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

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Abstract

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

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... Mollusk shells function primarily as a defense against predation, and changes in shell structure due to OA could alter species' vulnerabilities to predation, limit their capacity for inducible defenses by increased calcification, and favor species that are more resilient to OA. For example, the snail Nucella lapillus produces thinner and weaker shells with elevated CO 2 as both juveniles (Rühl et al. 2017) and adults (Queirós et al. 2015), which will likely make this species more vulnerable to predation. OA conditions can also reduce both behavioral responses (Jellison et al. 2016) and inducible defenses (Bibby et al. 2007) of herbivorous snails in the presence of crab predators. ...
... The goal of the present study was to measure the impact that elevated temperature and CO 2 could have on predator-prey relationships between native and invasive species. Recent studies (Ries 2011, Queirós et al. 2015, Harvey & Moore 2017, Rühl et al. 2017) have examined snail growth and feeding under a variety of future conditions and have predicted how species will respond to predators in a changing environment. The present study builds upon this work by comparing responses of 3 snail species with varying shell composition under future ocean conditions and testing varied responses of all 3 species to a globally abundant predatory crab. ...
... The last whorl of an individual of each species was ground and homogenized, then analyzed with Rietveld quantitative analysis (Bish & Howard 1988) to obtain accurate measures of calcite and aragonite ratios. Because shell composition can vary depending on the part of the shell examined (Harper 2000, Queirós et al. 2015, Rühl et al. 2017, the Rietveld analysis was only performed on the last whorl for all of the snail species (the newest part of the shell), and all snail individuals were of similar sizes to minimize any variability in age. The latitudinal cross-sections of the snail shells were also done on similarly sized individuals and were only used to observe relative thickness of calcite and aragonite layers; these were not used for any quantitative analysis. ...
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Predator–prey interactions often drive ecological patterns and are governed by factors including predator feeding rates, prey behavioral avoidance, and prey structural defenses. Invasive species can also play a large ecological role by disrupting food webs, driving local extinctions, and influencing evolutionary changes in prey defense mechanisms. This study documents a substantial reduction in the behavioral and morphological responses of multiple gastropod species (Nucella lapillus, N. ostrina, Urosalpinx cinerea) to an invasive predatory crab (green crab Carcinus maenas) under ocean acidification conditions. These results suggest that climate-related changes in ocean chemistry may diminish non-lethal effects of predators on prey responses including behavioral avoidance. While snails with varying shell mineralogies were similarly successful at deterring predation, those with primarily aragonitic shells were more susceptible to dissolution and erosion under high CO 2 conditions. The varying susceptibility to predation among species with similar ecological roles could indicate that the impacts of invasive species like green crabs could be modulated by the ability of native and invasive prey to withstand ocean acidification conditions. © J. Lord, E. Harper, J. Barry, Moravian College, Monterey Bay Aquarium Research Institute, Cambridge University 2019.
... Shell calcification processes in marine organisms can be affected by ocean acidification as has been documented for a number of molluscan taxa (Gaylord et al., 2011;Coleman et al., 2014;Queirós et al., 2015;Marshall et al., 2019;Doney et al., 2020). The scientific literature on the effects of ocean acidification on molluscs has expanded rapidly indicating a general trend of shell growth reduction under low pH conditions. ...
... The shell degradation of gastropods as a result of ocean acidification (low pH) has been documented in previous studies for several species (Bibby et al., 2007;Gazeau et al., 2013;Melatunan et al., 2013;Queirós et al., 2015;Chatzinikolaou et al., 2017;Harvey et al., 2018;Barclay et al., 2020). The capacity of marine organisms to construct calcium carbonate (CaCO 3 ) shells and skeletons can be impaired by reduced pH (Doney et al., 2009). ...
... A previous study also indicated that the maximum reduction (8%) estimated in the shell density of this species was in the apex, which is the oldest shell part and might often be eroded (Chatzinikolaou et al., 2017). Similarly in Nucella lapillus low pH caused apex dissolution and a 20-30% reduction in shell lip density (Queirós et al., 2015). Mytilus edulis shells grown under ocean acidification conditions displayed significant reductions in shell aragonite thickness, shell thickness index and changes to shell shape, making them more vulnerable to fracture (Fitzer et al., 2015). ...
Article
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The increased absorption of atmospheric CO 2 by the ocean reduces pH and affects the carbonate chemistry of seawater, thus interfering with the shell formation processes of marine calcifiers. The present study aims to examine the effects of ocean acidification and warming on the shell morphological properties of two intertidal gastropod species, Nassarius nitidus and Columbella rustica. The experimental treatments lasted for 3 months and combined a temperature increase of 3 • C and a pH reduction of 0.3 units. The selected treatments reflected the high emissions (RCP 8.5) "business as usual" scenario of the Intergovernmental Panel on Climate Change models for eastern Mediterranean. The morphological and architectural properties of the shell, such as density, thickness and porosity were examined using 3D micro-computed tomography, which is a technique giving the advantage of calculating values for the total shell (not only at specific points) and at the same time leaving the shells intact. Nassarius nitidus had a lower shell density and thickness and a higher porosity when the pH was reduced at ambient temperature, but the combination of reduced pH and increased temperature did not have a noticeable effect in comparison to the control. The shell of Columbella rustica was less dense, thinner and more porous under acidic and warm conditions, but when the temperature was increased under ambient pH the shells were thicker and denser than the control. Under low pH and ambient temperature, shells showed no differences compared to the control. The vulnerability of calcareous shells to ocean acidification and warming appears to be variable among species. Plasticity of shell building organisms as an acclimation action toward a continuously changing marine environment needs to be further investigated focusing on species or shell region specific adaptation mechanisms.
... Global atmospheric carbon dioxide (CO 2 ) levels have increased from about 280 ppm at Pre-industrial to about 400 ppm today (Queiros et al., 2015). The increasing levels of CO 2 in the atmosphere trap more and more heat, warming the atmosphere and the ocean. ...
... At the same time, the increase in atmospheric CO 2 has caused dissolution of 550 billion tons of atmospheric CO 2 in global oceans, resulting in an increase of ocean acidity (pH is reduced from 8.2 to 8.1) in a process known as ocean acidification (OA) (Fig. 1) (Sabine et al. 2011;Feely et al., 2009). It is predicted that by the end of the 21st century, the pH of the ocean will decrease by 0.32 units (Queiros et al., 2015), as a result, OA is considered to be one of the greatest threats to marine ecosystems in the near future Repeated episodes of bivalve massive mortalities (Tan and Zheng, 2019;Tan and Ransangan, 2019;Tan and Ransangan, 2016c;McFarland et al., 2016), deadly bivalve disease outbreaks (Asplund et al., 2014;Williams et al., 2014), growth and development retardations (Watson et al., 2009), bivalve shell dissolutions (Yang et al., 2017;Waldbusser et al., 2010) and bivalve byssus strength reductions O'Donnell et al., 2013), which involved all life stages of bivalve from larvae to juvenile and adults, have been reported worldwide. It is clear from the literature that the ocean warming and ocean acidification are the two main drivers of anthropogenic climate change and have deleterious effects on marine biological processes. ...
... 0.32 units in 2100, and this pH reduction is equivalent to a 150% increase in ocean acidity (IPCC, 2007). According to climate model projections, Queiros et al. (2015) and Gattuso and Hansson (2011) predicted that the ocean pH will reach pH 7.4 by 2300. Higherlatitude systems typically have low buffering capacity to resist acidity changes, therefore, OA has a greater impact on high latitude temperate waters than in tropical waters (Lough and Hobday, 2011;IPCC, 2007). ...
... Alternatively, metabolic depression may be the reason for reduced responsiveness under acidified conditions, as reported for Nassarius festivus, which was able to down-regulate its metabolic rate under low pH conditions (Leung et al., 2015). Nucella lapillus decreased foraging cost with increasing acidification, thus coping with the higher energetic expenditure that could have been triggered by limited chemo-sensory function under these conditions (Queirós et al., 2015). However, in other species, such as the whelk Tenguella marginalba, high pCO 2 increased the standard metabolic rate and energy requirements as the gastropods attempted to maintain homoeostasis, thus resulting in higher consumption of the prey Crassostrea gigas (Wright et al., 2018). ...
... Low pH had negative effects on the locomotion and sensory performance of Concholepas concholepas in the presence of a prey odour, thereby decreasing the overall ability of the gastropods to forage efficiently (Domenici et al., 2017). Queirós et al. (2015) note that low pH affected the predatory response of N. lapillus by increasing foraging distance, indicating a decreased ability to locate food. Although in our study very few adult and juvenile H. trunculus reached their food successfully at low pH, for the ones that eventually managed to do so, low pH did not affect any of the evaluated feeding behaviour parameters (response time, duration, speed, path index). ...
... Although in our study very few adult and juvenile H. trunculus reached their food successfully at low pH, for the ones that eventually managed to do so, low pH did not affect any of the evaluated feeding behaviour parameters (response time, duration, speed, path index). Similarly to our results, foraging duration did not increase with acidification in N. lapillus (Queirós et al., 2015). The opposite was recorded for C. concholepas, for which movement duration and decision time increased, while route finding was not as efficient under high PCO 2 (Domenici et al., 2017). ...
Article
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Increased atmospheric CO2 produced by anthropogenic activities will be absorbed by the oceans over the next century, causing ocean acidification and changes in seawater carbonate chemistry. Elevated CO2 causes sublethal physiological and behavioural responses of the locomotion and foraging behaviour of marine organisms. This study aims to investigate the independent and synergistic effects of long term exposure to low pH and increased temperature on the feeding behaviour of two gastropod species, Hexaplex trunculus and Nassarius nitidus, both in adults and juveniles. The gastropods were kept under controlled temperature (ambient = 20°C, increased = 23°C) and pH (ambient = 8, low = 7.6) conditions for 2.5 years. The percentage of animals that successfully reached their food, response time, total time required to reach their food (duration), and total distance covered, was measured. The speed and path index (i.e. how straightforward the movement is) were estimated as means of foraging efficiency. Increased temperature (under ambient pH) resulted in faster responses, a shorter duration until food was reached and a higher speed in H. trunculus adults. H. trunculus (both adults and juveniles) were less successful in reaching their food source under low pH and ambient temperature compared to all other treatments. The response time, duration, speed and path index were not affected by low pH (at ambient or increased temperature) for H. trunculus adults and juveniles, as well as for N. nitidus. The foraging performance of juveniles hatched and developed under low pH (either at ambient or increased temperature) was more effective than the performance of adults of the same species, thus indicating a degree of acclimation. Also, the scavenger N. nitidus was more successful and responded faster in reaching carrion than the predator H. trunculus, whereas no significant effects were observed for N. nitidus under low pH.
... Alternatively, metabolic depression may be the reason for reduced responsiveness under acidified conditions, as reported for Nassarius festivus, which was able to down-regulate its metabolic rate under low pH conditions (Leung et al., 2015). Nucella lapillus decreased foraging cost with increasing acidification, thus coping with the higher energetic expenditure that could have been triggered by limited chemo-sensory function under these conditions (Queirós et al., 2015). However, in other species, such as the whelk Tenguella marginalba, high pCO 2 increased the standard metabolic rate and energy requirements as the gastropods attempted to maintain homoeostasis, thus resulting in higher consumption of the prey Crassostrea gigas (Wright et al., 2018). ...
... Low pH had negative effects on the locomotion and sensory performance of Concholepas concholepas in the presence of a prey odour, thereby decreasing the overall ability of the gastropods to forage efficiently (Domenici et al., 2017). Queirós et al. (2015) note that low pH affected the predatory response of N. lapillus by increasing foraging distance, indicating a decreased ability to locate food. Although in our study very few adult and juvenile H. trunculus reached their food successfully at low pH, for the ones that eventually managed to do so, low pH did not affect any of the evaluated feeding behaviour parameters (response time, duration, speed, path index). ...
... Although in our study very few adult and juvenile H. trunculus reached their food successfully at low pH, for the ones that eventually managed to do so, low pH did not affect any of the evaluated feeding behaviour parameters (response time, duration, speed, path index). Similarly to our results, foraging duration did not increase with acidification in N. lapillus (Queirós et al., 2015). The opposite was recorded for C. concholepas, for which movement duration and decision time increased, while route finding was not as efficient under high PCO 2 (Domenici et al., 2017). ...
Article
Full-text available
Increased atmospheric CO2 produced by anthropogenic activities will be absorbed by the oceans over the next century causing ocean acidification and changes in the seawater carbonate chemistry. Elevated CO2 causes sublethal physiological and behavioural responses on the locomotion and foraging behaviour of marine organisms. This study aims to investigate the independent and synergistic effects of long term exposure to low pH and increased temperature on the feeding behaviour of two gastropod species, Hexaplex trunculus and Nassarius nitidus, both in adults and juveniles. Gastropods were maintained under controlled conditions of temperature (ambient = 20°C, increased = 23°C) and pH (ambient = 8, low = 7.6) for 2.5 years. The percentage of animals which successfully reached their food, the response time until gastropods began moving, the total duration until they reached food and the total distance covered, were measured. Speed and path index (i.e how straightforward the movement is) were estimated as means of foraging efficiency. Increased temperature (under ambient pH) resulted in faster responses, a shorter duration until food was reached and a higher speed in H. trunculus adults. H. trunculus (both adults and juveniles) were less successful in reaching their food source under low pH and ambient temperature in comparison to all other treatments. The response time, duration, speed and path index were not affected by low pH (at ambient or increased temperature) for H. trunculus adults and juveniles, as well as for N. nitidus. The foraging performance of juveniles hatched and developed under low pH (either at ambient or increased temperature) was more effective than adults of the same species, thus indicating a degree of acclimation. Also, the scavenger N. nitidus was more successful and responded faster in reaching carrion than the predator H. trunculus, whereas no significant effects were observed for N. nitidus under low pH.
... In intertidal zones in the California Current System of western North America, dogwhelks of the genus Nucella are important predators, consuming sedentary, foundational prey (West 1986). Nucella have direct-developing larvae and very low dispersal ability (Strathmann 1987), which gives them an increased ability to adapt to environmental conditions such as temperature and pH that affect foraging strategies (Yamane and Gilman 2009;Queirós et al. 2015;Cerny-Chipman 2016;King and Sebens 2018;Sadler et al. 2018). For example, populations of N. canaliculata exhibit local adaptation in mussel prey selectivity (Sanford et al. 2003;Sanford and Worth 2010). ...
... We predict that temperature will have important effects on size selectivity because it is known to influence Nucella foraging and ingestion rates (Largen 1967;Bayne and Scullard 1978;Sanford 2002;Yamane and Gilman 2009;Miller 2013;King and Sebens 2018). We further expect that pH will shape prey size selectivity because it affects prey detection and predation rate across a wide range of taxa (de la Haye et al. 2012;Pistevos et al. 2015;Watson et al. 2017), including other Nucella species (Queirós et al. 2015;Cerny-Chipman 2016;Sadler et al. 2018). We hypothesize that neutral genetic relatedness will not have a strong effect on size selectivity because Nucella populations have limited dispersal, providing ample opportunity for local adaptation and plasticity to modify feeding traits (Strathmann 1987;Marko 1998;Sanford et al. 2003;Dawson et al. 2014). ...
... Temperature and pH are two important stressors that alter Nucella foraging behavior and can influence size selectivity via risk of prolonged handling. Acidified seawater increases handling time (Queirós et al. 2015;Cerny-Chipman 2016) and causes shell dissolution in Nucella (Nienhuis et al. 2010), so dogwhelks exposed to low pH face a tradeoff between foraging and hiding from their own predators. This tradeoff could lead dogwhelks in lower pH to choose smaller mussels with shorter handling times. ...
Article
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Trait variation among populations is important for shaping ecological dynamics. In marine intertidal systems, seawater temperature, low tide emersion temperature, and pH can drive variation in traits and affect species interactions. In western North America, Nucella dogwhelks are intertidal drilling predators of the habitat-forming mussel Mytilus californianus. Nucella exhibit local adaptation, but it is not known to what extent environmental factors and genetic structure contribute to variation in prey selectivity among populations. We surveyed drilled mussels at sites across Oregon and California, USA, and used multiple regression and Mantel tests to test the effects of abiotic factors and Nucella neutral genetic relatedness on the size of mussels drilled across sites. Our results show that Nucella at sites characterized by higher and less variable temperature and pH drilled larger mussels. Warmer temperatures appear to induce faster handling time, and more stable pH conditions may prolong opportunities for active foraging by reducing exposure to repeated stressful conditions. In contrast, there was no significant effect of genetic relatedness on prey size selectivity. Our results emphasize the role of climate in shaping marine predator selectivity on a foundation species. As coastal climates change, predator traits will respond to localized environmental conditions, changing ecological interactions.
... In intertidal zones in the California Current System of western North America, dogwhelks of the genus Nucella are important predators, consuming sedentary, foundational prey (West 1986). Nucella have direct-developing larvae and very low dispersal ability (Strathmann 1987), which gives them an increased ability to adapt to environmental conditions such as temperature and pH that affect foraging strategies (Yamane and Gilman 2009;Queirós et al. 2015;Cerny-Chipman 2016;King and Sebens 2018;Sadler et al. 2018). For example, populations of N. canaliculata exhibit local adaptation in mussel prey selectivity (Sanford et al. 2003;Sanford and Worth 2010). ...
... We predict that temperature will have important effects on size selectivity because it is known to influence Nucella foraging and ingestion rates (Largen 1967;Bayne and Scullard 1978;Sanford 2002;Yamane and Gilman 2009;Miller 2013;King and Sebens 2018). We further expect that pH will shape prey size selectivity because it affects prey detection and predation rate across a wide range of taxa (De la Haye et al. 2012;Pistevos et al. 2015;Watson et al. 2017), including other Nucella species (Queirós et al. 2015;Cerny-Chipman 2016;Sadler et al. 2018). We hypothesize that neutral genetic relatedness will not have a strong effect on size selectivity because Nucella populations have limited dispersal, providing ample opportunity for local adaptation and plasticity to modify feeding traits (Strathmann 1987;Marko 1998;Sanford et al. 2003;Dawson et al. 2014). ...
... Temperature and pH are two important stressors that alter Nucella foraging behavior and can influence size selectivity via risk of prolonged handling. Acidified seawater increases handling time (Queirós et al. 2015;Cerny-Chipman 2016) and causes shell dissolution in Nucella (Nienhuis et al. 2010), so dogwhelks exposed to low pH face a tradeoff between foraging and hiding from their own predators. This tradeoff could lead dogwhelks in lower pH to choose smaller mussels with shorter handling times. ...
Thesis
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Recent research highlights the prevalence of intraspecific trait variation, even in relatively open ocean habitats. The ecological importance of intraspecific trait variation, however, while shown in freshwater and terrestrial ecosystems, remains unexplored in marine systems. While climates change rapidly and differentially across marine environments at scales within species ranges, population-level trait variation in response to abiotic drivers is inevitable. It is therefore timely and important to explore the climate drivers of intraspecific trait changes and their ecological consequences in marine systems. In this dissertation, I explore these dynamics in a model predator-prey system. The predators, Nucella ostrina-emarginata dogwhelks, exhibit low population connectivity and gene flow due to their life history. The prey, Mytilus californianus, the California mussel, is a foundational mussel that supports high intertidal diversity. These species exist throughout a mosaic of climate conditions in the California Current System, setting the stage for local scale climate effects on Nucella predation that have community consequences. In Chapter 1, I examine climate drivers of population-level variation in size selectivity of Nucella on Mytilus. I find that abiotic variables such as temperature and pH are the strongest drivers of Nucella prey size selectivity rather than neutral genetic relationships among populations, which have no effect. In Chapter 2, I test for population-level differences in the responses to acute exposure to acidified seawater on Nucella size selectivity and consumption time. I find that populations are affected differently by acidification, showing that climate change can affect Nucella predation on local scales. In Chapter 3, I test for community effects of population-level differences in Nucella predation on mussel beds in the field. I find that Nucella predation affects mussel bed size structure and in turn, size structure affects community composition, showing differential predation on a foundation species can alter communities. My dissertation links climate change, trait variation, and community ecology, demonstrating how climate can indirectly alter communities by shaping predator traits on local scales, and expanding the study of population-level trait variation into marine ecosystems.
... The resulting integrated SS-DBEM (size-spectrum and DBEM model) projected slower fish species shifts than in models that did not account for energy limitation. The SS-DBEM has also been applied to several conservation issues (Jones et al., 2013;Queirós et al., 2015) as well as socio-economic assessments in the North-East Atlantic (Fernandes et al., 2017;Mullon et al., 2016;Queirós et al., 2016) and developing countries . However, projections of future species and fishery distributions at local scales are uncertain Frölicher, Rodgers, Stock, & Cheung, 2016;Payne et al., 2016). ...
... Existing studies have made projections of fish distribution and abundance at scales ranging from higher resolution (Burrows et al., 2014;Cheung et al., 2011;Molinos et al., 2016) to higher level of aggregation Mullon et al., 2016). Many studies, including those for fisheries management, only need information at relatively low level of aggregation such as LMEs, FAO areas, seas, ICES areas, EEZs or subregions Fernandes et al., 2016;Mullon et al., 2016;Queirós et al., 2015;Queirós, Fernandes, Genevier, & Lynam, 2018). Applications not dependent on high resolution data are often used for management and economic research, linked with long-term scenarios (Mullon et al., 2016;Queirós et al., 2018) or ecological studies looking at overall impacts on specific species or habitats (Queirós et al., 2015). ...
... Many studies, including those for fisheries management, only need information at relatively low level of aggregation such as LMEs, FAO areas, seas, ICES areas, EEZs or subregions Fernandes et al., 2016;Mullon et al., 2016;Queirós et al., 2015;Queirós, Fernandes, Genevier, & Lynam, 2018). Applications not dependent on high resolution data are often used for management and economic research, linked with long-term scenarios (Mullon et al., 2016;Queirós et al., 2018) or ecological studies looking at overall impacts on specific species or habitats (Queirós et al., 2015). However, other studies and management activities require a finer level of detail, for instance cells of 0.5 × 0.5° or smaller as in the case of ICES rectangles, or even 1 × 1 km 2 for local marine spatial planning or studies of shifts in species abundance centroids (Coccoli et al., 2018;Fernandes et al., 2017;Jones et al., 2013;Queirós et al., 2016). ...
Article
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Large‐scale and long‐term changes in fish abundance and distribution in response to climate change have been simulated using both statistical and process‐based models. However, national and regional fisheries management requires also shorter term projections on smaller spatial scales, and these need to be validated against fisheries data. A 26‐year time series of fish surveys with high spatial resolution in the North‐East Atlantic provides a unique opportunity to assess the ability of models to correctly simulate the changes in fish distribution and abundance that occurred in response to climate variability and change. We use a dynamic bioclimate envelope model forced by physical–biogeochemical output from eight ocean models to simulate changes in fish abundance and distribution at scales down to a spatial resolution of 0.5°. When comparing with these simulations with annual fish survey data, we found the largest differences at the 0.5° scale. Differences between fishery model runs driven by different biogeochemical models decrease dramatically when results are aggregated to larger scales (e.g. the whole North Sea), to total catches rather than individual species or when the ensemble mean instead of individual simulations are used. Recent improvements in the fidelity of biogeochemical models translate into lower error rates in the fisheries simulations. However, predictions based on different biogeochemical models are often more similar to each other than they are to the survey data, except for some pelagic species. We conclude that model results can be used to guide fisheries management at larger spatial scales, but more caution is needed at smaller scales. A 26‐year time series of fish surveys with high spatial resolution in the North‐East Atlantic provides a unique opportunity to assess the ability of models to correctly simulate the changes in fish distribution and abundance that occurred in response to climate.
... As ocean acidification primarily affects the saturation of carbonate ions in seawater (Morse et al., 2007;Ries et al., 2009), it is possible that decreases in carbonate saturation itself may not affect calcification in organisms that are biologically able to control the site of calcification (DeCarlo et al., 2018;Findlay et al., 2011;Leung et al., 2020Leung et al., , 2019Roleda et al., 2012), although hypercapnia would likely still be an added stress (Byrne and Fitzer, 2019). However, existing calcified structures are still susceptible to dissolution if seawater becomes undersaturated in carbonates (Chatzinikolaou et al., 2017;Harvey et al., 2018;Nienhuis et al., 2010;Queirós et al., 2015). Therefore the ability to maintain existing structures or respond plastically to changes in carbonate saturation may become critical for molluscs as carbonate saturation decreases (Duquette et al., 2017;Langer et al., 2014;Leung et al., 2017b;Ries, 2011a), particularly as shells provide protection against predators, parasites, and may also provide temperature regulation (Geller, 1982a;Vermeij, 1993). ...
... The results of the previous study match the prediction that shells with greater aragonite and organic content (e. g., Tegula) experience greater costs with respect to shell production and maintenance under decreased seawater pH than those with more calcite. Other studies of Nucella species have also found that calcification is maintained under decreased pH, and have found that the shells experienced dissolution (Nienhuis et al., 2010;Queirós et al., 2015;Rühl et al., 2017). However, without examining the shell structure of both species in more detail, it is unclear how these responses manifested in either of the species studied, or whether reductions in strength are due to dissolution. ...
... As dissolution was expected to occur on the external surface, for each selected slice, a band of the external shell surface was selected for analysis. Each image was edited in ImageJ, where a 15 pixel-wide band was created along the external edge of the shell to be used in analyses, following the methods of Queirós et al. (2015) (Fig. 1), except in the case of the lip area, where the shell was thin enough to include the entire section. The mean greyscale values of each section were then determined using the binary threshold module (1-255 greyscale histogram range) in the program CTAnalyser (Sky-Scan™), with higher greyscale values indicating greater density of the shell materials. ...
Article
Organisms, such as molluscs, that produce their hard parts from calcium carbonate are expected to show increased difficulties growing and maintaining their skeletons under ocean acidification (OA). Any loss of shell integrity increases vulnerability, as shells provide protection against predation, desiccation, and disease. Not all species show the same responses to OA, which may be due to the composition and microstructural arrangement of their shells. We explore the role of shell composition and microstructure in resisting dissolution caused by decreases in seawater pH using a combination of microCT scans, XRD analysis, and SEM imaging. Two gastropods with different shell compositions and microstructure, Tegula funebralis and Nucella ostrina, were exposed to simulated ocean acidification conditions for six months. Both species showed signs of dissolution on the exterior of their shells, but changes in density were significantly more pronounced in T. funebralis. XRD analysis indicated that the exterior layer of both shell types was made of calcite. T. funebralis may be more prone to dissolution because their outer fibrous calcite layer has more crystal edges and faces exposed, potentially increasing the surface area on which dissolution can occur. These results support a previous study where T. funebralis showed significant decreases in both shell growth and strength, but N. ostrina only showed slight reductions in shell strength, and unaffected growth. We suggest that microstructural arrangement of shell layers in molluscs, more so than their composition alone, is critical for determining the vulnerability of mollusc shells to OA.
... CO 2 was mixed into the airlines supplying the 2 treatment tanks following methods described by Findlay et al. (2008) and gas concentrations were logged and recorded continuously using a CO 2 gas analyser (Licor LI-820). All treatment tanks contained submersible pumps to aid water circulation and mixing, and all were tightly covered with 2mm thick clear PVC sheeting to allow the headspace to equilibrate and to reduce gas exchange with the atmosphere (Queirós et al., 2015). Measurements of seawater pH were recorded daily, and alkalinity samples (A T , 250ml) were taken weekly. ...
... DBEMs calculate changes in growth and other life history traits in response to changes in environmental conditions based on algorithms derived from empirical growth and metabolic functions (Fernandes et al., 2013). It can be argued that the predictions made by these DBEMs are likely to be more robust than those made by other SDMs due to the fact that projected species' distributions are limited by more than just the distribution of suitable habitat (Queirós et al., 2015). However, they are still limited by the fact that they do not incorporate a fully mechanistic physiological understanding of species' responses to changing conditions. ...
... By describing water transports and mixing as well ecosystem processes, it captures essential rates and flows of matter and energy in space and time and links them to the environment and biota. This allows it to project the bulk properties of ecosystems into the future and the past (Allen et al., 2010;Queirós et al., 2015). The large-scale environmental patterns projected by NEMO-ERSEM can be used to force a DEB model, which consequently makes predictions for ecologically relevant endpoints e.g. ...
Thesis
Predicting how marine communities will be affected by environmental change is one of the most significant challenges facing researchers today. In order to tackle this challenge, a mechanistic understanding of climate impacts at the individual level is necessary, as variations in species physiological responses are often reflected in patterns at higher organisational levels such as populations and communities. In order to explore the relationship between individual physiology and higher-level dynamics more fully, the swimming crab Liocarcinus depurator (Linnaeus, 1758) was selected as a model species for experimental work in which whole organism responses (growth, respiration and allocation to reproduction) to climate drivers were investigated using a bio-energetic modelling approach. This species was selected as a model organism after analysis of epibenthic time-series from the Western English Channel monitoring Station L4 revealed that decapod crustaceans played a key role in structuring the benthic community, and that L. depurator was one of the most dominant species in the area, in terms of both abundance and biomass. A bio-energetic approach was used as the same time-series analysis identified water temperature and seasonal phytodetrital input (e.g. food) as the predominant drivers of variation in benthic community wet biomass at L4, with the two drivers appearing to primarily influence community biomass at different times of the year. It is possible that warmer water temperatures in the autumn trigger gonad development and a consequent increase in reproductive biomass, while the sedimentation of the spring phytoplankton bloom drives an increase in somatic biomass. This time-series analysis clearly highlighted the role of organism energetics, and the environmental conditions that influence energy allocation, in structuring benthic communities. Further work elucidated the relationship between environmental variables and individual energy budgets. L. depurator responses to climate drivers (temperature, hypoxia and ocean acidification) were tested experimentally, and a mechanistic Dynamic Energy Budget (DEB) model was parameterised to describe the life history characteristics of crustaceans. At an individual level the model was able to accurately describe and predict observed responses to environmental drivers, both in isolation and in multiple stressor scenarios. Experimental results suggested that L. depurator was broadly tolerant of those climate drivers tested in the short term. Over the longer term however, model scenarios suggested that OA and the combined stressors may have an adverse effect on growth. When the multi-stressor model was forced with environmental projections from a coupled hydrodynamic-biogeochemical model (NEMO-ERSEM), it could be used to make predictions regarding ultimate carbon mass, age-at-maturity and cumulative allocation to reproduction, which were used to infer possible population level effects such as species distributions and population viability. Model scenarios suggested that, in the future, the optimum settlement time for juvenile L. depurator would shift forward across the north-west European shelf, and that this crustacean species may be able to expand its range further into the northern North Sea. The DEB model presented here can provide a mechanistic underpinning of observed species responses to climate drivers, and more broadly, the thesis demonstrates how multi-stressor models can be built from data collected in single stressor experiments, thereby providing a way of synthesising single stressor data into a modelling environment. This approach allows us to simulate more complex, ecologically relevant conditions. At a broader scale, the coupled DEB-ERSEM model showed that it can provide insight into why changes in species’ distributions are predicted, as these distributions are an emergent property of the processes being modelled.
... to predation themselves under rising ocean acidification (Queirós et al., 2015). ...
... Expanding the scope from an individual to a community level is a highly complex undertaking. An generalization approach was made by Queirós et al. (2015), who modeled the distribution of the marine gastropod Nucella lapillus under three IPCC scenarios. Based on the results of a lab experiment, they concluded that spatial distribution cannot be accurately predicted from single-species experiments. ...
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Most intertidal rocky systems are exposed to severe tidal, diurnal, and seasonal changes in environmental parameters. In addition, they show extreme vulnerability to anthropogenic impacts. Research on multiple drivers is therefore crucial to understand the complexity of their potential interactions. Here, we first give an overview of the natural environment and impacts of climate change on rocky shore intertidal systems, and then focus on the impacts of multiple drivers. We further provide a summary of existing multiple driver studies in the literature with the aim for a better understanding of multiple driver interactions. As multiple drivers can affect rocky shore intertidal systems at different spatial and temporal scales, and the outcome of their effects are still more of an “ecological surprise,” we recommend a more widespread assessment of the environmental and biological context. We propose a new, integrated approach based on existing literature: this complements previous frameworks but with an improved understanding of co-occurring multiple driver systems of the rocky intertidal, in order to find management solutions based on accurate and informed predictions in these times of global change.
... Experiments seeking to understand the changes to biotic interactions caused by acidification have grown in recent years and have illustrated a variety of potential ways that acidification may impact these relationships. These include: 1) Temporal and spatial shifts in interactions (e.g., shifts in the timing of development relative to food availability) (Lord et al., 2017); 2) Physiological (metabolism, acid-base balance, calcification) changes that influence feeding rates or increase susceptibility to predation (Queirós et al., 2015;Lord et al., 2017;Sadler et al., 2018); 3) Neurological and behavioral impairments in key functional groups that reduce effectiveness of prey detection and capture ability (Dixson et al., 2015;Queirós et al., 2015;Glaspie et al., 2017;Yu et al., 2017); and 4) Limitations of food resources required to fulfill high energetic demands required to cope with acidification (Saba et al., 2012;Seibel et al., 2012;Ramajo et al., 2016;Hurst et al., 2017). ...
... Experiments seeking to understand the changes to biotic interactions caused by acidification have grown in recent years and have illustrated a variety of potential ways that acidification may impact these relationships. These include: 1) Temporal and spatial shifts in interactions (e.g., shifts in the timing of development relative to food availability) (Lord et al., 2017); 2) Physiological (metabolism, acid-base balance, calcification) changes that influence feeding rates or increase susceptibility to predation (Queirós et al., 2015;Lord et al., 2017;Sadler et al., 2018); 3) Neurological and behavioral impairments in key functional groups that reduce effectiveness of prey detection and capture ability (Dixson et al., 2015;Queirós et al., 2015;Glaspie et al., 2017;Yu et al., 2017); and 4) Limitations of food resources required to fulfill high energetic demands required to cope with acidification (Saba et al., 2012;Seibel et al., 2012;Ramajo et al., 2016;Hurst et al., 2017). ...
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The estuaries and continental shelf system of the United States Mid-Atlantic are subject to ocean acidification driven by atmospheric CO2, and coastal acidification caused by nearshore and land-sea interactions that include biological, chemical, and physical processes. These processes include freshwater and nutrient input from rivers and groundwater; tidally-driven outwelling of nutrients, inorganic carbon, alkalinity; high productivity and respiration; and hypoxia. Hence, these complex dynamic systems exhibit substantial daily, seasonal, and interannual variability that is not well captured by current acidification research on Mid-Atlantic organisms and ecosystems. We present recommendations for research priorities that target better understanding of the ecological impacts of acidification in the U. S. Mid-Atlantic region. Suggested priorities are: 1) Determining the impact of multiple stressors on our resource species as well as the magnitude of acidification; 2) Filling information gaps on major taxa and regionally important species in different life stages to improve understanding of their response to variable temporal scales and sources of acidification; 3) Improving experimental approaches to incorporate realistic environmental variability and gradients, include interactions with other environmental stressors, increase transferability to other systems or organisms, and evaluate community and ecosystem response; 4) Determining the capacity of important species to acclimate or adapt to changing ocean conditions; 5) Considering multi-disciplinary, ecosystem-level research that examines acidification impacts on biodiversity and biotic interactions; and 6) Connecting potential acidification-induced ecological impacts to ecosystem services and the economy. These recommendations, while developed for the Mid-Atlantic, can be applicable to other regions will help align research towards knowledge of potential larger-scale ecological and economic impacts.
... This research brings together a range of methods: experimental manipulations in the laboratory and with mesocosms (e.g. [2,3]), observations of 'natural laboratories' such as low-pH vent sites (e.g. [4,5]), and analysis of large databases (e.g. ...
... The scale of the problem is especially challenging: marine global change involves multiple stressors acting on hundreds of thousands of species, throughout the entire ocean and over decades to centuries of evolutionary time (figure 1). Experimental manipulations, and even natural laboratories, can only provide information at small spatial and temporal scales, and in many cases in controlled but artificial conditions [3,8]. Outcomes may depend on the rate of change [9,10], making extrapolations to longer timescales more difficult. ...
Article
Conservation of marine species requires the ability to predict the effects of climate-related stressors in an uncertain future. Experiments and observations in modern settings provide crucial information, but lack temporal scale and cannot anticipate emergent effects during ongoing global change. By contrast, the deep-time fossil record contains the long-term perspective at multiple global change events that can be used, at a broad scale, to test hypothesized effects of climate-related stressors. For example, geologically rapid carbon cycle disruption has often caused crises in reef ecosystems, and selective extinctions support the hypothesis that greater activity levels promote survival. Geographical patterns of extinction and extirpation were more variable than predicted from modern physiology, with tropical and temperate extinction peaks observed at different ancient events. Like any data source, the deep-time record has limitations but also provides opportunities that complement the limitations of modern and historical data. In particular, the deep-time record is the best source of information on actual outcomes of climate-related stressors in natural settings and over evolutionary timescales. Closer integration of modern and deep-time evidence can expand the types of hypotheses testable with the fossil record, yielding better predictions of extinction risk as climate-related stressors continue to intensify in future oceans. This article is part of a discussion meeting issue ‘The past is a foreign country: how much can the fossil record actually inform conservation?’
... Once in the mesocosm laboratory at Plymouth Marine Laboratory, the core tubes were randomly 192 allocated to two 1-m 3 mesocosm tanks. The laboratory is a temperature controlled room where air 193 temperature is maintained such that aquarium water in the room follows the seasonal cycle of 194 bottom water at the L4 station (Findlay et al., 2008, Queirós et al., 2015. The 1-m 3 mesocosm 195 tanks were used as water baths to ensure that base temperature and light (absence of) conditions 196 experienced by each core tube were as similar as possible during laboratory exposures, and water 197 was not circulated between individual (microcosm) sediment cores. ...
... reproduction, growth, calcification, locomotion) may 573 determine their survival under a changing climate . Increase of food 574 uptake is a strategy that has been observed across taxa, and reflects higher metabolic costs to the 575 individual associated with stress response pathways (Queirós et al., 2015, Thomsen et al., 2013. 576 ...
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Anthropogenic stressors can alter the structure and functioning of infaunal communities, which are key drivers of the carbon cycle in marine soft sediments. Nonetheless, the compounded effects of anthropogenic stressors on carbon fluxes in soft benthic systems remain largely unknown. Here, we investigated the cumulative effects of ocean acidification and hypoxia on the organic carbon fate in marine sediments, through a mesocosm experiment. Isotopically-labelled macroalgal detritus (13C) was used as a tracer to assess carbon incorporation in faunal tissue and in sediments under different experimental conditions. In addition, labelled macroalgae (13C), previously exposed to elevated CO2, were also used to assess the organic carbon uptake by fauna and sediments, when both sources and consumers were exposed to elevated CO2. At elevated CO2, infauna increased the uptake of carbon, likely as compensatory response to the higher energetic costs faced under adverse environmental conditions. By contrast, there was no increase in carbon uptake by fauna exposed to both stressors in combination, indicating that even a short-term hypoxic event may weaken the ability of marine invertebrates to withstand elevated CO2 conditions. In addition, both hypoxia and elevated CO2 increased organic carbon burial in the sediment, potentially affecting sediment biogeochemical processes. Since hypoxia and ocean acidification are predicted to increase in the face of climate change, our results suggest that local reduction of hypoxic events may mitigate the impacts of global climate change on marine soft-sediment systems.
... . Temperature changes throughout this range are likely to engender changes in community structure and functioning (Schindler, 1990), including shifts in population dynamics and species abundances in temperate marine communities (Hale, Calosi, McNeill, Mieszkowska, & Widdicombe, 2011;Queirós et al., 2015). ...
... The goal of this study, however, was not to study stress response, but rather to determine whether increases in near-future mean temperatures would affect E. elongata, intertidal habitat builders, and whether different portions of the environmental mosaic within the intertidal habitat would be differentially affected based on their organisms' environmental history. Such ecophysiological responses may not be tied directly to metabolic rates and photosynthesis, but to population-and community-level changes in competition or ecosystem structure due to range shifts of focal or interacting species (Burrows et al., 2014;Queirós et al., 2015;Vergés et al., 2014). Other studies have found growth sensitivity to nonstressful temperature treatments, though they do not appear tied to differences in photosynthetic activity (Clark, Poore, Ralph, & Doblin, 2013). ...
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Abstract Organisms inhabiting the intertidal zone have been used to study natural ecophysiological responses and adaptations to thermal stress because these organisms are routinely exposed to high‐temperature conditions for hours at a time. While intertidal organisms may be inherently better at withstanding temperature stress due to regular exposure and acclimation, they could be more vulnerable to temperature stress, already living near the edge of their thermal limits. Strong gradients in thermal stress across the intertidal zone present an opportunity to test whether thermal tolerance is a plastic or canalized trait in intertidal organisms. Here, we studied the intertidal pool‐dwelling calcified alga, Ellisolandia elongata, under near‐future temperature regimes, and the dependence of its thermal acclimatization response on environmental history. Two timescales of environmental history were tested during this experiment. The intertidal pool of origin was representative of long‐term environmental history over the alga's life (including settlement and development), while the pool it was transplanted into accounted for recent environmental history (acclimation over many months). Unexpectedly, neither long‐term nor short‐term environmental history, nor ambient conditions, affected photosynthetic rates in E. elongata. Individuals were plastic in their photosynthetic response to laboratory temperature treatments (mean 13.2°C, 15.7°C, and 17.7°C). Further, replicate ramets from the same individual were not always consistent in their photosynthetic performance from one experimental time point to another or between treatments and exhibited no clear trend in variability over experimental time. High variability in climate change responses between individuals may indicate the potential for resilience to future conditions and, thus, may play a compensatory role at the population or species level over time.
... At current CO 2 emission rates, by 2100 atmospheric CO 2 concentrations are predicted to exceed 1000 ppm in some instances (Stocker et al., 2013), leading to reductions in pH of between 0.3 and 0.4 units from today's conditions (Doney et al., 2009). For marine life, changes in pH can have significant, detrimental effects on their structure and functioning (Lemasson et al., 2017a including abnormal larval development (Kurihara et al., 2007;Kurihara, 2008), increased pressure on acid-base regulatory mechanisms (Lindinger et al., 1984;Thomsen and Melzner, 2010;Scanes et al., 2017) and changes to behavior (Queiros et al., 2015;Sadler et al., 2018). For calcifying species, OA can be especially problematic due to disruption of the biomineralization process that is so crucial to the development of shells, exoskeletons and tests. ...
... (e.g., Carcinus maenas, Nucella lapillus) typically use crushing or boring to penetrate the shell (Elner, 1978;Sadler et al., 2018). Furthermore, predators of bivalve mollusks may themselves become susceptible to OA through impacts on fitness (Whiteley, 2011;Landes and Zimmer, 2012;Sadler et al., 2018), feeding structures (Landes and Zimmer, 2012) and predatory behavior (Dixson et al., 2010;Queiros et al., 2015;Sadler et al., 2018). ...
Article
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Negative impacts of global climate change are predicted for a range of taxa. Projections predict marked increases in sea surface temperatures and ocean acidification (OA), arguably placing calcifying organisms at most risk. While detrimental impacts of environmental change on the growth and ultrastructure of bivalve mollusk shells have been shown, rapid and diel fluctuations in pH typical of coastal systems are often not considered. Mytilus edulis, an economically important marine calcifier vulnerable to climate change, were exposed to current and future OA (380 and 1000 ppm pCO2), warming (17 and 20°C), and ocean acidification and warming (OAW) scenarios in a seawater system incorporating natural fluctuations in pH. Both macroscopic morphometrics (length, width, height, volume) and microscopic changes in the crystalline structure of shells (ultrastructure) using electron backscatter diffraction (EBSD) were measured over time. Increases in seawater temperature and OAW scenarios led to increased and decreased shell growth respectively and on marginal changes in cavity volumes. Shell crystal matrices became disordered shifting toward preferred alignment under elevated temperatures indicating restricted growth, whereas Mytilus grown under OAW scenarios maintained single crystal fabrics suggesting OA may ameliorate some of the negative consequences of temperature increases. However, both elevated temperature and OAW led to significant increases in crystal size (grain area and diameter) and misorientation frequencies, suggesting a propensity toward increased shell brittleness. Results suggest adult Mytilus may become more susceptible to biological determinants of survival in the future, altering ecosystem structure and functioning.
... The response of organisms to changing environmental conditions is, therefore, governed by cumulative effects across multiple biological levels, from molecular pathways to whole-organism processes. Ultimately, changes in organism fitness will affect population composition, ecosystem structure, and evolutionary potential (Munday et al., 2013;Queirós et al., 2014). ...
... In contrast, an impact with weaker or less direct links across levels would be a reduction in metabolic efficiency, which represents a malleable response with largely indirect effects on individual fitness through energy reallocation (Cao et al., 2018). Connections across biological levels, therefore, require further consideration as, despite our limited understanding, they are recognized to play a key influence in the overall response of mollusks (Queirós et al., 2014) and other calcifying taxa to environmental change (Calosi et al., 2013;Palumbi et al., 2014). ...
Article
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Biological fitness relies on processes acting at various levels of organization, all of which can be modified by environmental change. Application of synthesis frameworks, such as the Adverse Outcome Pathway (AOP), can enhance our understanding of the responses to stressors identified in studies at each level, as well as the links among them. However, the use of such frameworks is often limited by a lack of data. In order to identify contexts with sufficient understanding to apply the AOP framework, we conducted a meta-analysis of studies considering ocean acidification effects on calcifying mollusks. Our meta-analysis identified that most studies considered the adult life history stage, bivalve taxonomic group, individual-level changes, and growth- and metabolism-related responses. Given the characteristics of the published literature, we constructed an AOP for the effects of ocean acidification on calcification in an adult bivalve, specifically the Pacific oyster (Magallana gigas). By structuring results within the AOP framework, we identify that, at present, the supported pathways by which ocean acidification affects oyster calcification are via the downregulation of cavortin and arginine kinase transcription. Such changes at the molecular level can prompt changes in cellular and organ responses, including altered enzyme activities, lipid peroxidation, and regulation of acid–base status, which have impacts on organism level metabolic rate and, therefore, calcification. Altered calcification may then impact organism mortality and population sizes. We propose that when developed and incorporated in future studies, the AOP framework could be used to investigate sources of complexity including varying susceptibility within and among species, feedback mechanisms, exposure duration and magnitude, and species interactions. Such applications of the AOP framework will allow more effective reflections of the consequences of environmental change, such as ocean acidification, on all levels of biological organization.
... Testing these stressors in isolation and together revealed that only elevated CO 2 was driving behavioral responses when both stressors were present. A similar pattern was found in dogwhelks (Queirós et al., 2015) and sharks , where only elevated CO 2 disrupted chemosensory perception of prey while warming had no effect on these behaviors. Identifying multiple stressor effects on a single sensory modality can allow the correct identification of specific stressor effects on different senses [e.g., chemosensation , vision (Ferrari et al., 2012), and audition (Simpson et al., 2011) in coral reef fish]. ...
Article
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Ecosystems are shaped by complex interactions between species and their environment. However, humans are rapidly changing the environment through increased carbon dioxide (CO2) emissions, creating global warming and elevated CO2 levels that affect ecological communities through multiple processes. Understanding community responses to climate change requires examining the consequences of changing behavioral interactions between species, such as those affecting predator and prey. Understanding the underlying sensory process that govern these interactions and how they may be affected by climate change provides a predictive framework, but many studies examine behavioral outcomes only. This review summarizes the current knowledge of global warming and elevated CO2 impacts on predator-prey interactions with respect to the relevant aspects of sensory ecology, and we discuss the potential consequences of these effects. Our specific questions concern how climate change affects the ability of predators and prey to collect information and how this affects predator-prey interactions. We develop a framework for understanding how warming and elevated CO2 can alter behavioral interactions by examining how the processes (steps) of sensory cue (or signal) production, transmission and reception may change. This includes both direct effects on cue production and reception resulting from changes in organismal physiology, but also effects on cue transmission resulting from modulation of the physical environment via physical and biotic changes. We suggest that some modalities may be particularly prone to disruption, and that aquatic environments may suffer more serious disruptions as a result of elevated CO2 and warming that collectively affect all steps of the signaling process. Temperature by itself may primarily operate on aspects of cue generation and transmission, implying that sensory-mediated disruptions in terrestrial environments may be less severe. However, significant biases in the literature in terms of modalities (chemosensation), taxa (fish), and stressors (elevated CO2) examined currently prevents accurate generalizations. Significant issues such as multimodal compensation and altered transmission or other environmental effects remain largely unaddressed. Future studies should strive to fill these knowledge gaps in order to better understand and predict shifts in predator-prey interactions in a changing climate.
... Thus, overall organisms, populations, communities, and entire ecosystems are predicted to have their physiological and ecological potential reduced across multiple abiotic (and consequently biotic) dimensions and traits, which will lead to pronounced impacts on both non-vital and vital functions, severely compromising organism, population, community, and ecosystem viability. It is important to retain that these alterations are ongoing and that field data already reveals significant alterations in community dynamics and species distribution (Breitburg et al. 2018;Queirós et al. 2015;Queiroz et al. 2016;Stramma et al. 2012). Ocean deoxygenation, warming, and acidification alter biogeochemical cycles, climate-regulating processes, heat distribution, wind regimes, and ecosystem services for the human population (Breitburg et al. 2018;IPCC 2013;Kroeker et al. 2012). ...
... All aquaria were gently topped up using local seawater on deck, and were individually aerated using diffusing air stones and shaded during the 4-6 h transport to the PML mesocosm laboratory. This is a temperature-controlled room where the air temperature is regulated on a monthly basis so that aquaria in the room follow the seasonal temperature cycle of bottom seawater at Station L4 (Findlay et al. 2008, Queir os et al. 2015a). On arrival to the mesocosm laboratory, the temperature of seawater overlying each core was recorded again, verifying that it had not risen by more than 2°C. ...
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Macroalgae drive the largest CO 2 flux fixed globally by marine macrophytes. Most of the resulting biomass is exported through the coastal ocean as detritus and yet almost no field measurements have verified its potential net sequestration in marine sediments. This gap limits the scope for the inclusion of macroalgae within blue carbon schemes that support ocean carbon sequestration globally, and the understanding of the role their carbon plays within distal food webs. Here, we pursued three lines of evidence (eDNA sequencing, Bayesian Stable Isotope Mixing Modeling, and benthic-pelagic process measurements) to generate needed, novel data addressing this gap. To this end, a 13-month study was undertaken at a deep coastal sedimentary site in the English Channel, and the surrounding shoreline of Plymouth, UK. The eDNA sequencing indicated that detritus from most macroalgae in surrounding shores occurs within deep, coastal sediments, with detritus supply reflecting the seasonal ecology of individual species. Bayesian stable isotope mixing modeling [C and N] highlighted its vital role in supporting the deep coastal benthic food web (22-36% of diets), especially when other resources are seasonally low. The magnitude of detritus uptake within the food web and sediments varies seasonally, with an average net sedimentary organic macroalgal carbon sequestration of 8.75 g CÁm À2 Áyr À1. The average net sequestration of particulate organic carbon in sediments is 58.74 g CÁm À2 Áyr À1 , the two rates corresponding to 4-5% and 26-37% of those associated with mangroves, salt marshes, and seagrass beds, systems more readily identified as blue carbon habitats. These novel data provide important first estimates that help to contextualize the importance of macroalgal-sedimentary connectivity for deep coastal food webs, and measured fluxes help constrain its role within global blue carbon that can support policy development. At a time when climate change mitigation is at the foreground of environmental policy development, embracing the full potential of the ocean in supporting climate regulation via CO 2 sequestration is a necessity.
... The impact of OA on shell microstructure of several species, maintained in experimental OA for various lengths of time has been investigated (Table 1). For the ecologically important species, Nucella lapillus and Nassarius nitidus, the reduction in shell density determined using μCT was marked (20-50%), while that for Columbella rustica was much less (0.8-8%) (Queirós et al., 2015;Chatzinikolaou et al., 2017). This shows differences in vulnerability of shells to OA between closely related gastropod species. ...
Article
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Ocean acidification (OA), from seawater uptake of anthropogenic CO2, has a suite of negative effects on the ability of marine invertebrates to produce and maintain their skeletons. Increased organism pCO2 causes hypercapnia, an energetically costly physiological stress. OA alters seawater carbonate chemistry, limiting the carbonate available to form the calcium carbonate (CaCO3) minerals used to build skeletons. The reduced saturation state of CaCO3 also causes corrosion of CaCO3 structures. Global change is also accelerating coastal acidification driven by land-run off (e.g. acid soil leachates, tannic acid). Building and maintaining marine biomaterials in the face of changing climate will depend on the balance between calcification and dissolution. Overall, in response to environmental acidification, many calcifiers produce less biomineral and so have smaller body size. Studies of skeleton development in echinoderms and molluscs across life stages show the stunting effect of OA. For corals, linear extension may be maintained, but at the expense of less dense biomineral. Conventional metrics used to quantify growth and calcification need to be augmented by characterisation of the changes to biomineral structure and mechanical integrity caused by environmental acidification. Scanning electron microscopy and microcomputed tomography of corals, tube worms and sea urchins exposed to experimental (laboratory) and natural (vents, coastal run off) acidification show a less dense biomineral with greater porosity and a larger void space. For bivalves, CaCO3 crystal deposition is more chaotic in response to both ocean and coastal acidification. Biomechanics tests reveal that these changes result in weaker, more fragile skeletons, compromising their vital protective roles. Vulnerabilities differ among taxa and depend on acidification level. Climate warming has the potential to ameliorate some of the negative effects of acidification but may also make matters worse. The integrative morphology-ecomechanics approach is key to understanding how marine biominerals will perform in the face of changing climate.
... software. Ten vertical 2-D cross sections were randomly selected throughout the length of each shell, and the total shell density was measured by tracing a ROI around the perimeter of the shell (Chatzinikolaou et al., 2017;Queirós et al., 2015). To compare the density of the periostracum (rough outer layer of the shell exposed to the water) to the overall shell density, 15 pixels were selected as the ROI along the outside of the shell (Papageorgiou and Schmidbaur, 2014;Rühl et al., 2017). ...
... Many studies have found that the change in hydrogen ion concentration that underpins reduced pH can cause physiological challenges that require additional energy consumption to rectify/moderate (e.g. oxygen consumption - Queirós et al., 2015;heart rate -Lim and Harley, 2018). Analogous research conducted on the lecithotrophic lifestages of non-parasitic organisms has repeatedly shown that exposure to reduced pH and associated acid-base challenges has resulted in increased energetic expenditure (Dupont et al., 2010a). ...
Article
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 (OAW), and have the potential to substantially alter the physiological performance of many marine organisms. It is vital that the effects of OAW on marine organisms are explored so that we can predict how marine communities may change in future. In particular, the effect of OAW 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 OAW 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.
... Thus, overall organisms, populations, communities, and entire ecosystems are predicted to have their physiological and ecological potential reduced across multiple abiotic (and consequently biotic) dimensions and traits, which will lead to pronounced impacts on both non-vital and vital functions, severely compromising organism, population, community, and ecosystem viability. It is important to retain that these alterations are ongoing and that field data already reveals significant alterations in community dynamics and species distribution (Breitburg et al. 2018;Queirós et al. 2015;Queiroz et al. 2016;Stramma et al. 2012). Ocean deoxygenation, warming, and acidification alter biogeochemical cycles, climate-regulating processes, heat distribution, wind regimes, and ecosystem services for the human population (Breitburg et al. 2018;IPCC 2013;Kroeker et al. 2012). ...
... Previous studies on M. edulis found that thread production increases as temperature ranges from 2 • C to 18 • C Young, 1985). The results reported here suggest that 25 • C exceeds the point where temperature has a positive influence on thread production for M. trossulus and the temperature optimum for thread production in this species likely lies between 18 • C and 25 • C. Mussels used in this study were collected from a location where temperatures range as high as 25 • C on warm summer days (Nishizaki & Carrington, 2014), suggesting that temperature effects on structural integrity could play a role in defining species distribution limits (Queirós et al., 2015;Kroeker et al., 2016). We note, however, that temperatures only reach 25 • C on the hottest days in the summer, and it is worth exploring further how the interplay of warming level, duration, and frequency in relation to hydrodynamic loading determines when and where temperature will become too high to support viable mussel attachment. ...
Article
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Predicting how combinations of stressors will affect failure risk is a key challenge for the field of ecomechanics and, more generally, ecophysiology. Environmental conditions often influence the manufacture and durability of biomaterials, inducing structural failure that potentially compromises organismal reproduction, growth, and survival. Species known for tight linkages between structural integrity and survival include bivalve mussels, which produce numerous byssal threads to attach to hard substrate. Among the current environmental threats to marine organisms are ocean warming and acidification. Elevated pCO2 exposure is known to weaken byssal threads by compromising the strength of the adhesive plaque. This study uses structural analysis to evaluate how an additional stressor, elevated temperature, influences byssal thread quality and production. Mussels (Mytilus trossulus) were placed in controlled temperature and pCO2 treatments, and then, newly produced threads were counted and pulled to failure to determine byssus strength. The effects of elevated temperature on mussel attachment were dramatic; mussels produced 60% weaker and 65% fewer threads at 25°C in comparison to 10°C. These effects combine to weaken overall attachment by 64–88% at 25°C. The magnitude of the effect of pCO2 on thread strength was substantially lower than that of temperature and, contrary to our expectations, positive at high pCO2 exposure. Failure mode analysis localized the effect of temperature to the proximal region of the thread, whereas pCO2 affected only the adhesive plaques. The two stressors therefore act independently, and because their respective target regions are interconnected (resisting tension in series), their combined effects on thread strength are exactly equal to the effect of the strongest stressor. Altogether, these results show that mussels, and the coastal communities they support, may be more vulnerable to the negative effects of ocean warming than ocean acidification.
... However, reduced preference for algae with lower nutritional content results in contrastingly decreased consumption in multiple choice feeding assays (Borell et al., 2013;Duarte et al., 2016), highlighting the role that foraging behavior plays in mediating rates of herbivory. Foraging behavior may also be negatively impacted by ocean acidification through the impaired ability of grazers to orient to the chemical cues produced by their food, which has been documented in urchins , molluscs (Queirós et al., 2015;Horwitz et al., 2020), and crustaceans . Therefore, predicting the effects of ocean acidification on urchin herbivory is complicated by the interaction of complex urchin and macroalgae physiology and the resulting impacts to multiple facets of their trophic interaction. ...
Article
When predicting the response of marine ecosystems to climate change, it is increasingly recognized that understanding the indirect effects of ocean acidification on trophic interactions is as important as studying direct effects on organism physiology. Furthermore, comprehensive studies that examine these effects simultaneously are needed to identify and link the underlying mechanisms driving changes in species interactions. Using an onshore ocean acidification simulator system, we investigated the direct and indirect effects of elevated seawater pCO2 on the physiology and trophic interaction of fleshy macroalgae and the grazing sea urchin Lytechinus variegatus. Macroalgal (Dictyota spp.) biomass increased despite decreased photosynthetic rates after two-week exposure to elevated pCO2. Algal tissue carbon content remained constant, suggesting the use of alternative carbon acquisition pathways beneficial to growth under acidification. Higher C:N ratios driven by a slight reduction in N content in algae exposed to elevated pCO2 suggest a decrease in nutritional content under acidification. Urchin (L. variegatus) respiration, biomass, and righting time did not change significantly after six-week exposure to elevated pCO2, indicating that physiological stress and changes in metabolism are not mechanisms through which the trophic interaction was impacted. Correspondingly, urchin consumption rates of untreated macroalgae (Caulerpa racemosa) were not significantly affected by pCO2. In contrast, exposure of urchins to elevated pCO2 significantly reduced the number of correct foraging choices for ambient macroalgae (Dictyota spp.), indicating impairment of urchin chemical sensing under acidification. However, exposure of algae to elevated pCO2 returned the number of correct foraging choices in similarly exposed urchins to ambient levels, suggesting alongside higher C:N ratios that algal nutritional content was altered in a way detectable by the urchins under acidification. These results highlight the importance of studying the indirect effects of acidification on trophic interactions simultaneously with direct effects on physiology. Together, these results suggest that changes to urchin chemical sensing and algal nutritional quality are the driving mechanisms behind surprisingly unaltered urchin foraging behavior for fleshy macroalgae under joint exposure to ocean acidification. Consistent foraging behavior and consumption rates suggest that the trophic interaction between L. variegatus and fleshy macroalgae may be sustained under future acidification. However, increases in fleshy macroalgal biomass driven by opportunistic carbon acquisition strategies have the potential to cause ecological change, depending on how grazer populations respond. Additional field research is needed to determine the outcome of these results over time and under a wider range of environmental conditions.
... Due to anthropogenic activities, largely increased amount of atmospheric carbon dioxide (CO 2 ) sinks into ocean, causing significant alterations in oceanic carbon chemistry (Barnett et al., 2005;Feely et al., 2009;Hoegh-Guldberg et al., 2014). The increased dissolution of CO 2 into ocean lowered the average pH level of global ocean by reducing the carbonate ions, while the acidity of ocean is expected to be dropped by approximately 0.4 pH units in the year 2100 (Caldeira et al., 2005;Cao et al., 2007;Bopp et al., 2013), and anticipated to cause adverse effects on biological and physiological processes in organisms which inhabit in marine ecosystems (Riebesell et al., 2004;Raven et al., 2005;Doney et al., 2009;Kroeker et al., 2013;Queirós et al., 2014). Although the potential effect of ocean acidification (OA) was widely studied, most researches mainly focused on the physiological effects in calcifying _______________________ * Corresponding authors. ...
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Ocean acidification (OA) is caused by alteration of global ocean carbon chemistry due to the increased pCO2 in the atmosphere and caused deleterious impacts on the marine ecosystem. Although various detrimental effects of OA were reported in marine organisms, the potential impact of OA on aquatic invertebrates still remain largely unknown. Here, we examined changes in life parameters and antioxidant system in response to low pH (7.5 and 7) in the marine copepod Tigriopus japonicus. Exposure of lower pHs (pH 7.5 and 7.0) to copepod resulted in reduction in the developmental time with decreased fecundity and body length. Also, they showed the increased reactive oxygen species with the enhanced glutathione S-transferase and glutathione reductase activities but showed the decreased glutathione peroxidase and superoxide dismutase activities in pH-dependent manner, indicating that OA exposure caused disturbance of the redox system in T. japonicus. Among several oxidative stress-related genes, GSTs2b was significantly up-regulated in response to OA. These findings will be helpful for a better understanding on the potential impact of OA on life parameters and antioxidant system in the marine copepod T. japonicus.
... following a 12 month exposure (Tambutté et al., 2015). For the gastropods Nassarius nitidus and Nucella lapillus, μCT images revealed lower shell density and abnormal thinning and cracks after a year in pH NBS 7.6 (Queirós et al., 2015;Chatzinikolaou et al., 2017). ...
Article
The article is available for open view until January 7, 2020 at https://authors.elsevier.com/c/1a3tS51aUZHW~ Anthropogenic CO2–driven ocean acidification (OA) is causing a decrease in seawater pH and the saturation state of calcium carbonate minerals, compromising the ability of calcifying species to produce and maintain their skeletons. Sea urchins are ecologically important calcifying species and we investigated the impacts of long-term (9 month) exposure to near-future OA (Ambient–pHNBS 8.01; OA–pHNBS 7.6) on the skeleton microstructure ofHeliocidaris erythrogramma using scanning electron microscopy (SEM), micro-computed tomography (μCT) and nanoindentation. SEM revealed that the youngest plates (apical plates) which had likely grown in experimental conditions had larger pores in the OA group (pore surface area ~ 72% larger) compared with those of urchins maintained in ambient pH. High-resolution,μCT 3-D reconstructions of the apical plates revealed that the experimental OA treatment urchins had a ~14% greater porosity and ~17% less biomineral, suggesting an inability to finely regulate skeletogenesis. The mid-test ambital plates established prior to this study did not show any OA associated change in porosity. Nanoindentation of the apical plates indicated that OA reduced skeletal hardness and elasticity. Stereom pore size is a key trait of the sea urchin endoskeleton and increased porosity in H. erythrogrammais likely to impact its biological functions as well as its biomechanical capacity to defend against predation and physical disturbances
... We exposed P. oceanica meadows, at both ambient and low pH, to different levels of nutrient enrichment (control, moderate, and high) for 16 months. Under OA scenario, food availability seems to play a critical role for marine invertebrates, by providing the energy required to support physiological responses to pH stress ( Thomsen et al. 2013;Queiros et al. 2015;Ramajo et al. 2016). Under these circumstances, a moderate increase in nutrient availability could have positive effects on meiobenthos at low pH, possibly increasing the consumption of more bioavailable food (Danovaro 1996;Antón et al. 2011). ...
Article
Seagrass meadows are an important organic matter (OM) reservoir but, are currently being lost due to global and regional stressors. Yet, there is limited research investigating the cumulative impacts of anthropogenic stressors on the structure and functioning of seagrass benthic assemblages, key drivers of OM mineralization and burial. Here, using a 16‐month field experiment, we assessed how meiobenthic assemblages and extracellular enzymatic activities (as a proxy of OM degradation) in Posidonia oceanica sediments responded to ocean acidification (OA) and nutrient loadings, at CO2 vents. P. oceanica meadows were exposed to three nutrient levels (control, moderate, and high) at both ambient and low pH sites. OA altered meiobenthic assemblage structure, resulting in increased abundance of annelids and crustaceans, along with a decline in foraminifera. In addition, low pH enhanced OM degradation rates in seagrass sediments by enhancing extracellular enzymatic activities, potentially decreasing the sediment carbon storage capacity of seagrasses. Nutrient enrichment had no effect on the response variables analyzed, suggesting that, under nutrient concentration unlikely to cause N or P imitation, a moderate increase of dissolved nutrients in the water column had limited influence on meiobenthic assemblages. These findings show that OA can significantly alter meiobenthic assemblage structure and enhance OM degradation rates in seagrass sediments. As meiofauna are ubiquitous key actors in the functioning of benthic ecosystems, we postulated that OA, altering the structure of meiobenthic assemblages and OM degradation, could affect organic carbon sequestration over large spatial scales.
... Those that have, have focussed on gastropod shells, the outer surfaces of which undergo passive dissolution in ways that often correlate with reductions in seawater pH and aragonite saturation (Ω ara ) (Bednaršek et al., 2012a, b, 2014, Marshall et al., 2008. Many studies have used gastropod shells to understand ecological impacts of ocean acidification Garilli et al., 2015;Hall-Spencer et al., 2008), but these have largely concerned shell calcification and energetics (Chen et al., 2015;Connell et al., 2017;Harvey et al., 2016Harvey et al., , 2018Spalding et al., 2017), to appreciate the constraints on building shells under carbonate-undersaturated conditions Duquette et al., 2017;Queirós et al., 2015;Rodolfo-Metalpa et al., 2011). Gastropod biomonitoring studies have specifically considered pelagic, oceanic pteropods (Bednaršek et al., 2012a(Bednaršek et al., , b, 2014, and estuarine and rocky shore animals (Marshall et al., 2008(Marshall et al., , 2019. ...
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The rapidly changing marine environmental chemistry associated with growing industrialisation, urban population expansion, and the unabated rise in atmospheric CO2 necessitates monitoring. Traditional approaches using metres, dataloggers, and buoys to monitor marine acidification have limited application in coastal oceans and intertidal zones subjected to direct wave action. The present study trialled a system to biomonitor coastal acidification (carbonate ion and pH) based on the dissolution of living gastropod shells. We extended on an approach that ranked shell erosion (SER) in Nerita chamaeleon (Nc) in environments where such erosion was found to correlate with exposure to acidified water. We assessed the spatial scale at which the Nc-SER marker could detect change in acidification along rocky shores, and whether snail body size affected this marker. We found that proportional and unique Nc-SERs not only varied between acidified and non-acidified reference shores at a coarse spatial scale (10 km), but also in predictable ways at fine scales (metres), vertically and horizontally within a shore. Differences between acidified and reference shores in the relationship for snail size and Nc-SER were accentuated by less weathered shells at reference localities, highlighting the value of including small, juvenile snails in monitoring protocols. Gastropod shells are shown to be useful for assessing point sources of acidification and the spatial area of affected coastal zones. This cost-effective and easy-to-use approach (potentially even by citizen-scientists) offers an early warning system of acidification of rocky shore ecosystems, where the deployment of instruments is precluded.
... • Weakly acidic environments impaired the ability of juvenile salmon to detect and respond to alarm cues [100]. • Littorinid snails on rocky shores switched from thickening shells in the presence of predators to avoidance behavior when exposed to OA [101], while the predatory snail, Nucella, showed decreased chemosensory abilities, taking longer to locate food [102]. • Impacts on chemosensory function when exposed to acidification have also been observed in some crabs [103] and fish (e.g., Refs. ...
Chapter
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Oceans have the capacity to absorb large amounts of carbon dioxide (CO2) because CO2 dissolves and reacts in seawater to form bicarbonate (HCO3⁻) and protons (H⁺). About a quarter to a third of the CO2 emitted into the atmosphere from the burning of fossil fuels, cement manufacturing, and land use changes has been absorbed by the oceans. Over thousands of years, the changes in pH have been buffered by bases, such as carbonate ions (CO3²⁻). However, the rate at which CO2 is currently being absorbed into the oceans is too rapid to be buffered sufficiently to prevent substantial changes in ocean pH and CO3²⁻. As a consequence, the relative seawater concentrations of CO2, HCO3–, CO3²⁻, and pH have been altered. Since preindustrial times, the ocean pH has decreased by a global average of 0.1. It is estimated that unmitigated CO2 emissions will cause ocean pH to decrease by as much as 0.4 by 2100 and 0.77 by 2300. These will be the most rapid and greatest changes in ocean carbonate chemistry experienced by marine organisms over the past tens of millions of years. Laboratory experiments, field observations of natural CO2-rich seawater “hot spots,” and studies of previous ocean acidification in Earth's history indicate that these changes are a threat to the survival of many marine organisms but particularly organisms that use CaCO3 to produce shells, tests, and skeletons. The only way of reducing the impacts of ocean acidification on a global scale is through urgent and substantial reductions in anthropogenic CO2 emissions. Ocean acidification is a key argument for united global societal action in ongoing climate change negotiations.
... It was demonstrated that short-term exposures to low pH values could lead to hypercapnic conditions in sea urchin coelomic fluid (Miles et al. 2007;Spicer et al. 2011;Dupont and Thorndyke 2012;Spicer and Widdicombe 2012;Stumpp et al. 2012;Holtmann et al. 2013;Kurihara et al. 2013). This effect could represent a "shock" response (Byrne 2012;Queirós et al. 2015) possibly related to both the absence of respiratory pigments and low capability of sea urchins to regulate ions. Generally, in most species, with few exceptions (Kurihara et al. 2013), the acid-base balance is recovered in some days or weeks (Calosi et al. 2013;Dupont and Thorndyke 2012;Stumpp et al. 2012;Holtman et al. 2013;Moulin et al. 2014). ...
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Seawater pH lowering, known as ocean acidification, is considered among the major threats to marine environment. In this study, post-spawning adults of the sea urchin Paracentrotus lividus were maintained at three pH values (8.0, 7.7, 7.4) for 60 days. Physiological, biochemical, cellular, behavioural and reproductive responses were evaluated in males and females. Significant differences between sexes were observed, with higher ammonia excretion and lower catalase activity in males. Respiration rate (after 21 days), catalase activity in gonads and total coelomocyte count showed the same increasing trend in males and females under low pH. Ammonia excretion, gonadosomatic index and lysozyme activity exhibited opposite responses to low pH, with an increasing trend in males and decreasing in females. Results demonstrated that exposure to low pH could result in different response strategies of male and female sea urchins at a physiological, biochemical and immunological level. Reduced female gonadosomatic index under low pH suggested decreased energy investment in reproduction.
... In parallel, many seabed species have limited ability to track climatedriven changes in habitat distributions due to limited movement and dispersal ability (Hiddink et al., 2015). This is exacerbated by individual-level trade-offs between energetically costly stress response pathways triggered by environmental change and processes supporting dispersal potential (Calosi et al., 2013;Queirós et al., 2015). Such impacts likely further affect important benthic-pelagic coupling processes mediated by benthic communities, indicating that CC may in this way indirectly affect broader ocean functioning in the region (Snelgrove et al., 2018). ...
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Marine spatial planning that addresses ocean climate-driven change (‘climate-smart MSP’) is a global aspiration to support economic growth, food security and ecosystem sustainability. Ocean climate change (‘CC’) modelling may become a key decision-support tool for MSP, but traditional modelling analysis and communication challenges prevent their broad uptake. We employed MSP-specific ocean climate modelling analyses to inform a real-life MSP process; addressing how nature conservation and fisheries could be adapted to CC. We found that the currently planned distribution of these activities may become unsustainable during the policy's implementation due to CC, leading to a shortfall in its sustainability and blue growth targets. Significant, climate-driven ecosystem-level shifts in ocean components underpinning designated sites and fishing activity were estimated, reflecting different magnitudes of shifts in benthic versus pelagic, and inshore versus offshore habitats. Supporting adaptation, we then identified: CC refugia (areas where the ecosystem remains within the boundaries of its present state); CC hotspots (where climate drives the ecosystem towards a new state, inconsistent with each sectors’ present use distribution); and for the first time, identified bright spots (areas where oceanographic processes drive range expansion opportunities that may support sustainable growth in the medium term). We thus create the means to: identify where sector-relevant ecosystem change is attributable to CC; incorporate resilient delivery of conservation and sustainable ecosystem management aims into MSP; and to harness opportunities for blue growth where they exist. Capturing CC bright spots alongside refugia within protected areas may present important opportunities to meet sustainability targets while helping support the fishing sector in a changing climate. By capitalizing on the natural distribution of climate resilience within ocean ecosystems, such climate-adaptive spatial management strategies could be seen as nature-based solutions to limit the impact of CC on ocean ecosystems and dependent blue economy sectors, paving the way for climate-smart MSP.
... Finally, artificial reefs, like natural reefs, are being subjected to a warmer and acidified marine environment. The combination of acidification and warming leads to substantial, non-additive and complex changes in community dynamics (Queirós et al., 2015), affects pelagic and benthic nutrient cycling (Braeckman et al., 2014), and alters the mechanism behind predator-prey interactions (Draper and Weissburg, 2019). Thus, current understanding of the artificial reef effect in OWFs must be considered within a modern changing environment. ...
Article
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Offshore wind farms (OWFs) are proliferating globally. The submerged parts of their structures act as artificial reefs, providing new habitats and likely affecting fisheries resources. While acknowledging that the footprints of these structures may result in loss of habitat, usually soft sediment, we focus on how the artificial reefs established by OWFs affect ecosystem structure and functioning. Structurally, the ecological response begins with high diversity and biomass in the flora and fauna that gradually colonize the complex hard substrate habitat. The species may include nonindigenous ones that are extending their spatial distributions and/or strengthening populations, locally rare species (e.g., hard substrate-associated fish), and habitat-forming species that further increase habitat complexity. Functionally, the response begins with dominant suspension feeders that filter organic matter from the water column. Their fecal deposits alter the surrounding seafloor communities by locally increasing food availability, and higher trophic levels (fish, birds, marine mammals) also profit from locally increased food availability and/or shelter. The structural and functional effects extend in space and time, impacting species differently throughout their life cycles. Effects must be assessed at those larger spatiotemporal scales.
... The flumes were then left to settle for 7 days in darkened and aerated conditions at L4 bottom water temperatures. The mesocosm laboratory is an environment controlled room where, in addition to other controls, air, and thereby also water, temperature is regulated on a monthly basis to follow the seasonal cycle of bottom water temperature at L4 (Queirós et al., 2015a). ...
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In coastal temperate environments, many processes known to affect the exchange of particulate and dissolved matter between the seafloor and the water column follow cyclical patterns of intra-annual variation. This study assesses the extent to which these individual short term temporal variations affect specific direct drivers of seafloor-water exchanges, how they interact with one another throughout the year, and what the resulting seasonal variation in the direction and magnitude of benthic-pelagic exchange. Existing data from a multidisciplinary long-term time-series from the Western Channel Observatory, UK, were combined with new experimental and in-situ data collected throughout a full seasonal cycle. These data were used to define an average year, split into five ‘periods’ (winter, pre-bloom, bloom, post-bloom and autumn) based around the known importance of pelagic primary production and hydrodynamically active phases of the year. Multivariate analyses were used to identify specific sub-sets of parameters that described the various direct drivers of seafloor-water exchanges. Both dissolved and particulate exchange showed three distinct periods of significant flux during the year, although the specific timings of these periods and the cause-effect relationships to the direct and indirect drivers differed between the two types of flux. Dissolved matter exchange was dominated by an upward flux in the pre-bloom period driven by diffusion, then a biologically induced upward flux during the bloom and an autumn downward flux. The latter was attributable to the interactions of hydrodynamic and biological activity on the seafloor. Particulate matter exchanges exhibited a strongly hydrologically influenced upward flux during the winter, followed by a biologically induced downward flux during the bloom and a second period of downward flux throughout post-bloom and autumn periods. This was driven primarily through interactions between biological activity, and physical and meteorological drivers. The integrated, holistic and quantitative data-based analysis of intra-annual variability in benthic/pelagic fluxes presented in this study in a representative temperate coastal environment, demonstrates not only the various process’ inter-connectivity, but also their relative importance to each other. Future investigations or modelling efforts of similar systems will benefit greatly from the relationships and baseline rules established in this study.
... Increased atmospheric CO 2 concentrations contribute to both a decrease in global ocean pH (i.e. ocean acidification) as well as an increase in ocean temperatures [5][6][7] . Sea surface temperatures have increased on average by 0.6 °C over the past 100 years and models suggest that the deep-sea is warming in some areas at a rate of 0.01-0.1 °C per decade 4,8,9 . ...
Article
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There are numerous studies highlighting the impacts of direct and indirect stressors on marine organisms, and multi-stressor studies of their combined effects are an increasing focus of experimental work. Lophelia pertusa is a framework-forming cold-water coral that supports numerous ecosystem services in the deep ocean. These corals are threatened by increasing anthropogenic impacts to the deep-sea, such as global ocean change and hydrocarbon extraction. This study implemented two sets of experiments to assess the effects of future conditions (temperature: 8 °C and 12 °C, pH: 7.9 and 7.6) and hydrocarbon exposure (oil, dispersant, oil + dispersant combined) on coral health. Phenotypic response was assessed through three independent observations of diagnostic characteristics that were combined into an average health rating at four points during exposure and recovery. In both experiments, regardless of environmental condition, average health significantly declined during 24-hour exposure to dispersant alone but was not significantly altered in the other treatments. In the early recovery stage (24 hours), polyp health returned to the pre-exposure health state under ambient temperature in all treatments. However, increased temperature resulted in a delay in recovery (72 hours) from dispersant exposure. These experiments provide evidence that global ocean change can affect the resilience of corals to environmental stressors and that exposure to chemical dispersants may pose a greater threat than oil itself.
... Using a simple size-spectrum approach based on temperature and primary production (Jennings et al. 2008), an increase of 2°C in temperature (and at the same primary production level) can trigger a 20% decrease in total biomass, but an increase of smaller size fish abundance and biomass . This is consistent with higher trophic and benthic species projected to decrease as a result of warming and ocean acidification in southern areas of the NEA (Queirós et al. 2015;Fernandes et al. 2017;Lotze et al. 2019). The two biogeochemical models show agreement on the main trends and areas of impacts. ...
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Small- and intermediate-size pelagic fisheries are highly impacted by environmental variability and climate change. Their wide geographical distribution and high mobility makes them more likely to shift their distribution under climate change. Here, we explore the potential impact of different climate change scenarios on the four main commercial pelagic species in the North-East Atlantic (NEA): Atlantic mackerel (Scomber scombrus), European sprat (Sprattus sprattus), Atlantic herring (Clupea harengus) and blue whiting (Micromesistius poutassou). We used a process-based fisheries model (SS-DBEM), where all the target species were exploited at their maximum sustainable yield (MSY), to project future potential catches under a high- and low-future-greenhouse-gas scenario (RCP 2.6 and 8.5, respectively). Two ocean biogeochemical models (GDFL and MEDUSA) were used to force the environmental conditions. Mackerel and sprat are projected to have increases in a potential catch under both scenarios. Herring and blue whiting are projected to increase under the RCP2.6, but future projections under RCP8.5 show mixed responses with decreases or no changes forecasted. Overall, the potential catch is projected to increase in the northern area of the NEA but is projected to decrease in the southern area. These projected changes are mainly driven by changes in temperature and primary production. Shifts in the distribution of pelagic resources may destabilize existing international agreements on sharing of straddling resources as exemplified by the dispute in sharing of quota for Atlantic mackerel. Novel climate-ready policy approaches considering full species distribution are needed to complement current stock-based approaches.
... OA-W has a significant impact on marine invertebrates and the ecosystems they inhabit [14][15][16] . Beyond the physical properties, these phenomena alter seawater carbonate chemistry affecting both carbonate secretion by marine calcifiers such as gastropod molluscs [17][18][19] , and its dissolution, with increased vulnerability of younger shells usually composed of more soluble calcium carbonate (CaCO 3 ) polymorphs 20,21 . ...
Article
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Ocean acidification and warming (OA-W) result mainly from the absorption of carbon dioxide and heat by the oceans, altering its physical and chemical properties and affecting carbonate secretion by marine calcifiers such as gastropods. These processes are ongoing, and the projections of their aggravation are not encouraging. This work assesses the concomitant effect of the predicted pH decrease and temperature rise on early life stages of the neogastropod Tritia reticulata (L.), a common scavenger of high ecological importance on coastal ecosystems of the NE Atlantic. Veligers were exposed for 14 days to 12 OA-W experimental scenarios generated by a factorial design of three pH levels (targeting 8.1, 7.8 and 7.5) at four temperatures (16, 18, 20 and 22 °C). Results reveal effects of both pH and temperature (T °C) on larval development, growth, shell integrity and survival, individually or interactively at different exposure times. All endpoints were initially driven by pH, with impaired development and high mortalities being recorded in the first week, constrained by the most acidic scenarios (pHtarget 7.5). Development was also significantly driven by T °C, and its acceleration with warming was observed for the remaining exposure time. Still, by the end of this 2-weeks trial, larval performance and survival were highly affected by the interaction between pH and T °C: growth under warming was evident but only for T °C ≤ 20 °C and carbonate saturation (pHtarget ≥ 7.8). In fact, carbonate undersaturation rendered critical larval mortality (100%) at 22 °C, and the occurrence of extremely vulnerable, unshelled specimens in all other tested temperatures. As recruitment cohorts are the foundation for future populations, our results point towards the extreme vulnerability of this species in case tested scenarios become effective that, according to the IPCC, are projected for the northern hemisphere, where this species is ubiquitous, by the end of the century. Increased veliger mortality associated with reduced growth rates, shell dissolution and loss under OA-W projected scenarios will reduce larval performance, jeopardizing T. reticulata subsistence.
... The effects of global climate change have been extensively documented, revealing impacts to both abiotic and biotic processes that are essential to community structure and ecosystem function (Queirós et al., 2014;Ruckelshaus et al., 2013;Wernberg et al., 2011). Indicators or metrics of physiological state can serve as powerful tools to assess the vulnerability of ecologically important organisms under current and future scenarios (Gunderson et al., 2016). ...
Article
The effects of climate change and associated increases in temperature on organisms are a major focus of scientific research, but how these impacts play out within ecological contexts is complex and hence often ignored. For example, the influence of predation risk (nonconsumptive effects, NCEs) can alter behavior, creating scenarios where individual physiological responses depend on the interaction between abiotic conditions, such as temperature, and the presence of risk in the environment. Yet a mechanistic understanding of how the interplay among abiotic and biotic stressors, especially NCEs, shapes the short-term physiological performance of intertidal organisms remains limited. From both physiological and biochemical perspectives, we explored the short-term interaction between temperature, feeding history, and predation risk from a predatory snail (Nucella lapillus) on the intertidal mussel (Mytilus edulis). We measured heart rate, key aerobic (citrate synthase) and anaerobic (cytosolic malate dehydrogenase) metabolic enzymes, and total antioxidant capacity to elucidate metabolic strategies utilized by mussels in short-term, multi-stressor events. After 60 min of continuous exposure to increased temperature and predation risk, heart rate and aerobic capacity were primarily impacted by temperature, whereas total cytosolic malate dehydrogenase activity displayed an antagonistic relationship in response to the combined effects of feeding history and predation risk. In contrast, total antioxidant capacity displayed a three-way interaction among all treatments (feeding history, temperature and predation risk), driven by opposing thermal responses between fed and starved mussels in the absence of risk. Our results suggest that although mussels are fairly tolerant of acute stress events, the interaction of feeding history and predation risk may prevent them from launching a coordinated stress response when thermal stress is high.
... Multiple shallow-water bivalve species exhibit reduced growth and/or survival under either low oxygen or low pH conditions; together, these stressors often induce additive or synergistic negative effects (Figure 4), although high tolerance may exist in some species or life stages (Gobler and Baumann, 2016). Elevated CO 2 typically raises energetic requirements of invertebrates, inducing greater OC consumption (Thomsen et al., 2013;Queirós et al., 2015); the addition of hypoxia causes metabolic depression, potentially impairing how sediment fauna cope with acidification (Ravaglioli et al., 2019). Studies of the combined effects of reduced oxygen and elevated CO 2 suggest slower microbial degradation and enhanced carbon burial, although additional studies of coupled abiotic stressors involving oxygen are needed (Sampaio et al., 2021). ...
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Benthic animals profoundly influence the cycling and storage of carbon and other elements in marine systems, particularly in coastal sediments. Recent climate change has altered the distribution and abundance of many seafloor taxa and modified the vertical exchange of materials between ocean and sediment layers. Here, we examine how climate change could alter animal-mediated biogeochemical cycling in ocean sediments.The fossil record shows repeated major responses from the benthos during mass extinctions and global carbon perturbations, including reduced diversity, dominance of simple trace fossils, decreased burrow size and bioturbation intensity, and nonrandom extinction of trophic groups. The broad dispersal capacity of many extant benthic species facilitates poleward shifts corresponding to their environmental niche as overlying water warms. Evidence suggests that locally persistent populations will likely respond to environmental shifts through either failure to respond or genetic adaptation rather than via phenotypic plasticity. Regional and global ocean models insufficiently integrate changes in benthic biological activity and their feedbacks on sedimentary biogeochemical processes. The emergence of bioturbation, ventilation, and seafloor-habitat maps and progress in our mechanistic understanding of organism-sediment interactions enable incorporation of potential effects of climate change on benthic macrofaunal mediation of elemental cycles into regional and global ocean biogeochemical models.
... Behavioral alterations at elevated CO 2 have also been demonstrated in a variety of marine invertebrates, including cnidaria, polychaetes, echinoderms, arthropods and mollusks, from a range of environments, including the intertidal zone, coastal and offshore waters and the deep-sea (Clements and Hunt, 2015;Nagelkerken and Munday, 2015;Wang and Wang, 2019). Marine invertebrates exhibit alterations in a range of behaviors at elevated CO 2 , including activity levels (Rosa and Seibel, 2008;Ellis et al., 2009;Spady et al., 2014), feeding rates (Saba et al., 2012;Vargas et al., 2014), settlement and metamorphosis behaviors (Albright et al., 2010;Doropoulos et al., 2012;Guo et al., 2015), burrowing behaviors (Green et al., 2013;Clements and Hunt, 2014), shelter selection (de la Haye et al., 2011), predatory behaviors (behaviors related to finding and eating prey) (Kim et al., 2015;Queirós et al., 2015;Spady et al., 2018) and predator avoidance (Bibby et al., 2007;Manríquez et al., 2013Manríquez et al., , 2014aSpady et al., 2014;Watson et al., 2014). Behavioral categorization is often ambiguous as one behavior may actually include multiple behaviors or decisionmaking processes. ...
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... Population-level variation can be an important source of variation in the responses of organisms to climate change (Barton 2011, Fryxell & Palkovacs 2017, but population-level responses are rarely considered in ocean acidification studies (Munday et al. 2009, Barton et al. 2012, Kroeker et al. 2014, Queirós et al. 2015, Lord et al. 2019. We found population-specific differences in how acute seawater acidification affected Nucella consumption time. ...
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Most research on Ocean Acidification (OA) has largely focused on the process of calcification and the physiological trade-offs employed by calcifying organisms to support the building of calcium carbonate structures. However, there is growing evidence that OA can also impact upon other key biological processes such as survival, growth and behaviour. On wave-swept rocky shores the ability of gastropods to self-right after dislodgement, and rapidly return to normal orientation, reduces the risk of predation. The impacts of OA on this self-righting behaviour and other important parameters such as growth, survival, shell dissolution and shell deposition in Concholepas concholepas (loco) were investigated under contrasting pCO2 levels. Although no impacts of OA on either growth or net shell calcification were found, the results did show that OA can significantly affect self-righting behaviour during the early ontogeny of this species with significantly faster righting times recorded for individuals of C. concholepas reared under increased average pCO2 concentrations (± SE) (716±12 and 1036±14 µatm CO2) compared to those reared at concentrations equivalent to those presently found in the surface ocean (388±8 µatm CO2). When loco were also exposed to the predatory crab Acanthocyclus hassleri, righting times were again increased by exposure to elevated CO2, although self-righting times were generally twice as fast as those observed in the absence of the crab. These results suggest that self-righting in the early ontogeny of C. concholepas will be positively affected by pCO2 levels expected by the end of the 21st century and beginning of the next one. However, as the rate of self-righting is an adaptive trait evolved to reduce lethal predatory attacks, our result also suggest that OA may disrupt prey responses to predators in nature.
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Climate change has already altered the distribution of marine fishes. Future predictions of fish distributions and catches based on bioclimate envelope models are available, but to date they have not considered inter-specific interactions. We address this by combining the species-based Dynamic Bioclimate Envelope Model (DBEM) with a size-based trophic model. The new approach provides spatially and temporally resolved predictions of changes in species' size, abundance and catch potential that account for the effects of ecological interactions. Predicted latitudinal shifts are, on average, reduced by 20% when species interactions are incorporated, compared to DBEM predictions, with pelagic species showing the greatest reductions. Goodness-of-fit to biomass data from fish stock assessments in the North Atlantic between 1991 and 2003 is improved slightly by including species interactions. The differences between predictions from the two models may be relatively modest because, at the North Atlantic basin scale, (1) predators and competitors may respond to climate change together; (2) existing parameterization of the DBEM might implicitly incorporate trophic interactions; and/or (3) trophic interactions might not be the main driver of responses to climate. Future analyses using ecologically-explicit models and data will improve understanding of the effects of inter-specific interactions on responses to climate change, and better inform managers about plausible ecological and fishery consequences of a changing environment. This article is protected by copyright. All rights reserved.
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