‘Ten Years After’—a long-term settlement and bioerosion experiment in an Arctic rhodolith bed (Mosselbukta,Svalbard)

Rhodolith beds and bioherms formed by ecosystem engineering crustose coralline algae support the northernmost centres of carbonate production, referred to as polar cold-water carbonate factories. Yet, little is known about biodiversity and recruitment of these hard-bottom communities or the bioeroders degrading them, and there is a demand for carbonate budgets to include respective rates of polar carbonate build-up and bioerosion. To address these issues, a 10-year settlement and bioerosion experiment was carried out at the Arctic Svalbard archipelago in and downslope of a rhodolith bed. The calcifiers recorded on experimental settlement tiles (56 taxa) were dominated by bryozoans, serpulids and foraminiferans. The majority of the bioerosion traces (30 ichnotaxa) were microborings, followed by attachment etchings and grazing traces. Biodiversity metrics show that calcifier diversity and bioerosion ichnodiversity are both elevated in the rhodolith bed, if compared to adjacent aphotic waters, but these differences are statistically insignificant. Accordingly, there were only low to moderate dissimilarities in the calcifier community structure and bioerosion trace assemblages between the two depth stations (46 and 127 m), substrate orientations (up- and down-facing) and substrate types (PVC and limestone), in that order of relevance. In contrast, surface coverage as well as the carbonate accretion and bioerosion rates were all significantly elevated in the rhodolith bed, reflecting higher abundance or size of calcifiers and bioerosion traces. All three measures were highest for up-facing substrates at 46 m, with a mean coverage of 78.2% (on PVC substrates), a mean accretion rate of 24.6 g m^?2  year^?1 (PVC), and a mean bioerosion rate of ?35.1 g m^?2 year^?1 (limestone). Differences in these metrics depend on the same order of factors than the community structure. Considering all limestone substrates of the two platforms, carbonate accretion and bioerosion were nearly in balance at a net rate of ?2.5 g m^?2 year^?1. A latitudinal comparison with previous settlement studies in the North Atlantic suggests that despite the harsh polar environment there is neither a depletion in the diversity of hard-bottom calcifier communities nor in the ichnodiversity of grazing traces, attachment etchings and microborings formed by organotrophs. In contrast, microborings produced by phototrophs are strongly depleted because of limitations in the availability of light (condensed photic zonation, polar night, shading by sea ice). Also, macroborings were almost absent, surprisingly. With respect to carbonate production, the Svalbard carbonate factory marks the low end of a latitudinal gradient while bioerosion rates are similar or even higher than at comparable depth or photic regime at lower latitudes, although this might not apply to shallow euphotic waters (not covered in our experiment), given the observed depletion in bioeroding microphytes and macroborers. While echinoid grazing is particularly relevant for the bioerosion in the rhodolith bed, respective rates are far lower than those reported from tropical shallow-water coral reefs. The slow pace of carbonate production but relatively high rates of bioerosion (both promoted by low carbonate supersaturation states in Arctic waters), in concert with high retention of skeletal carbonates on the seafloor and no calcite cements forming in open pore space created by microborers, suggest a low fossilisation potential for polar carbonates, such as those formed in the Mosselbukta rhodolith beds.     

Colonisation and bioerosion of experimental substrates by benthic foraminiferans from euphotic to aphotic depths (Kosterfjord, SW Sweden)

In the cold-temperate setting of the Swedish Kosterfjord area, experimental carbonate and PVC substrates were deployed for a 6, 12 and 24-month duration along a transect from euphotic to aphotic depths in order to study bioerosion and carbonate accretion patterns. Among the organisms that contribute to the latter by secreting calcareous skeletons, epibenthic foraminiferans represent a major component, both in terms of diversity (a dozen species) as well as in the number of individuals (exceeding 50,000 individuals per m² at certain depths). The by far dominating species were found to be Cibicides lobatulus and the agglutinating Lituotuba lituiformis, along with smaller numbers of Planorbulina mediterranensis, Tholosina vesicularis and Nubecularia lucifuga. The foraminiferal distribution exhibits a pronounced abundance maximum in shallow waters at 7 and especially 15 m and a maximum in diversity at 15-50 m water depth. Some of the foraminiferans encountered, such as Cibicides lobatulus and the rare Gypsina vesicularis, were found to contribute also to the bioerosion of the calcareous substrates by etching shallow attachment scars. These prominent traces witness the former presence of benthic foraminiferans on fossil to Recent hardgrounds, inferring a potential applicability as an in situ proxy where tests are not preserved. Estimated minimum carbonate production rates for the dominant Cibicides lobatulus reach a maximum of 0.326 g/m²/year with the highest rates occurring at 7 to 30 m water depth. Carbonate production rates are up to two magnitudes higher on the PVC (0-0.326 g/m²/year) than on the carbonate substrates (0-0.010 g/m²/year) and are considerably higher than estimates previously reported from the western Baltic.     

The infaunal echinoid Micraster: Taphonomic pathways indicated by sclerozoan trace and body fossils from the Upper Cretaceous of northern Spain

Micraster echinoid fossils are common in the Upper Cretaceous Olazagutía Formation of northern Spain. Tests frequently record sclerobiont signatures (including bioerosion and encrustation), left by reaction and/or coaction phenomena. Among bioerosion structures, Oichnus simplex, O. paraboloides, O. ichnosp. A, Trypanites solitarius, Rogerella ichnosp. indet., Centrichnus cf. eccentricus, Maeandropolydora ichnosp. indet. and fungal microborings are found, as well as pits and fractures. As for sclerozoan body fossils, bivalves (Dimyidae, Anomiidae, Plicatulidae and other Pectinacea), polychaete annelids (serpulids and spirorbids), lituolid foraminiferans (Haddoniidae and Coscinophragmatidae) and bryozoans (cheilostomate ones and others undetermined), as well as other less common groups, have been identified. Taphonomic paths followed by Micraster tests are analysed, based on conservation state and degree of colonisation and sedimentary filling. This allows to distinguish between accumulated fossils and non accumulated ones (including resedimented and reelaborated ones).     

Sexual Reproduction of the Foraminiferan Planoglabratella opercularis (d'Orbigny, 1839) in Nature

The taxonomic composition of the entire foraminiferan assemblage in the southern part of Peter the Great Bay (Sea of Japan) was studied during the summer 1999 expedition. Along with other foraminiferans, several offspring-filled tests of Planoglabratella opercularis (d'Orbigny, 1839) were found in surface sediment samples. Based on the extensive literature on the reproductive cycle of foraminiferans and indirect evidence, it was found that each of the parent individuals of this species was a partner in a plastogamous pair, thus being a gamont, while their offspring were agamonts. The gamogony stage, contrary to the generally accepted opinion, took place under natural conditions in summer (not in fall or winter) at a bottom water temperature of 15–18°C. The results of this research may be important in paleogeographical and paleoecological reconstructions.     

Nautilus pompilius life history and demographics at the Osprey Reef Seamount, Coral Sea, Australia

Nautiloids are the subject of speculation as to their threatened status arising from the impacts of targeted fishing for the ornamental shell market. Life history knowledge is essential to understand the susceptibility of this group to overfishing and to the instigation of management frameworks. This study provides a comprehensive insight into the life of Nautilus in the wild. At Osprey Reef from 1998–2008, trapping for Nautilus was conducted on 354 occasions, with 2460 individuals of one species, Nautilus pompilius, captured and 247 individuals recaptured. Baited remote underwater video systems (BRUVS) were deployed on 15 occasions and six remotely operated vehicle (ROV) dives from 100–800 m were conducted to record Nautilus presence and behavior. Maturity, sex and size data were recorded, while measurements of recaptured individuals allowed estimation of growth rates to maturity, and longevity beyond maturity. We found sexual dimorphism in size at maturity (males: 131.9±SD = 2.6 mm; females: 118.9±7.5 mm shell diameter) in a population dominated by mature individuals (58%). Mean growth rates of 15 immature recaptured animals were 0.061±0.023 mm day⁻¹ resulting in an estimate of around 15.5 years to maturation. Recaptures of mature animals after five years provide evidence of a lifespan exceeding 20 years. Juvenile Nautilus pompilius feeding behavior was recorded for the first time within the same depth range (200–610 m) as adults. Our results provide strong evidence of a K-selected life history for Nautilus from a detailed study of a ‘closed’ wild population. In conjunction with population size and density estimates established for the Osprey Reef Nautilus, this work allows calculations for sustainable catch and provides mechanisms to extrapolate these findings to other extant nautiloid populations (Nautilus and Allonautilus spp.) throughout the Indo-Pacific.     

Effects of Water Depth,Seasonal Exposure,and Substrate Orientation on Microbial Bioerosion in the Ionian Sea (Eastern Mediterranean)

The effects of water depth, seasonal exposure, and substrate orientation on microbioerosion were studied by means of a settlement experiment deployed in 15, 50, 100, and 250 m water depth south-west of the Peloponnese Peninsula (Greece). At each depth, an experimental platform was exposed for a summer period, a winter period, and about an entire year. On the up- and down-facing side of each platform, substrates were fixed to document the succession of bioerosion traces, and to measure variations in bioerosion and accretion rates. In total, 29 different bioerosion traces were recorded revealing a dominance of microborings produced by phototrophic and organotrophic microendoliths, complemented by few macroborings, attachment scars, and grazing traces. The highest bioerosion activity was recorded in 15 m up-facing substrates in the shallow euphotic zone, largely driven by phototrophic cyanobacteria. Towards the chlorophyte-dominated deep euphotic to dysphotic zones and the organotroph-dominated aphotic zone the intensity of bioerosion and the diversity of bioerosion traces strongly decreased. During summer the activity of phototrophs was higher than during winter, which was likely stimulated by enhanced light availability due to more hours of daylight and increased irradiance angles. Stable water column stratification and a resulting nutrient depletion in shallow water led to lower turbidity levels and caused a shift in the photic zonation that was reflected by more phototrophs being active at greater depth. With respect to the subordinate bioerosion activity of organotrophs, fluctuations in temperature and the trophic regime were assumed to be the main seasonal controls. The observed patterns in overall bioeroder distribution and abundance were mirrored by the calculated carbonate budget with bioerosion rates exceeding carbonate accretion rates in shallow water and distinctly higher bioerosion rates at all depths during summer. These findings highlight the relevance of bioerosion and accretion for the carbonate budget of the Ionian Sea.     

Foraminiferan (Protozoa) epizoites on deep-water arcturids (Crustacea,Isopoda)

Foraminiferan (Protozoa) epizoites were examined on two deep-water isopods, Neastacilla sp. and Pleuroprion hystrix (Valvifera, Arcturidae), from the north Atlantic Ocean and the Nordic Seas. Most foraminiferans belonged to genus Cibicides and occurred on 27% of Neastacilla and 23% of Pleuroprion hystrix. The pattern of foraminiferan epibiosis was similar in both species, with a concentration on the dorsal body and on the posterior pereopods. The arcturids showed an increase in number of foraminiferans with increased body size, suggesting that available time span for settling on the younger stages (mancas) was relatively short. There was a trend towards larger numbers of foraminiferans occurring on larger arcturid species, suggesting that available space is the factor that determines the total number of foraminiferans for epibenthic arcturid isopod species.     

Chemico-structural phylogeny of the discinoid brachiopod shell

Stratiform shells of living discinids are composed of membranous laminae and variously aggregated, protein-coated granules of apatitic francolite supported by proteinaceous and chitinous nets in glycosaminoglycans (GAGs) to form laminae in rhythmic sets. The succession is like that of living lingulids but differs significantly in the structure of the periostracum, the nature of baculate sets and in its organic composition. In particular, discinids have a higher level of amino acids although with relatively lower acidic and higher basic concentrations; and their overall lower organic content is owing to lower levels of hydrophilic components, like GAGs and chitin. The organic constituents are not completely degraded during fossilization; but data are presently too meagre to distinguish between linguloid and discinoid ancestries. Many differences among three of the four described extant genera emanate from transformations with a long geological history. Pelagodiscus is characterized by regular, concentric rheomorphic folding (fila) of the flexible periostracum and the plastic primary layer and by sporadically developed hemispherical imprints of periostracal vesicles. Both features are more strikingly developed in Palaeozoic discinids. In the oldest discinid, the Ordovician Schizotreta, and the younger Orbiculoidea and related genera, vesicles were persistent, hexagonal close-packed arrays fading out over fila. They must have differed in composition, however, as the larger vesicles of Schizotreta were simple (possibly mucinous), whereas the smaller vesicles of Orbiculoidea and younger genera were composites of thickly coated spheroids, possibly of lipoproteins, which survive as disaggregated relicts in Pelagodiscus. Baculate sets within the secondary layer are also less well developed in living discinids, being incipient in Pelagodiscus and restricted to the dorsal valve of Discinisca. The trellised rods (baculi) with proteinaceous cores are composed of pinacoids or prisms of apatite, depending on whether they are supported by chitinous nets or proteinaceous strands in GAGs. This differentiation occurred in Schizotreta but in that stock (and Trematis) the baculate set is symmetrical with baculi subtended between compact laminae, whereas in younger and post-Palaeozoic species the outer bounding lamina(e) of the set is normally membranous and/or stratified. The most striking synapomorphy of living discinids is the intravesicular secretion of organsiliceous tablets with a crystalline habit within the larval outer epithelium and their exocytosis as a close- or open-packed, transient, biomineral cover for larvae. Canals, on the other hand, are homologous with those pervading lingulid shells. Both systems interconnect with chitinous and proteinaceous sets and have probably always served as vertical struts in an organic scaffolding supporting the stratiform successions. A phylogenetic analysis based mainly on shell structure confirms the discinoids as the sister group of the linguloids but, contrary to current taxonomic practice, also supports the inclusion of acrotretoids within a ''discinoid'' clade as a sister group to the discinids.     

The bivalve boring Cuenulites amygdaloides nov. isp. in siliceous sponges from the Upper Cretaceous of Germany

Lower Campanian siliceous sponges from epicontinental deposits of the Subhercynian Cretaceous Basin in Germany contain amygdaloidal depressions which are distinguished as a new ichnospecies of the ichnogenus Cuenulites. These bioerosion traces are interpreted as borings of semi-endolithic bivalves, produced without significant rotation movement, probably mostly by chemical action. As there are no signs of tissue reaction in the bored sponges, the structure is considered to be produced post mortem to the sponge, probably in a foreshore-shoreface setting, with redeposition offshore.     

Sponge bioerosion accelerated by ocean acidification across species and latitudes?

In many marine biogeographic realms, bioeroding sponges dominate the internal bioerosion of calcareous substrates such as mollusc beds and coral reef framework. They biochemically dissolve part of the carbonate and liberate so-called sponge chips, a process that is expected to be facilitated and accelerated in a more acidic environment inherent to the present global change. The bioerosion capacity of the demosponge Cliona celata Grant, 1826 in subfossil oyster shells was assessed via alkalinity anomaly technique based on 4 days of experimental exposure to three different levels of carbon dioxide partial pressure (pCO₂) at ambient temperature in the cold-temperate waters of Helgoland Island, North Sea. The rate of chemical bioerosion at present-day pCO₂ was quantified with 0.08–0.1 kg m^?2 year^?1. Chemical bioerosion was positively correlated with increasing pCO₂, with rates more than doubling at carbon dioxide levels predicted for the end of the twenty-first century, clearly confirming that C. celata bioerosion can be expected to be enhanced with progressing ocean acidification (OA). Together with previously published experimental evidence, the present results suggest that OA accelerates sponge bioerosion (1) across latitudes and biogeographic areas, (2) independent of sponge growth form, and (3) for species with or without photosymbionts alike. A general increase in sponge bioerosion with advancing OA can be expected to have a significant impact on global carbonate (re)cycling and may result in widespread negative effects, e.g. on the stability of wild and farmed shellfish populations, as well as calcareous framework builders in tropical and cold-water coral reef ecosystems.     

Monitoring the Reductive and Oxidative Half-Reactions of a Flavin-Dependent Monooxygenase using Stopped-Flow Spectrophotometry

Aspergillus fumigatus siderophore A (SidA) is an FAD-containing monooxygenase that catalyzes the hydroxylation of ornithine in the biosynthesis of hydroxamate siderophores that are essential for virulence (e.g. ferricrocin or N'',N",N''''''-triacetylfusarinine C)¹. The reaction catalyzed by SidA can be divided into reductive and oxidative half-reactions (Scheme 1). In the reductive half-reaction, the oxidized FAD bound to Af SidA, is reduced by NADPH^2,3. In the oxidative half-reaction, the reduced cofactor reacts with molecular oxygen to form a C4a-hydroperoxyflavin intermediate, which transfers an oxygen atom to ornithine. Here, we describe a procedure to measure the rates and detect the different spectral forms of SidA using a stopped-flow instrument installed in an anaerobic glove box. In the stopped-flow instrument, small volumes of reactants are rapidly mixed, and after the flow is stopped by the stop syringe (Figure 1), the spectral changes of the solution placed in the observation cell are recorded over time. In the first part of the experiment, we show how we can use the stopped-flow instrument in single mode, where the anaerobic reduction of the flavin in Af SidA by NADPH is directly measured. We then use double mixing settings where Af SidA is first anaerobically reduced by NADPH for a designated period of time in an aging loop, and then reacted with molecular oxygen in the observation cell (Figure 1). In order to perform this experiment, anaerobic buffers are necessary because when only the reductive half-reaction is monitored, any oxygen in the solutions will react with the reduced flavin cofactor and form a C4a-hydroperoxyflavin intermediate that will ultimately decay back into the oxidized flavin. This would not allow the user to accurately measure rates of reduction since there would be complete turnover of the enzyme. When the oxidative half-reaction is being studied the enzyme must be reduced in the absence of oxygen so that just the steps between reduction and oxidation are observed. One of the buffers used in this experiment is oxygen saturated so that we can study the oxidative half-reaction at higher concentrations of oxygen. These are often the procedures carried out when studying either the reductive or oxidative half-reactions with flavin-containing monooxygenases. The time scale of the pre-steady-state experiments performed with the stopped-flow is milliseconds to seconds, which allow the determination of intrinsic rate constants and the detection and identification of intermediates in the reaction⁴. The procedures described here can be applied to other flavin-dependent monooxygenases.^5,6     

Bioerosion along a bathymetric gradient in a cold-temperate setting (Kosterfjord, SW Sweden): an experimental study

In the cold-temperate setting of the Swedish Kosterfjord, a 2-year experiment was launched in order to assess bioerosion rates and to investigate the endolithic borer communities in relation to light availability (relative bathymetry), hydrography and exposure time. The inventory of microendolithic traces, studied by SEM analysis of epoxy resin casts of planted bivalve shells, yields diverse ichnocoenoses comprising a total of 21 traces produced by boring cyanobacteria (7), chlorophytes (4), fungi (6) and traces of uncertain affinity (4). The link between the endoliths (biotaxa) and the traces they leave (ichnotaxa) is evaluated by the study of the boring organisms in situ by transmission light microscopy of planted Iceland spar and bivalve shells. Additionally, the activity of various macroborers (foraminiferans, polychaetes, echinoids, gastropods and sponges) is documented, adding to a distinct diversity maximum at 7 m water depth. A highly condensed photic zonation, due to the high latitude (59°) and eutrophic conditions, is recorded by the measurement of the Photosynthetically Active Radiation (PAR) and is confirmed by the bathymetric range of the photic related ichnocoenoses. At 1 m water depth, a mature shallow euphotic ichnocoenosis dominated by cyanobacteria and at 7 m, a deep euphotic ichnocoenosis dominated by chlorophytes, respectively, is developed after as little as 12 months exposure. With the vanishing light availability from 15 m downwards, the ichnocoenoses development is significantly slowed and only immature dysphotic and aphotic borer communities (dominated by fungi) are encountered. Strong fluctuations of salinity (down to 8%) and temperature (0–20°C) in the euphotic zone indicate most phototrophs present to be considerably euryhaline and eurytherm, while most endolithic fungi appear preferentially in the deeper, more stable marine waters.     

Hidden impacts of ocean acidification to live and dead coral framework

Cold-water corals, such as Lophelia pertusa, are key habitat-forming organisms found throughout the world''s oceans to 3000 m deep. The complex three-dimensional framework made by these vulnerable marine ecosystems support high biodiversity and commercially important species. Given their importance, a key question is how both the living and the dead framework will fare under projected climate change. Here, we demonstrate that over 12 months L. pertusa can physiologically acclimate to increased CO₂, showing sustained net calcification. However, their new skeletal structure changes and exhibits decreased crystallographic and molecular-scale bonding organization. Although physiological acclimatization was evident, we also demonstrate that there is a negative correlation between increasing CO₂ levels and breaking strength of exposed framework (approx. 20–30% weaker after 12 months), meaning the exposed bases of reefs will be less effective ‘load-bearers’, and will become more susceptible to bioerosion and mechanical damage by 2100.     

Macrobioerosion in Porites corals in subtropical northern South China Sea: a limiting factor for high-latitude reef framework development

Bioerosion is an important limiting factor in carbonate accretion and reef framework development; however, few studies have quantified the direct impact of macroborers on high-latitude coral communities, which are viewed as potential refuge during a period of global warming. In this study, internal macrobioerosion of Porites corals was examined at Daya Bay, subtropical northern South China Sea. The principal borers were the bivalve Lithophaga spp. and the sponges Cliona spp. and Cliothosa spp. (≥80 %), while sipunculid and polychaete worms and barnacles accounted for small amounts of bioerosion (≤20 %). Porites corals were heavily bioeroded in areas impacted by aquacultural and urban activities (10.34–27.55 %) compared with corals in relatively unpolluted areas (2.18–6.76 %). High levels of bioerosion, especially boring bivalve infestation, significantly weaken the corals and increase their susceptibility to dislodgement and fragmentation in typhoons, limiting accumulation of limestone framework. This study implies that carbonate accretion and reef development for high-latitude coral communities may be limited in future high-CO₂ and eutrophication-stressed environments.     

Paleoecology of the marine endobenthos

Endobenthic animals, which reside within the sea bottom, include stationary suspension feeders, mobile deposit feeders and both stationary and mobile carnivores. Their activities, especially with regard to dwelling, feeding, walking/crawling and resting/nesting, are recorded as trace fossils.Abundance, diversity and density of some kinds of trace fossils allow interpretation of the population strategies of the trace-makers in terms of opportunistic (r-selected) and equilibrium (K-selected) strategies. Opportunistic ichnotaxa tend to be faciesbreaking traces, which are highly localized in low-diversity, high-density trace fossil associations in rocks representing environmental extremes (e.g., variable salinities, harsh temperatures, low oxygen levels or shifting substrates). Equilibrium ichnotaxa usually are restricted to particular sedimentary facies and are characteristic of high-diversity, low-dominance trace fossil associations in sediments reflecting stable, predictable environmental conditions.The most important environmental factors influencing the composition of trace fossil assemblages in marine settings are bathymetry, substrate, oxygen and hydrodynamic energy. The four factors are closely interrelated, because as water depth increases, there is a general decrease in sediment grain size and hydrodynamic energy of the depositional environment. As depth below the water—sediment interface increases, the firmness of the sediment (due to compaction and dewatering) increases and the oxygen content of interstitial waters drops drastically.Marine ichnofacies are largely substrate-controlled. Soupgrounds are water-saturated, incompetent substrates typified by highly compressed and usually unidentifiable burrows. Softgrounds commonly contain numerous distinctive burrows and are zoned bathymetrically by the Skolithos, Cruziana, Zoophycos and Nereites Ichnofacies. Firmgrounds are characterized by stiff, compacted sediments, in which traces of the Glossifungites Ichnofacies are excavated. Hardgrounds are cemented substrates, in which bioerosion traces of the Trypanites Ichnofacies are bored. Woodgrounds are woody materials that have been exposed to the sea and bored by bivalves, which produce characteristic traces of the Teredolites Ichnofacies. Tiering of endobenthic communities is common and is related to substrate preference of the burrowers and oxygen stratification of interstitial waters.     

Spectroscopic studies for identifying the chemical states of the periostracum of the Corbicula species in Lake Biwa

Corbicula clam shells consist of thin periostracum and calcareous layers made of calcium carbonate (CaCO₃). Depending on habitat conditions, the shell exhibits various colorations, such as yellow, brown, and black. The chemical state of the periostracum of the Corbicula species in Lake Biwa was studied by X-ray absorption fine structure (XAFS) and Raman scattering spectroscopies. Fe K-edge X-ray absorption near edge structure (XANES) revealed that the Fe³⁺ intensity increases as the color of the shell changes from yellow to black. Raman spectra suggested that quinone-based polymers cover the yellow shell, and the black shell is further covered by dihydroxyphenylalanine (DOPA) rings of amino acid derivatives. From Fe K-edge extended X-ray absorption fine structure (EXAFS), it was found that Fe³⁺ in the periostracum was surrounded by five to six oxygen atoms with an average Fe-O ligand distance of 2.0 Å. Accordingly, a tris-DOPA-Fe³⁺ complex is formed, which is responsible for the periostracum’s black color.     

A Novel μCT Analysis Reveals Different Responses of Bioerosion and Secondary Accretion to Environmental Variability

Corals build reefs through accretion of calcium carbonate (CaCO₃) skeletons, but net reef growth also depends on bioerosion by grazers and borers and on secondary calcification by crustose coralline algae and other calcifying invertebrates. However, traditional field methods for quantifying secondary accretion and bioerosion confound both processes, do not measure them on the same time-scale, or are restricted to 2D methods. In a prior study, we compared multiple environmental drivers of net erosion using pre- and post-deployment micro-computed tomography scans (μCT; calculated as the % change in volume of experimental CaCO₃ blocks) and found a shift from net accretion to net erosion with increasing ocean acidity. Here, we present a novel μCT method and detail a procedure that aligns and digitally subtracts pre- and post-deployment μCT scans and measures the simultaneous response of secondary accretion and bioerosion on blocks exposed to the same environmental variation over the same time-scale. We tested our method on a dataset from a prior study and show that it can be used to uncover information previously unattainable using traditional methods. We demonstrated that secondary accretion and bioerosion are driven by different environmental parameters, bioerosion is more sensitive to ocean acidity than secondary accretion, and net erosion is driven more by changes in bioerosion than secondary accretion.