Conflicting effects of predator cue and ocean acidification on the mussel Mytilus coruscus byssus production

Understanding the impact of ocean acidification and warming on communities and ecosystems is a researcher priority. This can only be achieved through a combination of experimental and field approaches that would allow developing a mechanistic understanding of impacts across level of biological organizations. Surprisingly, most published studies are still focusing on single species responses with little consideration for interspecific interactions. In this study, the impacts of a 3 days exposure to three parameters (temperature, pH, and presence/absence of the predator cue of the crab Charybdis japonica) and their interactions on an ecologically important endpoint were evaluated: the byssus production of the mussel Mytilus coruscus. Tested temperatures (25°C and 30°C) were within the present range of natural variability whereas pH (8.1, 7.7, and 7.4) covered present as well as near-future natural variability. As expected, the presence of the crab cue induced an antipredator response in Mytilus coruscus (significant 10% increase in byssus secretion rate, 22% increase in frequency of shed byssus, and 30% longer byssus). Decreased pH but not temperature had a significant negative impact on the same endpoints (up to a 17% decrease in byssus secretion rate, 40% decrease in frequency of shed byssus, and 10% shorter byssus at pH 7.3 as compared with pH 8.1) with no significant interactions between the three tested parameters. In this study, it has been hypothesized that pH and predator cue have different modes of action and lead to conflicting functional responses (escape response versus stronger attachment). Functional consequences for ecosystem dynamics still need to be investigated.

Sampling sites, sampled birds and plants of young larch plantations in Tokachi district, eastern Hokkaido, northern Japan, 2011

Models and data used to describe species-area relationships confound sampling with ecological process as they fail to acknowledge that estimates of species richness arise due to sampling. This compromises our ability to make ecological inferences from and about species-area relationships. We develop and illustrate hierarchical community models of abundance and frequency to estimate species richness. The models we propose separate sampling from ecological processes by explicitly accounting for the fact that sampled patches are seldom completely covered by sampling plots and that individuals present in the sampling plots are imperfectly detected. We propose a multispecies abundance model in which community assembly is treated as the summation of an ensemble of species-level Poisson processes and estimate patch-level species richness as a derived parameter. We use sampling process models appropriate for specific survey methods. We propose a multispecies frequency model that treats the number of plots in which a species occurs as a binomial process. We illustrate these models using data collected in surveys of early-successional bird species and plants in young forest plantation patches. Results indicate that only mature forest plant species deviated from the constant density hypothesis, but the null model suggested that the deviations were too small to alter the form of species-area relationships. Nevertheless, results from simulations clearly show that the aggregate pattern of individual species density-area relationships and occurrence probability-area relationships can alter the form of species-area relationships. The plant community model estimated that only half of the species present in the regional species pool were encountered during the survey. The modeling framework we propose explicitly accounts for sampling processes so that ecological processes can be examined free of sampling artefacts. Our modeling approach is extensible and could be applied to a variety of study designs and allows the inclusion of additional environmental covariates.

Growth and carbon metabolism of the seaweed Fucus vesiculosus in the western Baltic Sea ("Kiel Outdoor Benthocosms")

Warming and acidification of the oceans as a consequence of increasing CO2-concentrations occur at large scales. Numerous studies have shown the impact of single stressors on individual species. However, studies on the combined effect of multiple stressors on a multi-species assemblage, which is ecologically much more realistic and relevant, are still scarce. Therefore, we orthogonally crossed the two factors warming and acidification in mesocosm experiments and studied their single and combined impact on the brown alga Fucus vesiculosus associated with its natural community (epiphytes and mesograzers) in the Baltic Sea in all seasons (from April 2013 to April 2014). We superimposed our treatment factors onto the natural fluctuations of all environmental variables present in the Benthocosms in so-called delta-treatments. Thereby we compared the physiological responses of F. vesiculosus (growth and metabolites) to the single and combined effects of natural Kiel Fjord temperatures and pCO2 conditions with a 5 °C temperature increase and/or pCO2 increase treatment (1100 ppm in the headspace above the mesocosms). Responses were also related to the factor photoperiod which changes over the course of the year. Our results demonstrate complex seasonal pattern. Elevated pCO2 positively affected growth of F. vesiculosus alone and/or interactively with warming. The response direction (additive, synergistic or antagonistic), however, depended on season and daylength. The effects were most obvious when plants were actively growing during spring and early summer. Our study revealed for the first time that it is crucial to always consider the impact of variable environmental conditions throughout all seasons. In summary, our study indicates that in future F. vesiculosus will be more affected by detrimental summer heat-waves than by ocean acidification although the latter consequently enhances growth throughout the year. The mainly negative influence of rising temperatures on the physiology of this keystone macroalga may alter and/or hamper its ecological functions in the shallow coastal ecosystem of the Baltic Sea.

Abundance of calcareous nannofossils across the K/T boundary at ODP Hole 113-690C (Table 1)

A biostratigraphically continuous, but intensely bioturbated, Cretaceous/Tertiary boundary sequence was cored during Ocean Drilling Program (ODP) Leg 113 on Maud Rise (65°S) in the Weddell Sea off East Antarctica. This interval is the first recovered by ODP/DSDP in the South Atlantic sector of the Southern Ocean and offers a unique opportunity to study the nannofossil sequences leading up to and beyond the terminal Cretaceous event at a high southern latitude. The K/T boundary lies just within Chron 29R and is placed at ODP Sample 113-690C-15X-4, 41.5 cm. An iridium anomaly was independently noted at about this level as well. Upper Maestrichtian-lower Paleocene sediments consist mostly of light-colored nannofossil chalks. Dark brown sediments at the base of the Danian (Zone CPla) are characterized by an increased clay content attributed to a drop in calcareous microplankton productivity following the terminal Cretaceous event. Although delineation of the boundary is hampered by intense bioturbation, the sharp color contrast between overlying clay-rich, dark brown chalks of the Tertiary and light cream colored chalks of the Cretaceous aids in the selection of the K/T horizon. Several dark colored burrows sampled at intervals as far as 1.3 m below the boundary and within the light colored Cretaceous chalk were found to contain up to 17% Tertiary nannofossils. Calcareous nannofossils from the boundary interval were divided into three groups for quantitative study. The three groups, "Cretaceous," "Tertiary," and "Survivor," exhibit a sequential change across the boundary with the Cretaceous forms giving way to a Survivor-dominated assemblage beginning at the boundary followed shortly thereafter by the appearance of the Tertiary taxa, Cruciplacolithus and Hornibrookina. The species, H. edwardsii, comprises nearly 50% of the assemblage just above the Zone CPla/CPlb boundary, an abundance not reported elsewhere at this level. Calculation of individual species abundances reveals several additional differences between this K/T boundary interval and those studied from middle and low latitude sections. The percentage of Thoracosphaera is much lower at the boundary in this section and a small form, Prediscosphaera stoveri, is extremely abundant in Cretaceous sediments just below the boundary.

Maestrichtian calcareous nannofossils of MAud Rise, Weddell Sea

Recovery of an essentially complete upper Maestrichtian/lower Paleocene interval on Maud Rise at 65 °S latitude in the Weddell Sea during Ocean Drilling Program Leg 113 marks the first time that this interval has been cored at these high latitudes. The entire interval was missing at all Falkland Plateau sites drilled during DSDP Legs 36 and 71. Maestrichtian nannofossil assemblages in sediments from Sites 689 and 690, therefore, provide the basis for a needed revision of Maestrichtian coccolith zonation schemes for high southern latitudes. Three zones and two new subzones are described: the uppermost Maestrichtian Nephrolithus frequens Zone, which is subdivided into the Cribrosphaerella daniae Subzone and the underlying N. corystus Subzone, and the Biscutum magnum and B. coronum Zones. A complete calcareous nannofossil biostratigraphy based on the proposed scheme is given including a description of individual species abundance, preservation, and stratigraphic distribution. At this site, the southernmost carbonate site yet drilled by DSDP/ODP, it is evident that the Falkland Plateau Nannofossil Biogeographic Province can be extended to the margins of Antarctica. In addition, the biogeographic ranges of many calcareous nannofossils can likewise be extended. Last, we hypothesize that Nephrolithus frequens evolved from N. corystus prior to its dispersal to the lower latitudes where it is an important zonal marker. Three new taxa, Neocrepidolithus watkinsii n. sp., Nephrolithus frequens miniporus emend, n. comb, and Psyktosphaera firthii n. gen., n. sp. are described.

Geographic distribution of dinoflagellate cysts in surface sediments

Dinoflagellate cysts are useful for reconstructing upper water conditions. For adequate reconstructions detailed information is required about the relationship between modern day environmental conditions and the geographic distribution of cysts in sediments. This Atlas summarises the modern global distribution of 71 organicwalled dinoflagellate cyst species. The synthesis is based on the integration of literature sources together with data of 2405 globally distributed surface sediment samples that have been preparedwith a comparable methodology and taxonomy. The distribution patterns of individual cyst species are being comparedwith environmental factors that are knownto influence dinoflagellate growth, gamete production, encystment, excystment and preservation of their organic-walled cysts: surface water temperature, salinity, nitrate, phosphate, chlorophyll-a concentrations and bottom water oxygen concentrations. Graphs are provided for every species depicting the relationship between seasonal and annual variations of these parameters and the relative abundance of the species. Results have been compared with previously published records; an overview of the ecological significance as well as information about the seasonal production of each individual species is presented. The relationship between the cyst distribution and variation in the aforementioned environmental parameters was analysed by performing a canonical correspondence analysis. All tested variables showed a positive relationship on the 99% confidence level. Sea-surface temperature represents the parameter corresponding to the largest amount of variance within the dataset (40%) followed by nitrate, salinity, phosphate and bottom-water oxygen concentration, which correspond to 34%, 33%, 25% and 24% of the variance, respectively. Characterisations of selected environments as well as a discussion about how these factors could have influenced the final cyst yield in sediments are included.

Dinoflagellates of multicorer surface sediments

Only very few studies focus on recent calcareous dinoflagellate cyst diversity, geographic distribution and ecology, so that information on the distribution patterns and environmental affinities of individual cyst species is extremely limited. This information is, however, essential if we want to use calcareous dinoflagellate cysts for palaeoenvironmental reconstruction. Surface sediment samples from the generally oligotrophic western equatorial Atlantic Ocean, offshore northeast Brazil, were therefore quantitatively analysed for their calcareous dinoflagellate cyst content, including the calcareous vegetative coccoid Thoracosphaera heimii. Seven calcareous dinoflagellate cyst species/morphotypes and T. heimii were encountered in high concentrations throughout the area. Substantial differences in the distribution patterns were observed. The highest concentrations of cysts are found in sediments of the more oligotrophic, oceanic regions, beyond the influence of Amazon River discharge waters. Dinoflagellates producing calcareous cysts thus appear to be capable of surviving low nutrient concentrations and produce large numbers of cysts in relatively stable and predictable environments affected by minimal seasonality. To test for the environmental affinities of individual species, distribution patterns in surface sediments were compared with temperature, salinity, density and stratification gradients within the upper water column (0-100 m) over different times of the year, using principal components analysis and redundancy analysis. T. heimii and four of the seven encountered cyst species (Sphaerodinella? albatrosiana, two morphotypes of Sphaerodinella? tuberosa and Scrippsiella regalis) relate to these parameters significantly and the variations in the cyst associations appear to be associated with the different surface water currents characterising the area. The results imply that calcareous dinoflagellate cyst distributions can potentially be used to distinguish between different open oceanic environments and they could, therefore, be useful in tracing water mass movements throughout the late Quaternary.

Relative abundance of dinoflagellate cysts in surface sediments and dinoflagellate counts from three sediment cores from the Atlantic sector of the Southern Ocean

In this study we investigate the potential of organic-walled dinoflagellate cysts (dinocysts) as tools for quantifying past sea-surface temperatures (SST) in the Southern Ocean. For this purpose, a dinocyst reference dataset has been formed, based on 138 surface sediment samples from different circum-Antarctic environments. The dinocyst assemblages of these samples are composed of phototrophic (gonyaulacoid) and heterotrophic (protoperidinioid) species that provide a broad spectrum of palaeoenvironmental information. The relationship between the environmental parameters in the upper water column and the dinocyst distribution patterns of individual species has been established using the statistical method of Canonical Correspondence Analysis (CCA). Among the variables tested, summer SST appeared to correspond to the maximum variance represented in the dataset. To establish quantitative summer SST reconstructions, a Modern Analogue Technique (MAT) has been performed on data from three Late Quaternary dinocyst records recovered from locations adjacent to prominent oceanic fronts in the Atlantic sector of the Southern Ocean. These dinocyst time series exhibit periodic changes in the dinocyst assemblage during the last two glacial/interglacial-cycles. During glacial conditions the relative abundance of protoperidinioid cysts was highest, whereas interglacial conditions are characterised by generally lower cyst concentrations and increased relative abundance of gonyaulacoid cysts. The MAT palaeotemperature estimates show trends in summer SST changes following the global oxygen isotope signal and a strong correlation with past temperatures of the last 140,000 years based on other proxies. However, by comparing the dinocyst results to quantitative estimates of summer SSTs based on diatoms, radiolarians and foraminifer-derived stable isotope records it can be shown that in several core intervals the dinocyst-based summer SSTs appeared to be extremely high. In these intervals the dinocyst record seems to be highly influenced by selective degradation, leading to unusual temperature ranges and to unrealistic palaeotemperatures. We used the selective degradation index (kt-index) to determine those intervals that have been biased by selective degradation in order to correct the palaeotemperature estimates. We show that after correction the dinocyst based SSTs correspond reasonably well with other palaeotemperature estimates for this region, supporting the great potential of dinoflagellate cysts as a basis for quantitative palaeoenvironmental studies.

Stable oxygen isotope composition of planktonic foraminifera in the Canary Islands region

Seasonal depth stratified plankton tows, sediment traps and core tops taken from the same stations along a transect at 29°N off NW Africa are used to describe the seasonal succession, the depth habitats and the oxygen isotope ratios (delta18O(shell)) of five planktic foraminiferal species. Both the delta18O(shell) and shell concentration profiles show variations in seasonal depth habitats of individual species. None of the species maintain a specific habitat depth exclusively within the surface mixed layer (SML), within the thermocline, or beneath the thermocline. Globigerinoides ruber (white) and (pink) occur with moderate abundance throughout the year along the transect, with highest abundances in the winter and summer/fall season, respectively. The average delta18O(shell) of G. ruber (w) from surface sediments is similar to the delta18O(shell) values measured from the sediment-trap samples during winter. However, the delta18O(shell) of G. ruber (w) underestimates sea surface temperature (SST) by 2 °C in winter and by 4 °C during summer/fall indicating an extension of the calcification/depth habitat into colder thermocline waters. Globigerinoides ruber (p) continues to calcify below the SML as well, particularly in summer/fall when the chlorophyll maximum is found within the thermocline. Its vertical distribution results in delta18O(shell) values that underestimate SST by 2 °C. Shell fluxes of Globigerina bulloides are highest in summer/fall, where it lives and calcifies in association with the deep chlorophyll maximum found within the thermocline. Pulleniatina obliquiloculata and Globorotalia truncatulinoides, dwelling and calcifying a part of their lives in the winter SML, record winter thermocline (~180 m) and deep surface water (~350 m) temperatures, respectively. Our observations define the seasonal and vertical distribution of multiple species of foraminifera and the acquisition of their delta18O(shell).

Benthic foraminifera groups of ODP Site 162-982

The pulsed decline and eventual extinction of 51 species of elongate, cylindrical deep-sea benthic foraminifera (Stilostomellidae, Pleurostomellidae, and some Nodosariidae) occurred at intermediate water depths (1145-2168 m, Sites 980 and 982) in the northern North Atlantic during the mid-Pleistocene transition (MPT, 1.2-0.6 Ma). In the early Pleistocene, prior to their disappearance, these species comprised up to 20% of the total abundance of the benthic foraminiferal assemblage at 2168 m, but up to only 2% at 1145 m. The MPT extinction of 51 species represents ?20% of the total benthic foraminiferal diversity at bathyal depths in the North Atlantic (excluding the myriad of small unilocular forms). The extinction rate during the MPT was approximately 10 species per 0.1 myr, being one or two orders of magnitude greater than normal background turnover rates of deep-sea benthic foraminifera. Comparison of the precise timings of declines and disappearances (= highest occurrences) of each species shows that they were often diachronous between the two depths. The last of these species to disappear in the North Atlantic was Pleurostomella alternans at ~0.679 and ~0.694 Ma in Sites 980 and 982, respectively, which is in good agreement with the previously documented global "Stilostomella extinction" datum within the period 0.7-0.58 Ma. Comparison with similar studies in intermediate depth waters in the Southwest Pacific Gateway indicates that ~61% of the extinct species were common to both regions, and that although the pattern of pulsed decline was similar, the precise order and timing of the extinction of individual species were mostly different on opposite sides of the world. Previous studies have indicated that this extinct group of elongate, cylindrical foraminifera lived infaunally and had their greatest abundances in poorly ventilated, lower oxygen environments. This is supported by our study where there is a strong positive correlation (r = ~+ 0.8) between the flux of the extinction group and low-oxygen/high organic input species (such as Uvigerina, Bulimina and Bolivina) during the MPT, suggesting a close relationship with lower oxygen levels and high food supply to the sea floor. The absolute abundance, flux, and number of the extinction group of species show a progressive withdrawal pattern with major decreases occurring in cold periods with high d13C values. This might be related to increasing chemical ventilation of glacial intermediate water.

Seawater carbonate chemistry and cell numbers during experiments with dinoflagellates, 2007

The effects of dissolved inorganic carbon (DIC) on the growth of 3 red-tide dinoflagellates (Ceratium lineatum, Heterocapsa triquetra and Prorocentrum minimum) were studied at pH 8.0 and at higher pH levels, depending upon the pH tolerance of the individual species. The higher pH levels chosen for experiments were 8.55 for C. lineatum and 9.2 for the other 2 species. At pH 8.0, which approximates the pH found in the open sea, the maximum growth in all species was maintained until the total DIC concentration was reduced below ~0.4 and 0.2 mM for C. lineatum and the other 2 species, respectively. Growth compensation points (concentration of inorganic carbon needed for maintenance of cells) were reached at ~0.18 and 0.05 mM DIC for C. lineatum and the other 2 species, respectively. At higher pH levels, maximum growth rates were lower compared to growth at pH 8, even at very high DIC concentrations, indicating a direct pH effect on growth. Moreover, the concentration of bio-available inorganic carbon (CO2 + HCO3-) required for maintenance as well as the half-saturation constants were increased considerably at high pH compared to pH 8.0. Experiments with pH-drift were carried out at initial concentrations of 2.4 and 1.2 mM DIC to test whether pH or DIC was the main limiting factor at a natural range of DIC. Independent of the initial DIC concentrations, growth rates were similar in both incubations until pH had increased considerably. The results of this study demonstrated that growth of the 3 species was mainly limited by pH, while inorganic carbon limitation played a minor role only at very high pH levels and low initial DIC concentrations.

Seawater carbonate chemistry and growth rate during experiments with dinoflagellates, 2007

The effects of dissolved inorganic carbon (DIC) on the growth of 3 red-tide dinoflagellates (Ceratium lineatum, Heterocapsa triquetra and Prorocentrum minimum) were studied at pH 8.0 and at higher pH levels, depending upon the pH tolerance of the individual species. The higher pH levels chosen for experiments were 8.55 for C. lineatum and 9.2 for the other 2 species. At pH 8.0, which approximates the pH found in the open sea, the maximum growth in all species was maintained until the total DIC concentration was reduced below ~0.4 and 0.2 mM for C. lineatum and the other 2 species, respectively. Growth compensation points (concentration of inorganic carbon needed for maintenance of cells) were reached at ~0.18 and 0.05 mM DIC for C. lineatum and the other 2 species, respectively. At higher pH levels, maximum growth rates were lower compared to growth at pH 8, even at very high DIC concentrations, indicating a direct pH effect on growth. Moreover, the concentration of bio-available inorganic carbon (CO2 + HCO3-) required for maintenance as well as the half-saturation constants were increased considerably at high pH compared to pH 8.0. Experiments with pH-drift were carried out at initial concentrations of 2.4 and 1.2 mM DIC to test whether pH or DIC was the main limiting factor at a natural range of DIC. Independent of the initial DIC concentrations, growth rates were similar in both incubations until pH had increased considerably. The results of this study demonstrated that growth of the 3 species was mainly limited by pH, while inorganic carbon limitation played a minor role only at very high pH levels and low initial DIC concentrations.