Population structure, growth and production of the surf clam Donax hanleyanus (Bivalvia: Donacidae) from northern Argentinean beaches

The surf clams Mesodesma mactroides Reeve, 1854 and Donax hanleyanus Philippi, 1847 are the two dominating species in macrobenthic communities of sandy beaches off northern Argentina, with the latter now surpassing M. mactroides populations in abundance and biomass. Before stock decimation caused by exploitation (during the 1940s and 1950s) and mass mortality events (1995, 1999 and 2007) M. mactroides was the prominent primary consumer in the intertidal ecosystem and an important economic resource in Argentina. Since D. hanleyanus was not commercially fished and not affected by mass mortality events, it took over as the dominant species, but did never reach the former abundance of M. mactroides. Currently abundance and biomass of both surf clams are a multiple smaller than those of forty years ago, indicating the conservation status of D. hanleyanus and M. mactroides as endangered. Therefore the aim of this study is to analyse the population dynamics (population structure, growth and reproductive biology) of D. hanleyanus and M. mactroides, and to compare the results with historical data in order to detect possible differences within surf clam populations forty years ago and at present.

Growth increments of the recent brachiopod Magellania venosa mechanically marked in Paso Comau and Comau Fjord, Chile, 2011/2012

Magellania venosa, the largest recent brachiopod, occurs in clusters and banks in population densities of up to 416 ind/m**2 in Comau Fjord, Northern Chilean fjord region. Below 15 m, it co-occurs with the mytilid Aulacomya atra and it dominates the benthic community below 20 m. To determine the question of why M. venosa is a successful competitor, the in situ growth rate of the brachiopod was studied and its overall growth performance compared with that of other brachiopods and mussels. The growth in length was measured between February 2011 and March 2012 after mechanical tagging and calcein staining. Settlement and juvenile growth were determined from recruitment tiles installed in 2009 and from subsequent photocensus. Growth of M. venosa is best described by the general von Bertalanffy growth function, with a maximum shell length (Linf) of 71.53 mm and a Brody growth constant (K) of 0.336/year. The overall growth performance (OGP index = 5.1) is the highest recorded for a rynchonelliform brachiopod and in the range of that for Mytilus chilensis (4.8-5.27), but lower than that of A. atra (5.74). The maximal individual production (PInd) is 0.29 g AFDM/ind/year at 42 mm shell length and annual production ranges from 1.28 to 89.25 g AFDM/year/m**2 (1-57% of that of A. atra in the respective fjords). The high shell growth rate of M. venosa, together with its high overall growth performance may explain the locally high population density of this brachiopod in Comau Fjord. However, the production per biomass of the population (P/B-ratio) is low (0.535) and M. venosa may play only a minor role in the food chain. Settling dynamics indicates that M. venosa is a pioneer species with low juvenile mortality. The coexistence of the brachiopod and bivalve suggests that brachiopod survival is affected by neither the presence of potential brachiopod predators nor that of space competitors (i.e. mytilids).

Ocean acidification alleviates low-temperature effects on growth and photosynthesis of the red alga Neosiphonia harveyi (Rhodophyta)

This study aimed to examine interactive effects between ocean acidification and temperature on the photosynthetic and growth performance of Neosiphonia harveyi. N. harveyi was cultivated at 10 and 17.5 °C at present (~380 µatm), expected future (~800 µatm), and high (~1500 µatm) pCO2. Chlorophyll a fluorescence, net photosynthesis, and growth were measured. The state of the carbon-concentrating mechanism (CCM) was examined by pH-drift experiments (with algae cultivated at 10 °C only) using ethoxyzolamide, an inhibitor of external and internal carbonic anhydrases (exCA and intCA, respectively). Furthermore, the inhibitory effect of acetazolamide (an inhibitor of exCA) and Tris (an inhibitor of the acidification of the diffusive boundary layer) on net photosynthesis was measured at both temperatures. Temperature affected photosynthesis (in terms of photosynthetic efficiency, light saturation point, and net photosynthesis) and growth at present pCO2, but these effects decreased with increasing pCO2. The relevance of the CCM decreased at 10 °C. A pCO2 effect on the CCM could only be shown if intCA and exCA were inhibited. The experiments demonstrate for the first time interactions between ocean acidification and temperature on the performance of a non-calcifying macroalga and show that the effects of low temperature on photosynthesis can be alleviated by increasing pCO2. The findings indicate that the carbon acquisition mediated by exCA and acidification of the diffusive boundary layer decrease at low temperatures but are not affected by the cultivation level of pCO2, whereas the activity of intCA is affected by pCO2. Ecologically, the findings suggest that ocean acidification might affect the biogeographical distribution of N. harveyi.

Juvenile sea stars exposed to acidification decrease feeding and growth with no acclimation potential

Ocean acidification has the potential to affect growth and calcification of benthic marine invertebrates, particularly during their early life history. We exposed field-collected juveniles of Asterias rubens from Kiel Fjord (western Baltic Sea) to 3 seawater CO2 partial pressure (pCO2) levels (ranging from around 650 to 3500 µatm) in a long-term (39 wk) and a short-term (6 wk) experiment. In both experiments, survival and calcification were not affected by elevated pCO2. However, feeding rates decreased strongly with increasing pCO2, while aerobic metabolism and NH4+ excretion were not significantly affected by CO2 exposure. Consequently, high pCO2 reduced the scope for growth in A. rubens. Growth rates decreased substantially with increasing pCO2 and were reduced even at pCO2 levels occurring in the habitat today (e.g. during upwelling events). Sea stars were not able to acclimate to higher pCO2, and growth performance did not recover during the long-term experiment. Therefore, the top-down control exerted by this keystone species may be diminished during periods of high environmental pCO2 that already occur occasionally and will be even higher in the future. However, some individuals were able to grow at high rates even at high pCO2, indicating potential for rapid adaption. The selection of adapted specimens of A. rubens in this seasonally acidified habitat may lead to higher CO2 tolerance in adult sea stars of this population compared to the juvenile stage. Future studies need to address the synergistic effects of multiple stressors such as acidification, warming and reduced salinity, which will simultaneously impact the performance of sea stars in this habitat.

Experiment: Combined effects of CO2, temperature, irradiance and time on the physiological performance of Chondrus crispus (Rhodophyta)

In natural environments, marine biotas are exposed to a variety of simultaneously acting abiotic factors. Among these, temperature, irradiance and CO2 availability are major factors influencing the physiological performance of marine macroalgae. To test whether elevated levels of CO2 may remediate the otherwise reduced performance of uncalcified seaweeds under the influence of other stressful abiotic factors, we performed multifactorial experiments with the red alga Chondrus crispus from Helgoland (North Sea) with two levels of CO2, temperature and irradiance: low and high pCO2 levels were tested in combination with either (1) optimal and low irradiances or (2) optimal and sub-lethal high temperatures for growth. Performance of C. crispus was evaluated as biomass increase and relative growth rates (RGR), gross photosynthesis and pigment content. Acclimations of growth and photosynthesis were measured after 4 and 8 days. Acclimation time was crucial for elucidating single or combined CO2 effects on growth and photosynthesis. Signifi- cant CO2 effects became evident only in combination with either elevated temperature or reduced irradiance. Growth and photosynthesis had divergent patterns: RGR and biomass significantly increased only under a combination of high pCO2 and elevated temperature; gross photosynthesis was significantly reduced under high pCO2 conditions at low irradiance. Pigment content varied in response to irradiance and temperature, but was independent of pCO2.

Water temperature measured in Independence Bay, Pisco, Peru

Independencia Bay can be considered as one of the most productive invertebratc fishing grounds worldwide. One of the most important exploited species is the scallop (Argopecten purpuratus) with strong catching fluctuations related to El Nino and La Nina events and to inadequate Management strategies. During strong warming periods annual landings reach up to 50000 t in an area of about 150 km**2 and during cold years they remain around 500 to 1000 t. This study analyses the changes in scallop landings at Independencia Bay observed during the last two decades and discusses the main factors affecting the scallop proliferations during the EI Nino events. In this way data on landings, sea surface temperature and those related to growth, reproduction, predation, mean density and oxygen concentration from published and unpublished Papers are used. The relationship between annual catches and average water temperature over the preceding reproductive period of the scallop over the past 20 year's period, showed that scallop production is affected positively only with strong EI Nino such as those of 1983 and 1998. Our review showed that the scallop stock proliferation can be traced to the combined effect of (1) an increase in reproductive output through an acceleration of gonad maturation and a higher spawning frequency; (2) a shortening of the larval period and an increase in larval survival; (3) an increase in the individual growth performance; (4) an increase in the juvenile and adult survival through reduction of predator biomass; (5) an increase in carrying capacity of the scallop banks due to elevated oxygen levels.

(Table 1) Egg size, clutch asymmetry, hatch date and breeding success in south polar skua, brown skua and mixed pairs on King George Island

We studied how environmental conditions affect reproduction in sympatric skua species that differ in their reliance on marine resources: the exclusively marine foraging south polar skua Catharacta maccormicki, the terrestrially foraging brown skua C. antarctica lonnbergi and mixed species pairs with an intermediate diet. Egg size, clutch asymmetry and hatching dates varied between species and years without consistent patterns. In the south polar skuas, 12 to 38% of the variation in these parameters was explained by sea surface temperature, sea ice cover and local weather. In mixed species pairs and brown skuas, the influence of environmental factors on variation in clutch asymmetry and hatching date decreased to 10-29%, and no effect on egg size was found. Annual variation in offspring growth performance also differed between species with variable growth in chicks of south polar skuas and mixed species pairs, and almost uniform growth in brown skuas. Additionally, the dependency on oceanographic and climatic factors, especially local wind conditions, decreased from south polar skuas to brown skua chicks. Consistent in all species, offspring were more sensitive to environmental conditions during early stages; during the late chick stage (>33 d) chick growth was almost independent of environmental conditions. The net breeding success could not be predicted by any environmental factor in any skua species, suggesting it may not be a sensitive indicator of environmental conditions. Hence, the sensitivity of skuas to environmental conditions varied between species, with south polar skuas being more sensitive than brown skuas, and between breeding periods, with the egg parameters being more susceptible to oceanographic conditions. However, during offspring development, local climatic conditions became more important. We conclude that future climate change in the Maritime Antarctic will affect reproduction of skuas more strongly through changes in sea ice cover and sea surface temperature (and the resulting alterations to the marine food web) than through local weather conditions.

Habitat characteristics and energy budget of the ocean quahog (Arctica islandica)

We compared lifetime and population energy budgets of the extraordinary long-lived ocean quahog Arctica islandica from 6 different sites - the Norwegian coast, Kattegat, Kiel Bay, White Sea, German Bight, and off northeast Iceland - covering a temperature and salinity gradient of 4-10°C (annual mean) and 25-34, respectively. Based on von Bertalanffy growth models and size-mass relationships, we computed organic matter production of body (PSB) and of shell (PSS), whereas gonad production (PG) was estimated from the seasonal cycle in mass. Respiration (R) was computed by a model driven by body mass, temperature, and site. A. islandica populations differed distinctly in maximum life span (40 y in Kiel Bay to 197 y in Iceland), but less in growth performance (phi' ranged from 2.41 in the White Sea to 2.65 in Kattegat). Individual lifetime energy throughput, as approximated by assimilation, was highest in Iceland (43,730 kJ) and lowest in the White Sea (313 kJ). Net growth efficiency ranged between 0.251 and 0.348, whereas lifetime energy investment distinctly shifted from somatic to gonad production with increasing life span; PS/PG decreased from 0.362 (Kiel Bay, 40 y) to 0.031 (Iceland, 197 y). Population annual energy budgets were derived from individual budgets and estimates of population mortality rate (0.035/y in Iceland to 0.173/y in Kiel Bay). Relationships between budget ratios were similar on the population level, albeit with more emphasis on somatic production; PS/ PG ranged from 0.196 (Iceland) to 2.728 (White Sea), and P/B ranged from 0.203-0.285/y. Life span is the principal determinant of the relationship between budget parameters, whereas temperature affects net growth efficiency only. In the White Sea population, both growth performance and net growth efficiency of A. islandica were lowest. We presume that low temperature combined with low salinity represent a particularly stressful environment for this species.

Time series of seawater carbonate chemistry calculated throughout incubation periods of Boreogadus saida and Gadus morhua during exposure to different CO2 and temperature conditions

Oceans are experiencing increasing acidification in parallel to a distinct warming trend in consequence of ongoing climate change. Rising seawater temperatures are mediating a northward shift in distribution of Atlantic cod (Gadus morhua), into the habitat of polar cod (Boreogadus saida), that is associated with retreating cold water masses. This study investigates the competitive strength of the co-occurring gadoids under ocean acidification and warming (OAW) scenarios. Therefore, we incubated specimens of both species in individual tanks for 4 months, under different control and projected temperatures (polar cod: 0, 3, 6, 8 °C, Atlantic cod: 3, 8, 12, 16 °C) and PCO2 conditions (390 and 1170 µatm) and monitored growth, feed consumption and standard metabolic rate. Our results revealed distinct temperature effects on both species. While hypercapnia by itself had no effect, combined drivers caused nonsignificant trends. The feed conversion efficiency of normocapnic polar cod was highest at 0 °C, while optimum growth performance was attained at 6 °C; the long-term upper thermal tolerance limit was reached at 8 °C. OAW caused only slight impairments in growth performance. Under normocapnic conditions, Atlantic cod consumed progressively increasing amounts of feed than individuals under hypercapnia despite maintaining similar growth rates during warming. The low feed conversion efficiency at 3 °C may relate to the lower thermal limit of Atlantic cod. In conclusion, Atlantic cod displayed increased performance in the warming Arctic such that the competitive strength of polar cod is expected to decrease under future OAW conditions.

Light-modulated responses of growth and photosynthetic performance to ocean acidification in the model diatom Phaeodactylum tricornutum

Ocean acidification (OA) due to atmospheric CO2 rise is expected to influence marine primary productivity. In order to investigate the interactive effects of OA and light changes on diatoms, we grew Phaeodactylum tricornutum, under ambient (390 ppmv; LC) and elevated CO2 (1000 ppmv; HC) conditions for 80 generations, and measured its physiological performance under different light levels (60 µmol/m**2/s, LL; 200 µmol/m**2/s, ML; 460 µmol/m**2/s, HL) for another 25 generations. The specific growth rate of the HC-grown cells was higher (about 12-18%) than that of the LC-grown ones, with the highest under the ML level. With increasing light levels, the effective photochemical yield of PSII (Fv'/Fm') decreased, but was enhanced by the elevated CO2, especially under the HL level. The cells acclimated to the HC condition showed a higher recovery rate of their photochemical yield of PSII compared to the LC-grown cells. For the HC-grown cells, dissolved inorganic carbon or CO2 levels for half saturation of photosynthesis (K1/2 DIC or K1/2 CO2) increased by 11, 55 and 32%, under the LL, ML and HL levels, reflecting a light dependent down-regulation of carbon concentrating mechanisms (CCMs). The linkage between higher level of the CCMs down-regulation and higher growth rate at ML under OA supports the theory that the saved energy from CCMs down-regulation adds on to enhance the growth of the diatom.

Effects of ocean acidification on the photosynthetic performance, carbonic anhydrase activity and growth of the giant kelp Macrocystis pyrifera

Under ocean acidification (OA), the 200 % increase in CO2(aq) and the reduction of pH by 0.3-0.4 units are predicted to affect the carbon physiology and growth of macroalgae. Here we examined how the physiology of the giant kelp Macrocystis pyrifera is affected by elevated pCO2/low pH. Growth and photosynthetic rates, external and internal carbonic anhydrase (CA) activity, HCO3 (-) versus CO2 use were determined over a 7-day incubation at ambient pCO2 400 µatm/pH 8.00 and a future OA treatment of pCO2 1200 µatm/pH 7.59. Neither the photosynthetic nor growth rates were changed by elevated CO2 supply in the OA treatment. These results were explained by the greater use of HCO3 (-) compared to CO2 as an inorganic carbon (Ci) source to support photosynthesis. Macrocystis is a mixed HCO3 (-) and CO2 user that exhibits two effective mechanisms for HCO3 (-) utilization; as predicted for species that possess carbon-concentrating mechanisms (CCMs), photosynthesis was not substantially affected by elevated pCO2. The internal CA activity was also unaffected by OA, and it remained high and active throughout the experiment; this suggests that HCO3 (-) uptake via an anion exchange protein was not affected by OA. Our results suggest that photosynthetic Ci uptake and growth of Macrocystis will not be affected by elevated pCO2/low pH predicted for the future, but the combined effects with other environmental factors like temperature and nutrient availability could change the physiological response of Macrocystis to OA. Therefore, further studies will be important to elucidate how this species might respond to the global environmental change predicted for the ocean.

Environmental characteristics, and growth traits and leaf chemistry of tundra plants in a warming experiment at Alexandra Fiord

Understanding plant trait responses to elevated temperatures in the Arctic is critical in light of recent and continuing climate change, especially because these traits act as key mechanisms in climate-vegetation feedbacks. Since 1992, we have artificially warmed three plant communities at Alexandra Fiord, Nunavut, Canada (79°N). In each of the communities, we used open-top chambers (OTCs) to passively warm vegetation by 1-2 °C. In the summer of 2008, we investigated the intraspecific trait responses of five key species to 16 years of continuous warming. We examined eight traits that quantify different aspects of plant performance: leaf size, specific leaf area (SLA), leaf dry matter content (LDMC), plant height, leaf carbon concentration, leaf nitrogen concentration, leaf carbon isotope discrimination (LCID), and leaf d15N. Long-term artificial warming affected five traits, including at least one trait in every species studied. The evergreen shrub Cassiope tetragona responded most frequently (increased leaf size and plant height/decreased SLA, leaf carbon concentration, and LCID), followed by the deciduous shrub Salix arctica (increased leaf size and plant height/decreased SLA) and the evergreen shrub Dryas integrifolia (increased leaf size and plant height/decreased LCID), the forb Oxyria digyna (increased leaf size and plant height), and the sedge Eriophorum angustifolium spp. triste (decreased leaf carbon concentration). Warming did not affect d15N, leaf nitrogen concentration, or LDMC. Overall, growth traits were more sensitive to warming than leaf chemistry traits. Notably, we found that responses to warming were sustained, even after many years of treatment. Our work suggests that tundra plants in the High Arctic will show a multifaceted response to warming, often including taller shoots with larger leaves.

Seawater carbonate chemistry and maximum growth rates of Skeletonema marinoi and Alexandrium ostenfeldii, toxin composition of Alexandrium ostenfeldii in a laboratory experiment

Phytoplankton populations can display high levels of genetic diversity that, when reflected by phenotypic variability, may stabilize a species response to environmental changes. We studied the effects of increased temperature and CO2 availability as predicted consequences of global change, on 16 genetically different isolates of the diatom Skeletonema marinoi from the Adriatic Sea and the Skagerrak (North Sea), and on eight strains of the PST (paralytic shellfish toxin)-producing dinoflagellate Alexandrium ostenfeldii from the Baltic Sea. Maximum growth rates were estimated in batch cultures of acclimated isolates grown for five to 10 generations in a factorial design at 20 and 24 °C, and present day and next century applied atmospheric pCO2, respectively. In both species, individual strains were affected in different ways by increased temperature and pCO2. The strongest response variability, buffering overall effects, was detected among Adriatic S. marinoi strains. Skagerrak strains showed a more uniform response, particularly to increased temperature, with an overall positive effect on growth. Increased temperature also caused a general growth stimulation in A. ostenfeldii, despite notable variability in strain-specific response patterns. Our data revealed a significant relationship between strain-specific growth rates and the impact of pCO2 on growth-slow growing cultures were generally positively affected, while fast growing cultures showed no or negative responses to increased pCO2. Toxin composition of A. ostenfeldii was consistently altered by elevated temperature and increased CO2 supply in the tested strains, resulting in overall promotion of saxitoxin production by both treatments. Our findings suggest that phenotypic variability within populations plays an important role in the adaptation of phytoplankton to changing environments, potentially attenuating short-term effects and forming the basis for selection. In particular, A. ostenfeldii blooms may expand and increase in toxicity under increased water temperature and atmospheric pCO2 conditions, with potentially severe consequences for the coastal ecosystem.

Seawater carbonate chemistry, the abiotic conditions in the fluid surrounding the embryo, growth, calcification of the cuttlefish Sepia officinalis in a laboratory experiment

This study investigated the effects of seawater pH (i.e., 8.10, 7.85 and 7.60) and temperature (16 and 19 °C) on (a) the abiotic conditions in the fluid surrounding the embryo (viz. the perivitelline fluid), (b) growth, development and (c) cuttlebone calcification of embryonic and juvenile stages of the cephalopod Sepia officinalis. Egg swelling increased in response to acidification or warming, leading to an increase in egg surface while the interactive effects suggested a limited plasticity of the swelling modulation. Embryos experienced elevated pCO2 conditions in the perivitelline fluid (>3-fold higher pCO2 than that of ambient seawater), rendering the medium under-saturated even under ambient conditions. The growth of both embryos and juveniles was unaffected by pH, whereas 45Ca incorporation in cuttlebone increased significantly with decreasing pH at both temperatures. This phenomenon of hypercalcification is limited to only a number of animals but does not guarantee functional performance and calls for better mechanistic understanding of calcification processes.