vulnerable to changing environmental conditions. We simultaneously exposed encapsulated molluscan embryos to three abiotic stressors—acidified conditions, elevated temperate, and solar UV radiation in large outdoor water tables in a multifactorial design. Solar UV radiation was modified with plastic filters, while levels of the other factors reflected IPCC predictions for near-future change. We quantified mortality and the rate of embryonic development for a mid-shore littorinid, Bembicium nanum, and low-shore opisthobranch, Dolabrifera brazieri. Outcomes were consistent for these model species with embryos faring significantly better at 26°C than 22°C. Mortality sharply increased at the lowest temperature (22°C) and lowest pH (7.6) examined, producing a significant interaction. Under these conditions mortality approached 100% for each species, representing a 2- to 4-fold increase in mortality relative to warm (26°C) non-acidified conditions. Predictably, development was more rapid at the highest temperature but this again interacted with acidified conditions. Development was slowed under acidified conditions at the lowest temperature. The presence of UV radiation had minimal impact on the outcomes, only slowing development for the littorinid and not interacting with the other factors. Our findings suggest that a warming ocean, at least to a threshold, may compensate for the effects of decreasing pH for some species. It also appears that stressors will interact in complex and unpredictable ways in a changing climate.     

The effect of ocean acidification and temperature on the fertilization and embryonic development of the Sydney rock oyster Saccostrea glomerata (Gould 1850)

This study investigated the synergistic effects of ocean acidification (caused by elevations in the partial pressure of carbon dioxide pCO₂) and temperature on the fertilization and embryonic development of the economically and ecologically important Sydney rock oyster, Saccostrea glomerata (Gould 1850). As pCO₂ increased, fertilization significantly decreased. The temperature of 26 °C was the optimum temperature for fertilization, as temperature increased and decreased from this optimum, fertilization decreased. There was also an effect of pCO₂ and temperature on embryonic development. Generally as pCO₂ increased, the percentage and size of D‐veligers decreased and the percentage of D‐veligers that were abnormal increased. The optimum temperature was 26 °C and embryonic development decreased at temperatures that were above and below this temperature. Abnormality of D‐veligers was greatest at 1000 ppm and 18 and 30 °C (≥90%) and least at 375 ppm and 26 °C (≤4%). Finally prolonged exposure of elevated pCO₂ and temperature across early developmental stages led to fewer D‐veligers, more abnormality and smaller sizes in elevated CO₂ environments and may lead to lethal effects at suboptimal temperatures. Embryos that were exposed to the pCO₂ and temperature treatments for fertilization and embryonic development had fewer D‐veligers, greater percentage of abnormality and reduced size than embryos that were exposed to the treatments for embryonic development only. Further at the elevated temperature of 30 °C and 750–1000 ppm, there was no embryonic development. The results of this study suggest that predicted changes in ocean acidification and temperature over the next century may have severe implications for the distribution and abundance of S. glomerata as well as possible implications for the reproduction and development of other marine invertebrates.     

Interactive effects of near-future temperature increase and ocean acidification on physiology and gonad development in adult Pacific sea urchin,Echinometra sp. A

Increased atmospheric CO₂ will have a twofold impact on future marine ecosystems, increasing global sea surface temperatures and uptake of CO₂ (Ocean Acidification). Many experiments focus on the investigation of one of these stressors, but under realistic future climate predictions, these stressors may have interactive effects on individuals. Here, we investigate the effect of warming and acidification in combination. We test for interactive effects of potential near-future (2100) temperature (+2 to 3 °C) and pCO₂ (~860–940 μAtm) levels on the physiology of the tropical echinoid Echinometra sp. A. The greatest reduction in growth was under simultaneous temperature and pH/pCO₂ stress (marginally significant temperature × pH/pCO₂ interaction). This was mirrored by the physiological data, with highest metabolic activity (measured as respiration and ammonium excretion) occurring at the increased temperature and pCO₂ treatment, although this was not significant for excretion. The perivisceral coelomic fluid pH was ~7.5–7.6, as typical for echinoids, and showed no significant changes between treatments. Indicative of active calcification, internal magnesium and calcium concentrations were reduced compared to the external medium, but were not different between treatments. Gonad weight was lower at the higher temperature, and this difference was more distinct and statistically significant for males. The condition of the gonads assessed by histology declined in increased temperature and low pH treatments. The Echinometra grew in all treatments indicating active calcification of their magnesium calcite tests even as carbonate mineral saturation decreased. Our results indicate that the interactive temperature and pH effects are more important for adult echinoids than individual stressors. Although adult specimens grow and survive in near-future conditions, higher energy demands may influence gonad development and thus population maintenance.     

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.     

Ocean Warming,More than Acidification,Reduces Shell Strength in a Commercial Shellfish Species during Food Limitation

Ocean surface pH levels are predicted to fall by 0.3–0.4 pH units by the end of the century and are likely to coincide with an increase in sea surface temperature of 2–4°C. The combined effect of ocean acidification and warming on the functional properties of bivalve shells is largely unknown and of growing concern as the shell provides protection from mechanical and environmental challenges. We examined the effects of near-future pH (ambient pH –0.4 pH units) and warming (ambient temperature +4°C) on the shells of the commercially important bivalve, Mytilus edulis when fed for a limited period (4–6 h day⁻¹). After six months exposure, warming, but not acidification, significantly reduced shell strength determined as reductions in the maximum load endured by the shells. However, acidification resulted in a reduction in shell flex before failure. Reductions in shell strength with warming could not be explained by alterations in morphology, or shell composition but were accompanied by reductions in shell surface area, and by a fall in whole-body condition index. It appears that warming has an indirect effect on shell strength by re-allocating energy from shell formation to support temperature-related increases in maintenance costs, especially as food supply was limited and the mussels were probably relying on internal energy reserves. The maintenance of shell strength despite seawater acidification suggests that biomineralisation processes are unaffected by the associated changes in CaCO₃ saturation levels. We conclude that under near-future climate change conditions, ocean warming will pose a greater risk to shell integrity in M. edulis than ocean acidification when food availability is limited.     

The reef-building coral Siderastrea siderea exhibits parabolic responses to ocean acidification and warming

Anthropogenic increases in atmospheric CO₂ over this century are predicted to cause global average surface ocean pH to decline by 0.1–0.3 pH units and sea surface temperature to increase by 1–4°C. We conducted controlled laboratory experiments to investigate the impacts of CO₂-induced ocean acidification (pCO₂ = 324, 477, 604, 2553 µatm) and warming (25, 28, 32°C) on the calcification rate of the zooxanthellate scleractinian coral Siderastrea siderea, a widespread, abundant and keystone reef-builder in the Caribbean Sea. We show that both acidification and warming cause a parabolic response in the calcification rate within this coral species. Moderate increases in pCO₂ and warming, relative to near-present-day values, enhanced coral calcification, with calcification rates declining under the highest pCO₂ and thermal conditions. Equivalent responses to acidification and warming were exhibited by colonies across reef zones and the parabolic nature of the corals'' response to these stressors was evident across all three of the experiment''s 30-day observational intervals. Furthermore, the warming projected by the Intergovernmental Panel on Climate Change for the end of the twenty-first century caused a fivefold decrease in the rate of coral calcification, while the acidification projected for the same interval had no statistically significant impact on the calcification rate—suggesting that ocean warming poses a more immediate threat than acidification for this important coral species.