Seawater carbonate chemistry and reproduction of the two calanoid copepods Centropages typicus and Temora longicornis in a laboratory experiment

Some planktonic groups suffer negative effects from ocean acidification (OA), although copepods might be less sensitive. We investigated the effect of predicted CO2 levels (range 480-750 ppm), on egg production and hatching success of two copepod species, Centropages typicus and Temora longicornis. In these short-term incubations there was no significant effect of high CO2 on these parameters. Additionally a very high CO2 treatment, (CO2 = 9830 ppm), representative of carbon capture and storage scenarios, resulted in a reduction of egg production rate and hatching success of C. typicus, but not T. longicornis. In conclusion, reproduction of C. typicus was more sensitive to acute elevated seawater CO2 than that of T. longicornis, but neither species was affected by exposure to CO2 levels predicted for the year 2100. The duration and seasonal timing of exposures to high pCO2, however, might have a significant effect on the reproduction success of calanoid copepods.

Seawater carbonate chemistry and reproduction of a coral reef fish in a laboratory experiment

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

Environment and habitat characteristics of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.

Characteristics of eggs and female adults of Calanus finmarchicus, C. glacialis and C. hyperboreus from Disko Bay, western Greenland

This is the first high temporal-resolution study in Disko Bay covering population dynamics, grazing, reproduction, and biochemical composition of 3 dominating copepod species (Calanus finmarchicus, C. glacialis and C. hyperboreus) from late winter to midsummer in 2008. C. finmarchicus and C. glacialis ascended to the surface layer at the onset of the spring phytoplankton bloom, followed by C. hyperboreus 2 wk later. C. finmarchicus spawning occurred during the bloom and postbloom period, partially fueled by wax esters. C. glacialis commenced spawning before the bloom, yet it was greatly stimulated when food became available. However, feeding and reproduction was terminated after the main bloom despite the presence of food. In terms of feeding, this was also the strategy for C. hyperboreus. Between pre-bloom and post-bloom, C. finmarchicus showed an increase in carbon, nitrogen, and phospholipid content but a decrease in total lipid content. This was likely the result of protein synthesis, oocyte maturation, and spawning fueled by wax esters and by feeding. C. glacialis showed a similar pattern, although with an increasing total lipid content from pre-bloom to post-bloom, and an increasing wax ester and decreasing phospholipid content after reproduction was terminated. C. hyperboreus showed greatly increased content of carbon, nitrogen, and all lipid classes between the pre- and post-bloom periods. Hence, C. finmarchicus commenced feeding and spawning at the onset of the bloom and continued throughout the remaining study period. Both C. glacialis and C. hyperboreus females refueled their storage lipids (wax esters) during the bloom and post-bloom period, suggesting that they may spawn in an additional year.

Seasonal fatty acid and fatty alcohol composition of Pseudocalanus minutus and Oithona similis in Kongsfjorden (Svalbard)

Seasonal lipid dynamics of various developmental stages were investigated in Pseudocalanus minutus and Oithona similis. For P. minutus, the dominance of 16:1(n?7), 16:4(n?3) and 20:5(n?3) fatty acids indicated a diatom-based nutrition in spring, whereas 22:6(n?3), 16:0, 18:2(n?6) and 18:1(n?9) pointed to a flagellate-based diet during the rest of the year as well as omnivorous/carnivorous low-level feeding during winter. The shorter-chain fatty alcohols 14:0 and 16:0 prevailed, also reflecting biosynthetic processes typical of omnivores or carnivores. Altogether, the lipid signatures characterized P. minutus as an opportunistic feeder. In contrast, O. similis had consistently high amounts of the 18:1(n?9) fatty acid in all stages and during all seasons pointing to a generally omnivorous/carnivorous/detritivorous diet. Furthermore, the fatty alcohol 20:1(n?9) reached high percentages especially in adult females and males, and feeding on Calanus faecal pellets is suggested. Fatty alcohols, as wax ester moieties, revealed significant seasonal variations in O. similis and a seasonal trend towards wax ester accumulation in autumn in P. minutus. P. minutus utilized its lipid deposits for development in the copepodite stages III and IV and for gonad maturation in CV and females during the dark season. However, CVs and females depended on the spring phytoplankton bloom for final maturation processes and reproduction. O. similis fueled gonad maturation and egg production for reproduction in June by wax esters, whereas reproduction in August/September co-occurred with the accumulation of new depot lipids. Both species revealed significantly higher wax ester levels in deeper (>50 m) as compared to surface (0-50 m) dwelling individuals related to a descent prior to overwintering.

Effects of ocean acidification on population dynamics and community structure of crustose coralline algae

Calcification and growth of crustose coralline algae (CCA) are affected by elevated seawater pCO2 and associated changes in carbonate chemistry. However, the effects of ocean acidification (OA) on population and community-level responses of CCA have barely been investigated. We explored changes in community structure and population dynamics (size structure and reproduction) of CCA in response to OA. Recruited from an experimental flow-through system, CCA settled onto the walls of plastic aquaria and developed under exposure to one of three pCO2 treatments (control [present day, 389±6 ppm CO2], medium [753±11 ppm], and high [1267±19 ppm]). Elevated pCO2 reduced total CCA abundance and affected community structure, in particular the density of the dominant species Pneophyllum sp. and Porolithon onkodes. Meanwhile, the relative abundance of P. onkodes declined from 24% under control CO2 to 8.3% in high CO2 (65% change), while the relative abundance of Pneophyllum sp. remained constant. Population size structure of P. onkodes differed significantly across treatments, with fewer larger individuals under high CO2. In contrast, the population size structure and number of reproductive structures (conceptacles) per crust of Pneophyllum sp. was similar across treatments. The difference in the magnitude of the response of species abundance and population size structure between species may have the potential to induce species composition changes in the future. These results demonstrate that the impacts of OA on key coral reef builders go beyond declines in calcification and growth, and suggest important changes to aspects of population dynamics and community ecology.

Effect of simulated ocean acidification on the acute toxicity of Cu and Cd to Tigriopus japonicus

Heavy metals pollution in marine environments has caused great damage to marine biological and ecological systems. Heavy metals accumulate in marine creatures, after which they are delivered to higher trophic levels of marine organisms through the marine food chain, which causes serious harm to marine biological systems and human health. Additionally, excess carbon dioxide in the atmosphere has caused ocean acidification. Indeed, about one third of the CO2 released into the atmosphere by anthropogenic activities since the beginning of the industrial revolution has been absorbed by the world's oceans, which play a key role in moderating climate change. Modeling has shown that, if current trends in CO2 emissions continue, the average pH of the ocean will reach 7.8 by the end of this century, corresponding to 0.5 units below the pre-industrial level, or a three-fold increase in H+ concentration. The ocean pH has not been at this level for several millions of years. Additionally, these changes are occurring at speeds 100 times greater than ever previously observed. As a result, several marine species, communities and ecosystems might not have time to acclimate or adapt to these fast changes in ocean chemistry. In addition, decreasing ocean pH has the potential to seriously affect the growth, development and reproduction reproductive processes of marine organisms, as well as threaten normal development of the marine ecosystem. Copepods are an important part of the meiofauna that play an important role in the marine ecosystem. Pollution of the marine environment can influence their growth and development, as well as the ecological processes they are involved in. Accordingly, there is important scientific value to investigation of the response of copepods to ocean acidification and heavy metals pollution. In the present study, we evaluated the effects of simulated future ocean acidification and the toxicological interaction between ocean acidity and heavy metals of Cu and Cd on T. japonicus. To accomplish this, harpacticoids were exposed to Cu and Cd concentration gradient seawater that had been equilibrated with CO2 and air to reach pH 8.0, 7.7, 7.3 and 6.5 for 96 h. Survival was not significantly suppressed under single sea water acidification, and the final survival rates were greater than 93% in both the experimental groups and the controls. The toxicity of Cu to T. japonicus was significantly affected by sea water acidification, with the 96h LC50 decreasing by nearly threefold from 1.98 to 0.64 mg/L with decreasing pH. The 96 h LC50 of Cd decreased with decreasing pH, but there was no significant difference in mortality among pH treatments. The results of the present study demonstrated that the predicted future ocean acidification has the potential to negatively affect survival of T. japonicus by exacerbating the toxicity of Cu. The calculated safe concentrations of Cu were 11.9 (pH 7.7) and 10.5 (pH 7.3) µg/L, which were below the class I value and very close to the class II level of the China National Quality Standard for Sea Water. Overall, these results indicate that the Chinese coastal sea will face a

Increased feeding and nutrient excretion of adult antarctic krill, Euphausia superba, exposed to enhanced carbon dioxide (CO2)

Ocean acidification has a wide-ranging potential for impacting the physiology and metabolism of zooplankton. Sufficiently elevated CO2 concentrations can alter internal acid-base balance, compromising homeostatic regulation and disrupting internal systems ranging from oxygen transport to ion balance. We assessed feeding and nutrient excretion rates in natural populations of the keystone species Euphausia superba (Antarctic krill) by conducting a CO2 perturbation experiment at ambient and elevated atmospheric CO2 levels in January 2011 along the West Antarctic Peninsula (WAP). Under elevated CO2 conditions (~672 ppm), ingestion rates of krill averaged 78 µg C/individual/d and were 3.5 times higher than krill ingestion rates at ambient, present day CO2 concentrations. Additionally, rates of ammonium, phosphate, and dissolved organic carbon (DOC) excretion by krill were 1.5, 1.5, and 3.0 times higher, respectively, in the high CO2 treatment than at ambient CO2 concentrations. Excretion of urea, however, was ~17% lower in the high CO2 treatment, suggesting differences in catabolic processes of krill between treatments. Activities of key metabolic enzymes, malate dehydrogenase (MDH) and lactate dehydrogenase (LDH), were consistently higher in the high CO2 treatment. The observed shifts in metabolism are consistent with increased physiological costs associated with regulating internal acid-base equilibria. This represents an additional stress that may hamper growth and reproduction, which would negatively impact an already declining krill population along the WAP.

Temperature and acidification variability reduce physiological performance in the intertidal zone porcelain crab Petrolisthes//cinctipes

We show here that increased variability of temperature and pH synergistically negatively affects the energetics of intertidal zone crabs. Under future climate scenarios, coastal ecosystems are projected to have increased extremes of low tide-associated thermal stress and ocean acidification-associated low pH, the individual or interactive effects of which have yet to be determined. To characterize energetic consequences of exposure to increased variability of pH and temperature, we exposed porcelain crabs, Petrolisthes cinctipes, to conditions that simulated current and future intertidal zone thermal and pH environments. During the daily low tide, specimens were exposed to no, moderate or extreme heating, and during the daily high tide experienced no, moderate or extreme acidification. Respiration rate and cardiac thermal limits were assessed following 2.5 weeks of acclimation. Thermal variation had a larger overall effect than pH variation, though there was an interactive effect between the two environmental drivers. Under the most extreme temperature and pH combination, respiration rate decreased while heat tolerance increased, indicating a smaller overall aerobic energy budget (i.e. a reduced O2 consumption rate) of which a larger portion is devoted to basal maintenance (i.e. greater thermal tolerance indicating induction of the cellular stress response). These results suggest the potential for negative long-term ecological consequences for intertidal ectotherms exposed to increased extremes in pH and temperature due to reduced energy for behavior and reproduction.

Diatom abundance and fatty acid composition of ice algae and of Calanus glacialis in samples obtained in 2008 in the eastern Beaufort Sea

The copepod Calanus glacialis plays a key role in the lipid-based energy flux in Arctic shelf seas. By utilizing both ice algae and phytoplankton, this species is able to extend its growth season considerably in these seasonally ice-covered seas. This study investigated the impacts of the variability in timing and extent of the ice algal bloom on the reproduction and population success of C. glacialis. The vertical distribution, reproduction, amount of storage lipids, stable isotopes, fatty acid and fatty alcohol composition of C. glacialis were assessed during the Circumpolar Flaw Lead System Study. Data were collected in the Amundsen Gulf, south-eastern Beaufort Sea, from January to July 2008 with the core-sampling from March to April. The reduction in sea ice thickness and coverage observed in the Amundsen Gulf in 2007 and 2008 affected the life strategy and reproduction of C. glacialis. Developmental stages CIII and CIV dominated the overwintering population, which resulted in the presence of very few CV and females during spring 2008. Spawning began at the peak of the ice algal bloom that preceded the precocious May ice break-up. Although the main recruitment may have occurred later in the season, low abundance of females combined with a potential mismatch between egg production/development to the first feeding stage and phytoplankton bloom resulted in low recruitment of C. glacialis in the early summer of 2008.

Data compilation of marine pelagic organism biomasses, swimming velocities, clearance and respiration rates

Jellyfishes have functionally replaced several overexploited commercial stocks of planktivorous fishes. This is paradoxical, because they use a primitive prey capture mechanism requiring direct contact with the prey, whereas fishes use more efficient visual detection. We have compiled published data to show that, in spite of their primitive life-style, jellyfishes exhibit similar instantaneous prey clearance and respiration rates as their fish competitors and similar potential for growth and reproduction. To achieve this production, they have evolved large, water-laden bodies that increase prey contact rates. Although larger bodies are less efficient for swimming, optimization analysis reveals that large collectors are advantageous if they move through the water sufficiently slowly.

Follow-up of the sponge and asteroid populations at the station Larsen A south (PS69/724-1 and PS77/253-1), Antarctic Peninsula, between 2007 and 2011

Over 30% of the Antarctic continental shelf is permanently covered by floating ice shelves, providing aphotic conditions for a depauperate fauna sustained by laterally advected food. In much of the remaining Antarctic shallows (<300 m depth), seasonal sea-ice melting allows a patchy primary production supporting rich megabenthic communities dominated by glass sponges (Porifera, Hexactinellida). The catastrophic collapse of ice shelves due to rapid regional warming along the Antarctic Peninsula in recent decades has exposed over 23,000 km**2 of seafloor to local primary production. The response of the benthos to this unprecedented flux of food is, however, still unknown. In 2007, 12 years after disintegration of the Larsen A ice shelf, a first biological survey interpreted the presence of hexactinellids as remnants of a former under-ice fauna with deep-sea characteristics. Four years later, we revisited the original transect, finding 2- and 3-fold increases in glass sponge biomass and abundance, respectively, after only two favorable growth periods. Our findings, along with other long-term studies, suggest that Antarctic hexactinellids, locked in arrested growth for decades, may undergo boom-and-bust cycles, allowing them to quickly colonize new habitats. The cues triggering growth and reproduction in Antarctic glass sponges remain enigmatic.

Seawater carbonate chemistry and Celleporella hyalina biological processes during experiments, 2011

Increased anthropogenic CO2 emissions in the last two centuries have lead to rising sea surface temperature and falling ocean pH, and it is predicted that current global trends will worsen over the next few decades. There is limited understanding of how genetic variation among individuals will influence the responses of populations and species to these changes. A microcosm system was set up to study the effects of predicted temperature and CO2 levels on the bryozoan Celleporella hyalina. In this marine species, colonies grow by the addition of male, female and feeding modular individuals (zooids) and can be physically subdivided to produce a clone of genetically identical colonies. We studied colony growth rate (the addition of zooids), reproductive investment (the ratio of sexual to feeding zooids) and sex ratio (male to female zooids) in four genetically distinct clonal lines. There was a significant effect of clone on growth rate, reproductive investment and sex ratio, with clones showing contrasting responses to the various temperature and pH combinations. Overall, decreasing pH and increasing temperature caused reduction of growth, and eventual cessation of growth was often observed at the highest temperature, especially during the latter half of the 15-day trials. Reproductive investment increased with increasing temperature and decreasing pH, varying more widely with temperature at the lowest pH. The increased production of males, a general stress response of the bryozoan, was seen upon exposure to reduced pH, but was not expressed at the highest temperature tested, presumably due to the frequent cessation of growth. Further to the significant effect of pH on the measured whole-colony parameters, observation by scanning electron microscopy revealed surface pitting of the calcified exoskeleton in colonies that were exposed to increased acidity. Studying ecologically relevant processes of growth and reproduction, we demonstrate the existence of relevant levels of variation among genetic individuals which may enable future adaptation via non-mutational natural selection to falling pH and rising temperature.

Seawater carbonate chemistry and severe tissue damage in Atlantic cod larvae under increasing ocean acidification, 2012

Ocean acidification, caused by increasing atmospheric concentrations of CO2, is one of the most critical anthropogenicthreats to marine life. Changes in seawater carbonate chemistry have the potential to disturb calcification, acid-base regulation, blood circulation and respiration, as well as the nervous system of marine organisms, leading to long-term effects such as reduced growth rates and reproduction. In teleost fishes, early life-history stages are particularly vulnerable as they lack specialized internal pH regulatory mechanisms. So far, impacts of relevant CO2concentrations on larval fish have been found in behaviour and otolith size, mainly in tropical, non-commercial species. Here we show detrimental effects of ocean acidification on the development of a mass-spawning fish species of high commercial importance. We reared Atlantic cod larvae at three levels of CO2, (1) present day, (2) end of next century and (3) an extreme, coastal upwelling scenario, in a long-term ( 2.5 1/2 months) mesocosm experiment. Exposure to CO2 resulted in severe to lethal tissue damage in many internal organs, with the degree of damage increasing with CO2 concentration. As larval survival is the bottleneck to recruitment, ocean acidification has the potential to act as an additional source of natural mortality, affecting populations of already exploited fish stocks.