Interactive effects of ocean acidification and rising sea temperatures alter predation rate and predator selectivity in reef fish communities

Ocean warming and acidification are serious threats to marine life. While each stressor alone has been studied in detail, their combined effects on the outcome of ecological interactions are poorly understood. We measured predation rates and predator selectivity of two closely related species of damselfish exposed to a predatory dottyback. We found temperature and CO2 interacted synergistically on overall predation rate, but antagonistically on predator selectivity. Notably, elevated CO2 or temperature alone reversed predator selectivity, but the interaction between the two stressors cancelled selectivity. Routine metabolic rates of the two prey showed strong species differences in tolerance to CO2 and not temperature, but these differences did not correlate with recorded mortality. This highlights the difficulty of linking species-level physiological tolerance to resulting ecological outcomes. This study is the first to document both synergistic and antagonistic effects of elevated CO2 and temperature on a crucial ecological process like predator-prey dynamics.

The population dynamics of the Galapagos Flightless Cormorant Phalacrocorax harrisi in relation to sea temperature

The Flightless Cormorant Phalacrocorax harrisi is restricted to c. 400 km of the western coastline of the Galápagos archipelago coinciding with the local occurrence of seasonal upwelling of oceanic currents. Individuals frequently make more than one breeding attempt per year, usually change mates, and when juveniles are raised, females desert them to the further care of their mates who complete the rearing alone. Here we report data from a ten-year historical study of a colony stretching c.2 km along the coast-line and representing c. 12% of the total population of the species. The number of clutches laid and juveniles fledged were linked to the occurrence of cold water in off-shore foraging grounds. Most Flightless Cormorants have attachments to local stretches of coastline several hundred metres long. However, a few birds travelled many kilometres, including between colonies, sometimes over open sea. We show that males invest more in nest-building and feeding of the offspring than their mates, and we relate this to the (presumed) in-bred nature of the colony and to male and female reproductive strategies. Our data validate a published demographic model of the species (Valle 1995).

From the river to the open sea - a critical life phase of young Atlantic salmon migrating from the Simojoki river

Long-term monitoring data collected from wild smolts of Atlantic salmon (Salmo salar) in the Simojoki river, northern Finland, were used in studying the relationships between the smolt size and age, smolt and postsmolt migration, environmental conditions and postsmolt survival. The onset of the smolt run was significantly dependent on the rising water temperature and decreasing discharge of the river in the spring. The mean length of smolts migrating early in the season was commonly higher and the mean age always older than among smolts migrating later. Many of the smolts migrating early in the season and almost all smolts migrating later had started their new growth in spring in the river before their sea entry. Among postsmolts, the time required for emigration from the estuary was dependent on the sea surface temperature (SST) off the river, being significantly shorter in years with warm than cold sea temperatures. After leaving the estuary, the postsmolts migrated southwards along the eastern coast of the northern Gulf of Bothnia, the geographical distribution of the tag recoveries coinciding with the warm thermal zone in spring in the coastal area. After arriving in the southern Gulf of Bothnia in late summer the postsmolts mostly migrated near the western coast, reaching the Baltic Main Basin in late autumn. Until the early 1990s there was only a weak positive association between smolt length and postsmolt survival. However, following a subsequent decrease in the mean smolt size, a significant positive dependence was observed between smolt size and the reported recapture rate of tagged salmon. The differences in recapture rates between smolts tagged during the first and second half of the annual migration season were insignificant, indicating that the seasonal variation in smolt size and age seem to be too small to affect survival. Among the climatic factors examined, the summer SST in the Gulf of Bothnia was most clearly related to the survival of the wild postsmolts. Postsmolt survival appeared to be highest in years when the SST in June in the Bothnian Bay varied between 9 and 12 ºC. In addition, the survival of wild postsmolts showed a significant positive dependence on the SST in July in the Bothnian Sea, but not on the abundance of the prey fish (0+ herring, Clupea harengus and sprat, Sprattus sprattus) in the Bothnian Sea and in the Baltic Main Basin. The results suggest, that if the incidence of extreme weather conditions were to increase due to climatic changes, it would probably reduce the postsmolt survival of wild salmon populations. For improving the performance of hatchery-reared smolts, it could be useful to examine opportunities to produce smolts that are in their smolt traits and abilities more similar to the wild smolts described in this thesis.

Diurnal and nocturnal temperatures for Atlantic salmon postsmolts (Salmo salar L.) during their early marine life

Data storage tags (DSTs) were applied to Atlantic salmon (Salmo salar L.) smolts during their seaward migration in the spring of 2002 at a fish counting fence on Campbellton River, Newfoundland. Our objectives were to discover whether or not salmon smolts could carry DSTs and survive, whether or not useful data on thermal habitat could be obtained and interpreted, and whether or not salmon smolts moved vertically in the water column. Data were downloaded from 15 of the recovered tags and revealed the hourly water temperatures experienced by the fish for periods of 3 to 71 days. The data on the DSTs were analyzed for temperature patterns in relation to migration behavior and diurnal movement of the fish. While in the sea, the DSTs recorded night temperatures of 12.5°C, which were higher than day temperatures of 11.6°C; the record from moored recorders, however, indicated that sea temperatures actually declined at night. It is hypothesized that posts-molts avoid avian predators during daylight hours by positioning themselves deeper in the water column and that they were pursuing prey during the deeper vertical descents or ascents noted during the periods of more rapid changes in temperature.

Have jellyfish in the Irish Sea benefited from climate change and overfishing?

Climate change and overfishing may lead to ecosystem instability and may benefit nonexploited organisms such as jellyfish. In the Irish Sea, an increase in jellyfish abundance was evident (r2=0.29, P=0.03) in a 16-year time-series (1994–2009) collected during juvenile fish surveys. Jellyfish abundance correlated positively with sea surface temperature (SST) over the preceding 18 months (r=0.65, pACF<0.001) and copepod biomass in the previous year (r=0.56, pACF=0.03) and negatively with spring (February–May) precipitation (r=−0.57, pACF=0.02). Principal components regression indicated that climatic indices explained 68% of the interannual variability in jellyfish abundance (P=0.003), where the components were based on the North Atlantic Oscillation Index, SST and precipitation. The frequency of cnidarian material present in Continuous Plankton Recorder (CPR) samples has also increased since 1970, with a period of frequent outbreaks between 1982 and 1991. Before this period, the herring stock in the northern Irish Sea declined rapidly to a low level, potentially stimulating structural change in the ecosystem. In 1985, there was a step decrease in CPR copepod biomass and in 1989, a step increase in the phytoplankton colour index, suggesting a cascading regime shift during the 1980s. Subsequent overexploitation of gadids, coupled with warm temperatures and the poor recruitment of cod, led to the rapid decline in cod biomass from 1990. While the biomass of sprat has decreased in the last decade, the herring stock has recovered partially. Reductions in demersal fishing pressure since 2000, intended to stimulate cod recovery, appear to have facilitated further rises in haddock biomass. Since the 1980s regime shift, sea temperatures have increased, the fish community has altered and jellyfish abundance has risen such that jellyfish and haddock may now play an increasingly important role in the ecosystem.

Western Arabian Sea SST estimation

Millennial-scale records of planktonic foraminiferal Mg/Ca, bulk sediment UK37', and planktonic foraminiferal d18O are presented across the last two deglaciations in sediment core NIOP929 from the Arabian Sea. Mg/Ca-derived temperature variability during the penultimate and last deglacial periods falls within the range of modern day Arabian Sea temperatures, which are influenced by monsoon-driven upwelling. The UK37'-derived temperatures in MIS 5e are similar to modern intermonsoon values and are on average 3.5°C higher than the Mg/Ca temperatures in the same period. MIS 5e UK37' and Mg/Ca temperatures are 1.5°C warmer than during the Holocene, while the UK37'-Mg/Ca temperature difference was about twice as large during MIS 5e. This is surprising as, nowadays, both proxy carriers have a very similar seasonal and depth distribution. Partial explanations for the MIS 5e UK37'-Mg/Ca temperature offset include carbonate dissolution, the change in dominant alkenone-producing species, and possibly lateral advection of alkenone-bearing material and a change in seasonal or depth distribution of proxy carriers. Our findings suggest that (1) Mg/Ca of G. ruber documents seawater temperature in the same way during both studied deglaciations as in the present, with respect to, e.g., season and depth, and (2) UK37'-based temperatures from MIS 5 (or older) represent neither upwelling SST nor annual average SST (as it does in the present and the Holocene) but a higher temperature, despite alkenone production mainly occurring in the upwelling season. Further we report that at the onset of the deglacial warming, the Mg/Ca record leads the UK37' record by 4 ka, of which a maximum of 2 ka may be explained by postdepositional processes. Deglacial warming in both temperature records leads the deglacial decrease in the d18O profile, and Mg/Ca-based temperature returns to lower values before d18O has reached minimum interglacial values. This indicates a substantial lead in Arabian Sea warming relative to global ice melting.

Impacts of the Oceans on Climate Change

The oceans play a key role in climate regulation especially in part buffering (neutralising) the effects of increasing levels of greenhouse gases in the atmosphere and rising global temperatures. This chapter examines how the regulatory processes performed by the oceans alter as a response to climate change and assesses the extent to which positive feedbacks from the ocean may exacerbate climate change. There is clear evidence for rapid change in the oceans. As the main heat store for the world there has been an accelerating change in sea temperatures over the last few decades, which has contributed to rising sea‐level. The oceans are also the main store of carbon dioxide (CO2), and are estimated to have taken up ∼40% of anthropogenic-sourced CO2 from the atmosphere since the beginning of the industrial revolution. A proportion of the carbon uptake is exported via the four ocean ‘carbon pumps’ (Solubility, Biological, Continental Shelf and Carbonate Counter) to the deep ocean reservoir. Increases in sea temperature and changing planktonic systems and ocean currents may lead to a reduction in the uptake of CO2 by the ocean; some evidence suggests a suppression of parts of the marine carbon sink is already underway. While the oceans have buffered climate change through the uptake of CO2 produced by fossil fuel burning this has already had an impact on ocean chemistry through ocean acidification and will continue to do so. Feedbacks to climate change from acidification may result from expected impacts on marine organisms (especially corals and calcareous plankton), ecosystems and biogeochemical cycles. The polar regions of the world are showing the most rapid responses to climate change. As a result of a strong ice–ocean influence, small changes in temperature, salinity and ice cover may trigger large and sudden changes in regional climate with potential downstream feedbacks to the climate of the rest of the world. A warming Arctic Ocean may lead to further releases of the potent greenhouse gas methane from hydrates and permafrost. The Southern Ocean plays a critical role in driving, modifying and regulating global climate change via the carbon cycle and through its impact on adjacent Antarctica. The Antarctic Peninsula has shown some of the most rapid rises in atmospheric and oceanic temperature in the world, with an associated retreat of the majority of glaciers. Parts of the West Antarctic ice sheet are deflating rapidly, very likely due to a change in the flux of oceanic heat to the undersides of the floating ice shelves. The final section on modelling feedbacks from the ocean to climate change identifies limitations and priorities for model development and associated observations. Considering the importance of the oceans to climate change and our limited understanding of climate-related ocean processes, our ability to measure the changes that are taking place are conspicuously inadequate. The chapter highlights the need for a comprehensive, adequately funded and globally extensive ocean observing system to be implemented and sustained as a high priority. Unless feedbacks from the oceans to climate change are adequately included in climate change models, it is possible that the mitigation actions needed to stabilise CO2 and limit temperature rise over the next century will be underestimated.

Predicting future thermal habitat suitability of competing native and invasive fish species: from metabolic scope to oceanographic modelling

Global increase in sea temperatures has been suggested to facilitate the incoming and spread of tropical invaders. The increasing success of these species may be related to their higher physiological performance compared with indigenous ones. Here, we determined the effect of temperature on the aerobic metabolic scope (MS) of two herbivorous fish species that occupy a similar ecological niche in the Mediterranean Sea: the native salema (Sarpa salpa) and the invasive marbled spinefoot (Siganus rivulatus). Our results demonstrate a large difference in the optimal temperature for aerobic scope between the salema (21.8°C) and the marbled spinefoot (29.1°C), highlighting the importance of temperature in determining the energy availability and, potentially, the distribution patterns of the two species. A modelling approach based on a present-day projection and a future scenario for oceanographic conditions was used to make predictions about the thermal habitat suitability (THS, an index based on the relationship between MS and temperature) of the two species, both at the basin level (the whole Mediterranean Sea) and at the regional level (the Sicilian Channel, a key area for the inflow of invasive species from the Eastern to the Western Mediterranean Sea). For the present-day projection, our basin-scale model shows higher THS of the marbled spinefoot than the salema in the Eastern compared with the Western Mediterranean Sea. However, by 2050, the THS of the marbled spinefoot is predicted to increase throughout the whole Mediterranean Sea, causing its westward expansion. Nevertheless, the regional-scale model suggests that the future thermal conditions of Western Sicily will remain relatively unsuitable for the invasive species and could act as a barrier for its spread westward. We suggest that metabolic scope can be used as a tool to evaluate the potential invasiveness of alien species and the resilience to global warming of native species.

Marine foundation species shifting ranges and genetic baselines

This thesis revealed the most importance factors shaping the distribution, abundance and genetic diversity of four marine foundation species. Environmental conditions, particularly sea temperatures, nutrient availability and ocean waves, played a primary role in shaping the spatial distribution and abundance of populations, acting on scales varying from tens of meters to hundreds of kilometres. Furthermore, the use of Species Distribution Models (SDMs) with biological records of occurrence and high-resolution oceanographic data, allowed predicting species distributions across time. This approach highlighted the role of climate change, particularly when extreme temperatures prevailed during glacial and interglacial periods. These results, when combined with mtDNA and microsatellite genetic variation of populations allowed inferring for the influence of past range dynamics in the genetic diversity and structure of populations. For instance, the Last Glacial Maximum produced important shifts in species ranges, leaving obvious signatures of higher genetic diversities in regions where populations persisted (i.e., refugia). However, it was found that a species’ genetic pool is shaped by regions of persistence, adjacent to others experiencing expansions and contractions. Contradicting expectations, refugia seem to play a minor role on the re(colonization) process of previously eroded populations. In addition, the available habitat area for expanding populations and the inherent mechanisms of species dispersal in occupying available habitats were also found to be fundamental in shaping the distributions of genetic diversity. However, results suggest that the high levels of genetic diversity in some populations do not rule out that they may have experienced strong genetic erosion in the past, a process here named shifting genetic baselines. Furthermore, this thesis predicted an ongoing retraction at the rear edges and extinctions of unique genetic lineages, which will impoverish the global gene pool, strongly shifting the genetic baselines in the future.

A surface energy perspective on climate change

A surface forcing response framework is developed that enables an understanding of time-dependent climate change from a surface energy perspective. The framework allows the separation of fast responses that are unassociated with global-mean surface air temperature change (ΔT), which is included in the forcing, and slow feedbacks that scale with ΔT. The framework is illustrated primarily using 2 × CO2 climate model experiments and is robust across the models. For CO2 increases, the positive downward radiative component of forcing is smaller at the surface than at the tropopause, and so a rapid reduction in the upward surface latent heat (LH) flux is induced to conserve the tropospheric heat budget; this reduces the precipitation rate. Analysis of the time-dependent surface energy balance over sea and land separately reveals that land areas rapidly regain energy balance, and significant land surface warming occurs before global sea temperatures respond. The 2 × CO2 results are compared to a solar increase experiment and show that some fast responses are forcing dependent. In particular, a significant forcing from the fast hydrological response found in the CO2 experiments is much smaller in the solar experiment. The different fast response explains why previous equilibrium studies found differences in the hydrological sensitivity between these two forcings. On longer time scales, as ΔT increases, the net surface longwave and LH fluxes provide positive and negative surface feedbacks, respectively, while the net surface shortwave and sensible heat fluxes change little. It is found that in contrast to their fast responses, the longer-term response of both surface energy fluxes and the global hydrological cycle are similar for the different forcing agents.

Impact of atmospheric forcing on Antarctic continental shelf water masses

The Antarctic continental shelf seas feature a bimodal distribution of water mass temperature, with the Amundsen and Bellingshausen Seas flooded by Circumpolar Deep Water that is several degrees Celsius warmer than the cold shelf waters prevalent in the Weddell and Ross Seas. This bimodal distribution could be caused by differences in atmospheric forcing, ocean dynamics, ocean and ice feedbacks, or some combination of these factors. In this study, a highly simplified coupled sea ice–mixed layer model is developed to investigate the physical processes controlling this situation. Under regional atmospheric forcings and parameter choices the 10-yr simulations demonstrate a complete destratification of the Weddell Sea water column in winter, forming cold, relatively saline shelf waters, while the Amundsen Sea winter mixed layer remains shallower, allowing a layer of deep warm water to persist. Applying the Weddell atmospheric forcing to the Amundsen Sea model destratifies the water column after two years, and applying the Amundsen forcing to the Weddell Sea model results in a shallower steady-state winter mixed layer that no longer destratifies the water column. This suggests that the regional difference in atmospheric forcings alone is sufficient to account for the bimodal distribution in Antarctic shelf-sea temperatures. The model prediction of mixed layer depth is most sensitive to the air temperature forcing, but a switch in all forcings is required to prevent destratification of the Weddell Sea water column.