Biomineralization in bryozoans: present,past and future

Many animal phyla have the physiological ability to produce biomineralized skeletons with functional roles that have been shaped by natural selection for more than 500 million years. Among these are bryozoans, a moderately diverse phylum of aquatic invertebrates with a rich fossil record and importance today as bioconstructors in some shallow‐water marine habitats. Biomineralizational patterns and, especially, processes are poorly understood in bryozoans but are conventionally believed to be similar to those of the related lophotrochozoan phyla Brachiopoda and Mollusca. However, bryozoan skeletons are more intricate than those of these two phyla. Calcareous skeletons have been acquired independently in two bryozoan clades – Stenolaemata in the Ordovician and Cheilostomata in the Jurassic – providing an evolutionary replicate. This review aims to highlight the importance of biomineralization in bryozoans and focuses on their skeletal ultrastructures, mineralogy and chemistry, the roles of organic components, the evolutionary history of bimineralization in bryozoans with respect to changes in seawater chemistry, and the impact of contemporary global changes, especially ocean acidification, on bryozoan skeletons. Bryozoan skeletons are constructed from three different wall types (exterior, interior and compound) differing in the presence/absence and location of organic cuticular layers. Skeletal ultrastructures can be classified into wall‐parallel (i.e. laminated) and wall‐perpendicular (i.e. prismatic) fabrics, the latter apparently found in only one of the two biomineralizing clades (Cheilostomata), which is also the only clade to biomineralize aragonite. A plethora of ultrastructural fabrics can be recognized and most occur in combination with other fabrics to constitute a fabric suite. The proportion of aragonitic and bimineralic bryozoans, as well as the Mg content of bryozoan skeletons, show a latitudinal increase into the warmer waters of the tropics. Responses of bryozoan mineralogy and skeletal thickness to oscillations between calcite and aragonite seas through geological time are equivocal. Field and laboratory studies of living bryozoans have shown that predicted future changes in pH (ocean acidification) combined with global warming are likely to have detrimental effects on calcification, growth rate and production of polymorphic zooids for defence and reproduction, although some species exhibit reasonable levels of resilience. Some key questions about bryozoan biomineralization that need to be addressed are identified.     

Calcite and aragonite distributions in the skeletons of bimineralic bryozoans as revealed by Raman spectroscopy

Abstract. Bryozoans are among a diverse range of invertebrates capable of secreting calcium carbonate skeletons. Relatively little is known about biomineralization in bryozoans, despite the importance of understanding biomineralization processes for nanotechnology and the threats imposed by ocean acidification on organisms having calcareous skeletons. Ten species of cheilostome bryozoans that are reported to have bimineralic skeletons of calcite and aragonite are studied here using Raman spectroscopy. This technique allowed identification of the two mineral phases at submicron spatial resolution, allowing the distributions of calcite and aragonite within bryozoan skeletons to be determined with unprecedented precision. Confirming previous findings based on the use of chemical stains, most of the bimineralic species analyzed exhibited a calcitic skeletal framework, composed of basal, vertical, and inner frontal walls, having aragonite deposited subsequently onto the outer surfaces of the frontal walls. In one species ( Odontionella cyclops ), aragonite formed the superstructure above the autozooids, and in two others, traces of aragonite were detected on the undersides of the frontal shields. Using Raman spectroscopy, it was possible for the first time to determine the mineralogy of small-scale structures, including orificial rims, condyles and hinge teeth, avicularian pivotal bars and rostra, and ascopore rims and sieve plates. Even when surrounded by aragonitic frontal shields, these structures were found typically to be calcitic, the two exceptions being the aragonitic avicularia of Stylopoma inchoans and O. cyclops . Unexpectedly, the first-formed part of the basal wall at the distalmost growing edge of Pentapora foliacea was found to consist mainly of aragonite. This may point to a precursory phase of biomineralization comparable with the unusual mineralogies identified previously in the earliest-formed skeletons of members of some other invertebrate phyla.     

Patterns of benthic mega-invertebrate habitat associations in the Pacific Northwest continental shelf waters: a reassessment

As human impacts and demands for ocean space increase (fisheries, aquaculture, marine reserves, renewable energy), identification of marine habitats hosting sensitive biological assemblages has become a priority. Epifaunal invertebrates, especially the structure-forming species, are an increasing conservation concern as many traditional (bottom-contact fishing) and novel (marine renewable energy) ocean uses have the potential to displace or otherwise impact these slow-growing organisms. The differences in mega-invertebrate species assemblages between high-relief rocks and low-relief sediments are well documented and likely hold for most marine environments. In anticipation of potential development of marine renewable energy faculties off Oregon and Washington (USA), a survey of the benthic invertebrate assemblages and habitats was conducted on three rocky reefs on the continental shelf of the Pacific Northwest, using video footage collected by remotely operated vehicle, to more finely characterize these assemblage–habitat associations. Benthic assemblages appeared to first group by depth (50–80 vs. 100–120 m), then by relief (consolidated rocks vs. unconsolidated rocks and soft sediments). Consolidated rocks were characterized at each site by a combination of various sponges, gorgonians, sea anemones and echinoderms; unconsolidated rocks were characterized at Grays Bank by sea anemones and burrowing brittle stars, and at Bandon-Arago by sponges and echinoderms; soft sediments were characterized at Grays Bank and Siltcoos Reef by sea whips and burrowing brittle stars, as well as pink shrimps and sea stars at Siltcoos Reef, and at Bandon-Arago by sponges, gorgonians and echinoderms. The results of this study will help classify and map the seafloor in a way that represents benthic habitats reflective of biological species assemblage distributions, rather than solely geological features, and support conservation and management planning.     

Coralline algae (Rhodophyta) in a changing world: integrating ecological,physiological, and geochemical responses to global change

Coralline algae are globally distributed benthic primary producers that secrete calcium carbonate skeletons. In the context of ocean acidification, they have received much recent attention due to the potential vulnerability of their high‐Mg calcite skeletons and their many important ecological roles. Herein, we summarize what is known about coralline algal ecology and physiology, providing context to understand their responses to global climate change. We review the impacts of these changes, including ocean acidification, rising temperatures, and pollution, on coralline algal growth and calcification. We also assess the ongoing use of coralline algae as marine climate proxies via calibration of skeletal morphology and geochemistry to environmental conditions. Finally, we indicate critical gaps in our understanding of coralline algal calcification and physiology and highlight key areas for future research. These include analytical areas that recently have become more accessible, such as resolving phylogenetic relationships at all taxonomic ranks, elucidating the genes regulating algal photosynthesis and calcification, and calibrating skeletal geochemical metrics, as well as research directions that are broadly applicable to global change ecology, such as the importance of community‐scale and long‐term experiments in stress response.     

Mineralogy of Arctic bryozoan skeletons in a global context

Bryozoans are major carbonate producers in some ancient and Recent benthic environments, including parts of the Arctic Ocean. Seventy-six species of bryozoans from within the Arctic Circle have been studied using XRD to determine their carbonate mineralogies and the Mg content of the calcite. The majority of species were found to be calcitic, only four having bimineralic skeletons that combined calcite and aragonite, and none being entirely aragonitic. In almost all species, the calcite was of the low- (<4 mol% MgCO₃) or intermediate-Mg (4–11.99 mol% MgCO₃) varieties. Previous regional studies of bryozoan biomineralogy have found higher proportions of bimineralic and/or aragonitic species in New Zealand and the Mediterranean, with a greater number of calcitic species employing intermediate- and high-Mg calcite. The Antarctic bryozoan fauna, however, has a similar mineralogical composition to the Arctic. The lesser solubility of low-Mg calcite compared to both Mg calcite and aragonite in cold polar waters is most likely responsible for this latitudinal pattern. However, it is unknown to what extent environmental factors drive the pattern directly through eliciting an ecophenotypic response from the bryozoans concerned or the pattern reflects genetic adaptations by particular bryozoan clades.     

The origins of graptolites and other pterobranchs: a journey from ‘Polyzoa’

The graptolites, known only from fossils, have been convincingly allied to the pterobranch hemichordates, a group of tiny, mostly colonial marine invertebrates bearing feeding arms. The phylogenetic position of pterobranchs has been subject to debate and revision for over a century. Their colonial lifestyle and feeding arms were originally seen as evidence placing them among the bryozoans, until later and more careful anatomical studies revealed more characters in common with acorn worms. Pterobranchs and acorn worms are now grouped as the phylum Hemichordata. For many decades, it was thought that pterobranchs were closer to the ancestral form of hemichordates, particularly because ‘lophophorate’ invertebrates also possess feeding arms, notably the phoronids and bryozoans, as do crinoid echinoderms. This traditional view has been challenged by recent molecular evidence. First, there is strong molecular evidence to indicate that lophophorates are very distant from hemichordates and echinoderms, in a different major branch of the animal phylogenetic tree. Therefore, similarities between the feeding structures must be due to convergent evolution. Second, there is strong evidence that hemichordates and echinoderms form a clade (Ambulacraria) within the deuterostomes, rather than hemichordates being closer to chordates. Third, there is weaker evidence that pterobranchs may be derived from acorn worms, and hence that the vermiform body plan may be ancestral within hemichordates. This suggestion warrants further testing. Here we review the evidence for these conclusions, highlight strengths and weaknesses in the data and analyses, and consider the implications for the origins of pterobranchs and graptolites.     

Effects of reduced seawater pH on fertilisation, embryogenesis and larval development in the Antarctic seastar Odontaster validus

The effects of ocean acidification will be pronounced in high-latitude marine communities, although little is known on how reproduction in free-spawning polar invertebrates will respond. Using the circum-Antarctic sea star Odontaster validus, we examined fertilisation, larval survival and development under a controlled seawater treatment (temperature = ?0.5 °C, pH 8.1, pCO₂(aq) = 326.6 μatm, TA = 2,274.2 μmol kg soln^?1), two near-future pH treatments (pH 7.8 and 7.6) and an extreme treatment (pH 7.0). At a sperm concentration of 3.5 × 10⁵ sperm ml^?1, percentage of fertilisation was 98–90 % across a pH 8.1–7.0 range. At near-future pH ranges (pH 7.8 and 7.6), fertilisation was not significantly lower than in the control pH 8.1 except at the lowest sperm concentration (2.2 × 10³ sperm ml^?1) where fertilisation was reduced to 60 and 61 % in pH 7.6 and 7.8, respectively. Larval survival was not significantly affected by a decrease in pH of 0.3 units, but at pH 7.6 survival was significantly reduced. This difference was apparent at 9 days, and at the end of the experiment at 58 days, survival was 55 % compared with 85 % in the ambient treatment. Near-future changes to pH yielded smaller larvae, a result of both subtle differences in their morphology and slowed development rates, while larvae at pH 7.0 showed evidence of abnormal development. O. validus fertilisation and larval success declines in seawater pH conditions expected in coastal Antarctica over the coming decades, although the responses observed are within the range observed in warmer-water echinoderms.     

Giant Clams and Rising CO2: Light May Ameliorate Effects of Ocean Acidification on a Solar-Powered Animal

Global climate change and ocean acidification pose a serious threat to marine life. Marine invertebrates are particularly susceptible to ocean acidification, especially highly calcareous taxa such as molluscs, echinoderms and corals. The largest of all bivalve molluscs, giant clams, are already threatened by a variety of local pressures, including overharvesting, and are in decline worldwide. Several giant clam species are listed as ‘Vulnerable’ on the IUCN Red List of Threatened Species and now climate change and ocean acidification pose an additional threat to their conservation. Unlike most other molluscs, giant clams are ‘solar-powered’ animals containing photosynthetic algal symbionts suggesting that light could influence the effects of ocean acidification on these vulnerable animals. In this study, juvenile fluted giant clams Tridacna squamosa were exposed to three levels of carbon dioxide (CO₂) (control ~400, mid ~650 and high ~950 μatm) and light (photosynthetically active radiation 35, 65 and 304 μmol photons m^-2 s^-1). Elevated CO₂ projected for the end of this century (~650 and ~950 μatm) reduced giant clam survival and growth at mid-light levels. However, effects of CO₂ on survival were absent at high-light, with 100% survival across all CO₂ levels. Effects of CO₂ on growth of surviving clams were lessened, but not removed, at high-light levels. Shell growth and total animal mass gain were still reduced at high-CO₂. This study demonstrates the potential for light to alleviate effects of ocean acidification on survival and growth in a threatened calcareous marine invertebrate. Managing water quality (e.g. turbidity and sedimentation) in coastal areas to maintain water clarity may help ameliorate some negative effects of ocean acidification on giant clams and potentially other solar-powered calcifiers, such as hard corals.     

The effects of changing climate on faunal depth distributions determine winners and losers

Changing climate is predicted to impact all depths of the global oceans, yet projections of range shifts in marine faunal distributions in response to changing climate seldom evaluate potential shifts in depth distribution. Marine ectotherms' thermal tolerance is limited by their ability to maintain aerobic metabolism (oxygen‐ and capacity‐limited tolerance), and is functionally associated with their hypoxia tolerance. Shallow‐water (<200 m depth) marine invertebrates and fishes demonstrate limited tolerance of increasing hydrostatic pressure (pressure exerted by the overlying mass of water), and hyperbaric (increased pressure) tolerance is proposed to depend on the ability to maintain aerobic metabolism, too. Here, we report significant correlation between the hypoxia thresholds and the hyperbaric thresholds of taxonomic groups of shallow‐water fauna, suggesting that pressure tolerance is indeed oxygen limited. Consequently, it appears that the combined effects of temperature, pressure and oxygen concentration constrain the fundamental ecological niches (FENs) of marine invertebrates and fishes. Including depth in a conceptual model of oxygen‐ and capacity‐limited FENs' responses to ocean warming and deoxygenation confirms previous predictions made based solely on consideration of the latitudinal effects of ocean warming (e.g. Cheung et al., 2009), that polar taxa are most vulnerable to the effects of climate change, with Arctic fauna experiencing the greatest FEN contraction. In contrast, the inclusion of depth in the conceptual model reveals for the first time that temperate fauna as well as tropical fauna may experience substantial FEN expansion with ocean warming and deoxygenation, rather than FEN maintenance or contraction suggested by solely considering latitudinal range shifts.     

Perspectives in coral reef hydrodynamics

Knowledge of skeletogenesis in scleractinian corals is central to reconstructing past ocean and climate histories, assessing and counteracting future climate and ocean acidification impacts upon coral reefs, and determining the taxonomy and evolutionary path of the Scleractinia. To better understand skeletogenesis and mineralogy in extant scleractinian corals, we have investigated the nature of the initial calcium carbonate skeleton deposited by newly settling coral recruits. Settling Acropora millepora larvae were sampled daily for 10 days from initial attachment, and the carbonate mineralogy of their newly deposited skeletons was investigated. Bulk analyses using Raman and infrared spectroscopic methods revealed that the skeletons were predominantly comprised of aragonite, with no evidence of calcite or an amorphous precursor phase, although presence of the latter cannot be discounted. Sensitive selected area electron diffraction analyses of sub-micron areas of skeletal regions further consolidated these data. These findings help to address the uncertainty surrounding reported differences in carbonate mineralogy between larval and adult extant coral skeletons by indicating that skeletons of new coral recruits share the same aragonitic mineralogy as those of their mature counterparts. In this respect, we can expect that skeletogenesis in both larval and mature growth stages of scleractinian corals will be similarly affected by ocean acidification and predicted environmental changes.     

Carbonate mineralogy of a tropical bryozoan biota and its vulnerability to ocean acidification

Decreasing pH levels in the world’s oceans are widely recognized as a threat to marine life. Bryozoans are among several phyla that produce calcium carbonate skeletons potentially affected by ocean acidification (OA). Depending on species, bryozoan skeletons can consist of calcite, aragonite or have a bimineralic combination of these two minerals. Aragonite is generally more soluble in seawater than calcite, making aragonitic species more vulnerable to OA. Here, for the first time we use Raman spectroscopy to determine the mineral composition of a tropical bryozoan biota. Compared with bryozoan biotas from higher latitudes in which calcite predominates, aragonite was found to occur in a much higher proportion of the 22 cheilostome bryozoan species collected from the shorelines of Penang and Langkawi in Malaysia, where 46% of species are calcitic, 41% aragonitic and 13% bimineralic. All but one of the aragonitic or bimineralic species belong to the ascophorans, whereas calcitic skeletons characterized most of the anascans, many of which are primitive ‘weedy’ malacostegines. These results suggest a relatively high vulnerability of tropical bryozoan faunas to OA, with the weedier taxa likely to be least impacted.     

Detrimental effects of ocean acidification on the economically important Mediterranean red coral (Corallium rubrum)

The mean predicted decrease of 0.3–0.4 pH units in the global surface ocean by the end of the century has prompted urgent research to assess the potential effects of ocean acidification on the marine environment, with strong emphasis on calcifying organisms. Among them, the Mediterranean red coral (Corallium rubrum) is expected to be particularly susceptible to acidification effects, due to the elevated solubility of its Mg‐calcite skeleton. This, together with the large overexploitation of this species, depicts a bleak future for this organism over the next decades. In this study, we evaluated the effects of low pH on the species from aquaria experiments. Several colonies of C. rubrum were long‐term maintained for 314 days in aquaria at two different pH levels (8.10 and 7.81, pH_T). Calcification rate, spicule morphology, major biochemical constituents (protein, carbohydrates and lipids) and fatty acids composition were measured periodically. Exposure to lower pH conditions caused a significant decrease in the skeletal growth rate in comparison with the control treatment. Similarly, the spicule morphology clearly differed between both treatments at the end of the experiment, with aberrant shapes being observed only under the acidified conditions. On the other hand, while total organic matter was significantly higher under low pH conditions, no significant differences were detected between treatments regarding total carbohydrate, lipid, protein and fatty acid composition. However, the lower variability found among samples maintained in acidified conditions relative to controls, suggests a possible effect of pH decrease on the metabolism of the colonies. Our results show, for the first time, evidence of detrimental ocean acidification effects on this valuable and endangered coral species.     

Multiple phases of mg‐calcite in crustose coralline algae suggest caution for temperature proxy and ocean acidification assessment: lessons from the ultrastructure and biomineralization in Phymatolithon (Rhodophyta,Corallinales)1

Magnesium content, strongly correlated with temperature, has been developed as a climate archive for the late Holocene without considering anatomical controls on Mg content. In this paper, we explore the ultrastructure and cellular scale Mg‐content variations within four species of North Atlantic crust‐forming Phymatolithon. The cell wall has radial grains of Mg‐calcite, whereas the interfilament (middle lamella) has grains aligned parallel to the filament axis. The proportion of interfilament and cell wall carbonate varies by tissue and species. Three distinct primary phases of Mg‐calcite were identified: interfilament Mg‐calcite (mean 8.9 mol% MgCO₃), perithallial cell walls Mg‐calcite (mean 13.4 mol% MgCO₃), and hypothallium Mg‐calcite (mean 17.1 mol% MgCO₃). Magnesium content for the bulk crust, an average of all phases present, showed a strongly correlated (R² = 0.975) increase of 0.31 mol% MgCO₃ per °C. Of concern for climate reconstructions is the potential for false warming signals from undetected postgrazing wound repair carbonate that is substantially enriched in Mg, unrelated to temperature. Within a single crust, Mg‐content of component carbonates ranged from 8 to 20 mol% MgCO₃, representing theoretical thermodynamic stabilities from aragonite‐equivalent to unstable higher‐Mg‐calcite. It is unlikely that existing current predictions of ocean acidification impact on coralline algae, based on saturation states calculated using average Mg contents, provide an environmentally relevant estimate.     

Completeness of a fossil record: the Pleistocene echinoids of the Antilles

Diversity of Pleistocene marine invertebrates with single component, robust, easily identifiable skeletons shows a good correspondence to that of the Recent when compared between similar environments in the same region. Such comparisons are less easily made for multi-element skeletons. In the Antilles there are 24 extant species of echinoids in 0–30  m water depth. Considering all available fossils - complete tests, disarticulated spines, and test fragments and ossicles - the Pleistocene echinoids of the Antillean region have a high similarity with the extant shallow water fauna (c. 63% specific and 72% generic similarity). Many of the extant, shallow water, regular echinoid taxa are known from the Pleistocene, although irregular echinoids are less well represented. These results are comparable with those determined for coeval benthic invertebrates from this and other geographic regions, and indicate a high degree of similarity from a survey which is so far limited to only a few of the islands. Not all Pleistocene units considered herein were deposited in shallow water, but they include autochthonous specimens of taxa whose depth range extends beyond shallow water at the present day or presumed allochthonous specimens derived by downslope transport from shallower water, or both.     

海洋酸化效应对海水鱼类的综合影响评述

人类活动引起的大气CO2浓度的升高,除了使全球温度升高外,导致的另一个严重生态问题——海洋酸化(Ocean acidification,OA),受到社会各界包括科研界的高度重视,该领域的大部分研究结果都是在近十年才发表出来的,目前还有很多需要解决的问题。海洋酸化的研究涉及到很多学科的交叉包括化学、古生物学、生态学、生物地球化学等等。在生物学领域,海洋酸化主要围绕敏感物种,例如由碳酸钙形成贝壳或外骨骼的贝类,珊瑚礁群体等。鱼类作为海洋脊椎动物的代表生物类群,自身具有一定的酸碱平衡调节能力,但相关海洋酸化方向的研究并不是很多。尽管人们对于海洋酸化对鱼类的影响了解甚少,这并不说明海洋酸化对鱼类没有作用或者效应小,在相关研究逐步展开的同时,发现鱼类同样受到海洋酸化的危害,几乎涉及到鱼类整个生活史和几乎大部分生理过程,尤其是早期生活史的高度敏感。因此就目前国内外对此领域研究结果做综述,以期待业界同行能够对海水鱼类这个大的类群引起重视。     

Gene expression changes in the coccolithophore Emiliania huxleyi after 500 generations of selection to ocean acidification

Coccolithophores are unicellular marine algae that produce biogenic calcite scales and substantially contribute to marine primary production and carbon export to the deep ocean. Ongoing ocean acidification particularly impairs calcifying organisms, mostly resulting in decreased growth and calcification. Recent studies revealed that the immediate physiological response in the coccolithophore Emiliania huxleyi to ocean acidification may be partially compensated by evolutionary adaptation, yet the underlying molecular mechanisms are currently unknown. Here, we report on the expression levels of 10 candidate genes putatively relevant to pH regulation, carbon transport, calcification and photosynthesis in E. huxleyi populations short-term exposed to ocean acidification conditions after acclimation (physiological response) and after 500 generations of high CO₂ adaptation (adaptive response). The physiological response revealed downregulation of candidate genes, well reflecting the concomitant decrease of growth and calcification. In the adaptive response, putative pH regulation and carbon transport genes were up-regulated, matching partial restoration of growth and calcification in high CO₂-adapted populations. Adaptation to ocean acidification in E. huxleyi likely involved improved cellular pH regulation, presumably indirectly affecting calcification. Adaptive evolution may thus have the potential to partially restore cellular pH regulatory capacity and thereby mitigate adverse effects of ocean acidification.     

The palaeobiology of belemnites – foundation for the interpretation of rostrum geochemistry

Belemnites are an extinct group of Mesozoic coleoid cephalopods with a fossil record ranging from the early Late Triassic [about 240 million years ago (Mya)] to the Cretaceous/Palaeogene boundary (65 Mya). Belemnites were widely distributed, highly abundant and diverse, and an important component of Mesozoic marine food webs. Their internal shells, specifically their low‐Mg calcite rostra, have been used as palaeoenvironmental carbonate archives for the last 70 years. This is primarily due to the assumption that the rostrum calcite formed in equilibrium with the oxygen isotope composition of ambient sea water. Of prime importance for the reliable interpretation of isotope data derived from these biogenic carbonates is a robust reconstruction of the palaeobiology of their producers. Here we provide a critical assessment of published reconstructions of belemnite soft‐body organization and their lifestyle and habitats. Different lines of evidence, including sedimentological, geochemical, morphological, and biomechanical data, point towards an outer shelf habitat of belemnites, for some taxa also including the littoral area. Belemnite habitat temperatures, oxygen content, salinities, and life span are constrained based on observations of the ecology and life history of modern coleoids. Belemnite habitat depth might have been largely controlled by food and temperature, with a temperature optimum between 10°C and 30°C. The distribution of modern coleoids is for most species restricted to well‐oxygenated water masses and a salinity between 27 and 37 psu. The trophic position of belemnites as both predators and prey is documented by unique fossil finds of stomach contents and soft tissue preservation, such as jaws, hooks, and ink sacs. Belemnites were medium‐sized predators in the epipelagic zone (not deeper than ～200 m) hunting for crustaceans, other cephalopods, and fishes. Taxa with elongated rostra probably were fast and highly manoeuvrable swimmers. Forms with conical rostra represent slow but highly manoeuvrable swimmers, and forms with depressed rostra likely had a bottom‐related life habit. Predators of adult belemnites were sharks, bony fishes, and marine reptiles. Belemnites, like most of the modern coleoids, were relatively short lived, most likely living only for 1–2 years. Understanding the biomineralization of belemnite rostra is highly relevant for an improved interpretation of their geochemistry. Here we confirm that belemnite rostra are composed of low Mg‐calcite fibres, but they do not contain distinct types of laminae. These fibres are composed of two distinct calcite phases. One phase is a filigree network of tetrahedral organic‐rich calcite and the second phase is represented by organic‐poor calcite.     

Effects of seawater acidification on a coral reef meiofauna community

Despite the increasing risk that ocean acidification will modify benthic communities, great uncertainty remains about how this impact will affect the lower trophic levels, such as members of the meiofauna. A mesocosm experiment was conducted to investigate the effects of water acidification on a phytal meiofauna community from a coral reef. Community samples collected from the coral reef subtidal zone (Recife de Fora Municipal Marine Park, Porto Seguro, Bahia, Brazil), using artificial substrate units, were exposed to a control pH (ambient seawater) and to three levels of seawater acidification (pH reductions of 0.3, 0.6, and 0.9 units below ambient) and collected after 15 and 30 d. After 30 d of exposure, major changes in the structure of the meiofauna community were observed in response to reduced pH. The major meiofauna groups showed divergent responses to acidification. Harpacticoida and Polychaeta densities did not show significant differences due to pH. Nematoda, Ostracoda, Turbellaria, and Tardigrada exhibited their highest densities in low-pH treatments (especially at the pH reduction of 0.6 units, pH 7.5), while harpacticoid nauplii were strongly negatively affected by low pH. This community-based mesocosm study supports previous suggestions that ocean acidification induces important changes in the structure of marine benthic communities. Considering the importance of meiofauna in the food web of coral reef ecosystems, the results presented here demonstrate that the trophic functioning of coral reefs is seriously threatened by ocean acidification.     

海洋酸化生态学研究进展

工业革命以来,人类排放的大量二氧化碳引起温室效应的同时,也被海洋吸收使得全球海洋出现了严重的酸化。海洋酸化及伴随的海水碳酸盐化学体系的变化对海洋生物产生深远的影响。以海洋酸化对钙化作用和光合作用的影响为重点,总结了近年来关于海洋酸化的研究,介绍了海洋中不同生态系统对海洋酸化的响应。一方面,海水中CO23-浓度和碳酸钙饱和度的降低对海洋钙化生物造成严重损害,生活在高纬的冷水珊瑚和翼足目等文石生产者是最早的受害者;贝类和棘皮动物在钙化早期对海洋酸化尤其敏感,其幼体存活率受到海洋酸化的严重制约。另一方面,CO2浓度的增加能促进海洋植物的光合作用和生长,增加初级生产力,改变浮游植物的群落组成。此外,海洋酸化可以促进固氮和脱氮作用同时削弱硝化作用,改变溶氧浓度分布和金属的生物可利用性,从而对海洋生物产生间接影响。海洋酸化对海洋生态系统的影响机制复杂,影响程度深远。为了能准确的评估海洋酸化的生态学效应,需要更全面深入的研究。     

Effects of Ocean Acidification on Juvenile Red King Crab (Paralithodes camtschaticus) and Tanner Crab (Chionoecetes bairdi) Growth,Condition, Calcification,and Survival

Ocean acidification, a decrease in the pH in marine waters associated with rising atmospheric CO₂ levels, is a serious threat to marine ecosystems. In this paper, we determine the effects of long-term exposure to near-future levels of ocean acidification on the growth, condition, calcification, and survival of juvenile red king crabs, Paralithodes camtschaticus, and Tanner crabs, Chionoecetes bairdi. Juveniles were reared in individual containers for nearly 200 days in flowing control (pH 8.0), pH 7.8, and pH 7.5 seawater at ambient temperatures (range 4.4–11.9 °C). In both species, survival decreased with pH, with 100% mortality of red king crabs occurring after 95 days in pH 7.5 water. Though the morphology of neither species was affected by acidification, both species grew slower in acidified water. At the end of the experiment, calcium concentration was measured in each crab and the dry mass and condition index of each crab were determined. Ocean acidification did not affect the calcium content of red king crab but did decrease the condition index, while it had the opposite effect on Tanner crabs, decreasing calcium content but leaving the condition index unchanged. This suggests that red king crab may be able to maintain calcification rates, but at a high energetic cost. The decrease in survival and growth of each species is likely to have a serious negative effect on their populations in the absence of evolutionary adaptation or acclimatization over the coming decades.