Isotopes Trace Biogeochemistry and Sources of Cu and Zn in an intertidal soil

River floodplain soils are sinks and potential sources for toxic trace metals like Cu and Zn. We hypothesize that stable Cu and Zn isotope ratios reflect both the mobilization and the sources of metals. We determined the soil properties, the concentrations and partitioning of Cu and Zn, and variations in δ65Cu and δ66Zn values in a core obtained from an Aquic Udifluvent developed on a freshwater intertidal mudflat of the River Elbe, Germany. The core was sampled at 2 cm intervals to a depth of 34 cm, which corresponds to approximately 9 yr of sedimentation. Elevated concentrations of Cu (up to 320 μg g−1) and Zn (up to 2080 μg g−1) indicated anthropogenic pollution. At the time of sampling the redox conditions changed from oxic (Eh 200 to 400 mV, above 22 cm deep) to strongly anoxic conditions (-100 to -200 mV, below 22 cm deep). The δ65Cu values varied systematically with depth (from -0.02 to 0.16‰) and were correlated with the Fe, C, and N concentrations. Although pre-depositional variations cannot be ruled out, the systematic variation with depth suggests post-sedimentation fractionation of δ65Cu in response to seasonally variable organic matter deposition and redox conditions. In contrast, the δ66ZnIRMM values were uniform (from -0.07 to 0.01‰) throughout the core, indicating that the Zn isotopes did not significantly fractionate after deposition and that the Zn sources were homogeneous throughout the sedimentation.

Growth and Persistence of Diverse Intertidal Crusts: Survival of the Slow in a Fast-Paced World

Encrusting algae are conspicuous components of hard-substratum benthic communities in the photic zone despite being poor competitors and slow growers. Little is known about their growth rates or about mechanisms controlling key processes such as wound healing and surviving overgrowth. We manipulated 12 crustose species (including red and brown algae and a lichen) from the intertidal zone of Washington, USA, studying their varying responses to identical experimental conditions. Three of 8 crust species tested showed increased growth rates with size. Species healed from standardized wounds at different rates and using different mechanisms (e.g. lateral vs vertical regeneration) as seen in cross-sections. Three species showed altered growth rates at unwounded margins of wounded crusts, suggesting intrathallus communication. Year-long experiments involving simulated overgrowth showed that some species can maintain healthy tissue in a covered area, and one (the coralline Lithothamnion phymatodeum) even grew new tissue there. Other species gradually lost color, thickness, and area in covered areas. Hildenbrandia occidentalis survived remarkably well when covered, possibly due to its very slow growth and low metabolic demand. One suggested mechanism underlying the high variation in responses among crusts is the degree to which their thalli may be anatomically integrated by features such as cell fusions; physiological work testing translocation via these features is needed. Other mechanisms allowing persistence include rapid wound healing and frequent recruitment.

Physiological Responses to Environmental Variation in Intertidal Red Algae: Does Thallus Morphology Matter?

Morphological variation within and among many species of algae show correlated life history traits. The trade-offs of Life history traits among different morphs are presumed to be determined by morphology. Form-function hypotheses also predict that algae of different morphological groups exhibit different tolerances to physiological stress, whereas algae within a morphological group respond similarly to stress. We tested this hypothesis by comparing photosynthetic and respiratory responses to variation in season, light, temperature, desiccation and freezing among the morphologically similar fronds of Chondrus crispus and Mastocarpus stellatus and the alternate stage crust of M. stellatus. Physiological differences between fronds of the 2 species and crusts and fronds were consistent with their patterns of distribution and abundance in the intertidal zone. However, there was no clear relationship between algal morphology and physiological response to environmental variation. These results suggest that among macroalgae the correlation between Life history traits and morphology is not always causal. Rather, the link between life history traits and morphology is constrained by the extent to which physiological characteristics codetermine these features.

Two intertidal, non-calcifying macroalgae (Palmaria palmata and Saccharina latissima) show complex and variable responses to short-term CO2 acidification

Ocean acidification, the result of increased dissolution of carbon dioxide (CO2) in seawater, is a leading subject of current research. The effects of acidification on non-calcifying macroalgae are, however, still unclear. The current study reports two 1-month studies using two different macroalgae, the red alga Palmaria palmata (Rhodophyta) and the kelp Saccharina latissima (Phaeophyta), exposed to control (pHNBS = 8.04) and increased (pHNBS = 7.82) levels of CO2-induced seawater acidification. The impacts of both increased acidification and time of exposure on net primary production (NPP), respiration (R), dimethylsulphoniopropionate (DMSP) concentrations, and algal growth have been assessed. In P. palmata, although NPP significantly increased during the testing period, it significantly decreased with acidification, whereas R showed a significant decrease with acidification only. S. latissima significantly increased NPP with acidification but not with time, and significantly increased R with both acidification and time, suggesting a concomitant increase in gross primary production. The DMSP concentrations of both species remained unchanged by either acidification or through time during the experimental period. In contrast, algal growth differed markedly between the two experiments, in that P. palmata showed very little growth throughout the experiment, while S. latissima showed substantial growth during the course of the study, with the latter showing a significant difference between the acidified and control treatments. These two experiments suggest that the study species used here were resistant to a short-term exposure to ocean acidification, with some of the differences seen between species possibly linked to different nutrient concentrations between the experiments.

Barnacle recurit density, biomass, and reproductive output measured in rocky intertidal habitats in Glasgow Head, on the Atlantic coast of Nova Scotia, Canada

Through a field experiment, we show that a predator has negative nonconsumptive effects (NCEs) on different life-history stages of the same prey species. Shortly before the recruitment season of the barnacle Semibalanus balanoides (May-June), we established experimental cages in rocky intertidal habitats in Nova Scotia, Canada. The cages were used to manipulate the presence and absence of dogwhelks, Nucella lapillus, the main predators of barnacles. At the centre of each cage, we installed a tile where barnacle pelagic larvae could settle and the resulting recruits grow. Mesh prevented caged dogwhelks from accessing the tiles, but allowed waterborne dogwhelk cues to reach the tiles. Results in May indicated that barnacle larvae settled preferentially on tiles from cages without dogwhelks. In November, at the end of the dogwhelk activity period and once the barnacle recruits had grown to adult size, barnacle body mass was lower in the presence of dogwhelks. This limitation may have resulted from a lower barnacle feeding activity with nearby dogwhelks, as found by a previous study. The observed larval and adult responses in barnacles are consistent with attempts to decrease predation risk. November data also indicated that dogwhelk cues limited barnacle reproductive output, a possible consequence of the limited growth of barnacles. Overall, this study suggests that a predator species might influence trait evolution in a prey species through NCEs on different life-history stages.

Chlorophyll and true-color maps of sediments from the intertidal zones in Winyah Bay, South Carolina, USA and List, Sylt, Germany

We used hyperspectral imaging to study short-term effects of bioturbation by lugworms (Arenicola marina) on the surficial biomass of microphytobenthos (MPB) in permeable marine sediments. Within days to weeks after the addition of a lugworm to a homogenized and recomposed sediment, the average surficial MPB biomass and its spatial heterogeneity were, respectively, 150 - 250% and 280% higher than in sediments without lugworms. The surficial sediment area impacted by a single medium-sized lugworm (~4 g wet weight) over this time-scale was at least 340 cm**2. While sediment reworking was the primary cause of the increased spatial heterogeneity, experiments with lugworm-mimics together with modeling showed that bioadvective porewater transport from depth to the sediment surface, as induced by the lugworm ventilating its burrow, was the main cause of the increased surficial MPB biomass. Although direct measurements of nutrient fluxes are lacking, our present data show that enhanced advective supply of nutrients from deeper sediment layers induced by faunal ventilation is an important mechanism that fuels high primary productivity at the surface of permeable sediments even though these systems are generally characterized by low standing stocks of nutrients and organic material.

Community structure and biodiversity in marine benthic communities during experiments, 2011

A mesocosm experiment was conducted to quantify the effects of reduced pH and elevated temperature on an intact marine invertebrate community. Standardised faunal communities, collected from the extreme low intertidal zone using artificial substrate units, were exposed to one of eight nominal treatments (four pH levels: 8.0, 7.7, 7.3 and 6.7, crossed with two temperature levels: 12 and 16°C). After 60 days exposure communities showed significant changes in structure and lower diversity in response to reduced pH. The response to temperature was more complex. At higher pH levels (8.0 and 7.7) elevated temperature treatments contained higher species abundances and diversity than the lower temperature treatments. In contrast, at lower pH levels (7.3 and 6.7), elevated temperature treatments had lower species abundances and diversity than lower temperature treatments. The species losses responsible for these changes in community structure and diversity were not randomly distributed across the different phyla examined. Molluscs showed the greatest reduction in abundance and diversity in response to low pH and elevated temperature, whilst annelid abundance and diversity was mostly unaffected by low pH and was higher at the elevated temperature. The arthropod response was between these two extremes with moderately reduced abundance and diversity at low pH and elevated temperature. Nematode abundance increased in response to low pH and elevated temperature, probably due to the reduction of ecological constraints, such as predation and competition, caused by a decrease in macrofaunal abundance. This community-based mesocosm study supports previous suggestions, based on observations of direct physiological impacts, that ocean acidification induced changes in marine biodiversity will be driven by differential vulnerability within and between different taxonomical groups. This study also illustrates the importance of considering indirect effects that occur within multispecies assemblages when attempting to predict the consequences of ocean acidification and global warming on marine communities.

Data to investigate the role of Pomatoschistus microps in intertidal food webs, sampled off the island of Sylt

Ecological network analysis (ENA) was used to study the effects of Pomatoschistus microps on energy transport through the food web, its impact on other compartments and its possible role as a keystone species in the trophic webs of an Arenicola tidal flat ecosystem and a sparse Zostera noltii bed ecosystem. Three ENA models were constructed: (a) model 1 contains data of the original food web from prior research in the investigated area by Baird et al. (2007), (b) an updated model 2 which included biomass and diet data of P. microps from recent sampling, and (c) model 3 simulating a food web without P. microps. A comparison of energy transport between the different models revealed that more energy is transported from lower trophic levels up the food chain, in the presence of P. microps (models 1 and 2) than in its absence (model 3). Calculations of the keystone index (KSi) revealed the high overall impact (measured as eps_i) of this fish species on food webs. In model 1, P. microps was assigned a low KSi in the Arenicola flat and in the sparse Z. noltii bed. Calculations in model 2 ranked P. microps first for keystoneness and eps_i in both communities, the Arenicola flat and the sparse Z. noltii bed. Taken together, our results give insight into the role of P. microps when considering a whole food web and reveal direct and indirect trophic interactions of this small-sized fish species. These results might illustrate the impact and importance of abundant, widespread species in food webs and facilitate further investigations.

(Table 1) Grain size parameters of intertidal channel sand bars near Sylt, North-Frisian Wadden Sea

The aim of the present study is to investigate directional asymmetric properties and internal structures of the bedforms on the intertidal sand bars in comparison with the migration problems of the sand bodies developed in the channel systems of the tidal basin off the west coast of Schleswig-Holstein. The tidal channel sand bodies studied have 'V'-shaped outlines and are asymmetric in cross-section. Based on such knowledge it was hoped to understand and find possible factors for application to recent and ancient tidal depositional environments. The V-shaped intertidal channel sand bodies developed in the tidal environments between Sylt and Föhr Island are constantly migrating sand bars. The migration directions are in good agreement with the resultant vector mean directions of internal cross-stratification structures of asymmetric sedimentary bedforms. Finally, it is shown that the orientation of the apex of V-shaped sand bar as an equilibrium form alone can not indicate the migration direction, but that the orientation of the resultant vector mean of internal structures of sedimentary bedforms does indicate the migration direction. Based on the analyses of textural parameters of the migrating intertidal bar sands, it seems that sands of typical intertidal sand bars are negatively skewed and well sorted. The high rounding of quarz sand grains of these tidal channel sand bars seems to be an additional characteristical criterion for tidal depositional environments, as also indicated by Balazs and Klein (1972).

Effects of elevated CO2 and temperature on an intertidal meiobenthic community

In the near future, the marine environment is likely to be subjected to simultaneous increases in temperature and decreased pH. The potential effects of these changes on intertidal, meiofaunal assemblages were investigated using a mesocosm experiment. Artificial Substrate Units containing meiofauna from the extreme low intertidal zone were exposed for 60 days to eight experimental treatments (four replicates for each treatment) comprising four pH levels: 8.0 (ambient control), 7.7 & 7.3 (predicted changes associated with ocean acidification), and 6.7 (CO2 point-source leakage from geological storage), crossed with two temperatures: 12 °C (ambient control) and 16 °C (predicted). Community structure, measured using major meiofauna taxa was significantly affected by pH and temperature. Copepods and copepodites showed the greatest decline in abundance in response to low pH and elevated temperature. Nematodes increased in abundance in response to low pH and temperature rise, possibly caused by decreased predation and competition for food owing to the declining macrofauna density. Nematode species composition changed significantly between the different treatments, and was affected by both seawater acidification and warming. Estimated nematode species diversity, species evenness, and the maturity index, were substantially lower at 16 °C, whereas trophic diversity was slightly higher at 16 °C except at pH 6.7. This study has demonstrated that the combination of elevated levels of CO2 and ocean warming may have substantial effects on structural and functional characteristics of meiofaunal and nematode communities, and that single stressor experiments are unlikely to encompass the complexity of abiotic and biotic interactions. At the same time, ecological interactions may lead to complex community responses to pH and temperature changes in the interstitial environment.