Odor tracking in sharks is reduced under future ocean acidification conditions

Recent studies show that ocean acidification impairs sensory functions and alters the behavior of teleost fishes. If sharks and other elasmobranchs are similarly affected, this could have significant consequences for marine ecosystems globally. Here, we show that projected future CO₂ levels impair odor tracking behavior of the smooth dogfish (Mustelus canis). Adult M. canis were held for 5 days in a current‐day control (405 ± 26 μatm) and mid (741 ± 22 μatm) or high CO₂ (1064 ± 17 μatm) treatments consistent with the projections for the year 2100 on a ‘business as usual’ scenario. Both control and mid CO₂‐treated individuals maintained normal odor tracking behavior, whereas high CO₂‐treated sharks significantly avoided the odor cues indicative of food. Control sharks spent >60% of their time in the water stream containing the food stimulus, but this value fell below 15% in high CO₂‐treated sharks. In addition, sharks treated under mid and high CO₂ conditions reduced attack behavior compared to the control individuals. Our findings show that shark feeding could be affected by changes in seawater chemistry projected for the end of this century. Understanding the effects of ocean acidification on critical behaviors, such as prey tracking in large predators, can help determine the potential impacts of future ocean acidification on ecosystem function.     

Absence of cellular damage in tropical newly hatched sharks (<Emphasis Type="Italic">Chiloscyllium plagiosum</Emphasis>) under ocean acidification conditions

Sharks have maintained a key role in marine food webs for 400 million years and across varying physicochemical contexts, suggesting plasticity to environmental change. In this study, we investigated the biochemical effects of ocean acidification (OA) levels predicted for 2100 (pCO₂ ~?900 μatm) on newly hatched tropical whitespotted bamboo sharks (Chiloscyllium plagiosum). Specifically, we measured lipid, protein, and DNA damage levels, as well as changes in the activity of antioxidant enzymes and non-enzymatic ROS scavengers in juvenile sharks exposed to elevated CO₂ for 50 days following hatching. Moreover, we also assessed the secondary oxidative stress response, i.e., heat shock response and ubiquitin levels. Newly hatched sharks appear to cope with OA-related stress through a range of tissue-specific biochemical strategies, specifically through the action of antioxidant enzymatic compounds. Our findings suggest that ROS-scavenging molecules, rather than complex enzymatic proteins, provide an effective defense mechanism in dealing with OA-elicited ROS formation. We argue that sharks’ ancient antioxidant system, strongly based on non-enzymatic antioxidants (e.g., urea), may provide them with resilience towards OA, potentially beyond the tolerance of more recently evolved species, i.e., teleosts. Nevertheless, previous research has provided evidence of detrimental effects of OA (interacting with other climate-related stressors) on some aspects of shark biology. Moreover, given that long-term acclimation and adaptive potential to rapid environmental changes are yet experimentally unaccounted for, future research is warranted to accurately predict shark physiological performance under future ocean conditions.     

Elevated carbon dioxide alters the plasma composition and behaviour of a shark

Increased carbon emissions from fossil fuels are increasing the pCO₂ of the ocean surface waters in a process called ocean acidification. Elevated water pCO₂ can induce physiological and behavioural effects in teleost fishes, although there appear to be large differences in sensitivity between species. There is currently no information available on the possible responses to future ocean acidification in elasmobranch fishes. We exposed small-spotted catsharks (Scyliorhinus canicula) to either control conditions or a year 2100 scenario of 990 μatm pCO₂ for four weeks. We did not detect treatment effects on growth, resting metabolic rate, aerobic scope, skin denticle ultrastructure or skin denticle morphology. However, we found that the elevated pCO₂ group buffered internal acidosis via accumulation with an associated increase in Na⁺, indicating that the blood chemistry remained altered despite the long acclimation period. The elevated pCO₂ group also exhibited a shift in their nocturnal swimming pattern from a pattern of many starts and stops to more continuous swimming. Although CO₂-exposed teleost fishes can display reduced behavioural asymmetry (lateralization), the CO₂-exposed sharks showed increased lateralization. These behavioural effects may suggest that elasmobranch neurophysiology is affected by CO₂, as in some teleosts, or that the sharks detect CO₂ as a constant stressor, which leads to altered behaviour. The potential direct effects of ocean acidification should henceforth be considered when assessing future anthropogenic effects on sharks.     

CO2-induced fertilization impairment in Strongylocentrotus droebachiensis collected in the Arctic

Fertilization depends on distribution and aggregation patterns of sea urchins which influence gamete contact time and may potentially enhance their vulnerability to ocean acidification. In this study, we conducted fertilization experiments to assess the effects of selected pH scenarios on fertilization success of Strongylocentrotus droebachiensis, from Spitsbergen, Arctic. Acidification was achieved by aerating seawater with different CO₂ partial pressures to represent pre-industrial and present conditions (measured ~180–425 µatm) and future acidification scenarios (~550–800, ~1,300, ~2,000 µatm). Fertilization success was defined as the proportion of successful/unsuccessful fertilizations per treatment; eggs were classified according to features of their fertilization envelope (FE), hyaline layer (HL) and achievement of cellular division. The diagnostic findings of specific pathological aberrations were described in detail. We additionally measured intracellular pH changes in unfertilized eggs exposed for 1 h to selected acidification treatments using BCECF/AM. We conclude that (a) acidified conditions increase the proportion of eggs that failed fertilization, (b) acidification may increase the risk of polyspermy due to failures in the FE formation supported by the occasional observation of multiple sperms in the perivitelline space and (c) irregular formation of the embryo may arise due to impaired formation of the HL. The decrease in fertilization success could be also related to the observed changes in intracellular pH at pCO₂ ~ 1,000 μatm or higher.     

Ocean acidification and warming scenarios increase microbioerosion of coral skeletons

Biological mediation of carbonate dissolution represents a fundamental component of the destructive forces acting on coral reef ecosystems. Whereas ocean acidification can increase dissolution of carbonate substrates, the combined impact of ocean acidification and warming on the microbioerosion of coral skeletons remains unknown. Here, we exposed skeletons of the reef‐building corals, Porites cylindrica and Isopora cuneata, to present‐day (Control: 400 μatm – 24 °C) and future pCO₂–temperature scenarios projected for the end of the century (Medium: +230 μatm – +2 °C; High: +610 μatm – +4 °C). Skeletons were also subjected to permanent darkness with initial sodium hypochlorite incubation, and natural light without sodium hypochlorite incubation to isolate the environmental effect of acidic seawater (i.e., Ω_aragonite <1) from the biological effect of photosynthetic microborers. Our results indicated that skeletal dissolution is predominantly driven by photosynthetic microborers, as samples held in the dark did not decalcify. In contrast, dissolution of skeletons exposed to light increased under elevated pCO₂–temperature scenarios, with P. cylindrica experiencing higher dissolution rates per month (89%) than I. cuneata (46%) in the high treatment relative to control. The effects of future pCO₂–temperature scenarios on the structure of endolithic communities were only identified in P. cylindrica and were mostly associated with a higher abundance of the green algae Ostreobium spp. Enhanced skeletal dissolution was also associated with increased endolithic biomass and respiration under elevated pCO₂–temperature scenarios. Our results suggest that future projections of ocean acidification and warming will lead to increased rates of microbioerosion. However, the magnitude of bioerosion responses may depend on the structural properties of coral skeletons, with a range of implications for reef carbonate losses under warmer and more acidic oceans.     

Response to ocean acidification in larvae of a large tropical marine fish,Rachycentron canadum

Currently, ocean acidification is occurring at a faster rate than at any time in the last 300 million years, posing an ecological challenge to marine organisms globally. There is a critical need to understand the effects of acidification on the vulnerable larval stages of marine fishes, as there is potential for large ecological and economic impacts on fish populations and the human economies that rely on them. We expand upon the narrow taxonomic scope found in the literature today, which overlooks many life history characteristics of harvested species, by reporting on the larvae of Rachycentron canadum (cobia), a large, highly mobile, pelagic‐spawning, widely distributed species with a life history and fishery value contrasting other species studied to date. We raised larval cobia through the first 3 weeks of ontogeny under conditions of predicted future ocean acidification to determine effects on somatic growth, development, otolith formation, swimming ability, and swimming activity. Cobia exhibited resistance to treatment effects on growth, development, swimming ability, and swimming activity at 800 and 2100 μatm pCO₂. However, these scenarios resulted in a significant increase in otolith size (up to 25% larger area) at the lowest pCO₂ levels reported to date, as well as the first report of significantly wider daily otolith growth increments. When raised under more extreme scenarios of 3500 and 5400 μatm pCO₂, cobia exhibited significantly reduced size‐at‐age (up to 25% smaller) and a 2–3 days developmental delay. The robust nature of cobia may be due to the naturally variable environmental conditions this species currently encounters throughout ontogeny in coastal environments, which may lead to an increased acclimatization ability even during long‐term exposure to stressors.     

Effects of ocean acidification on the growth and biochemical composition of a green alga (Ulva fasciata) and its associated microbiota

In marine ecosystems, fluctuations in surface-seawater carbon dioxide (CO₂), significantly influence the whole metabolism of marine algae, especially during the early stages of macroalgal development. In this study, the response of the green alga Ulva fasciata for elevating ocean acidification was investigated using four levels of pCO₂ ~ 280, 550, 750 and 1050 µatm. Maximum growth rate (6.6% day⁻¹), protein (32.43 %DW) and pigment (2.9 mg/g) accumulation were observed at pCO₂-550 with an increase of ~2-fold compared to control. On the other hand, lipid and carbohydrate contents recorded their maximum production (4.23 and 46.96 %DW, respectively) at pCO₂-750 while control showed 3.70 and 42.37 %DW, respectively. SDS-PAGE showed the presence of unique bands in response to pCO_2, especially at 550 µatm. Dominant associated bacteria was shifted from Halomonas hydrothermalis of control to Vibrio toranzoniae at pCO₂-1050. These findings suggest that ocean acidification at 550 µatm might impose noticeable effects on growth, protein, pigments, and protein profile of U. fasciata, which could be a good source for fish farming. While, pCO₂-750 was recommended for energetic purpose, due to its high lipid and carbohydrate contents.     

Direct and indirect effects of high pCO2 on algal grazing by coral reef herbivores from the Gulf of Aqaba (Red Sea)

Grazing on marine macroalgae is a key structuring process for coral reef communities. However, ocean acidification from rising atmospheric CO₂ concentrations is predicted to adversely affect many marine animals, while seaweed communities may benefit and prosper. We tested how exposure to different pCO₂ (400, 1,800 and 4,000 μatm) may affect grazing on the green alga Ulva lactuca by herbivorous fish and sea urchins from the coral reefs in the northern Gulf of Aqaba (Red Sea), either directly, by changing herbivore behaviour, or indirectly via changes in algal palatability. We also determined the effects of pCO₂ on algal tissue concentrations of protein and the grazing-deterrent secondary metabolite dimethylsulfoniopropionate (DMSP). Grazing preferences and overall consumption were tested in a series of multiple-choice feeding experiments in the laboratory and in situ following exposure for 14 d (algae) and 28 d (herbivores). 4,000 μatm had a significant effect on the biochemical composition and palatability of U. lactuca. No effects were observed at 1,800 relative to 400 μatm (control). Exposure of U. lactuca to 4,000 μatm resulted in a significant decrease in protein and increase in DMSP concentration. This coincided with a reduced preference for these algae by the sea urchin Tripneustes gratilla and different herbivorous fish species in situ (Acanthuridae, Siganidae and Pomacanthidae). No feeding preferences were observed for the rabbitfish Siganus rivulatus under laboratory conditions. Exposure to elevated pCO₂ had no direct effect on the overall algal consumption by T. gratilla and S. rivulatus. Our results show that CO₂ has the potential to alter algal palatability to different herbivores which could have important implications for algal abundance and coral community structure. The fact that pCO₂ effects were observed only at a pCO₂ of 4,000 μatm, however, indicates that algal-grazer interactions may be resistant to predicted pCO₂ concentrations in the near future.     

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.     

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.     

Responses of a coral reef shark acutely exposed to ocean acidification conditions

Anthropogenic ocean acidification (OA) is a threat to coral reef fishes, but few studies have investigated responses of high-trophic-level predators, including sharks. We tested the effects of 72-hr exposure to OA-relevant elevated partial pressures of carbon dioxide (pCO₂) on oxygen uptake rates, acid–base status, and haematology of newborn tropical blacktip reef sharks (Carcharhinus melanopterus). Acute exposure to end-of-century pCO₂ levels resulted in elevated haematocrit (i.e. stress or compensation of oxygen uptake rates) and blood lactate concentrations (i.e. prolonged recovery) in the newborns. Conversely, whole blood and mean corpuscular haemoglobin concentrations, blood pH, estimates of standard and maximum metabolic rates, and aerobic scope remained unaffected. Taken together, newborn blacktip reef sharks appear physiologically robust to end-of-century pCO₂ levels, but less so than other, previously investigated, tropical carpet sharks. Our results suggest peak fluctuating pCO₂ levels in coral reef lagoons could still physiologically affect newborn reef sharks, but studies assessing the effects of long-term exposure and in combination with other anthropogenic stressors are needed.

     

Ocean warming has a greater effect than acidification on the early life history development and swimming performance of a large circumglobal pelagic fish

Ocean warming and acidification are serious threats to marine life; however, their individual and combined effects on large pelagic and predatory fishes are poorly understood. We determined the effects of projected future temperature and carbon dioxide (CO₂) levels on survival, growth, morphological development and swimming performance on the early life stages of a large circumglobal pelagic fish, the yellowtail kingfish Seriola lalandi. Eggs, larvae and juveniles were reared in cross‐factored treatments of temperature (21 and 25°C) and pCO₂ (500 and 985 μatm) from fertilisation to 25 days post hatching (dph). Temperature had the greatest effect on survival, growth and development. Survivorship was lower, but growth and morphological development were faster at 25°C, with surviving fish larger and more developed at 1, 11 and 21 dph. Elevated pCO₂ affected size at 1 dph, but not at 11 or 21 dph, and did not affect survival or morphological development. Elevated temperature and pCO₂ had opposing effects on swimming performance at 21 dph. Critical swimming speed (U_crit) was increased by elevated temperature but reduced by elevated pCO₂. Additionally, elevated temperature increased the proportion of individuals that responded to a startle stimulus, reduced latency to respond and increased maximum escape speed, potentially due to the more advanced developmental stage of juveniles at 25°C. By contrast, elevated pCO₂ reduced the distance moved and average speed in response to a startle stimulus. Our results show that higher temperature is likely to be the primary driver of global change impacts on kingfish early life history; however, elevated pCO₂ could affect critical aspects of swimming performance in this pelagic species. Our findings will help parameterise and structure fisheries population dynamics models and improve projections of impacts to large pelagic fishes under climate change scenarios to better inform adaptation and mitigation responses.     

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.     

Shallow Water Marine Sediment Bacterial Community Shifts Along a Natural CO2 Gradient in the Mediterranean Sea Off Vulcano,Italy

The effects of increasing atmospheric CO₂ on ocean ecosystems are a major environmental concern, as rapid shoaling of the carbonate saturation horizon is exposing vast areas of marine sediments to corrosive waters worldwide. Natural CO₂ gradients off Vulcano, Italy, have revealed profound ecosystem changes along rocky shore habitats as carbonate saturation levels decrease, but no investigations have yet been made of the sedimentary habitat. Here, we sampled the upper 2 cm of volcanic sand in three zones, ambient (median pCO₂ 419 μatm, minimum Ω_arag 3.77), moderately CO₂-enriched (median pCO₂ 592 μatm, minimum Ω_arag 2.96), and highly CO₂-enriched (median pCO₂ 1611 μatm, minimum Ω_arag 0.35). We tested the hypothesis that increasing levels of seawater pCO₂ would cause significant shifts in sediment bacterial community composition, as shown recently in epilithic biofilms at the study site. In this study, 454 pyrosequencing of the V1 to V3 region of the 16S rRNA gene revealed a shift in community composition with increasing pCO₂. The relative abundances of most of the dominant genera were unaffected by the pCO₂ gradient, although there were significant differences for some 5 % of the genera present (viz. Georgenia, Lutibacter, Photobacterium, Acinetobacter, and Paenibacillus), and Shannon Diversity was greatest in sediments subject to long-term acidification (>100 years). Overall, this supports the view that globally increased ocean pCO₂ will be associated with changes in sediment bacterial community composition but that most of these organisms are resilient. However, further work is required to assess whether these results apply to other types of coastal sediments and whether the changes in relative abundance of bacterial taxa that we observed can significantly alter the biogeochemical functions of marine sediments.     

Contrasting effects of ocean acidification on reproduction in reef fishes

Differences in the sensitivity of marine species to ocean acidification will influence the structure of marine communities in the future. Reproduction is critical for individual and population success, yet is energetically expensive and could be adversely affected by rising CO₂ levels in the ocean. We investigated the effects of projected future CO₂ levels on reproductive output of two species of coral reef damselfish, Amphiprion percula and Acanthochromis polyacanthus. Adult breeding pairs were maintained at current-day control (446 μatm), moderate (652 μatm) or high CO₂ (912 μatm) for a 9-month period that included the summer breeding season. The elevated CO₂ treatments were consistent with CO₂ levels projected by 2100 under moderate (RCP6) and high (RCP8) emission scenarios. Reproductive output increased in A. percula, with 45–75 % more egg clutches produced and a 47–56 % increase in the number of eggs per clutch in the two elevated CO₂ treatments. In contrast, reproductive output decreased at high CO₂ in Ac. polyacanthus, with approximately one-third as many clutches produced compared with controls. Egg survival was not affected by CO₂ for A. percula, but was greater in elevated CO₂ for Ac. polyacanthus. Hatching success was also greater for Ac. polyacanthus at elevated CO₂, but there was no effect of CO₂ treatments on offspring size. Despite the variation in reproductive output, body condition of adults did not differ between control and CO₂ treatments in either species. Our results demonstrate different effects of high CO₂ on fish reproduction, even among species within the same family. A greater understanding of the variation in effects of ocean acidification on reproductive performance is required to predict the consequences for future populations of marine organisms.     

Increased CO2 stimulates reproduction in a coral reef fish

Ocean acidification is predicted to negatively impact the reproduction of many marine species, either by reducing fertilization success or diverting energy from reproductive effort. While recent studies have demonstrated how ocean acidification will affect larval and juvenile fishes, little is known about how increasing partial pressure of carbon dioxide (pCO₂) and decreasing pH might affect reproduction in adult fishes. We investigated the effects of near‐future levels of pCO₂ on the reproductive performance of the cinnamon anemonefish, Amphiprion melanopus, from the Great Barrier Reef, Australia. Breeding pairs were held under three CO₂ treatments [Current‐day Control (430 μatm), Moderate (584 μatm) and High (1032 μatm)] for a 9‐month period that included the summer breeding season. Unexpectedly, increased CO₂ dramatically stimulated breeding activity in this species of fish. Over twice as many pairs bred in the Moderate (67% of pairs) and High (55%) compared to the Control (27%) CO₂ treatment. Pairs in the High CO₂ group produced double the number of clutches per pair and 67% more eggs per clutch compared to the Moderate and Control groups. As a result, reproductive output in the High group was 82% higher than that in the Control group and 50% higher than that in the Moderate group. Despite the increase in reproductive activity, there was no difference in adult body condition among the three treatment groups. There was no significant difference in hatchling length between the treatment groups, but larvae from the High CO₂ group had smaller yolks than Controls. This study provides the first evidence of the potential effects of ocean acidification on key reproductive attributes of marine fishes and, contrary to expectations, demonstrates an initially stimulatory (hormetic) effect in response to increased pCO₂. However, any long‐term consequences of increased reproductive effort on individuals or populations remain to be determined.     

EVOLUTIONARY RESPONSES OF A COCCOLITHOPHORID GEPHYROCAPSA OCEANICA TO OCEAN ACIDIFICATION

The ongoing ocean acidification associated with a changing carbonate system may impose profound effects on marine planktonic calcifiers. Here, we show that a coccolithophore, Gephyrocapsa oceanica, evolved in response to an elevated CO₂ concentration of 1000 μatm (pH reduced to 7.8) in a long‐term (～670 generations) selection experiment. The high CO₂‐selected cells showed increases in photosynthetic carbon fixation, growth rate, cellular particulate organic carbon (POC) or nitrogen (PON) production, and a decrease in C:N elemental ratio, indicating a greater upregulation of PON than of POC production under the ocean acidification condition. Cells from the low CO₂ selection process shifted to high CO₂ exposure showed an enhanced cellular POC and PON production rates. Our data suggest that the coccolithophorid could adapt to ocean acidification with enhanced assimilations of carbon and nitrogen but decreased C:N ratios.     

Giant Clams and Rising CO2: Light May Ameliorate Effects of Ocean Acidification on a Solar-Powered Animal

Global climate change and ocean acidification pose a serious threat to marine life. Marine invertebrates are particularly susceptible to ocean acidification, especially highly calcareous taxa such as molluscs, echinoderms and corals. The largest of all bivalve molluscs, giant clams, are already threatened by a variety of local pressures, including overharvesting, and are in decline worldwide. Several giant clam species are listed as ‘Vulnerable’ on the IUCN Red List of Threatened Species and now climate change and ocean acidification pose an additional threat to their conservation. Unlike most other molluscs, giant clams are ‘solar-powered’ animals containing photosynthetic algal symbionts suggesting that light could influence the effects of ocean acidification on these vulnerable animals. In this study, juvenile fluted giant clams Tridacna squamosa were exposed to three levels of carbon dioxide (CO₂) (control ~400, mid ~650 and high ~950 μatm) and light (photosynthetically active radiation 35, 65 and 304 μmol photons m^-2 s^-1). Elevated CO₂ projected for the end of this century (~650 and ~950 μatm) reduced giant clam survival and growth at mid-light levels. However, effects of CO₂ on survival were absent at high-light, with 100% survival across all CO₂ levels. Effects of CO₂ on growth of surviving clams were lessened, but not removed, at high-light levels. Shell growth and total animal mass gain were still reduced at high-CO₂. This study demonstrates the potential for light to alleviate effects of ocean acidification on survival and growth in a threatened calcareous marine invertebrate. Managing water quality (e.g. turbidity and sedimentation) in coastal areas to maintain water clarity may help ameliorate some negative effects of ocean acidification on giant clams and potentially other solar-powered calcifiers, such as hard corals.     

Food availability outweighs ocean acidification effects in juvenile Mytilus edulis: laboratory and field experiments

Ocean acidification is expected to decrease calcification rates of bivalves. Nevertheless, in many coastal areas high pCO₂ variability is encountered already today. Kiel Fjord (Western Baltic Sea) is a brackish (12–20 g kg^?1) and CO₂ enriched habitat, but the blue mussel Mytilus edulis dominates the benthic community. In a coupled field and laboratory study we examined the annual pCO₂ variability in this habitat and the combined effects of elevated pCO₂ and food availability on juvenile M. edulis growth and calcification. In the laboratory experiment, mussel growth and calcification were found to chiefly depend on food supply, with only minor impacts of pCO₂ up to 3350 μatm. Kiel Fjord was characterized by strong seasonal pCO₂ variability. During summer, maximal pCO₂ values of 2500 μatm were observed at the surface and >3000 μatm at the bottom. However, the field growth experiment revealed seven times higher growth and calcification rates of M. edulis at a high pCO₂ inner fjord field station (mean pCO₂ ca. 1000 μatm) in comparison to a low pCO₂ outer fjord station (ca. 600 μatm). In addition, mussels were able to out‐compete the barnacle Amphibalanus improvisus at the high pCO₂ site. High mussel productivity at the inner fjord site was enabled by higher particulate organic carbon concentrations. Kiel Fjord is highly impacted by eutrophication, which causes bottom water hypoxia and consequently high seawater pCO₂. At the same time, elevated nutrient concentrations increase the energy availability for filter feeding organisms such as mussels. Thus, M. edulis can dominate over a seemingly more acidification resistant species such as A. improvisus. We conclude that benthic stages of M. edulis tolerate high ambient pCO₂ when food supply is abundant and that important habitat characteristics such as species interactions and energy availability need to be considered to predict species vulnerability to ocean acidification.