Shell Condition and Survival of Puget Sound Pteropods Are Impaired by Ocean Acidification Conditions

We tested whether the thecosome pteropod Limacina helicina from Puget Sound, an urbanized estuary in the northwest continental US, experiences shell dissolution and altered mortality rates when exposed to the high CO₂, low aragonite saturation state (Ω_a) conditions that occur in Puget Sound and the northeast Pacific Ocean. Five, week-long experiments were conducted in which we incubated pteropods collected from Puget Sound in four carbon chemistry conditions: current summer surface (∼460–500 µatm CO₂, Ω_a≈1.59), current deep water or surface conditions during upwelling (∼760 and ∼1600–1700 µatm CO₂, Ω_a≈1.17 and 0.56), and future deep water or surface conditions during upwelling (∼2800–3400 µatm CO₂, Ω_a≈0.28). We measured shell condition using a scoring regime of five shell characteristics that capture different aspects of shell dissolution. We characterized carbon chemistry conditions in statistical analyses with Ω_a, and conducted analyses considering Ω_a both as a continuous dataset and as discrete treatments. Shell dissolution increased linearly as aragonite saturation state decreased. Discrete treatment comparisons indicate that shell dissolution was greater in undersaturated treatments compared to oversaturated treatments. Survival increased linearly with aragonite saturation state, though discrete treatment comparisons indicated that survival was similar in all but the lowest saturation state treatment. These results indicate that, under starvation conditions, pteropod survival may not be greatly affected by current and expected near-future aragonite saturation state in the NE Pacific, but shell dissolution may. Given that subsurface waters in Puget Sound’s main basin are undersaturated with respect to aragonite in the winter and can be undersaturated in the summer, the condition and persistence of the species in this estuary warrants further study.     

Description and quantification of pteropod shell dissolution: a sensitive bioindicator of ocean acidification

Anthropogenic ocean acidification is likely to have negative effects on marine calcifying organisms, such as shelled pteropods, by promoting dissolution of aragonite shells. Study of shell dissolution requires an accurate and sensitive method for assessing shell damage. Shell dissolution was induced through incubations in CO₂‐enriched seawater for 4 and 14 days. We describe a procedure that allows the level of dissolution to be assessed and classified into three main types: Type I with partial dissolution of the prismatic layer; Type II with exposure of underlying crossed‐lamellar layer, and Type III, where crossed‐lamellar layer shows signs of dissolution. Levels of dissolution showed a good correspondence to the incubation conditions, with the most severe damage found in specimens held for 14 days in undersaturated condition (Ω ~ 0.8). This methodology enables the response of small pelagic calcifiers to acidified conditions to be detected at an early stage, thus making pteropods a valuable bioindicator of future ocean acidification.     

Evolution and biomineralization of pteropod shells

Shelled pteropods, known as sea butterflies, are a group of small gastropods that spend their entire lives swimming and drifting in the open ocean. They build thin shells of aragonite, a metastable polymorph of calcium carbonate. Pteropod shells have been shown to experience dissolution and reduced thickness with a decrease in pH and therefore represent valuable bioindicators to monitor the impacts of ocean acidification. Over the past decades, several studies have highlighted the striking diversity of shell microstructures in pteropods, with exceptional mechanical properties, but their evolution and future in acidified waters remains uncertain. Here, we revisit the body-of-work on pteropod biomineralization, focusing on shell microstructures and their evolution. The evolutionary history of pteropods was recently resolved, and thus it is timely to examine their shell microstructures in such context. We analyse new images of shells from fossils and recent species providing a comprehensive overview of their structural diversity. Pteropod shells are made of the crossed lamellar and prismatic microstructures common in molluscs, but also of curved nanofibers which are proposed to form a helical three-dimensional structure. Our analyses suggest that the curved fibres emerged before the split between coiled and uncoiled pteropods and that they form incomplete to multiple helical turns. The curved fibres are seen as an important trait in the adaptation to a planktonic lifestyle, giving maximum strength and flexibility to the pteropod thin and lightweight shells. Finally, we also elucidate on the candidate biomineralization genes underpinning the shell diversity in these important indicators of ocean health.     

Dissolution Dominating Calcification Process in Polar Pteropods Close to the Point of Aragonite Undersaturation

Thecosome pteropods are abundant upper-ocean zooplankton that build aragonite shells. Ocean acidification results in the lowering of aragonite saturation levels in the surface layers, and several incubation studies have shown that rates of calcification in these organisms decrease as a result. This study provides a weight-specific net calcification rate function for thecosome pteropods that includes both rates of dissolution and calcification over a range of plausible future aragonite saturation states (Ω_ar). We measured gross dissolution in the pteropod Limacina helicina antarctica in the Scotia Sea (Southern Ocean) by incubating living specimens across a range of aragonite saturation states for a maximum of 14 days. Specimens started dissolving almost immediately upon exposure to undersaturated conditions (Ω_ar∼0.8), losing 1.4% of shell mass per day. The observed rate of gross dissolution was different from that predicted by rate law kinetics of aragonite dissolution, in being higher at Ω_ar levels slightly above 1 and lower at Ω_ar levels of between 1 and 0.8. This indicates that shell mass is affected by even transitional levels of saturation, but there is, nevertheless, some partial means of protection for shells when in undersaturated conditions. A function for gross dissolution against Ω_ar derived from the present observations was compared to a function for gross calcification derived by a different study, and showed that dissolution became the dominating process even at Ω_ar levels close to 1, with net shell growth ceasing at an Ω_ar of 1.03. Gross dissolution increasingly dominated net change in shell mass as saturation levels decreased below 1. As well as influencing their viability, such dissolution of pteropod shells in the surface layers will result in slower sinking velocities and decreased carbon and carbonate fluxes to the deep ocean.     

Interannual pteropod variability in sediment traps deployed above and below the aragonite saturation horizon in the Sub-Antarctic Southern Ocean

Anthropogenic inputs of CO₂ are altering ocean chemistry and may alter the role of marine calcifiers in ocean ecosystems. Laboratory research and ocean models suggest calcifiers in polar waters are especially at risk, particularly pteropods: pelagic aragonite-shelled molluscs. However, baseline data for natural populations of pteropods are limited, especially for polar and sub-polar waters. In order to establish baseline data on diversity, preservation state and shell flux of in situ populations of Sub-Antarctic Southern Ocean pteropods, we deployed sediment traps above (1,000 m) and below (2,000 m) the aragonite saturation horizon (ASH) (currently at 1,200 m) from 1997 to 2006 at 47°S, 142°E. We identified seven pteropod taxa. We applied a shell opacity index to each shell collected and found 50% of shells collected above the ASH to be in pristine condition but only 3% of the shells collected below the ASH showed such a high degree of preservation. We estimated pteropod shell mass fluxes for the region (0.17–4.99 mg m⁻² day⁻¹), and we identified significant reductions in shell flux for Limacina helicina antarctica forma rangi and Clio recurva to the trap series above the ASH and for Limacina helicina antarctica forma rangi and Limacina helicina antarctica forma antarctica to the trap series below the ASH over the interval 1997–2006. Our data establish a temporal and vertical snapshot of the current Sub-Antarctic pelagic pteropod community and provide a baseline against which to monitor Southern Ocean pteropods responses, if any, to changing ocean conditions projected for the region in the coming decades.     

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 m^?2 and 12 to 38 ind m^?2, 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 μatm] 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.     

Near‐future pH conditions severely impact calcification,metabolism and the nervous system in the pteropod Heliconoides inflatus

Shelled pteropods play key roles in the global carbon cycle and food webs of various ecosystems. Their thin external shell is sensitive to small changes in pH, and shell dissolution has already been observed in areas where aragonite saturation state is ~1. A decline in pteropod abundance has the potential to disrupt trophic networks and directly impact commercial fisheries. Therefore, it is crucial to understand how pteropods will be affected by global environmental change, particularly ocean acidification. In this study, physiological and molecular approaches were used to investigate the response of the Mediterranean pteropod, Heliconoides inflatus, to pH values projected for 2100 under a moderate emissions trajectory (RCP6.0). Pteropods were subjected to pH_T 7.9 for 3 days, and gene expression levels, calcification and respiration rates were measured relative to pH_T 8.1 controls. Gross calcification decreased markedly under low pH conditions, while genes potentially involved in calcification were up‐regulated, reflecting the inability of pteropods to maintain calcification rates. Gene expression data imply that under low pH conditions, both metabolic processes and protein synthesis may be compromised, while genes involved in acid–base regulation were up‐regulated. A large number of genes related to nervous system structure and function were also up‐regulated in the low pH treatment, including a GABA_A receptor subunit. This observation is particularly interesting because GABA_A receptor disturbances, leading to altered behavior, have been documented in several other marine animals after exposure to elevated CO₂. The up‐regulation of many genes involved in nervous system function suggests that exposure to low pH could have major effects on pteropod behavior. This study illustrates the power of combining physiological and molecular approaches. It also reveals the importance of behavioral analyses in studies aimed at understanding the impacts of low pH on marine animals.     

Microstructural shell strength of the Subantarctic pteropod <Emphasis Type="Italic">Limacina helicina antarctica</Emphasis>

Anthropogenic inputs of CO₂ are changing ocean chemistry and will likely affect calcifying marine organisms, particularly aragonite producers such as pteropods. This work seeks to set a benchmark analysis of pteropod shell properties and variability using nanoindentation and electron microscopy to measure the structural and mechanical properties of Subantarctic pteropod shells (Limacina helicina antarctica) collected in 1998 and 2007. The 1998 shells were collected by a sediment trap deployed at 2000 m, 47°S, 142°E, and the 2007 shells were collected using nets from mixed-layer waters in the region (44°–54°S, 140°–155°E). Transmission electron microscopy revealed that the shells are composed of a polycrystalline structure, and no obvious porosity was visible. The hardness and modulus of the shells were measured using shell cross-section nanoindentation, across various regions of the shell from the inner to outer whorl. No change in mechanical properties was found with respect to the region of the shell cross-section probed. There was no statistically significant difference in the mean modulus or hardness of the shells between the 1998 and 2007 data sets. No major changes in the mechanical properties of these pteropod shells were detected between the 1998 and 2007 data sets, and we discuss the possible biases in the sampling techniques in complicating our analysis. However, quantifying the mechanical properties and microstructure of calcified may still provide insights into the responses of calcification to environmental changes, such as ocean acidification.     

Impact of aragonite saturation state changes on migratory pteropods

Thecosome pteropods play a key role in the food web of various marine ecosystems and they calcify, secreting the unstable CaCO(3) mineral aragonite to form their shell material. Here, we have estimated the effect of ocean acidification on pteropod calcification by exploiting empirical relationships between their gross calcification rates (CaCO(3) precipitation) and aragonite saturation state Ω(a), combined with model projections of future Ω(a). These were corrected for modern model-data bias and taken over the depth range where pteropods are observed to migrate vertically. Results indicate large reductions in gross calcification at temperate and high latitudes. Over much of the Arctic, the pteropod Limacina helicina will become unable to precipitate CaCO(3) by the end of the century under the IPCC SRES A2 scenario. These results emphasize concerns over the future of shelled pteropods, particularly L. helicina in high latitudes. Shell-less L. helicina are not known to have ever existed nor would we expect them to survive. Declines of pteropod populations could drive dramatic ecological changes in the various pelagic ecosystems in which they play a critical role.     

Late Quaternary millennial-scale variability in pelagic aragonite preservation off Somalia

In order to better understand Late Quaternary pelagic aragonite preservation in the western Arabian Sea we have investigated a high-resolution sediment core 905 off Somalia. Pteropod preservation is enhanced in times of reduced monsoon-driven productivity, indicated by low amounts of C_org and low barium to aluminium (Ba/Al) ratios. All periods corresponding to Heinrich events in the North Atlantic are represented by maxima in shell preservation of the common pteropod Limacina inflata (LDX values < 2, except for H5-equivalent with a poorer shell preservation, LDX > 2.66). Good shell preservation is also found during stadials at 52.1–53.2, 36, 33.2, and 31.9 ka. Relative abundance of pteropods and their fragments in the coarse fraction reaches maxima during Marine Isotope Stage (MIS) 5.2, during time-equivalents of Heinrich events 4–6 and in stadials at  53,  42.5, and 41.4 ka.On longer time scales, the pteropod abundance corresponds to the ‘Indo-Pacific carbonate preservation type’ with poor preservation during interglacials and better preservation during glacials. Late MIS 5 to early MIS 4 sections (84.1–64.8 ka) and the Late Holocene interval (6.5–0 ka) of core 905 contain only traces of pteropods. The early Holocene (9.2–6.5 ka) part is characterized by low pteropod amounts. Between 64.8 and 43.4 ka strong fluctuations occur and an intermediate average relative pteropod abundance is revealed. Between 43.4 and 9.2 ka the highest amounts in relative pteropod abundance in core 905 are observed. Besides the regional monsoonal influence on deepwater chemistry, changes in deepwater circulation occurring on glacial/interglacial and stadial/interstadial time scales might have affected pteropod preservation. However, it remains elusive whether 1) deep water formation in the Arabian Sea, 2) inflow of Glacial North Atlantic Intermediate Water or 3) change in water mass properties of the Circumpolar Deep Water (which is the water mass currently bathing this site) contributed to the observed pteropod preservation pattern.     

Trophodynamics of Southern Ocean pteropods on the southern Kerguelen Plateau

Pteropods are a group of small marine gastropods that are highly sensitive to multiple stressors associated with climate change. Their trophic ecology is not well studied, with most research having focused primarily on the effects of ocean acidification on their fragile, aragonite shells. Stable isotopes analysis coupled with isotope‐based Bayesian niche metrics is useful for characterizing the trophic structure of biological assemblages. These approaches have not been implemented for pteropod assemblages. We used isotope‐based Bayesian niche metrics to investigate the trophic relationships of three co‐occurring pteropod species, with distinct feeding behaviors, sampled from the Southern Kerguelen Plateau area in the Indian Sector of the Southern Ocean—a biologically and economically important but poorly studied region. Two of these species were gymnosomes (shell‐less pteropods), which are traditionally regarded as specialist predators on other pteropods, and the third species was a thecosome (shelled pteropod), which are typically generalist omnivores. For each species, we aimed to understand (a) variability and overlap among isotopic niches; and (b) whether there was a relationship between body size and trophic position. Observed isotopic niche areas were broadest for gymnosomes, especially Clione limacina antarctica, whose observed isotopic niche area was wider than expected on both δ¹³C and δ¹⁵N value axes. We also found that trophic position significantly increased with increasing body length for Spongiobranchaea australis. We found no indication of a dietary shift toward increased trophic position with increasing body size for Clio pyramidata f. sulcata. Trophic positions ranged from 2.8 to 3.5, revealing an assemblage composed of both primary and secondary consumer behaviors. This study provides a comprehensive comparative analysis on trophodynamics in Southern Ocean pteropod species, and supports previous studies using gut content, fatty acid and stable isotope analyses. Combined, our results illustrate differences in intraspecific trophic behavior that may be attributed to differential feeding strategies at species level.     

台湾海峡浮游翼足类群聚之季节动态

本研究利用水产试验所研究船“水试一号”于2006年冬(1月)夏(7月)两季在台湾海峡海域进行水文探测及浮游动物采样, 以探讨浮游翼足类群聚组成与丰度之时空分布及其与水文环境的相关性。结果发现浮游翼足类夏季的丰度及多样性指数明显较冬季高, 而种类数则差异不大; 两季间优势种类组成相似但优势排名略有不同, 夏季主要优势种为棒笔帽螺(Creseis clava)、尖笔帽螺(C. acicula)和马蹄螔螺(Limacina trochiformis), 合占所有翼足类丰度的97%, 其中仅棒笔帽螺就占了总丰度的62%; 冬季主要优势种为胖螔螺(Limacina inflata), 占翼足类丰度的72%。翼足类丰度在空间上的分布亦有明显的季节差异, 夏季时以台湾海峡北部较高, 冬季则以南部较高。种类数与物种多样性指数则均以海峡南部较高。由水文站群分析及各站群生物及水文特征结果显示, 台湾海峡季节性水团的消长与翼足类群聚分布有十分密切的关联。     

EVIDENCE FROM THE FOSSIL RECORD OF AN ANTIPREDATORY EXAPTATION: CONCHIOLIN LAYERS IN CORBULID BIVALVES

Conchiolin layers, organic-rich laminae, are characteristic of the shells of corbulid bivalves. The retention of these layers, despite their high metabolic cost, throughout the evolutionary history of Corbulidae has prompted the proposal of several adaptive scenarios to explain the origin and maintenance of these layers. The most widely held hypothesis contends that conchiolin layers are an adaptation for inhibiting drilling by predatory naticid gastropods. However, others suggest that the layers are adaptations to retard shell dissolution in waters undersaturated with calcium carbonate or to increase shell strength in the face of durophagous (shell crushing) predators. In this paper, experiments using recent Corbula (Varicorbula) gibba (Olivi) and observations of corbulids' present natural habitat demonstrate the current utility of conchiolin layers for all three functions: retardation of shell dissolution in waters undersaturated in calcium carbonate, increase of mechanical shell strength, and inhibition of drilling by predatory naticid gastropods. Earlier analyses of the extensive history of naticid predator-corbulid prey interactions suggested that conchiolin layers were an adaptation, a feature that promotes fitness and was built by selection for its current role, for deterring naticid predators. Not only are naticid drillholes widespread in fossil and recent corbulid shells, but an unusually large number of incomplete drillholes terminate unsuccessfully at conchiolin layers. In addition, a phylogenetic analysis of the origin of conchiolin layers and its function to deter naticid predators is consistent with a hypothesis of adaptation for this function. However, this hypothesis is rejected by an examination of fossil Jurassic Corbulomima. These oldest corbulids contained conchiolin layers before the evolution of naticid drilling during the Early Cretaceous. Therefore, conchiolin layers appear to be an exaptation, characters evolved for other usages and later “coopted” for their current role, for defense against drilling predators. The layers may in fact be an adaptation to resist durophagous predation.     

Diverse trends in shell weight of three Southern Ocean pteropod taxa collected with Polar Frontal Zone sediment traps from 1997 to 2007

The impact of ocean acidification on key ocean calcifiers is predicted to be imminent, particularly in high-latitude ecosystems. Long-term field observations are essential to ground truth predictions of change in regional ecosystems. Here, we report on aragonitic pteropods collected to sediment traps at 800 m depth at 54°S, 140°E in the Polar Frontal Zone (PFZ) of the Southern Ocean from 1997 to 2007. Statistically significant trends were not identified in either mass or number flux from 1997 to 2007; however, differences emerged in decadal trends seen in shell weight for each of the three common taxa collected: Limacina helicina antarctica forma antarctica shells became significantly lighter (P < 0.05), L. retroversa australis shells became significantly heavier (P < 0.05) and L. helicina antarctica forma rangi shells did not change significantly. These results suggest that factors other than ocean acidification affect pteropod population variations on decadal timescales, with the potential to either amplify or counter the impact of decreasing aragonite saturation state, at least in the short term. Comparison to sea surface temperature and chlorophyll biomass did not identify these as significant drivers of the observed changes, and attribution across these multiple variables requires better understanding of pteropod physiology and ecology. Our PFZ pelagic pteropod observations provide a reference for evaluation of southern polar pteropod responses to changing ocean conditions in coming decades. Importantly, these data also raise the issue of taxonomic care when monitoring the region for impacts of ocean acidification on calcifiers.     

Gelatinous invertebrate zooplankton of the South Adriatic: species composition and vertical distribution

The species composition and vertical distribution of gelatinousinvertebrate zooplankton were investigated from April 1993 toJune 1994 in the 0–1000 m water column at a deep-sea stationin the northern part of the South Adriatic Pit. Fifty-sevenspecies were identified: 11 hydromedusae, 13 calycophores, 3ctenophores, 3 heteropods, 10 pteropods, 8 polychaetes and 9chaetognaths. The pteropod Desmopterus papilio and the heteropodProtatlanta mediterranea were recorded for the first time inthe Adriatic Sea. Data from this study differed from those ofprevious investigations in the South Adriatic as regards thenumerically dominant polychaete and pteropod species. All investigatedgroups generally were more abundant in the upper 100 m and decreasedwith depth. Different vertical distributions of life stageswere observed for those species that occupy a wide depth range:Persa incolorata, Solmissus albescens, Limacina inflata, Cymbuliaperoni, Pelagobia longicirrata, Sagitta lyra and Sagitta decipiens.     

Late pleistocene pteropod-rich sediment layer in the northeast atlantic and protoconch variation of Clio pyramidata Linné 1767

Sediment samples from the northeastern Atlantic are compared with recent plankton samples from the same area. Special attention is given to the protoconch variation in the pteropod Clio pyramidata. The dominance/diversity ratio of pteropods and the variation in volume of protoconchae of Clio pyramidata prove that the upwelling off NW Africa in the Late Würm is present in the same region as today, and that temperatures were 4–6°C lower in this period than during recent times. Possible reasons for the formation of the thin sediment layer rich in pteropods, synchronous from Portugal to Senegal, are discussed. As an hypothesis, red tides are proposed to be responsible for mass mortality of pteropods (and benthonic molluscs), thus forming the pteropod-rich sediment layer.     

东海表层沉积中的翼足类及其地质意义

对东海84个表层沉积中翼足类进行了定量分析,共获得7属15种,其中Limacinainflata在陆架外缘和上陆坡为优势种,Limacinatrochi formis,Creseisacicula和Creseis virgule在中外陆架最为丰富。根据翼足类的深度分布确定东海现代文石补偿深度在约600m水深处。以冲绳海槽北部柱状样B3GC为例,说明翼足类是研究地质时期古海洋学变化的重要证据。     

Ocean Acidification Disrupts Prey Responses to Predator Cues but Not Net Prey Shell Growth in Concholepas concholepas (loco)

Background

Most research on Ocean Acidification (OA) has largely focused on the process of calcification and the physiological trade-offs employed by calcifying organisms to support the building of calcium carbonate structures. However, there is growing evidence that OA can also impact upon other key biological processes such as survival, growth and behaviour. On wave-swept rocky shores the ability of gastropods to self-right after dislodgement, and rapidly return to normal orientation, reduces the risk of predation.

Methodology/Principal Findings

The impacts of OA on this self-righting behaviour and other important parameters such as growth, survival, shell dissolution and shell deposition in Concholepas concholepas (loco) were investigated under contrasting pCO₂ levels. Although no impacts of OA on either growth or net shell calcification were found, the results did show that OA can significantly affect self-righting behaviour during the early ontogeny of this species with significantly faster righting times recorded for individuals of C. concholepas reared under increased average pCO₂ concentrations (± SE) (716±12 and 1036±14 µatm CO₂) compared to those reared at concentrations equivalent to those presently found in the surface ocean (388±8 µatm CO₂). When loco were also exposed to the predatory crab Acanthocyclus hassleri, righting times were again increased by exposure to elevated CO₂, although self-righting times were generally twice as fast as those observed in the absence of the crab.

Conclusions and Significance

These results suggest that self-righting in the early ontogeny of C. concholepas will be positively affected by pCO₂ levels expected by the end of the 21st century and beginning of the next one. However, as the rate of self-righting is an adaptive trait evolved to reduce lethal predatory attacks, our result also suggest that OA may disrupt prey responses to predators in nature.     

Oceanic dispersal barriers in a holoplanktonic gastropod

Pteropods, a group of holoplanktonic gastropods, are regarded as bioindicators of the effects of ocean acidification on open ocean ecosystems, because their thin aragonitic shells are susceptible to dissolution. While there have been recent efforts to address their capacity for physiological acclimation, it is also important to gain predictive understanding of their ability to adapt to future ocean conditions. However, little is known about the levels of genetic variation and large‐scale population structuring of pteropods, key characteristics enabling local adaptation. We examined the spatial distribution of genetic diversity in the mitochondrial cytochrome c oxidase I (COI) and nuclear 28S gene fragments, as well as shell shape variation, across a latitudinal transect in the Atlantic Ocean (35°N–36°S) for the pteropod Limacina bulimoides. We observed high levels of genetic variability (COI π = 0.034, 28S π = 0.0021) and strong spatial structuring (COI Φ_ST = 0.230, 28S Φ_ST = 0.255) across this transect. Based on the congruence of mitochondrial and nuclear differentiation, as well as differences in shell shape, we identified a primary dispersal barrier in the southern Atlantic subtropical gyre (15–18°S). This barrier is maintained despite the presence of expatriates, a gyral current system, and in the absence of any distinct oceanographic gradients in this region, suggesting that reproductive isolation between these populations must be strong. A secondary dispersal barrier supported only by 28S pairwise Φ_ST comparisons was identified in the equatorial upwelling region (between 15°N and 4°S), which is concordant with barriers observed in other zooplankton species. Both oceanic dispersal barriers were congruent with regions of low abundance reported for a similar basin‐scale transect that was sampled 2 years later. Our finding supports the hypothesis that low abundance indicates areas of suboptimal habitat that result in barriers to gene flow in widely distributed zooplankton species. Such species may in fact consist of several populations or (sub)species that are adapted to local environmental conditions, limiting their potential for adaptive responses to ocean changes. Future analyses of genome‐wide diversity in pteropods could provide further insight into the strength, formation and maintenance of oceanic dispersal barriers.     

Functional morphology and palaeontological significance of the conchiolin layers in corbulid pelecypods

The Corbulidae, which today are slow, cumbersome, very shallow burrowers, developed special morphological features by which they obtained an outstanding capability to withstand the physical and biological stresses characteristic of their preferred habitat. These features are: an inequivalve, globose shape, thick shells, and conchiolin layers (at least one) embedded within their valves in a unique way. These features enable the corbulids to close their valves tightly during the unfavourable environmental conditions (e.g. low salinity, low oxygen content) which may prevail in the marginal marine regions inhabited by several corbulid species. The conchiolin layers act as a barrier preventing all chemically boring organisms from penetrating into the bivalve shell, or shell dissolution by sea water undersaturated with respect to calcium carbonate. The layered conchiolin weakens the shell mechanically, however, especially during fossilization, when the conchiolin is decomposed. The valve splits apart into two shells so completely different in appearance that they may be attributed to different taxa. The conchiolin layers are therefore of great ecological and palaeontological significance. The nature of these conchiolin layers in Corbula (Varicorbula) gibba (Olivi) is described and illustrated and their functional significance discussed in relation to other living and fossil corbulid species.