ALBERT

Description: Anthropogenic carbon dioxide (CO 2 ) emissions simultaneously increase ocean temperatures and reduce ocean surface pH, a process termed ocean acidification (OA). OA is expected to negatively affect the growth and physiology of many calcified organisms, but the response of non-calcified (fleshy) organisms is less well understood. Rising temperatures and p CO 2 can enhance photosynthetic rates (within tolerance limits). Therefore, warming may interact with OA to alter biological responses of macroalgae in complicated ways. Beyond thresholds of physiological tolerance, however, rising temperatures could further exacerbate negative responses to OA. Many studies have investigated the effects of OA or warming independently of each other, but few studies have quantified the interactive effects of OA and warming on marine organisms. We conducted four short-term independent factorial CO 2 enrichment and warming experiments on six common species of calcified and fleshy macroalgae from southern California to investigate the independent and interactive effects of CO 2 and warming on growth, carbonic anhydrase (CA) enzyme activity, pigment concentrations, and photosynthetic efficiency. There was no effect of elevated p CO 2 on CA activity, pigment concentration, and photosynthetic efficiency in the macroalgal species studies. However, we found that calcareous algae suffered reduced growth rates under high p CO 2 conditions alone, although the magnitude of the effect varied by species. Fleshy algae had mixed responses of growth rates to high p CO 2 , indicating that the effects of p CO 2 enrichment are inconsistent across species. The combined effects of elevated p CO 2 and warming had a significantly negative impact on growth for both fleshy and calcareous algae; calcareous algae experienced five times more weight loss than specimens in ambient control conditions and fleshy growth was reduced by 76%. Our results demonstrate the need to study the interactive effects of multiple stressors associated with global change on marine communities.

Description: Ocean acidification (OA) is likely to differentially affect the biology and physiology of calcifying and non-calcifying taxa, thereby potentially altering key ecological interactions (e.g., facilitation, competition, predation) in ways that are difficult to predict from single-species experiments. We used a two-factor experimental design to investigate how multispecies benthic assemblages in southern California kelp forests respond to OA and grazing by the purple sea urchin, Strongylocentrotus purpuratus. Settlement tiles accrued natural mixed assemblages of algae and invertebrates in a kelp forest off San Diego, CA for one year before being exposed to OA and grazing in a laboratory experiment for two months. Space occupying organisms were identified and pooled into six functional groups: calcified invertebrates, non-calcified invertebrates, calcified algae, fleshy algae, sediment, and bare space for subsequent analyses of community structure. Interestingly, communities that developed on separate tile racks were unique, despite being deployed close in space, and further changes in community structure in response to OA and grazing depended on this initial community state. On Rack 1, we found significant effects of both pCO2 and grazing with elevated pCO2 increasing cover of fleshy algae, but sea urchin grazers decreasing cover of fleshy algae. On Rack 2, we found a 35% higher percent cover of sediment on tiles reared in ambient pCO2 but observed 27% higher cover of bare space in the high pCO2 conditions. On Rack 3, we found an average of 45% lower percent cover of calcified sessile invertebrates at ambient pCO2 than in high pCO2 treatments on Rack 3. Net community calcification was 137% lower in elevated pCO2 treatments. Kelp sporophyte densities on tiles without urchins were 74% higher than on tiles with urchins and kelp densities were highest in the elevated pCO2 treatment. Urchin growth and grazing rates were 49% and 126% higher under ambient than high pCO2 conditions. This study highlights consistent negative impacts of OA on community processes such as calcification and grazing rates, even though impacts on community structure were highly context-dependent.

Description: The emergent responses of vulnerable species to global change can vary depending on the relative quality of resources available to support their productivity under increased stress, as well as the biotic interactions with other species that may alter their access to these resources. This research tested how seawater pCO2 may interact with seasonal light availability to affect the photosynthesis and calcification of high-latitude coralline algae, and whether the responses of these calcified macroalgae are modified by physical association with a non-calcified seaweed. Through an in situ approach, our study first investigated how current seasonal environmental variation affects the growth of the understory coralline algae Crusticorallina spp. and Bossiella orbigniana in Southeast Alaska's kelp forests. We then experimentally manipulated pH to simulate end-of-century acidification scenarios, light regime to simulate seasonal light availability at the benthos, and pairings of coralline algal species with and without a fleshy red alga to examine the interactive effects of these variables on coralline productivity and calcification. Our results indicate that: 1) coralline species may face net dissolution under projected future winter pH and carbonate saturation state conditions, 2) differences in seasonal light availability in productive, high-latitude waters may not be distinct enough to modify coralline algal net calcification, and 3) association with a non-calcified red alga does not alter the response of these coralline algal species to ocean acidification scenarios. This research highlights the necessity of incorporating locally informed scenarios of environmental variability and community interactions when predicting species' vulnerability to global change.

Description: Rising anthropogenic CO2 in the atmosphere is accompanied by an increase in oceanic CO2 and a concomitant decline in seawater pH (ref. 1). This phenomenon, known as ocean acidification (OA), has been experimentally shown to impact the biology and ecology of numerous animals and plants2, most notably those that precipitate calcium carbonate skeletons, such as reef-building corals3. Volcanically acidified water at Maug, Commonwealth of the Northern Mariana Islands (CNMI) is equivalent to near-future predictions for what coral reef ecosystems will experience worldwide due to OA. We provide the first chemical and ecological assessment of this unique site and show that acidification-related stress significantly influences the abundance and diversity of coral reef taxa, leading to the often-predicted shift from a coral to an algae-dominated state4, 5. This study provides field evidence that acidification can lead to macroalgae dominance on reefs.

Description: Understanding species' responses to upwelling may be especially important in light of ongoing environmental change. Upwelling frequency and intensity are expected to increase in the future, while ocean acidification and deoxygenation are expected to decrease the pH and dissolved oxygen of upwelled waters. However, the acute effects of a single upwelling event and the integrated effects of multiple upwelling events on marine organisms are poorly understood. Here, we use in situ measurements of pH, temperature, and dissolved oxygen to characterize the covariance of environmental conditions within upwelling-dominated kelp forest ecosystems. We then test the effects of acute (0-3 days) and chronic (1-3 month) upwelling on the performance of two species of kelp forest grazers, the echinoderm, Mesocentrotus franciscanus, and the gastropod, Promartynia pulligo. We exposed organisms to static conditions in a regression design to determine the shape of the relationship between upwelling and performance and provide insights into the potential effects in a variable environment. We found that respiration, grazing, growth, and net calcification decline linearly with increasing upwelling intensity for M. francicanus over both acute and chronic timescales. Promartynia pulligo exhibited decreased respiration, grazing, and net calcification with increased upwelling intensity after chronic exposure, but we did not detect an effect over acute timescales or on growth after chronic exposure. Given the highly correlated nature of pH, temperature, and dissolved oxygen in the California Current, our results suggest the relationship between upwelling intensity and growth in the 3-month trial could potentially be used to estimate growth integrated over long-term dynamic oceanographic conditions for M. franciscanus. Together, these results indicate current exposure to upwelling may reduce species performance and predicted future increases in upwelling frequency and intensity could affect ecosystem function by modifying the ecological roles of key species.

Description: Variation in environmental conditions across a species' range can alter their responses to environmental change through local adaptation and acclimation. Evolutionary responses, however, may be challenged in ecosystems with tightly coupled environmental conditions, where changes in the covariance of environmental factors may make it more difficult for species to adapt to global change. Here, we conduct a 3-month-long mesocosm experiment and find evidence for local adaptation/acclimation in populations of red sea urchins, Mesocentrotus franciscanus, to multiple environmental drivers. Moreover, populations differ in their response to projected concurrent changes in pH, temperature, and dissolved oxygen. Our results highlight the potential for local adaptation/acclimation to multivariate environmental regimes but suggest that thresholds in responses to a single environmental variable, such as temperature, may be more important than changes to environmental covariance. Therefore, identifying physiological thresholds in key environmental drivers may be particularly useful for preserving biodiversity and ecosystem functioning.

Description: It is well understood that differences in the cues used by consumers and their resources in fluctuating environments can give rise to trophic mismatches governing the emergent effects of global change. Trophic mismatches caused by changes in consumer energetics during periods of low resource availability have received far less attention, although this may be common for consumers during winter when primary producers are limited by light. Even less is understood about these dynamics in marine ecosystems, where consumers must cope with energetically costly changes in CO2‐driven carbonate chemistry that will be most pronounced in cold temperatures. This may be especially important for calcified marine herbivores, such as the pinto abalone (Haliotis kamschatkana). H. kamschatkana are of high management concern in the North Pacific due to the active recreational fishery and their importance among traditional cultures, and research suggests they may require more energy to maintain their calcified shells and acid/base balance with ocean acidification. Here we use field surveys to demonstrate seasonal mismatches in the exposure of marine consumers to low pH and algal resource identity during winter in a subpolar, marine ecosystem. We then use these data to test how the effects of exposure to seasonally relevant pH conditions on H. kamschatkana are mediated by seasonal resource identity. We find that exposure to projected future winter pH conditions decreases metabolism and growth, and this effect on growth is pronounced when their diet is limited to the algal species available during winter. Our results suggest that increases in the energetic demands of pinto abalone caused by ocean acidification during winter will be exacerbated by seasonal shifts in their resources. These findings have profound implications for other marine consumers and highlight the importance of considering fluctuations in exposure and resources when inferring the emergent effects of global change.