Influence of sea-ice dynamics on coastal Antarctic benthos: A case study on lantern clams (Laternula elliptica) in Adélie Land.

Polar regions are warming faster than the world average and are profoundly affected by changes in the spatio-temporal dynamics of sea ice, with largely unknown repercussions on the functioning of marine ecosystems. Here, we investigated the impacts of interannual sea-ice variability on coastal benthic communities in Antarctica, focusing on a close-to-pristine area (Adélie Land). We investigated shell growth of the circum-Antarctic bivalve Laternula elliptica, considered a key species in these soft bottom benthic communities. Chondrophores of live-collected clams were prepared using standard sclerochronological methods to study the interannual variability of shell growth from 1996 to 2015. Our results show that the master chronology varied with sea-ice dynamics. When sea ice breaks up too early, sympagic algae do not have time to accumulate sufficiently high biomass, thus strongly limiting the energy input to the benthos. This negatively affects the physiological performance of L. elliptica, thereby altering their population dynamics and hence the functioning of these soft-bottom ecosystems.

Assessing the elemental fingerprints of cockle shells (Cerastoderma edule) to confirm their geographic origin from regional to international spatial scales.

Geographic origin is directly linked to the quality and commercial value of bivalves. The globalization of the seafood trade and the increasing number of fraudulent practices in the bivalves industry has prompted consumers to become increasingly aware on the geographic origin of the seafood they consume. To enhance consumers' confidence and allow authorities to effectively enforce regulations and contain risks that threaten public health, fast and accurate tools must be made available to confirm claims along the trade chain on the geographic origin of bivalves. In the present study the efficiency of using the elemental fingerprints of a small-homogenized subsample of the shell of common cockles (Cerastoderma edule) to confirm their harvesting location is evaluated at different spatial scales: i) regional (along the Galician coast (Spain) - Espasante, Barallobre, Rio Anllóns, Camariñas, Muros, Noia, Carril, Grove, Combarro, Placeres, Moaña, and Baiona), ii) national (along the Portuguese coast - Ria de Aveiro, Óbidos lagoon, Tagus estuary, Sado estuary and Ria Formosa), and iii) international (along the Northeast Atlantic coast - Hejeltefjorden (Norway), Nykobing Mors (Denmark), Sylt (Germany), Slikken van Viane (Netherlands), Roscoff (France), Plymouth (England), Swansea (Wales), Ria de Aveiro (Portugal) and Oualidia (Morocco). Results confirm that elemental fingerprints of bivalve shells are significantly different among locations and that they can be successfully used with high accuracy to discriminate the geographic origin of cockles at all spatial scales surveyed (97.2% at regional scale, 99.3% at national scale and 100% at international scale). Overall, elemental fingerprints of a small-homogenized subsample of the shell showed to be a replicable, low cost and fast tool to reliably trace the place of origin of cockles sampled at different spatial scales, with success rate of discrimination directly increasing with distance between collection sites.

Shells of the bivalve Astarte moerchi give new evidence of a strong pelagic-benthic coupling shift occurring since the late 1970s in the North Water polynya.

Climate changes in the Arctic may weaken the currently tight pelagic-benthic coupling. In response to decreasing sea ice cover, arctic marine systems are expected to shift from a 'sea-ice algae-benthos' to a 'phytoplankton-zooplankton' dominance. We used mollusc shells as bioarchives and fatty acid trophic markers to estimate the effects of the reduction of sea ice cover on the food exported to the seafloor. Bathyal bivalve Astarte moerchi living at 600 m depth in northern Baffin Bay reveals a clear shift in growth variations and Ba/Ca ratios since the late 1970s, which we relate to a change in food availability. Tissue fatty acid compositions show that this species feeds mainly on microalgae exported from the euphotic zone to the seabed. We, therefore, suggest that changes in pelagic-benthic coupling are likely due either to local changes in sea ice dynamics, mediated through bottom-up regulation exerted by sea ice on phytoplankton production, or to a mismatch between phytoplankton bloom and zooplankton grazing due to phenological change. Both possibilities allow a more regular and increased transfer of food to the seabed. This article is part of the theme issue 'The changing Arctic Ocean: consequences for biological communities, biogeochemical processes and ecosystem functioning'.

Does trace element composition of bivalve shells record utra-high frequency environmental variations?

Saint-Pierre and Miquelon (SPM) is a small archipelago where instrumental measures based on water column velocity and temperature profiles compiled comprehensive evidence for strong near-diurnal (25.8h) current and bottom temperature oscillations (up to 11.5 °C) which is possibly the largest ever observed - at any frequency - on a stratified mid-latitude continental shelf. The main objective of our study was to identify if Placopecten magellanicus can record on its shell these high frequency environmental variations. To this end, we have tried to identify proxies for water temperature and food availability through development of a new ultra-high resolution LA-ICPMS analyses method capable of resolving shell surface elemental composition with a 10 µm resolution. This method was applied on two shell fragments, both representing the third year of growth and 2015 annual growth period, respectively coming from two environmentally contrasted sites, more (30 m depth) or less (10 m depth) affected by high frequency thermal oscillations. Our results strongly suggest a relationship between phytoplankton biomass and barium incorporation into P. magellanicus shells at both sites. Even if P. magellanicus might present a physiological control of magnesium incorporation, the shape of the two Mg/Ca profiles seems to illustrate that temperature also exerts a control on magnesium incorporation in P. magellanicus shells from SPM. While U/Ca and Mg/Ca profiles show a strong positive correlation for 30 m site shell, suggesting that uranium incorporation in P. magellanicus shell is at least partially temperature dependent. The absence of such correlation for 10 m site shell suggests differences in uranium environmental availability or in P. magellanicus biomineralization between these two sites. The resolution of this new analytical method raises questions about such data interpretation related to P. magellanicus growth dynamics and physiology or individual scale based environmental measurements.

Bivalve trophic ecology in the Mediterranean: Spatio-temporal variations and feeding behavior.

The trophic ecology of two bivalves, the clam Callista chione and the cockle Glycymeris bimaculata was studied using environmental and biochemical variables of the suspended particulate matter and the sediment. Samples were collected from two shallow sites, Pag and Cetina, in the coastal oligotrophic Mediterranean Sea, during a 17 month period. The temporal variation of the particulate matter reflected a mixture between marine and terrestrial sources throughout the year, with a clear marine influence during summer and fall, and predominance of terrestrial inputs during spring and winter. The digestive gland was a useful rapid turnover tissue, where the carbon isotope signal was species-specific and the nitrogen isotope one was site-specific. FA markers in the digestive gland revealed a mixed diet where Callista chione fed more upon fresh material than G. bimaculata which relied largely on bacteria-derived detritus. Overall, little feeding niche overlap was observed between the two species during the year, indicating resource partitioning, expected for a food-limited system. The present trophic ecology study in co-occurring species allowed identifying species-specific feeding adaptations to environmental variability.

Ligament, hinge, and shell cross-sections of the Atlantic surfclam (Spisula solidissima): Promising marine environmental archives in NE North America.

The Atlantic surfclam (Spisula solidissima) is a commercially important species in North American waters, undergoing biological and ecological shifts. These are attributed, in part, to environmental modifications in its habitat and driven by climate change. Investigation of shell growth patterns, trace elements, and isotopic compositions require an examination of growth lines and increments preserved in biogenic carbonates. However, growth pattern analysis of S. solidissima is challenging due to multiple disturbance lines caused by environmental stress, erosion in umbonal shell regions, and constraints related to sample size and preparation techniques. The present study proposes an alternative method for describing chronology. First, we analyzed growth patterns using growth lines within the shell and hinge. To validate the assumption of annual periodicity of growth line formation, we analyzed the oxygen isotope composition of the outer shell layer of two specimens (46°54'20"N; 56°18'58"W). Maximum δ18Oshell values occurred at the exact same location as internal growth lines in both specimens, confirming that they are formed annually and that growth ceases during winter. Next, we used growth increment width data to build a standardized growth index (SGI) time-series (25-year chronology) for each of the three parts of the shell. Highly significant correlations were found between the three SGI chronologies (p < 0.001; 0.55 < τ < 0.68) of all specimens. Thus, ligament growth lines provide a new method of determining ontogenetic age and growth rate in S. solidissima. In a biogeographic approach, the shell growth performance of S. solidissima in Saint-Pierre and Miquelon was compared to those in other populations along its distribution range in order to place this population in a temporal and regional context.

Drivers of shell growth of the bivalve, Callista chione (L. 1758) - Combined environmental and biological factors.

Seasonal shell growth patterns were analyzed using the stable oxygen and carbon isotope values of live-collected specimens of the bivalve Callista chione from two sites in the Adriatic Sea (Pag and Cetina, Croatia). Micromilling was performed on the shell surface of three shells per site and shell oxygen isotopes of the powder samples were measured. The timing and rate of seasonal shell growth was determined by aligning the δ18Oshell-derived temperatures so that the best fit was achieved with the instrumental temperature curve. According to the data, shells grew only at very low rates or not at all during the winter months, i.e., between January and March. Shell growth slowdown/shutdown temperatures varied among sites, i.e., 13.6 °C at Pag and 16.6 °C at Cetina, indicating that temperature was not the only driver of shell growth. Likely, seasonal differences in seawater temperature and food supply were the major component explaining contrasting growth rates of C. chione at two study sites. Decreasing shell growth rates were also associated with the onset of gametogenesis suggesting a major energy reallocation toward reproduction rather than growth. These results highlight the need to combine sclerochronological analyses with ecological studies to understand life history traits of bivalves as archives of environmental variables.

Influence of riverine input on the growth of Glycymeris glycymeris in the Bay of Brest, North-West France.

A crossdated, replicated, chronology of 114 years (1901-2014) was developed from internal growth increments in the shells of Glycymeris glycymeris samples collected monthly from the Bay of Brest, France. Bivalve sampling was undertaken between 2014 and 2015 using a dredge. In total 401 live specimens and 243 articulated paired valves from dead specimens were collected, of which 38 individuals were used to build the chronology. Chronology strength, assessed as the Expressed Population Signal, was above 0.7 throughout, falling below the generally accepted threshold of 0.85 before 1975 because of reduced sample depth. Significant positive correlations were identified between the shell growth and the annual averages of rainfall (1975-2008; r = 0.34) and inflow from the river Elorn (1989-2009; r = 0.60). A significant negative correlation was identified between shell growth and the annual average salinity (1998-2014; r = -0.62). Analysis of the monthly averages indicates that these correlations are associated with the winter months (November-February) preceding the G. glycymeris growth season suggesting that winter conditions predispose the benthic environment for later shell growth. Concentration of suspended particulate matter within the river in February is also positively correlated with shell growth, leading to the conclusion that food availability is also important to the growth of G. glycymeris in the Bay of Brest. With the addition of principle components analysis, we were able to determine that inflow from the River Elorn, nitrite levels and salinity were the fundamental drivers of G. glycymeris growth and that these environmental parameters were all linked.

Transcriptomic responses of the endangered freshwater mussel Margaritifera margaritifera to trace metal contamination in the Dronne River, France.

The freshwater pearl mussel Margaritifera margaritifera is one of the most threatened freshwater bivalves worldwide. In this study, we aimed (i) to study the processes by which water quality might affect freshwater mussels in situ and (ii) to provide insights into the ecotoxicological significance of water pollution to natural populations in order to provide necessary information to enhance conservation strategies. M. margaritifera specimens were sampled in two close sites located upstream or downstream from an illegal dumping site. The renal transcriptome of these animals was assembled and gene transcription determined by RNA-seq. Correlations between transcription levels of each single transcript and the bioaccumulation of nine trace metals, age (estimated by sclerochronology), and condition index were determined in order to identify genes likely to respond to a specific factor. Amongst the studied metals, Cr, Zn, Cd, and Ni were the main factors correlated with transcription levels, with effects on translation, apoptosis, immune response, response to stimulus, and transport pathways. However, the main factor explaining changes in gene transcription appeared to be the age of individuals with a negative correlation with the transcription of retrotransposon-related genes. To investigate this effect further, mussels were classified into three age classes. In young, middle-aged and old animals, transcription levels were mainly explained by Cu, Zn and age, respectively. This suggests differences in the molecular responses of this species to metals during its lifetime that must be better assessed in future ecotoxicology studies.

Combined Use of Morphological and Molecular Tools to Resolve Species Mis-Identifications in the Bivalvia The Case of Glycymeris glycymeris and G. pilosa.

Morphological and molecular tools were combined to resolve the misidentification between Glycymeris glycymeris and Glycymeris pilosa from Atlantic and Mediterranean populations. The ambiguous literature on the taxonomic status of these species requires this confirmation as a baseline to studies on their ecology and sclerochronology. We used classical and landmark-based morphometric approaches and performed bivariate and multivariate analyses to test for shell character interactions at the individual and population level. Both approaches generated complementary information. The former showed the shell width to length ratio and the valve asymmetry to be the main discriminant characters between Atlantic and Mediterranean populations. Additionally, the external microsculpture of additional and finer secondary ribs in G. glycymeris discriminates it from G. pilosa. Likewise, landmark-based geometric morphometrics revealed a stronger opisthogyrate beak and prosodetic ligament in G. pilosa than G. glycymeris. Our Bayesian and maximum likelihood phylogenetic analyses based on COI and ITS2 genes identified that G. glycymeris and G. pilosa form two separate monophyletic clades with mean interspecific divergence of 11% and 0.9% for COI and ITS2, respectively. The congruent patterns of morphometric analysis together with mitochondrial and nuclear phylogenetic reconstructions indicated the separation of the two coexisting species. The intraspecific divergence occurred during the Eocene and accelerated during the late Pliocene and Pleistocene. Glycymeris pilosa showed a high level of genetic diversity, appearing as a more robust species whose tolerance of environmental conditions allowed its expansion throughout the Mediterranean.

Potential impacts of blooms of the toxic dinoflagellate Karenia brevis on the growth, survival and juvenile recruitment of the non-native green mussel Perna viridis in southeastern United States.

Red tide blooms formed by Karenia brevis are frequent along the Gulf coast of Florida and it is unclear what tolerance the green mussel Perna viridis, a recently introduced species to coastal waters, has toward these events. Established populations of P. viridis were monitored along the coastal waters of Estero Bay, Florida before, during and following two consecutive red tide blooms to assess the potential effects on growth, survival and juvenile recruitment. Upon onset of the bloom, growth rates fell from 6 to 10 mm month(-1) (March 2011-November 2011) to less than 3 mm month(-1). In the succeeding years, K. brevis blooms were present, and average growth of individually tagged mussels remained below 3 mm month(-1). During growth monitoring the use of calcein as an internal marker was tested with positive staining results and no observed effect on growth or survival. In March 2012, following the first red tide bloom, a population-wide mortality event was observed. Following this event, increased mortality rates were observed with peaks during onset of the bloom in the fall of 2012 and 2013. Juvenile recruitment was also limited during years in which blooms persisted into the spring spawning period suggesting gamete and/or larval sensitivity to K. brevis. Although it cannot be conclusively determined that the cause of reduced growth and survival is due to red tide events, the parallels observed suggest that K. brevis is a factor in the observed changes in population structure.

Circadian behaviour of Tectus (Trochus) niloticus in the southwest Pacific inferred from accelerometry.

BACKGROUND: Behaviour and time spent active and inactive are key factors in animal ecology, with important consequences for bioenergetics. For the first time, here, we equipped the gastropod Tectus (= Trochus) niloticus with accelerometers to describe activity rhythms at two sites in the Southwest Pacific with different temperature regimes: New Caledonia and Vanuatu. RESULTS: Based on a 24-hour cycle, T. niloticus activity began at dusk and gradually stopped during the night, before sunrise. This nocturnal behaviour was characterised by short (duration <30 s), low intensity (acceleration < 0.12 É¡) movements and was probably associated with foraging behaviour. We assumed that activity ceased once the animal was satiated. Our analysis of two size groups in Vanuatu (80-90 mm vs. 120-140 mm, basal shell diameter) revealed a size effect; smaller specimens displayed greater activity, reflected by more intense and longer movements while migrating at night toward the edge of the reef. This nocturnal behaviour is not uncommon for grazing gastropods and is mainly associated with attempting to avoid visual predators whilst feeding. CONCLUSIONS: The use of accelerometers coupled with light and temperature sensors provided detailed information on topshell behaviour and physiology under natural conditions. These data provide a foundation for identifying potential changes in the fine-scale behaviour of T. niloticus in response to environmental changes, which is essential in animal ecology and stock conservation.

Endoliths in Lithophaga lithophaga shells--Variation in intensity of infestation and species occurrence.

Pronounced differences with respect to the extent of infestation and the degree of Lithophaga lithophaga shell damage inflicted by euendolithic taxa at two sites in the Adriatic Sea representing different productivity conditions, are described. Shells collected from the eastern part of Kastela Bay, which is characterized by higher primary productivity, have significantly more shell damage then the shell collected from a site on the outer coast of the island of Ciovo exposed to the oligotrophic Adriatic Sea. The presence of endoliths and their perforations were detected in different layers of the shell, including solidly mineralized parts of the skeleton and within the organic lamellae incorporated into the shell. Phototrophic endoliths were not observed in the specimens. The most serious damage to L. lithophaga shells was the boring clionaid sponge Pione vastifica, which was more common in shells collected from Kastela.

Responses of two scleractinian corals to cobalt pollution and ocean acidification.

The effects of ocean acidification alone or in combination with warming on coral metabolism have been extensively investigated, whereas none of these studies consider that most coral reefs near shore are already impacted by other natural anthropogenic inputs such as metal pollution. It is likely that projected ocean acidification levels will aggravate coral reef health. We first investigated how ocean acidification interacts with one near shore locally abundant metal on the physiology of two major reef-building corals: Stylophora pistillata and Acropora muricata. Two pH levels (pHT 8.02; pCO2 366 µatm and pHT 7.75; pCO2 1140 µatm) and two cobalt concentrations (natural, 0.03 µg L-1 and polluted, 0.2 µg L-1) were tested during five weeks in aquaria. We found that, for both species, cobalt input decreased significantly their growth rates by 28% while it stimulated their photosystem II, with higher values of rETRmax (relative Electron Transport Rate). Elevated pCO2 levels acted differently on the coral rETRmax values and did not affect their growth rates. No consistent interaction was found between pCO2 levels and cobalt concentrations. We also measured in situ the effect of higher cobalt concentrations (1.06 ± 0.16 µg L-1) on A. muricata using benthic chamber experiments. At this elevated concentration, cobalt decreased simultaneously coral growth and photosynthetic rates, indicating that the toxic threshold for this pollutant has been reached for both host cells and zooxanthellae. Our results from both aquaria and in situ experiments, suggest that these coral species are not particularly sensitive to high pCO2 conditions but they are to ecologically relevant cobalt concentrations. Our study reveals that some reefs may be yet subjected to deleterious pollution levels, and even if no interaction between pCO2 levels and cobalt concentration has been found, it is likely that coral metabolism will be weakened if they are subjected to additional threats such as temperature increase, other heavy metals, and eutrophication.

Proteomic-based comparison between populations of the Great Scallop, Pecten maximus.

Comparing populations residing in contrasting environments is an efficient way to decipher how organisms modulate their physiology. Here we present the proteomic signatures of two populations in a non-model marine species, the great scallop Pecten maximus, living in the northern (Hordaland, Norway) and in the center (Brest, France) of this species' latitudinal distribution range. The results showed 38 protein spots significantly differentially accumulated in mantle tissues between the two populations. We could unambiguously identify 11 of the protein spots by Maldi TOF-TOF mass spectrometry. Eight proteins corresponded to different isoforms of actin, two were identified as filamin, another protein related to the cytoskeleton structure, and one was the protease elastase. Our results suggest that scallops from the two populations assayed may modulate their cytoskeleton structures through regulation of intracellular pools of actin and filamin isoforms to better adapt to their environment. BIOLOGICAL SIGNIFICANCE: Marine mollusks are non-model organisms that have been poorly studied at the proteomic level, and this article is the first studying the great scallop (P. maximus) at this level. Furthermore, it addresses population proteomics, a new promising field, especially in environmental sciences. This article is part of a Special Issue entitled: Proteomics of non-model organisms.

Variation in size and growth of the great scallop Pecten maximus along a latitudinal gradient.

Understanding the relationship between growth and temperature will aid in the evaluation of thermal stress and threats to ectotherms in the context of anticipated climate changes. Most Pecten maximus scallops living at high latitudes in the northern hemisphere have a larger maximum body size than individuals further south, a common pattern among many ectotherms. We investigated differences in daily shell growth among scallop populations along the Northeast Atlantic coast from Spain to Norway. This study design allowed us to address precisely whether the asymptotic size observed along a latitudinal gradient, mainly defined by a temperature gradient, results from differences in annual or daily growth rates, or a difference in the length of the growing season. We found that low annual growth rates in northern populations are not due to low daily growth values, but to the smaller number of days available each year to achieve growth compared to the south. We documented a decrease in the annual number of growth days with age regardless of latitude. However, despite initially lower annual growth performances in terms of growing season length and growth rate, differences in asymptotic size as a function of latitude resulted from persistent annual growth performances in the north and sharp declines in the south. Our measurements of daily growth rates throughout life in a long-lived ectothermic species provide new insight into spatio-temporal variations in growth dynamics and growing season length that cannot be accounted for by classical growth models that only address asymptotic size and annual growth rate.

Specific pathways for the incorporation of dissolved barium and molybdenum into the bivalve shell: an isotopic tracer approach in the juvenile Great Scallop (Pecten maximus).

Dissolved barium and molybdenum incorporation in the calcite shell was investigated in the Great Scallop Pecten maximus. Sixty six individuals were exposed for 16 days to two successive dissolved Ba and Mo concentrations accurately differentiated by two different isotopic enrichments (97Mo, 95Mo; ¹³5Ba, ¹³7Ba). Soft tissue and shell isotopic composition were determined respectively by quantitative ICP-MS (Inductively Coupled Plasma Mass Spectrometer) and laser ablation--ICP-MS. Results from Ba enrichment indicate the direct incorporation of dissolved Ba into the shell in proportion to the levels in the water in which they grew with a 6-8 day delay. The low spike contributions and the low partition coefficient (D(Mo) = 0.0049 ± 0.0013), show that neither the soft tissue nor the shell were significantly sensitive to Mo enrichment. These results eliminate direct Mo shell enrichment by the dissolved phase, and favour a trophic uptake that will be investigated using the successive isotopic enrichment approach developed in this study.