Seawater acidification affects the physiological energetics and spawning capacity of the Manila clam Ruditapes philippinarum during gonadal maturation

Ocean acidification is predicted to have widespread implications for marine bivalve mollusks. While our understanding of its impact on their physiological and behavioral responses is increasing, little is known about their reproductive responses under future scenarios of anthropogenic climate change. In this study, we examined the physiological energetics of the Manila clam Ruditapes philippinarum exposed to CO2-induced seawater acidification during gonadal maturation. Three recirculating systems filled with 600 L of seawater were manipulated to three pH levels (8.0, 7.7, and 7.4) corresponding to control and projected pH levels for 2100 and 2300. In each system, temperature was gradually increased ca. 0.3 °C per day from 10 to 20 °C for 30 days and maintained at 20 °C for the following 40 days. Irrespective of seawater pH levels, clearance rate (CR), respiration rate (RR), ammonia excretion rate (ER), and scope for growth (SFG) increased after a 30-day stepwise warming protocol. When seawater pH was reduced, CR, ratio of oxygen to nitrogen, and SFG significantly decreased concurrently, whereas ammonia ER increased. RR was virtually unaffected under acidified conditions. Neither temperature nor acidification showed a significant effect on food absorption efficiency. Our findings indicate that energy is allocated away from reproduction under reduced seawater pH, potentially resulting in an impaired or suppressed reproductive function. This interpretation is based on the fact that spawning was induced in only 56% of the clams grown at pH 7.4. Seawater acidification can therefore potentially impair the physiological energetics and spawning capacity of R. philippinarum.

Seawater carbonate chemistry and biological processes during experiments with early life stages of the blue mussel Mytilus edulis, 2010

Several experiments have shown a decrease of growth and calcification of organisms at decreased pH levels. There is a growing interest to focus on early life stages that are believed to be more sensitive to environmental disturbances such as hypercapnia. Here, we present experimental data, acquired in a commercial hatchery, demonstrating that the growth of planktonic mussel (Mytilus edulis) larvae is significantly affected by a decrease of pH to a level expected for the end of the century. Even though there was no significant effect of a 0.25-0.34 pH unit decrease on hatching and mortality rates during the first 2 days of development nor during the following 13-day period prior to settlement, final shells were respectively 4.5±1.3 and 6.0±2.3% smaller at pHNBS~7.8 (pCO2~1100-1200 µatm) than at a control pHNBS of ~8.1 (pCO2~460-640 µatm). Moreover, a decrease of 12.0±5.4% of shell thickness was observed after 15d of development. More severe impacts were found with a decrease of ~0.5 pHNBS unit during the first 2 days of development which could be attributed to a decrease of calcification due to a slight undersaturation of seawater with respect to aragonite. Indeed, important effects on both hatching and D-veliger shell growth were found. Hatching rates were 24±4% lower while D-veliger shells were 12.7±0.9% smaller at pHNBS~7.6 (pCO2~1900 µatm) than at a control pHNBS of ~8.1 (pCO2~540 µatm). Although these results show that blue mussel larvae are still able to develop a shell in seawater undersaturated with respect to aragonite, the observed decreases of hatching rates and shell growth could lead to a significant decrease of the settlement success. As the environmental conditions considered in this study do not necessarily reflect the natural conditions experienced by this species at the time of spawning, future studies will need to consider the whole larval cycle (from fertilization to settlement) under environmentally relevant conditions in order to investigate the potential ecological and economical losses of a decrease of this species fitness in the field.

Seawater carbonate chemistry and Pacific oyster (Crassostrea gigas) biological processes during experiments, 2011

Ocean acidification, due to anthropogenic CO2 absorption by the ocean, may have profound impacts on marine biota. Calcareous organisms are expected to be particularly sensitive due to the decreasing availability of carbonate ions driven by decreasing pH levels. Recently, some studies focused on the early life stages of mollusks that are supposedly more sensitive to environmental disturbances than adult stages. Although these studies have shown decreased growth rates and increased proportions of abnormal development under low pH conditions, they did not allow attribution to pH induced changes in physiology or changes due to a decrease in aragonite saturation state. This study aims to assess the impact of several carbonate-system perturbations on the growth of Pacific oyster (Crassostrea gigas) larvae during the first 3 days of development (until shelled D-veliger larvae). Seawater with five different chemistries was obtained by separately manipulating pH, total alkalinity and aragonite saturation state (calcium addition). Results showed that the developmental success and growth rates were not directly affected by changes in pH or aragonite saturation state but were highly correlated with the availability of carbonate ions. In contrast to previous studies, both developmental success into viable D-shaped larvae and growth rates were not significantly altered as long as carbonate ion concentrations were above aragonite saturation levels, but they strongly decreased below saturation levels. These results suggest that the mechanisms used by these organisms to regulate calcification rates are not efficient enough to compensate for the low availability of carbonate ions under corrosive conditions.

Synergistic effects of ocean acidification and warming on overwintering pteropods in the Arctic

Ocean acidification and warming will be most pronounced in the Arctic Ocean. Aragonite shell-bearing pteropods in the Arctic are expected to be among the first species to suffer from ocean acidification. Carbonate undersaturation in the Arctic will first occur in winter and because this period is also characterized by low food availability, the overwintering stages of polar pteropods may develop into a bottleneck in their life cycle. The impacts of ocean acidification and warming on growth, shell degradation (dissolution), and mortality of two thecosome pteropods, the polar Limacina helicina and the boreal L. retroversa, were studied for the first time during the Arctic winter in the Kongsfjord (Svalbard). The abundance of L. helicina and L. retroversa varied from 23.5 to 120 ind /m2 and 12 to 38 ind /m2, and the mean shell size ranged from 920 to 981 µm and 810 to 823 µm, respectively. Seawater was aragonite-undersaturated at the overwintering depths of pteropods on two out of ten days of our observations. A 7-day experiment [temperature levels: 2 and 7 °C, pCO2 levels: 350, 650 (only for L. helicina) and 880 ?atm] revealed a significant pCO2 effect on shell degradation in both species, and synergistic effects between temperature and pCO2 for L. helicina. A comparison of live and dead specimens kept under the same experimental conditions indicated that both species were capable of actively reducing the impacts of acidification on shell dissolution. A higher vulnerability to increasing pCO2 and temperature during the winter season is indicated compared with a similar study from fall 2009. Considering the species winter phenology and the seasonal changes in carbonate chemistry in Arctic waters, negative climate change effects on Arctic thecosomes are likely to show up first during winter, possibly well before ocean acidification effects become detectable during the summer season.

Data compilation on the biological response to ocean acidification: environmental and experimental context of data sets and related literature

The exponential growth of studies on the biological response to ocean acidification over the last few decades has generated a large amount of data. To facilitate data comparison, a data compilation hosted at the data publisher PANGAEA was initiated in 2008 and is updated on a regular basis (doi:10.1594/PANGAEA.149999). By January 2015, a total of 581 data sets (over 4 000 000 data points) from 539 papers had been archived. Here we present the developments of this data compilation five years since its first description by Nisumaa et al. (2010). Most of study sites from which data archived are still in the Northern Hemisphere and the number of archived data from studies from the Southern Hemisphere and polar oceans are still relatively low. Data from 60 studies that investigated the response of a mix of organisms or natural communities were all added after 2010, indicating a welcomed shift from the study of individual organisms to communities and ecosystems. The initial imbalance of considerably more data archived on calcification and primary production than on other processes has improved. There is also a clear tendency towards more data archived from multifactorial studies after 2010. For easier and more effective access to ocean acidification data, the ocean acidification community is strongly encouraged to contribute to the data archiving effort, and help develop standard vocabularies describing the variables and define best practices for archiving ocean acidification data.

Avaliação da variação sedimentológica em uma área da plataforma continental sob a atuação da indústria petrolífera

Petroleum exploration activity occurs on the offshore Potiguar Basin, from very shallow (2-3 m) until about 50 m water depth, extending from Alto de Touros (RN) to Alto de Fortaleza (CE). Take in account the biological importance and the heterogeneity of sediments on this area, it is necessary the understanding of the sedimentological dynamics, and mainly the changes generated by petroleum exploration to prevent possible damages to environment. Despite the intense activity of oil exploration in this area, research projects like these are still rare. In view to minimize this gap, this study was developed to evaluate sedimentological, mineralogical and geochemical changes in the vicinity of a exploration well, here designated as well A, located on the Middle continental shelf, near the transition to Outer shelf. The well selected for this study was the first one drilled with Riserless Mud Recovery technology (RMR) in Brazil. The main difference from this to the conventional method is the possibility of drilling phase I of the well with return of drilling material to the rig tank, minimizing fluid and gravel discharging around the vicinity, during this phase. Monitoring consisted of three surveys, first of them done before start drilling, the second one done 19 days after the end of drilling and the third one done one year after then. Comparison of the studied variables (calcium carbonate and organic matter content, sediment size, mineralogy and geochemistry) was done with their average, median and coefficient of variation values to understand the changes after drilling activity. Because operating company technical reasons, the well location was changed after the first survey (C1), resulting in a shift of the sampled area on the two last surveys (C2 e C3). Nevertheless, the acquired data presented a good correlation, with no loss to the mean goal of the study. The sedimentological, mineralogical and geochemical analyzes were done at Federal University of Rio Grande do Norte (UFRN). The results indicated a predominantly sandy environment along the three surveys. It was noticed that the first survey (C1), presented different values for all the studied variables than to the second (C2) and third (C3) surveys, which had similar values. Siliciclastic sediments are prevalent at all surveys, and quartz is the main component (more than 80%). Heavy minerals (garnet, turmaline, zircon and lmenite), rock fragments and mud aggregates also was described. Bioclastic sediments are dominated by coralline algae (more than 45%) and mollusks (more than 30%), followed by benthic foraminifera, bryozoans and worm tubes. More rarely was observed ostracoda and spike of calcareous sponge. Because the low changes of the sediments at the studied area and by the using of RMR method in the drilling, it was possible to conclude that drilling activity did not promote significant alteration on the local sediment cover. Changes in the studied variables before and after drilling activity could be influenced by the changing in the sampling area after survey 1 (C1).

Pesca artesanal entre mudanças socioambientais: estudo de caso na APA Bonfim-Guaraíra/RN-Brasil

Artisanal fishing is one of the important economic activities of Rio Grande do Norte (RN), becoming the main form of survival and income for many families. The southern coast of RN has strong fishing tradition, both in exploitation of fish as in the collection of mollusks on the banks of Guaraíra Lagoon (APA Bonfim-Guaraíra). In recent years, fishing communities have become more vulnerable to outside influences, since the socio-economic and environmental changes are exposed. These changes require adjustments to the fishermen to different environmental conditions, as the social and natural systems are driven by reorganization and upgrading processes. The living conditions in fishing communities are objective and necessary in different ways, according to a selection that is cultural, so adaptable. This study has the general objective to analyze the main environmental changes in fishing communities, based on the understanding of fishermen and mollusk fisherwomen about this problem. The specific objectives searched to: describe the socioeconomic profile, prospects and difficulties of fishermen and mollusk fisherwomen; investigate the major social and environmental changes in the fishing communities of Patané and Camocim (Arez/RN). The methodology was qualitative-quantitative with the techniques of observation in the field and open and semistructured interviews applied to representatives of Cologne fishing, fishermen and mollusk fisherwomen. Between the months of January to July 2014 were applied semistructured interviews with 41 interlocutors in Patané and 23 of Camocim; totaling 64 people. The results indicated that the interlocutors are aged 40 to 60 years; have only completed elementary school level; are married and have children; survive on less than a minimum wage income and they are not receiving employment insurance. The case study also revealed that the socioeconomic and environmental problems reflected in changes in reproduction, organization and social division of labor, which generates adaptive changes of families to external influences, resulting directly from other economic activities and market pressure and indirectly from tourism. Therefore, adaptations to changes scenario presented as favorable to economic aspects and unfavorable to the social and environmental aspects. In short, the fishery develops on technical, bureaucratic and financial obstacles, in the everyday needs of men and women is the determining factor in willingness to continue exercising fishing and shell fishing on Guaraíra Lagoon, space representing relations social, practices and customs transmitted in the course of local history through social memory of the oldest to the youngest.

Seawater carbonate chemistry, calcification and dissolution response, and skeletal mineralogy of benthic orhanisms during experiments, 2011

Increasing atmospheric pCO2 reduces the saturation state of seawater with respect to the aragonite, high-Mg calcite (Mg/Ca > 0.04), and low-Mg calcite (Mg/Ca < 0.04) minerals from which marine calcifiers build their shells and skeletons. Notably, these polymorphs of CaCO3 have different solubilities in seawater: aragonite is more soluble than pure calcite, and the solubility of calcite increases with its Mg-content. Although much recent progress has been made investigating the effects of CO2-induced ocean acidification on rates of biological calcification, considerable uncertainties remain regarding impacts on shell/skeletal polymorph mineralogy. To investigate this subject, eighteen species of marine calcifiers were reared for 60-days in seawater bubbled with air-CO2 mixtures of 409 ± 6, 606 ± 7, 903 ± 12, and 2856 ± 54 ppm pCO2, yielding aragonite saturation states of 2.5 ± 0.4, 2.0 ± 0.4, 1.5 ± 0.3, and 0.7 ± 0.2. Calcite/aragonite ratios within bimineralic calcifiers increased with increasing pCO2, but were invariant within monomineralic calcifiers. Calcite Mg/Ca ratios (Mg/CaC) also varied with atmospheric pCO2 for two of the five high-Mg-calcite-producing organisms, but not for the low-Mg-calcite-producing organisms. These results suggest that shell/skeletal mineralogy within some-but not all-marine calcifiers will change as atmospheric pCO2 continues rising as a result of fossil fuel combustion and deforestation. Paleoceanographic reconstructions of seawater Mg/Ca, temperature, and salinity from the Mg/CaC of well-preserved calcitic marine fossils may also be improved by accounting for the effects of paleo-atmospheric pCO2 on skeletal Mg-fractionation.

Fidelity bias in mollusk assemblages from coastal lagoons of Southern Brazil

The South America southern coast exhibits many outcrops with abundant shell beds, from the Pleistocene through the Recent. How much biological information is preserved within these shell beds? Or, what is the actual probability a living community has to leave a fossil record corresponding to these shell deposits? Although ecological and biogeographical aspects might had been pointed, considering these temporal scales, up to the moment there is no taphonomically-oriented studies available. Quantitative comparisons between living (LAs), death (DAs) and fossil assemblages (FAs) are important not only in strictly taphonomic studies, but have grown a leading tool for conservation paleobiology analysis. Comparing LAs, DAs and FAs from estuaries and lagoons in the Rio Grande do Sul Coastal Plain makes possible to quantitatively understand the nature and quantity of biological information preserved in fossil associations in Holocene lagoon facies. As already noted by several authors, spatial scale parts the analysis, but we detected that the FAs refl ects live ones, rather than dead ones, as previously not realized. The results herein obtained illustrates that species present in DA are not as good preserved in recent (Holocene) fossil record as originally thought. Strictly lagoon species are most prone to leave fossil record. The authors consider that the fi delity pattern here observed for estuarine mollusks to be driven by (i) high temporal and spatial variability in the LAs, (ii) spatial mixing in the DA and (iii) differential preservation of shells, due to long residence times in the taphonomically active zone.

Grain size distribution of the lagoonal deposits within the South Malé Atoll, Maldives, Indian Ocean

Seismic and multibeam data, as well as sediment samples were acquired in the South Malé Atoll in the Maldives archipelago in 2011 to unravel the stratigraphy and facies of the lagoonal deposits. Multichannel seismic lines show that the sedimentary succession locally reaches a maximum thickness of 15-20 m above an unconformity interpreted as the emersion surface which developed during the last glacial sea-level lowstand. Such depocenters are located in current-protected areas flanking the reef rim of the atoll or in infillings of karst dolinas. Much of the 50 m deep sea floor in the lagoon interior is current swept, and has no or very minor sediment cover. Erosive current moats line drowned patch reefs, whereas other areas are characterized by nondeposition. Karst sink holes, blue holes and karst valleys occur throughout the lagoon, from its rim to its center. Lagoonal sediments are mostly carbonate rubble and coarse-grained carbonate sands with frequent large benthic foraminifers, Halimeda flakes, red algal nodules, mollusks, bioclasts, and intraclasts, some of them glauconitic, as well as very minor ooids. Finer-grained deposits locally are deposited in current-protected areas behind elongated faros, i.e., small atolls which are part of the rim of South Malé Atoll. The South Malé Atoll is a current-flushed atoll, where water and sediment export with the open sea is facilitated by the multiple passes dissecting the atoll rim. With an elevated reef rim and tower-like reefs in the atoll interior it is an example of a leaky bucket atoll which shares characteristics of incipiently drowned carbonate banks or drowning sequences as known from the geological record.

Pollen and macrofossil profiles from six cores in Illinois, U.S.A

Pollen analysis of Wisconsinan sediments from eleven localities in northern and central Illinois, combined with the results of older studies, allows a first general survey of the vegetational changes in Illinois during the last glaciation. In the late Altonian (after 40,000 B.P.), pine was already the most prevalent tree type in northern Illinois. Probably because of the influence of the last Altonian ice advance to northern Illinois, pine migrated to the south and reached south-central Illinois, which was at that time a region of prairie, with oak and hickory trees in favorable sites. Likewise in the late Altonian, spruce appeared in northern Illinois. Spruce also expanded its area to the south during the Wisconsinan, reaching south-central Illinois only after 21,000 B.P., in the early Woodfordian. Deciduous trees (predominantly oak) were present in south-central Illinois throughout the Wisconsinan. Their prevalence decreased to the north. The vegetation during the different subdivisions of the last glacial period in Illinois was approximately as follows: Late Altonian: Pine/spruce forest with some deciduous trees in northern and central Illinois; prairie and oak/hickory stands in south-central Illinois; immigration of pine. Farmdalian: Pine/spruce forest in central Illinois; deciduous trees and pine in south-central Illinois, with areas of open vegetation, perhaps similar to the present-day transition of prairie to forest in the northern Great Plains. Woodfordian: Northern and central Illinois ice covered; in south central Illinois, spruce and oak as dominant tree types, but also pine and grassland. During the Woodfordian, pine and spruce disappeared again from south-central Illinois, and oak/hickory forest and prairie again prevailed. The ice-free areas of northern Illinois become populated temporarily with spruce, but later there is proof of deciduous forest in this region. Pollen investigations in south-central Illinois have shown convincingly that deciduous trees could survive relatively close (less than 60 km) to the ice margin. Therefore the frequently presented view that arctic climatic conditions prevailed in North America during the last glaciation far south of the ice margin can be refuted for the Illinois area, confirming the opinion of other authors resulting from investigations of fossil mollusks and frost-soil features. The small number of localities investigated still permits no complete reconstruction of the vegetation zones and their possible movements in Illinois. During the Altonian and Farmdalian in Illinois, a vegetational zonation probably existed similar to that of today in North America. As the ice pushed southward as far as 39° 20' N. lat in the early Woodfordian, this zonation was apparently broken up under the influence of a relatively moderate climate. In any case, the Vandalia area, which was only about 60 km south of the ice, was at that time neither in a tundra zone nor in a zone of boreal coniferous forest.

Seawater carbonate chemistry and biological processes during experiments with oyster Crassostrea gigas, 2009

An increasing number of studies are now reporting the effects of ocean acidification on a broad range of marine species, processes and systems. Many of these are investigating the sensitive early life-history stages that several major reviews have highlighted as being potentially most susceptible to ocean acidification. Nonetheless there remain few investigations of the effects of ocean acidification on the very earliest, and critical, process of fertilization, and still fewer that have investigated levels of ocean acidification relevant for the coming century. Here we report the effects of near-future levels of ocean acidification (?0.35 pH unit change) on sperm swimming speed, sperm motility, and fertilization kinetics in a population of the Pacific oyster Crassostrea gigas from western Sweden. We found no significant effect of ocean acidification - a result that was well-supported by power analysis. Similar findings from Japan suggest that this may be a globally robust result, and we emphasise the need for experiments on multiple populations from throughout a species' range. We also discuss the importance of sound experimental design and power analysis in meaningful interpretation of non-significant results.

Seawater carbonate chemistry and energy status in the periwinkle Littorina littorea during experiments, 2011

In the future, marine organisms will face the challenge of coping with multiple environmental changes associated with increased levels of atmospheric Pco2, such as ocean warming and acidification. To predict how organisms may or may not meet these challenges, an in-depth understanding of the physiological and biochemical mechanisms underpinning organismal responses to climate change is needed. Here, we investigate the effects of elevated Pco2 and temperature on the whole-organism and cellular physiology of the periwinkle Littorina littorea. Metabolic rates (measured as respiration rates), adenylate energy nucleotide concentrations and indexes, and end-product metabolite concentrations were measured. Compared with values for control conditions, snails decreased their respiration rate by 31% in response to elevated Pco2 and by 15% in response to a combination of increased Pco2 and temperature. Decreased respiration rates were associated with metabolic reduction and an increase in end-product metabolites in acidified treatments, indicating an increased reliance on anaerobic metabolism. There was also an interactive effect of elevated Pco2 and temperature on total adenylate nucleotides, which was apparently compensated for by the maintenance of adenylate energy charge via AMP deaminase activity. Our findings suggest that marine intertidal organisms are likely to exhibit complex physiological responses to future environmental drivers, with likely negative effects on growth, population dynamics, and, ultimately, ecosystem processes.

Seawater carbonate chemistry and biological parameters of Sepia officinalis during experiments, 2009

Low pO2 values have been measured in the perivitelline fluids (PVF) of marine animal eggs on several occasions, especially towards the end of development, when embryonic oxygen consumption is at its peak and the egg case acts as a massive barrier to diffusion. Several authors have therefore suggested that oxygen availability is the key factor leading to hatching. However, there have been no measurements of PVF pCO2 so far. This is surprising, as elevated pCO2 could also constitute a major abiotic stressor for the developing embryo. As a first attempt to fill this gap in knowledge, we measured pO2, pCO2 and pH in the PVF of late cephalopod (Sepia officinalis) eggs. We found linear relationships between embryo wet mass and pO2, pCO2 and pH. pO2 declined from >12 kPa to less than 5 kPa, while pCO2 increased from 0.13 to 0.41 kPa. In the absence of active accumulation of bicarbonate in the PVF, pH decreased from 7.7 to 7.2. Our study supports the idea that oxygen becomes limiting in cephalopod eggs towards the end of development; however, pCO2 and pH shift to levels that have caused significant physiological disturbances in other marine ectothermic animals. Future research needs to address the physiological adaptations that enable the embryo to cope with the adverse abiotic conditions in their egg environment.