Posts Tagged 'bryozoa'



Impacts of seawater saturation state (ΩA = 0.4 – 4.6) and temperature (10, 25 °C) on the dissolution kinetics of whole-shell biogenic carbonates

Anthropogenic increase of atmospheric pCO2 since the Industrial Revolution has caused seawater pH to decrease and seawater temperatures to increase—trends that are expected to continue into the foreseeable future. Myriad experimental studies have investigated the impacts of ocean acidification and warming on marine calcifiers’ ability to build protective shells and skeletons. No studies, however, have investigated the combined impacts of ocean acidification and warming on the whole-shell dissolution kinetics of biogenic carbonates. Here, we present the results of experiments designed to investigate the effects of seawater saturation state (ΩA = 0.4 – 4.6) and temperature (10, 25 °C) on gross rates of whole-shell dissolution for ten species of benthic marine calcifiers: the oyster Crassostrea virginicus, the ivory barnacle Balanus eburneus, the blue mussel Mytilus edulis, the conch Strombus alatus, the tropical coral Siderastrea siderea, the temperate coral Oculina arbuscula, the hard clam Mercenaria mercenaria, the soft clam Mya arenaria, the branching bryozoan Schizoporella errata, and the coralline red alga Neogoniolithon sp. These experiments confirm that dissolution rates of whole-shell biogenic carbonates decrease with calcium carbonate (CaCO3) saturation state, increase with temperature, and vary predictably with respect to the relative solubility of the calcifiers’ polymorph mineralogy [high-Mg calcite (mol% Mg > 4) > aragonite > low-Mg calcite (mol% Mg < 4)], consistent with prior studies on sedimentary and inorganic carbonates. Furthermore, the severity of the temperature effects on gross dissolution rates also varied with respect to carbonate polymorph solubility, with warming (10 – 25 °C) exerting the greatest effect on biogenic high-Mg calcite, an intermediate effect on biogenic aragonite, and the least effect on biogenic low-Mg calcite. These results indicate that both ocean acidification and warming will lead to increased dissolution of biogenic carbonates in future oceans, with shells/skeletons composed of the more soluble polymorphs of CaCO3 being the most vulnerable to these stressors. The effects of saturation state and temperature on gross shell dissolution rate were modelled with an exponential asymptotic function (y = B0 – B2· eB1·x) that appeals to the general Arrhenius-derived rate equation for mineral dissolution [r = (C · e-Ea/RT)(1-Ω)n]. Although the dissolution curves for the investigated biogenic CaCO3 exhibited exponential asymptotic trends similar to those of inorganic CaCO3, the observation that gross dissolution of whole-shell biogenic CaCO3 occurred (albeit at lower rates) even in treatments that were oversaturated (Ω > 1) with respect to both aragonite and calcite reveals fundamental differences between the dissolution kinetics of whole-shell CaCO3 and inorganic CaCO3. Thus, applying stoichiometric solubility products derived for inorganic CaCO3 to model gross dissolution of biogenic carbonates may substantially underestimate the impacts of ocean acidification on net calcification (gross calcification minus gross dissolution) of systems ranging in scale from individual organisms to entire ecosystems (i.e., net ecosystem calcification). Finally, these experiments permit rough estimation of the impact of CO2-induced ocean acidification on the gross calcification rates of various marine calcifiers, calculated as the difference between net calcification rates derived empirically in prior studies and gross dissolution rates derived from the present study. Organisms’ gross calcification responses to acidification were generally less severe than their net calcification response patterns, with aragonite mollusks (bivalves, gastropods) exhibiting the most negative gross calcification response to acidification, and photosynthesizing organisms, including corals and coralline red algae, exhibiting relative resilience.

Continue reading ‘Impacts of seawater saturation state (ΩA = 0.4 – 4.6) and temperature (10, 25 °C) on the dissolution kinetics of whole-shell biogenic carbonates’

Field-based experimental acidification alters fouling community structure and reduces diversity

1.Increasing levels of CO2 in the atmosphere are affecting ocean chemistry, leading to increased acidification (i.e., decreased pH) and reductions in calcium carbonate saturation state. 2.Many species are likely to respond to acidification, but the direction and magnitude of these responses will be based on interspecific and ontogenetic variation in physiology and the relative importance of calcification. Differential responses to ocean acidification among species will likely result in important changes in community structure and diversity. 3.To characterize potential impacts of ocean acidification on community composition and structure, we examined the response of a marine fouling community to experimental CO2 enrichment in field-deployed flow-through mesocosm systems. 4.Acidification significantly altered community structure by altering the relative abundances of species and reduced community variability, resulting in more homogenous biofouling communities from one experimental tile to the next both among and within the acidified mesocosms. Mussel (Mytilus trossulus) recruitment was reduced by over 30% in the elevated CO2 treatment compared to the ambient treatment by the end of the experiment. Strong differences in mussel cover (up to 40% lower in acidified conditions) developed over the second half of the 10-week experiment. Acidification did not appear to affect mussel growth, as average mussel sizes were similar between treatments at the end of the experiment. Hydroid (Obelia dichotoma) cover was significantly reduced in the elevated CO2 treatment after eight weeks. Conversely, the percent cover of bryozoan colonies (Mebranipora membranacea) was higher under acidified conditions with differences becoming apparent after six weeks. Neither recruitment nor final size of barnacles (Balanus crenatus) was affected by acidification. By the end of the experiment, diversity was 41% lower in the acidified treatment relative to ambient conditions. 5.Overall, our findings support the general expectation that OA will simplify marine communities by acting on important ecological processes that ultimately determine community structure and diversity.

Continue reading ‘Field-based experimental acidification alters fouling community structure and reduces diversity’

Depth patterns in Antarctic bryozoan skeletal Mg-calcite: Can they provide an analogue for future environmental changes?

Factors related to depth have the potential to provide an analogue for future changes in the skeletal mineralogy of calcifying marine organisms and communities, given that oceanic pH decreases with depth, with a minimum pH of <7.7, which corresponds to the predicted pH of shallow waters in the next 85 yr. Antarctic bryozoans are often characterized by surprisingly broad bathymetrical ranges, and thus have potential for the study of depth-related environmental changes. This study addressed depth-related changes in the levels of magnesium (Mg)-calcite in Antarctic bryozoan skeletons for the first time in order to facilitate predictions of ocean acidification effects. Specimens (n = 103) belonging to 4 bryozoan species (3 cheilostomes and 1 cyclostome) were collected at various depths in East Antarctica (Terre Adelie and George V Land) during the CEAMARC cruise (December 2007 to January 2008), and Mg-calcite contents from their calcareous skeletons were studied using X-ray diffraction. A dataset was compiled from existing environmental data for both sampling and neighboring sites. All 4 species were found to be entirely calcitic with low or intermediate Mg-levels. The predicted negative correlation between pH and Mg-calcite was not evident. Higher Mg levels were found in Fasciculipora ramosa from the George V Basin, suggesting that high salinity shelf water creates favorable conditions for this species, although alternative environmental and biological factors influencing Mg-calcite in skeletons are also discussed for this species.

Continue reading ‘Depth patterns in Antarctic bryozoan skeletal Mg-calcite: Can they provide an analogue for future environmental changes?’

Dissolution rates of biogenic carbonates in natural seawater at different pCO2 conditions: a laboratory study

The bulk dissolution rates of six biogenic carbonates (goose barnacle, benthic foraminifera, bryozoan, sea urchin, and two types of coralline algae) and a sample of mixed sediment from the Bermuda carbonate platform were measured in natural seawater at pCO2 values ranging from approximately 3000 to 5500 μatm. This range of pCO2 values encompassed values regularly observed in porewaters at a depth of a few cm in carbonate sediments at shallow water depths (<15 m) on the Bermuda carbonate platform. The biogenic carbonates included calcites of varying Mg content (2–17 mol%) and a range of specific surface areas (0.01–2.7 m2 g−1) as determined by BET gas adsorption. Measured rates of dissolution increased with increasing pCO2 treatment for all substrates and ranged from 2.5 to 18 μmol g−1 h−1. The highest rates of dissolution were observed for the bryozoans and the lowest rates for the goose barnacles. The relative ranking in dissolution rates between different substrates was consistent at all pCO2 levels, indicating that substrates dissolve sequentially and that some substrates will be more vulnerable than others to rising CO2 and ocean acidification. Furthermore, dissolution rates were found to increase with increasing Mg content, though the relative dissolution rates were observed to be a function of both Mg content and microstructure (surface area).

Continue reading ‘Dissolution rates of biogenic carbonates in natural seawater at different pCO2 conditions: a laboratory study’

Bryozoans in climate and ocean acidification research: A reappraisal of an under-used tool

Bryozoans are colonial animals that are widely distributed in marine benthic environments and play an important role in temperate and cold-water oceanic shelves as habitat providers. Morphologically and mineralogically diverse, bryozoans are important carbonate producers with an extensive fossil record, which makes them good indicators in environmental and (paleo) environmental research. Existing data, though insufficient, suggests that bryozoans can become a valuable tool in investigating present-day climate change. This paper reviews the major characteristics of bryozoans, their function in shallow oceanic areas worldwide, and their potential as proxy organisms in climate and ocean acidification research.

Continue reading ‘Bryozoans in climate and ocean acidification research: A reappraisal of an under-used tool’

Forever young… Colonization pattern of epibionts on Posidonia oceanica artificial leaves in relation to ocean acidification

Ocean acidification (OA) is today considered one of the most pervasive stressors for marine biota at the level of species, communities and ecosystems. Naturally acidified systems, such as the CO2 vents, represent suitable laboratories to study the effects of OA on benthic organisms. An analysis of the colonization pattern of epibionts settled on artificial leaves (mimics) of Posidonia oceanica in relation to ocean acidification at the shallow CO2 vents off the island of Ischia, is here presented. Mimics of Posidonia oceanica artificial leaves (dark green flexible PVC stripes 1 cm wide x 36 cm long) were placed from September 2009 to September 2010 along a gradient of OA of the Ischia vent’s system at six stations (3 on the south and 3 on the north side of the study area), located at extreme low pH (mean pH 7.5), low pH (7.8), and control, normal pH conditions (8.12). Six artificial leaves per station were collected every three months and analysed for taxa identification and estimates of coverage (algae and sessile clonal invertebrates) and number of individuals (not clonal taxa). Patterns of colonization in control stations showed a progressive increase in time in coverage values of many organisms, mainly calcifying forms as coralline algae, which represent the dominant taxon, spirorbids and bryozoans. Colonization of artificial leaves located in low pH stations followed a similar temporal pattern as control conditions, but with lower coverage and higher patchiness of calcareous forms at 12 months of colonization. Epibionts in extreme low pH conditions were dominated by filamentous green/brown algae, with the occurrence of a few coralline algae, spirorbids and bryozoans, especially in the early months of colonization (3 and 6 months). Colonization at 9 and 12 months showed the disappearance of even these rare calcareous organisms and occurrence only of filamentous turf and fleshy algae, with a very simplified epibiont assemblage, remaining at an early, young colonization stage. These results indicate a strong selection of calcareous forms and the lack of successional stages in extreme low pH conditions, while the few calcifiers settled at short exposure time (3-6 months) do not seem to survive at longer exposure to critical values of OA.

Continue reading ‘Forever young… Colonization pattern of epibionts on Posidonia oceanica artificial leaves in relation to ocean acidification’

Morphological plasticity in a calcifying modular organism: evidence from an in situ transplant experiment in a natural CO2 vent system

Understanding is currently limited of the biological processes underlying the responses of modular organisms to climate change and the potential to adapt through morphological plasticity related to their modularity. Here, we investigate the effects of ocean acidification and seawater warming on the growth, life history and morphological plasticity in the modular bryozoan Calpensia nobilis using transplantation experiments in a shallow Mediterranean volcanic CO2 vents system that simulates pH values expected for the year 2100. Colonies exposed at vent sites grew at approximately half the rate of those from the control site. Between days 34 and 48 of the experiment, they reached a possible ‘threshold’, due to the combined effects of exposure time and pH. Temperature did not affect zooid length, but longer zooids with wider primary orifices occurred in low pH conditions close to the vents. Growth models describing colony development under different environmental scenarios suggest that stressed colonies of C. nobilis reallocate metabolic energy to the consolidation and strengthening of existing zooids. This is interpreted as a change in life-history strategy to support persistence under unfavourable environmental conditions. Changes in the skeletal morphology of zooids evident in C. nobilis during short-time (87 days) exposure experiments reveal morphological plasticity that may indicate a potential to adapt to the more acidic Mediterranean predicted for the future.

Continue reading ‘Morphological plasticity in a calcifying modular organism: evidence from an in situ transplant experiment in a natural CO2 vent system’

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.

Continue reading ‘Biomineralization in bryozoans: present, past and future’

Variability in the skeletal mineralogy of temperate bryozoans: the relative influence of environmental and biological factors

Bryozoans exhibit a highly variable geochemistry within their calcium carbonate skeletons. Previous studies have predominantly attributed this variability to differences in seawater temperature influencing the relative deposition of aragonite and calcite, and the extent of magnesium incorporation into the calcite lattice. However, the patterns and scale of this variability have not been examined in detail. We conducted a high-replicate, multi-site study on the skeletal mineralogy of temperate Northern Hemisphere bryozoans to investigate the range of skeletal aragonite and Mg-calcite variability between species and the relative influence of environmental and biological factors on skeletal biogeochemistry. During a cruise in May 2012 in Scapa Flow, Orkney, Northeast Scotland, 480 specimens from 3 bryozoan species were collected by SCUBA diving. Samples were obtained from 5 study sites with similar depths and physical characteristics. All specimens were collected within the same week and were selected to be of similar size, age and breeding status. The results of X-ray diffraction analysis showed that wt% MgCO3 in calcite and wt% aragonite in total CaCO3 were statistically different between sites for all species. This may be explained by differential population connectivity between sites influenced by the tidal regimes of Scapa Flow. No temperate bryozoan species showed the expected positive trends of increasing wt% MgCO3 in calcite or wt% aragonite in total CaCO3 with seawater temperature. Based on the data generated in this study, we suggest that both environmental and biological factors are involved in the control of skeletal mineralogy in some temperate bryozoan species.

Continue reading ‘Variability in the skeletal mineralogy of temperate bryozoans: the relative influence of environmental and biological factors’

Settlement pattern of Posidonia oceanica epibionts along a gradient of ocean acidification: an approach with mimics

Effects of ocean acidification (OA on the colonization/settlement pattern of the epibiont community of the leaves and rhizomesof the Mediterranean seagrass,Posidoniaoceanica, have been studied at volcanic CO2vents off Ischia (Italy), using “mimics”as artificial substrates. The experiments were conducted in shallowPosidoniastands (2-3 m depth), in three stations on the northand three on the south sides of the study area, distributed along a pH gradient. At each station, 4 rhizome mimics and 6 artificialleaves were collected every three months (Sept 2009-Sept 2010). The epibionts on both leaf and rhizome mimics showed clearchanges along the pH gradient; coralline algae and calcareous invertebrates (bryozoans, serpulid polychaetes and barnacles) weredominant at control stations but progressively disappeared at the most acidified stations. In these extremely low pH sites theassemblage was dominated by filamentous algae and non calcareous taxa such as hydroids and tunicates. Settlement pattern onthe artificial leaves and rhizome mimics over time showed a consistent distribution pattern along the pH gradient and highlightedthe peak of recruitment of the various organisms in different periods according to their life history.Posidoniamimics at theacidified station showed a poor and very simplified assemblage where calcifying epibionts seemed less competitive for space. Thisprofound difference in epiphyte communities in low pH conditions suggests cascading effects on the food web of the meadow and,consequently, on the functioning of the system.
Continue reading ‘Settlement pattern of Posidonia oceanica epibionts along a gradient of ocean acidification: an approach with mimics’


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