Ocean acidification induces multi-generational decline in copepod naupliar production with possible conflict for reproductive resource allocation

Climate change, including ocean acidification (OA), presents fundamental challenges to marine biodiversity and sustained ecosystem health. We determined reproductive response (measured as naupliar production), cuticle composition and stage specific growth of the copepod Tisbe battagliai over three generations at four pH conditions (pH 7.67, 7.82, 7.95, and 8.06). Naupliar production increased significantly at pH 7.95 compared with pH 8.06 followed by a decline at pH 7.82. Naupliar production at pH 7.67 was higher than pH 7.82. We attribute the increase at pH 7.95 to an initial stress response which was succeeded by a hormesis-like response at pH 7.67. A multi-generational modelling approach predicted a gradual decline in naupliar production over the next 100 years (equivalent to approximately 2430 generations). There was a significant growth reduction (mean length integrated across developmental stage) relative to controls. There was a significant increase in the proportion of carbon relative to oxygen within the cuticle as seawater pH decreased. Changes in growth, cuticle composition and naupliar production strongly suggest that copepods subjected to OA-induced stress preferentially reallocate resources towards maintaining reproductive output at the expense of somatic growth and cuticle composition. These responses may drive shifts in life history strategies that favour smaller brood sizes, females and perhaps later maturing females, with the potential to profoundly destabilise marine trophodynamics.

The photo-physiological response of a model cnidarian-dinoflagellate symbiosis to CO2-induced acidification at the cellular level

We measured the relationship between CO2-induced seawater acidification, photo-physiological performance and intracellular pH (pHi) in a model cnidarian-dinoflagellate symbiosis - the sea anemone Aiptasia sp. -under ambient (289.94 ± 12.54 µatm), intermediate (687.40 ± 25.10 µatm) and high (1459.92 ± 65.51 µatm) CO2 conditions. These treatments represented current CO2 levels, in addition to CO2 stabilisation scenarios IV and VI provided by the Intergovernmental Panel on Climate Change (IPCC). Anemones were exposed to each treatment for two months and sampled at regular intervals. At each time-point we measured a series of physiological responses: maximum dark-adapted fluorescent yield of PSII (Fv/Fm), gross photosynthetic rate, respiration rate, symbiont population density, and light-adapted pHi of both the dinoflagellate symbiont and isolated host anemone cell. We observed increases in all but one photo-physiological parameter (Pgross:R ratio). At the cellular level, increases in light-adapted symbiont pHi were observed under both intermediate and high CO2 treatments, relative to control conditions (pHi 7.35 and 7.46 versus pHi 7.25, respectively). The response of light-adapted host pHi was more complex, however, with no change observed under the intermediate CO2 treatment, but a 0.3 pH-unit increase under the high CO2 treatment (pHi 7.19 and 7.48, respectively). This difference is likely a result of a disproportionate increase in photosynthesis relative to respiration at the higher CO2 concentration. Our results suggest that, rather than causing cellular acidosis, the addition of CO2 will enhance photosynthetic performance, enabling both the symbiont and host cell to withstand predicted ocean acidification scenarios.

Ingestion and clearance rates of Copepods

Feeding activity, selective grazing and the potential grazing impact of two dominant grazers of the Polar Frontal Zone, Calanus simillimus and Rhincalanus gigas, and of copepods < 2 mm were investigated with incubation experiments in the course of an iron fertilized diatom bloom in November 2000. All grazers were already actively feeding in the low chlorophyll waters prior to the onset of the bloom. C. simillimus maintained constant clearance rates and fed predominantly on diatoms. R. gigas and the small copepods strongly increased clearance and ingestion of diatoms in response to their enhanced availability. All grazers preyed on microzooplankton, most steadily on ciliates, confirming the view that pure herbivory appears to be the exception rather than the rule in copepod feeding. The grazers exhibited differences in feeding behavior based on selectivity indices. C. simillimus and R. gigas showed prey switching from dinoflagellates to diatoms in response to the phytoplankton bloom. All grazers most efficiently grazed on large diatoms leading to differences in daily losses for large and small species, e.g. Corethron sp. or Thalassionema nitzschioides. Species-specific diatom mortality rates due to grazing suggest that the high feeding activity of C. simillimus prior to and during the bloom played a role in shaping diatom population dynamics

Adaptive variability to low-pH river discharges in Acartia tonsa and stress responses to high PCO2 conditions

Environmental transitions leading to spatial physical-chemical gradients are of ecological and evolutionary interest because they are able to induce variations in phenotypic plasticity. Thus, the adaptive variability to low-pH river discharges may drive divergent stress responses [ingestion rates (IR) and expression of stress-related genes such as Heat shock protein 70 (Hsp70) and Ferritin] in the neritic copepod Acartia tonsa facing changes in the marine chemistry associated to ocean acidification (OA). These responses were tested in copepod populations inhabiting two environments with contrasting carbonate system parameters (an estuarine versus coastal area) in the Southern Pacific Ocean, and assessing an in situ and 96-h experimental incubation under conditions of high pressure of CO2 (PCO2 1200 ppm). Adaptive variability was a determining factor in driving variability of copepods' responses. Thus, the food-rich but colder and corrosive estuary induced a traits trade-off expressed as depressed IR under in situ conditions. However, this experience allowed these copepods to tolerate further exposure to high PCO2 levels better, as their IRs were on average 43% higher than those of the coastal individuals. Indeed, expression of both the Hsp70 and Ferritin genes in coastal copepods was significantly higher after acclimation to high PCO2 conditions. Along with other recent evidence, our findings confirm that adaptation to local fluctuations in seawater pH seems to play a significant role in the response of planktonic populations to OA-associated conditions. Facing the environmental threat represented by the inter-play between multiple drivers of climate change, this biological feature should be examined in detail as a potential tool for risk mitigation policies in coastal management arrangements.

Ingestion rate and egg production of copepods collected in the upper 20m in the eastern Mediterranean Sea in March and April 2008 during SES_GR1

The ingestion on ciliates and phytoplankton dataset is based on samples taken during April 2008 in Northern Aegean Sea, the area influenced by the Black Sea water outflow. A Lagrangian experiment was established and copepod ingestion was estimated from experiments performed at stations according to the different positions of drifters during the cruise. Copepods for the experiments were obtained with slow non-quantitative tows from the upper 20 m layer of the water column using 200 µm mesh size nets fitted with a large non-filtering cod end. For the grazing experiments we used the following copepod species: Centropages typicus and Calanus helgolandicus according to the relevant reference (Bamstedt et al. 2000). Copepod clearance rates on ciliates were calculated according to Frost equations (Frost 1972). Ingestion rates were calculated by multiplying clearance rates by the initial standing stocks (Bamstedt et al. 2000). The egg production dataset is based on samples taken during April 2008 in Northern Aegean Sea, the area influenced by the Black Sea water outflow. A Lagrangian experiment was established and copepod egg production was estimated from experiments performed at stations according to the different positions of drifters during the cruise. Egg production rates of the dominant calanoid copepods were determined by incubation of fertilised females (eggs female/day) collected in the 0-20m layer. Copepod egg production was measured for the copepods Centropages typicus, Calanus helgolandicus. On board experiments for the estimation of copepod egg production were taken place. For the estimation of copepod production (mgC/ m**2 /day), lengths (copepods and eggs) were converted to body carbon (Hopcroft et al., 1998) and production was estimated from biomass and weight-specific egg production rates, by assuming that those rates are representative for juvenile specific growth rates (Berggreen et al., 1988).

Ingestion rate and egg production of copepods collected in the upper 20m in the eastern Mediterranean Sea in Oktober 2008

This dataset based on samples taken during October 2008 in Dardanelles Straits, Marmara Sea and Bosporus Straits at the third priority stations. Copepods for the experiments were obtained with slow non-quantitative tows from the upper 50 m layer of the water column using 200 µm mesh size nets fitted with a large non-filtering cod end. For the grazing experiments we used the following copepod species: Oithona spp., Clausocalanus furcatus, Acartia clausi and Oncaea spp. and in one cladoceran species Penilia avirostris according to the relevant reference (Bamstedt et al. 2000). Copepod clearance rates on ciliates were calculated according to Frost equations (Frost 1972). Ingestion rates were calculated by multiplying clearance rates by the initial standing stocks (Bamstedt et al. 2000). Egg production rates of the dominant calanoid copepods were determined by incubation of fertilised females (eggs/female/day) collected in the 0-20m layer. Copepod egg production was measured for the copepods Clausocalanus furcatus, Paracalanus parvus,Acaria clausi. On board experiments for the estimation of copepod egg production were taken place. For the estimation of copepod production (mg/m**2/day), lengths (copepods and eggs) were converted to body carbon (Hopcroft et al., 1998) and production was estimated from biomass and weight-specific egg production rates, by assuming that those rates are representative for juvenile specific growth rates (Berggreen et al., 1988).

Ingestion rate and egg production of copepods collected in the upper 100m in the eastern Mediterranean Sea in March and April 2008 during SES_GR1

The phytoplankton dataset is based on samples taken during March-April 2008 in Libyan Sea, Southern Aegean Sea and Northern Aegean Sea. Ingestion rates were estimated from experiments performed at all the third priority stations during the cruise according to DoW of Sesame project. Copepods for the experiments were obtained with slow non-quantitative tows from the upper 100 m layer of the water column using 200 µm mesh size nets fitted with a large non-filtering cod end. For the grazing experiments we used the following copepod species: Calanus helgolandicus and Centropages typicus according to the relevant reference (Bamstedt et al. 2000). Copepod clearance rates on ciliates were calculated according to Frost equations (Frost 1972). Ingestion rates were calculated by multiplying clearance rates by the initial standing stocks (Bamstedt et al. 2000). Egg production rates of the dominant calanoid copepods were determined by incubation of fertilised females (eggs/female/day) collected in the 0-100m layer. Copepod egg production was measured for the copepods Eucalanus monachus, Centropages typicus and Calanus helgolandicus. On board experiments for the estimation of copepod egg production were taken place. For the estimation of copepod production (mg/m**2/day), lengths (copepods and eggs) were converted to body carbon (Hopcroft et al., 1998) and production was estimated from biomass and weight-specific egg production rates, by assuming that those rates are representative for juvenile specific growth rates (Berggreen et al., 1988).

Station characteristics and copepod fatty alcohol, fatty acid and lipid composition during MSM02/4 in Fram Strait

The biodiversity of pelagic deep-sea ecosystems has received growing scientific interest in the last decade, especially in the framework of international marine biodiversity initiatives, such as Census of Marine Life (CoML). While a growing number of deep-sea zooplankton species has been identified and genetically characterized, little information is available on the mechanisms minimizing inter-specific competition and thus allowing closely related species to co-occur in the deep-sea pelagic realm. Focussing on the two dominant calanoid copepod families Euchaetidae and Aetideidae in Fram Strait, Arctic Ocean, the present study strives to characterize ecological niches of co-occurring species, with regard to vertical distribution, dietary composition as derived from lipid biomarkers, and trophic level on the basis of stable isotope signatures. Closely related species were usually restricted to different depth layers, resulting in a multi-layered vertical distribution pattern. Thus, vertical partitioning was an important mechanism to avoid inter-specific competition. Species occurring in the same depth strata usually belonged to different genera. They differed in fatty acid composition and trophic level, indicating different food preferences. Herbivorous Calanus represent major prey items for many omnivorous and carnivorous species throughout the water column. The seasonal and ontogenetic vertical migration of Calanus acts as a short-cut in food supply for pelagic deep-sea ecosystems in the Arctic.

Mesozooplankton abundance data from Disko Bay, West Greenland, 2008

The study site was located in the Disko Bay off Qeqertarsuaq, western Greenland. Due to land-connected sea ice coverage during winter, 2 sampling sites were combined. At the first site in winter (21 February to 23 March 2008), sampling was conducted through a hole in the ice at ca. 65 to 160 m depth approximately 0.5 nautical mile (n mile) south of Qeqertarsuaq (69° 14' N, 53° 29' W). In spring and summer (9 April to 18 July), sampling was done at a monitoring station 1 n mile south from Qeqertarsuaq (69° 14' N, 53° 23' W) at 300 m depth. Sampling was carried out between 10:00 and 17:00 h. During sampling from the ice, mesozooplankton was collected using a modified WP-2 net (45 µm) equipped with a closing mechanism (Hydrobios). Samples were collected in 3 depth strata (0-50, 50-100, and 100-150 m). During ship-based sampling, mesozooplankton was collected with a multinet (50 µm) equipped with a flow meter (Multinet, Hydrobios type midi), and 2 additional depth strata (150-200m and 200-250 m) were included. In addition to the seasonal study one diurnal investigation with sampling every 6 h was conducted from 29 April at 12:00 h to 30 April 30 at 12:00 h. Samples were immediately preserved in buffered formalin (5% final concentration) for later analyses. Biomass values of the different copepod species were calculated based on measurements of prosome length, and length/weight relationships. Two regressions for Calanus spp. were established for biomass calculations: one applicable prior to and during the phytoplankton bloom until 4 May, and another from 9 May onwards.