Egg production, hatching, faecal pellet production of Calanus finmarchicus in the North Atlantic

The samples were collected using a T-80 net (375 µm mesh size) equipped with a non-filtering cod-end in the North Atlantic during the G.O. Sars Trans-Atlantic cruise in 2013. Within 15-30 minutes after the recovery, 20 Calanus finmarchicus females were sorted out under microscope in ice chilled petri dishes and incubated individually in 600 ml polycarbonate culture bottles resulting in 20 replicate measurements. The bottles were filled with 50 µm screened seawater originated from 6 m water depth. The samples were incubated upright in thermoroom for 24 hours at the surface temperature (3°C). After the samples had been filtered (40 µm filter), female prosome length, egg as well as pellet abundance were determined. Subsequently, eggs from six females were incubated in petri dishes at 5°C. After 4 days, the number of nauplii and eggs were counted in order to calculate hatching success.

Parental exposure to elevated pCO2 influences the reproductive success of copepods

Substantial variations are reported for egg production and hatching rates of copepods exposed to elevated carbon dioxide concentrations (pCO2). One possible explanation, as found in other marine taxa, is that prior parental exposure to elevated pCO2 (and/or decreased pH) affects reproductive performance. Previous studies have adopted two distinct approaches, either (1) expose male and female copepoda to the test pCO2/pH scenarios, or (2) solely expose egg-laying females to the tests. Although the former approach is more realistic, the majority of studies have used the latter approach. Here, we investigated the variation in egg production and hatching success of Acartia tonsa between these two experimental designs, across five different pCO2 concentrations (385-6000 µatm pCO2). In addition, to determine the effect of pCO2 on the hatching success with no prior parental exposure, eggs produced and fertilized under ambient conditions were also exposed to these pCO2 scenarios. Significant variations were found between experimental designs, with approach (1) resulting in higher impacts; here >20% difference was seen in hatching success between experiments at 1000 µatm pCO2 scenarios (2100 year scenario), and >85% at 6000 µatm pCO2. This study highlights the potential to misrepresent the reproductive response of a species to elevated pCO2 dependent on parental exposure.

Seawater carbonate chemistry and egg production, hatching and metabolic rates of a Mediterranean copepod species (Acartia clausi) in a laboratory experiment

This study includes the first information on the combined effect of low pH and raised temperature on egg production rate (EP), hatching success (HS), excretion and respiration of the Mediterranean copepod Acartia clausi. Adult individuals of A. clausi and fresh surface seawater were collected at a coastal station in Saronikos Gulf during April 2012. Four different conditions were applied: two different pH levels (present: 8.09 and future: 7.83) at two temperature values (present: 16°C and present+4 °C= 20°C). EP and HS success decreased significantly over the duration of exposure at future pH at both temperature conditions. However, the analysis of the combined effect of pH, T, chlorophyll a and the duration of the experiments on EP and HS revealed that ocean acidification had no discernible effect, whereas warming; food and the duration of exposure were more significant for the reproductive output of A. clausi. Temperature appeared to have a positive effect on respiration and excretion. Acidification had no clear effect on respiration, but a negative effect on the A. clausi excretion was observed. Acidification and warming resulted in the increase of the excretion rate and the increase was higher than that observed by warming only. Our findings showed that a direct effect of ocean acidification on copepod's vital rates was not obvious, except maybe in the case of excretion. Therefore, the combination of acidification with the ambient oligotrophic conditions and the warming could result in species being less able to allocate resources for coping with multiple stressors.

(Table 1) Reproductive performance of brown skua pairs with and without feeding territories at Potter Peninsula/King George Island from 1998/1999 to 2000/2001

In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.

Lack of evidence for elevated CO2-induced bottom-up effects on marine copepods: a dinoflagellate-calanoid prey-predator pair

Rising levels of atmospheric CO2 are responsible for a change in the carbonate chemistry of seawater with associated pH drops (acidification) projected to reach 0.4 units from 1950 to 2100. We investigated possible indirect effects of seawater acidification on the feeding, fecundity, and hatching success of the calanoid copepod Acartia grani, mediated by potential CO2-induced changes in the nutritional characteristics of their prey. We used as prey the autotrophic dinoflagellate Heterocapsa sp., cultured at three distinct pH levels (control: 8.17, medium: 7.96, and low: 7.75) by bubbling pure CO2 via a computer automated system. Acartia grani adults collected from a laboratory culture were acclimatized for 3 d at food suspensions of Heterocapsa from each pH treatment (ca. 500 cells/ml; 300 ?g C/l). Feeding and egg production rates of the preconditioned females did not differ significantly among the three Heterocapsa diets. Egg hatching success, monitored once per day for the 72 h, did not reveal significant difference among treatments. These results are in agreement with the lack of difference in the cellular stoichiometry (C : N, C : P, and N : P ratios) and fatty acid concentration and composition encountered between the three tested Heterocapsa treatments. Our findings disagree with those of other studies using distinct types of prey, suggesting that this kind of indirect influence of acidification on copepods may be largely associated with interspecific differences among prey items with regard to their sensitivity to elevated CO2 levels.

Contrasting effects of ocean acidification on reproduction in reef fishes

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

PeECE III - Pelagic Ecosystem CO2 Enrichment Study, Raunefjord, Bergen, Norway, 2005

The effects of CO2-induced seawater acidification on plankton communities were also addressed in a series of 3 mesocosm experiments, called the Pelagic Ecosystem CO2 Enrichment (PeECE I-III) studies, which were conducted in the Large-Scale Mesocosm Facilities of the University of Bergen, Norway in 2001, 2003 and 2005, respectively. Each experiment consisted of 9 mesocosms, in which CO2 was manipulated to initial concentrations of 190, 350 and 750 µatm in 2001 and 2003, and 350, 700 and 1050 µatm in 2005. The present dataset concerns PeECE III.

Stickleback F2 generation transgenerational plasticity experiment: egg and body size data

Nongenetic inheritance mechanisms such as transgenerational plasticity (TGP) can buffer populations against rapid environmental change such as ocean warming. Yet, little is known about how long these effects persist and whether they are cumulative over generations. Here, we tested for adaptive TGP in response to simulated ocean warming across parental and grandparental generations of marine sticklebacks. Grandparents were acclimated for two months during reproductive conditioning, whereas parents experienced developmental acclimation, allowing us to compare the fitness consequences of short-term vs. prolonged exposure to elevated temperature across multiple generations. We found that reproductive output of F1 adults was primarily determined by maternal developmental temperature, but carry-over effects from grandparental acclimation environments resulted in cumulative negative effects of elevated temperature on hatching success. In very early stages of growth, F2 offspring reached larger sizes in their respective paternal and grandparental environment down the paternal line, suggesting that other factors than just the paternal genome may be transferred between generations. In later growth stages, maternal and maternal granddam environments strongly influenced offspring body size, but in opposing directions, indicating that the mechanism(s) underlying the transfer of environmental information may have differed between acute and developmental acclimation experienced by the two generations. Taken together, our results suggest that the fitness consequences of parental and grandparental TGP are highly context dependent, but will play an important role in mediating some of the impacts of rapid climate change in this system.

Seawater carbonate chemistry and hatch rates of Antarctic krill

Marine ecosystems of the Southern Ocean are particularly vulnerable to ocean acidification. Antarctic krill (Euphausia superba; hereafter krill) is the key pelagic species of the region and its largest fishery resource. There is therefore concern about the combined effects of climate change, ocean acidification and an expanding fishery on krill and ultimately, their dependent predators-whales, seals and penguins. However, little is known about the sensitivity of krill to ocean acidification. Juvenile and adult krill are already exposed to variable seawater carbonate chemistry because they occupy a range of habitats and migrate both vertically and horizontally on a daily and seasonal basis. Moreover, krill eggs sink from the surface to hatch at 700-1,000 m, where the carbon dioxide partial pressure (pCO2) in sea water is already greater than it is in the atmosphere. Krill eggs sink passively and so cannot avoid these conditions. Here we describe the sensitivity of krill egg hatch rates to increased CO2, and present a circumpolar risk map of krill hatching success under projected pCO2 levels. We find that important krill habitats of the Weddell Sea and the Haakon VII Sea to the east are likely to become high-risk areas for krill recruitment within a century. Furthermore, unless CO2 emissions are mitigated, the Southern Ocean krill population could collapse by 2300 with dire consequences for the entire ecosystem.

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.

Ocean acidification affects the phyto-zoo plankton trophic transfer efficiency

The critical role played by copepods in ocean ecology and biogeochemistry warrants an understanding of how these animals may respond to ocean acidification (OA). Whilst an appreciation of the potential direct effects of OA, due to elevated pCO2, on copepods is improving, little is known about the indirect impacts acting via bottom-up(food quality) effects. We assessed, for the first time, the chronic effects of direct and/or indirect exposures to elevated pCO2 on the behaviour, vital rates, chemical and biochemical stoichiometry of the calanoid copepod Acartia tonsa. Bottom-up effects of elevated pCO2 caused species-specific biochemical changes to the phytoplanktonic feed, which adversely affected copepod population structure and decreased recruitment by 30 %. The direct impact of elevated pCO2 caused gender-specific respiratory responses in A.tonsa adults, stimulating an enhanced respiration rate in males (> 2-fold), and a suppressed respiratory response in females when coupled with indirect elevated pCO2 exposures. Under the combined indirect+direct exposure, carbon trophic transfer efficiency from phytoplankton-to-zooplankton declined to < 50 % of control populations, with a commensurate decrease in recruitment. For the first time an explicit role was demonstrated for biochemical stoichiometry in shaping copepod trophic dynamics. The altered biochemical composition of the CO2-exposed prey affected the biochemical stoichiometry of the copepods, which could have ramifications for production of higher tropic levels, notably fisheries. Our work indicates that the control of phytoplankton and the support of higher trophic levels involving copepods have clear potential to be adversely affected under future OA scenarios.