Showing papers on "Benthic zone published in 2014"

Ocean Acidification in the Coastal Zone from an Organism's Perspective: Multiple System Parameters, Frequency Domains, and Habitats

Abstract: Multiple natural and anthropogenic processes alter the carbonate chemistry of the coastal zone in ways that either exacerbate or mitigate ocean acidification effects. Freshwater inputs and multiple acid-base reactions change carbonate chemistry conditions, sometimes synergistically. The shallow nature of these systems results in strong benthic-pelagic coupling, and marine invertebrates at different life history stages rely on both benthic and pelagic habitats. Carbonate chemistry in coastal systems can be highly variable, responding to processes with temporal modes ranging from seconds to centuries. Identifying scales of variability relevant to levels of biological organization requires a fuller characterization of both the frequency and magnitude domains of processes contributing to or reducing acidification in pelagic and benthic habitats. We review the processes that contribute to coastal acidification with attention to timescales of variability and habitats relevant to marine bivalves.

Benthic Exchange and Biogeochemical Cycling in Permeable Sediments

Abstract: The sandy sediments that blanket the inner shelf are situated in a zone where nutrient input from land and strong mixing produce maximum primary production and tight coupling between water column and sedimentary processes. The high permeability of the shelf sands renders them susceptible to pressure gradients generated by hydrodynamic and biological forces that modulate spatial and temporal patterns of water circulation through these sediments. The resulting dynamic three-dimensional patterns of particle and solute distribution generate a broad spectrum of biogeochemical reaction zones that facilitate effective decomposition of the pelagic and benthic primary production products. The intricate coupling between the water column and sediment makes it challenging to quantify the production and decomposition processes and the resultant fluxes in permeable shelf sands. Recent technical developments have led to insights into the high biogeochemical and biological activity of these permeable sediments and their role in the global cycles of matter.

Ecological effects of ocean acidification and habitat complexity on reef-associated macroinvertebrate communities

Abstract: The ecological effects of ocean acidification (OA) from rising atmospheric carbon dioxide (CO2) on benthic marine communities are largely unknown. We investigated in situ the consequences of long-term exposure to high CO2 on coral-reef-associated macroinvertebrate communities around three shallow volcanic CO2 seeps in Papua New Guinea. The densities of many groups and the number of taxa (classes and phyla) of macroinvertebrates were significantly reduced at elevated CO2 (425–1100 µatm) compared with control sites. However, sensitivities of some groups, including decapod crustaceans, ascidians and several echinoderms, contrasted with predictions of their physiological CO2 tolerances derived from laboratory experiments. High CO2 reduced the availability of structurally complex corals that are essential refugia for many reef-associated macroinvertebrates. This loss of habitat complexity was also associated with losses in many macroinvertebrate groups, especially predation-prone mobile taxa, including crustaceans and crinoids. The transition from living to dead coral as substratum and habitat further altered macroinvertebrate communities, with far more taxa losing than gaining in numbers. Our study shows that indirect ecological effects of OA (reduced habitat complexity) will complement its direct physiological effects and together with the loss of coral cover through climate change will severely affect macroinvertebrate communities in coral reefs.

Sea Ice Ecosystems

Abstract: Polar sea ice is one of the largest ecosystems on Earth. The liquid brine fraction of the ice matrix is home to a diverse array of organisms, ranging from tiny archaea to larger fish and invertebrates. These organisms can tolerate high brine salinity and low temperature but do best when conditions are milder. Thriving ice algal communities, generally dominated by diatoms, live at the ice/water interface and in recently flooded surface and interior layers, especially during spring, when temperatures begin to rise. Although protists dominate the sea ice biomass, heterotrophic bacteria are also abundant. The sea ice ecosystem provides food for a host of animals, with crustaceans being the most conspicuous. Uneaten organic matter from the ice sinks through the water column and feeds benthic ecosystems. As sea ice extent declines, ice algae likely contribute a shrinking fraction of the total amount of organic matter produced in polar waters.

Investigating hypoxia in aquatic environments: diverse approaches to addressing a complex phenomenon

Abstract: In this paper we provide an overview of new knowledge on oxygen depletion (hypoxia) and related phenomena in aquatic systems resulting from the EU-FP7 project HYPOX (“In situ monitoring of oxygen depletion in hypoxic ecosystems of coastal and open seas, and landlocked water bodies”, www.hypox.net). In view of the anticipated oxygen loss in aquatic systems due to eutrophication and climate change, HYPOX was set up to improve capacities to monitor hypoxia as well as to understand its causes and consequences. Temporal dynamics and spatial patterns of hypoxia were analyzed in field studies in various aquatic environments, including the Baltic Sea, the Black Sea, Scottish and Scandinavian fjords, Ionian Sea lagoons and embayments, and Swiss lakes. Examples of episodic and rapid (hours) occurrences of hypoxia, as well as seasonal changes in bottom-water oxygenation in stratified systems, are discussed. Geologically driven hypoxia caused by gas seepage is demonstrated. Using novel technologies, temporal and spatial patterns of watercolumn oxygenation, from basin-scale seasonal patterns to meter-scale sub-micromolar oxygen distributions, were resolved. Existing multidecadal monitoring data were used to demonstrate the imprint of climate change and eutrophication on long-term oxygen distributions. Organic and inorganic proxies were used to extend investigations on past oxygen conditions to centennial and even longer timescales that cannot be resolved by monitoring. The effects of hypoxia on faunal communities and biogeochemical processes were also addressed in the project. An investigation of benthic fauna is presented as an example of hypoxia-devastated benthic communities that slowly recover upon a reduction in eutrophication in a system where naturally occurring hypoxia overlaps with anthropogenic hypoxia. Biogeochemical investigations reveal that oxygen intrusions have a strong effect on the microbially mediated redox cycling of elements. Observations and modeling studies of the sediments demonstrate the effect of seasonally changing oxygen conditions on benthic mineralization pathways and fluxes. Data quality and access are crucial in hypoxia research. Technical issues are therefore also addressed, including the availability of suitable sensor technology to resolve the gradual changes in bottom-water oxygen in marine systems that can be expected as a result of climate change. Using cabled observatories as examples, we show how the benefit of continuous oxygen monitoring can be maximized by adopting proper quality control. Finally, we discuss strategies for state-of-the-art data archiving and dissemination in compliance with global standards, and how ocean observations can contribute to global earth observation attempts.

The future of the northeast Atlantic benthic flora in a high CO2 world

Abstract: Seaweed and seagrass communities in the northeast Atlantic have been profoundly impacted by humans, and the rate of change is accelerating rapidly due to runaway CO2 emissions and mounting pressures on coastlines associated with human population growth and increased consumption of finite resources. Here, we predict how rapid warming and acidification are likely to affect benthic flora and coastal ecosystems of the northeast Atlantic in this century, based on global evidence from the literature as interpreted by the collective knowledge of the authorship. We predict that warming will kill off kelp forests in the south and that ocean acidification will remove maerl habitat in the north. Seagrasses will proliferate, and associated epiphytes switch from calcified algae to diatoms and filamentous species. Invasive species will thrive in niches liberated by loss of native species and spread via exponential development of artificial marine structures. Combined impacts of seawater warming, ocean acidification, and increased storminess may replace structurally diverse seaweed canopies, with associated calcified and noncalcified flora, with simple habitats dominated by noncalcified, turf-forming seaweeds.

Hypoxia in the Baltic Sea: Biogeochemical Cycles, Benthic Fauna, and Management

Abstract: Hypoxia has occurred intermittently over the Holocene in the Baltic Sea, but the recent expansion from less than 10 000 km 2 before 1950 to (60 000 km 2 since 2000 is mainly caused by enhanced nutrient inputs from land and atmosphere. With worsening hypoxia, the role of sediments changes from nitrogen removal to nitrogen release as ammonium. At present, denitrification in the water column and sediments is equally important. Phos- phorus is currently buried in sediments mainly in organic form, with an additional contribution of reduced Fe-phos- phate minerals in the deep anoxic basins. Upon the tran- sition to oxic conditions, a significant proportion of the organic phosphorus will be remineralized, with the phos- phorus then being bound to iron oxides. This iron-oxide bound phosphorus is readily released to the water column upon the onset of hypoxia again. Important ecosystems services carried out by the benthic fauna, including bio- geochemical feedback-loops and biomass production, are also lost with hypoxia. The results provide quantitative knowledge of nutrient release and recycling processes under various environmental conditions in support of decision support tools underlying the Baltic Sea Action Plan.

How much did Glacial North Atlantic Water shoal

Abstract: Observations of δ13C and Cd/Ca from benthic foraminifera have been interpreted to reflect a shoaling of northern source waters by about 1000 m during the Last Glacial Maximum, with the degree of shoaling being significant enough for the water mass to be renamed Glacial North Atlantic Intermediate Water. These nutrient tracers, however, may not solely reflect changes in water mass distributions. To quantify the distribution of Glacial North Atlantic Water, we perform a glacial water mass decomposition where the sparsity of data, geometrical constraints, and nonconservative tracer effects are taken into account, and the extrapolation for the unknown water mass end-members is guided by the modern-day circulation. Under the assumption that the glacial sources of remineralized material are similar to that of the modern day, we find a steady solution consistent with 241 δ13C, 87 Cd/Ca, and 174 δ18O observations and their respective uncertainties. The water mass decomposition indicates that the core of Glacial North Atlantic Water shoals and southern source water extends in greater quantities into the abyssal North Atlantic, as previously inferred. The depth of the deep northern-southern water mass interface and the volume of North Atlantic Water, however, are not grossly different from that of the modern day. Under this scenario, the vertical structure of glacial δ13C and Cd/Ca is primarily due to the greater accumulation of nutrients in lower North Atlantic Water, which may be a signal of the hoarding of excess carbon from the atmosphere by the glacial Atlantic.

Benthic Coral Reef Calcium Carbonate Dissolution in an Acidifying Ocean

Abstract: The authors of this Review argue that changes to carbonate dissolution in an acidifying ocean, which have been relatively overlooked, are potentially more important than calcification for the future accretion and survival of coral reef ecosystems.

Global reductions in seafloor biomass in response to climate change

Abstract: Seafloor organisms are vital for healthy marine ecosystems, contributing to elemental cycling, benthic remineralization, and ultimately sequestration of carbon. Deep-sea life is primarily reliant on the export flux of particulate organic carbon from the surface ocean for food, but most ocean biogeochemistry models predict global decreases in export flux resulting from 21st century anthropogenically induced warming. Here we show that decadal-to-century scale changes in carbon export associated with climate change lead to an estimated 5.2% decrease in future (2091–2100) global open ocean benthic biomass under RCP8.5 (reduction of 5.2 Mt C) compared with contemporary conditions (2006–2015). Our projections use multi-model mean export flux estimates from eight fully coupled earth system models, which contributed to the Coupled Model Intercomparison Project Phase 5, that have been forced by high and low representative concentration pathways (RCP8.5 and 4.5, respectively). These export flux estimates are used in conjunction with published empirical relationships to predict changes in benthic biomass. The polar oceans and some upwelling areas may experience increases in benthic biomass, but most other regions show decreases, with up to 38% reductions in parts of the northeast Atlantic. Our analysis projects a future ocean with smaller sized infaunal benthos, potentially reducing energy transfer rates though benthic multicellular food webs. More than 80% of potential deep-water biodiversity hotspots known around the world, including canyons, seamounts, and cold-water coral reefs, are projected to experience negative changes in biomass. These major reductions in biomass may lead to widespread change in benthic ecosystems and the functions and services they provide.

Global warming and mass mortalities of benthic invertebrates in the Mediterranean Sea.

Abstract: Satellite data show a steady increase, in the last decades, of the surface temperature (upper few millimetres of the water surface) of the Mediterranean Sea. Reports of mass mortalities of benthic marine invertebrates increased in the same period. Some local studies interpreted the two phenomena in a cause-effect fashion. However, a basin-wide picture of temperature changes combined with a systematic assessment on invertebrate mass mortalities was still lacking. Both the thermal structure of the water column in the Mediterranean Sea over the period 1945–2011 and all documented invertebrate mass mortality events in the basin are analysed to ascertain if: 1- documented mass mortalities occurred under conditions of positive temperature trends at basin scale, and 2- atypical thermal conditions were registered at the smaller spatial and temporal scale of mass mortality events. The thermal structure of the shallow water column over the last 67 years was reconstructed using data from three public sources: MEDAR-MEDATLAS, World Ocean Database, MFS-VOS programme. A review of the mass mortality events of benthic invertebrates at Mediterranean scale was also carried out. The analysis of in situ temperature profiles shows that the Mediterranean Sea changed in a non-homogeneous fashion. The frequency of mass mortalities is increasing. The areas subjected to these events correspond to positive thermal anomalies. Statistically significant temperature trends in the upper layers of the Mediterranean Sea show an increase of up to 0.07°C/yr for a large fraction of the basin. Mass mortalities are consistent with both the temperature increase at basin scale and the thermal changes at local scale, up to 5.2°C. Our research supports the existence of a causal link between positive thermal anomalies and observed invertebrate mass mortalities in the Mediterranean Sea, invoking focused mitigation initiatives in sensitive areas.

Trophic interactions of fish communities at midwater depths enhance long-term carbon storage and benthic production on continental slopes

Abstract: Biological transfer of nutrients and materials between linked ecosystems influences global carbon budgets and ecosystem structure and function. Identifying the organisms or functional groups that are responsible for nutrient transfer, and quantifying their influence on ecosystem structure and carbon capture is an essential step for informed management of ecosystems in physically distant, but ecologically linked areas. Here, we combine natural abundance stable isotope tracers and survey data to show that mid-water and bentho-pelagic-feeding demersal fishes play an important role in the ocean carbon cycle, bypassing the detrital particle flux and transferring carbon to deep long-term storage. Global peaks in biomass and diversity of fishes at mid-slope depths are explained by competitive release of the demersal fish predators of mid-water organisms, which in turn support benthic fish production. Over 50% of the biomass of the demersal fish community at depths between 500 and 1800 m is supported by biological rather than detrital nutrient flux processes, and we estimate that bentho-pelagic fishes from the UK–Irish continental slope capture and store a volume of carbon equivalent to over 1 million tonnes of CO2 every year.

Explaining bathymetric diversity patterns in marine benthic invertebrates and demersal fishes: physiological contributions to adaptation of life at depth

Abstract: Bathymetric biodiversity patterns of marine benthic invertebrates and demersal fishes have been identified in the extant fauna of the deep continental margins. Depth zonation is widespread and evident through a transition between shelf and slope fauna from the shelf break to 1000 m, and a transition between slope and abyssal fauna from 2000 to 3000 m; these transitions are characterised by high species turnover. A unimodal pattern of diversity with depth peaks between 1000 and 3000 m, despite the relatively low area represented by these depths. Zonation is thought to result from the colonisation of the deep sea by shallow-water organisms following multiple mass extinction events throughout the Phanerozoic. The effects of low temperature and high pressure act across hierarchical levels of biological organisation and appear sufficient to limit the distributions of such shallow-water species. Hydrostatic pressures of bathyal depths have consistently been identified experimentally as the maximum tolerated by shallow-water and upper bathyal benthic invertebrates at in situ temperatures, and adaptation appears required for passage to deeper water in both benthic invertebrates and demersal fishes. Together, this suggests that a hyperbaric and thermal physiological bottleneck at bathyal depths contributes to bathymetric zonation. The peak of the unimodal diversity–depth pattern typically occurs at these depths even though the area represented by these depths is relatively low. Although it is recognised that, over long evolutionary time scales, shallow-water diversity patterns are driven by speciation, little consideration has been given to the potential implications for species distribution patterns with depth. Molecular and morphological evidence indicates that cool bathyal waters are the primary site of adaptive radiation in the deep sea, and we hypothesise that bathymetric variation in speciation rates could drive the unimodal diversity–depth pattern over time. Thermal effects on metabolic-rate-dependent mutation and on generation times have been proposed to drive differences in speciation rates, which result in modern latitudinal biodiversity patterns over time. Clearly, this thermal mechanism alone cannot explain bathymetric patterns since temperature generally decreases with depth. We hypothesise that demonstrated physiological effects of high hydrostatic pressure and low temperature at bathyal depths, acting on shallow-water taxa invading the deep sea, may invoke a stress–evolution mechanism by increasing mutagenic activity in germ cells, by inactivating canalisation during embryonic or larval development, by releasing hidden variation or mutagenic activity, or by activating or releasing transposable elements in larvae or adults. In this scenario, increased variation at a physiological bottleneck at bathyal depths results in elevated speciation rate. Adaptation that increases tolerance to high hydrostatic pressure and low temperature allows colonisation of abyssal depths and reduces the stress–evolution response, consequently returning speciation of deeper taxa to the background rate. Over time this mechanism could contribute to the unimodal diversity–depth pattern.

Deglacial whole-ocean δ13C change estimated from 480 benthic foraminiferal records

Abstract: Terrestrial carbon storage is dramatically decreased during glacial periods due to cold temperatures, increased aridity, and the presence of large ice sheets on land. Most of the carbon released by the terrestrial biosphere is stored in the ocean, where the light isotopic signature of terrestrial carbon is observed as a 0.32–0.7‰ depletion in benthic foraminiferal δ13C. The wide range in estimated δ13C change results from the use of different subsets of benthic δ13C data and different methods of weighting the mean δ13C by volume. We present a more precise estimate of glacial-interglacial δ13C change of marine dissolved inorganic carbon using benthic Cibicidoides spp. δ13C records from 480 core sites (more than 3 times as many sites as previous studies). We divide the ocean into eight regions to generate linear regressions of regional δ13C versus depth for the Late Holocene (0–6 ka) and Last Glacial Maximum (19–23 ka) and estimate a mean δ13C decrease of 0.38 ± 0.08‰ (2σ) for 0.5–5 km. Estimating large uncertainty ranges for δ13C change in the top 0.5 km, below 5 km, and in the Southern Ocean, we calculate a whole-ocean change of 0.34 ± 0.19‰. This implies a terrestrial carbon change that is consistent with recent vegetation model estimates of 330–694 Gt C. Additionally, we find that a well-constrained surface ocean δ13C change is essential for narrowing the uncertainty range of estimated whole-ocean δ13C change.

Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance

Abstract: A number of mechanisms result in a feedback between water clarity and macrophytes and, consequently, the occurrence of alternative stable states in shallow lakes. We hypothesize that bottom-up mechanisms and interactions within the benthic food web are more important in a charophyte-dominated clear-water state, while top-down mechanism and interactions in the planktonic food web prevail at angiosperm dominance. Charophytes, which dominate at lower nutrient concentrations and develop higher densities than most angiosperms, can have a higher influence on sedimentation, resuspension, and water column nutrients. During dominance of dense submerged vegetation like charophytes, zooplankton can be hampered by low food quality and quantity and by high predation pressure from juvenile fish, which in turn are favoured by the high refuge potential of this vegetation. Grazing pressure from zooplankton on phytoplankton can therefore be low in charophytes, but the main feedback in angiosperm-dominated ecosystems. Charophytes offer a higher surface than most angiosperms to periphyton, which favors benthic invertebrates. These support macrophytes by grazing periphyton and constitute a central link in a trophic cascade from fish to periphyton and macrophytes. To test these hypotheses, more experiments and field measurements comparing the effect of charophytes and angiosperms on water clarity are needed.

Influence of benthic macrofauna community shifts on ecosystem functioning in shallow estuaries

Abstract: We identify how ecosystem functioning in shallow estuaries is affected by shifts in benthic fauna communities. We use the shallow estuary, Odense Fjord, Denmark, as a case study to test our hypotheses that (1) shifts in benthic fauna composition and species functional traits affect biogeochemical cycling with cascading effects on ecological functioning, which may (2) modulate pelagic primary productivity with feedbacks to the benthic system. Odense Fjord is suitable because it experienced dramatic shifts in benthic fauna community structure from 1998 to 2008. We focused on infaunal species with emphasis on three dominating burrow-dwelling polychaetes: the native Nereis (Hediste) diversicolor and Arenicola marina, and the invasive Marenzelleria viridis. The impact of functional traits in the form of particle reworking and ventilation on biogeochemical cycles, i.e. sediment metabolism and nutrient dynamics, was determined from literature data. Historical records of summer nutrient levels in the water column of the inner Odense Fjord show elevated concentrations of NH4+ and NO3- (DIN) during the years 2004-2006, exactly when the N. diversicolor population declined and A. marina and M. viridis populations expanded dramatically. In support of our first hypothesis, we show that excess NH4+ delivery from the benthic system during the A. marina and M. viridis expansion period enriched the overlying water in DIN and stimulated phytoplankton concentration. The altered benthic-pelagic coupling and stimulated pelagic production may, in support of our second hypothesis, have feedback to the benthic system by changing the deposition of organic material. We therefore advice to identify the exact functional traits of the species involved in a community shift before studying its impact on ecosystem functioning. We also suggest studying benthic community shifts in shallow environments to obtain knowledge about the drivers and controls before exploring deep-water environments.

Lake benthic algae: toward an understanding of their ecology

Abstract: Lake phytoplankton studies outnumber studies on lake periphyton by an order of magnitude, and most periphyton research has been done in streams. Most benthic algal taxa found in lakes also can be found in lotic systems, but assemblages and taxa differ in a number of ways. The ecological characteristics of some lake benthic algae reflect habitat coupling. Littoral zones (benthic areas above the light compensation depth) are areas of high productivity and biodiversity. The proportion of benthic and planktonic primary production (autotrophic structure) is a key ecosystem property. The distribution of lake benthic algae is markedly influenced by the depth gradient and substratum, and assemblages change with depth from epilithic and epiphytic rheobionts to epipsammic and epipelic limnobionts. At shallow depths, periphyton must cope with the effects of high radiation, water-level fluctuations, wave action (e.g., desiccation, ultraviolet radiation exposure, shear stress), and seasonal shifts in temperatu...

Modelling Community Structure in Freshwater Ecosystems

Abstract: This volume presents approaches and methodologies for predicting the structure and diversity of key aquatic communities (namely, diatoms, benthic macroinvertebrates and fish), under natural conditions and under man-made disturbance. The intent is to offer an organized means for modeling, evaluating and restoring freshwater ecosystems.

Exploring local adaptation and the ocean acidification seascape - studies in the California Current Large Marine Ecosystem

Abstract: . The California Current Large Marine Ecosystem (CCLME), a temperate marine region dominated by episodic upwelling, is predicted to experience rapid environmental change in the future due to ocean acidification. The aragonite saturation state within the California Current System is predicted to decrease in the future with near-permanent undersaturation conditions expected by the year 2050. Thus, the CCLME is a critical region to study due to the rapid rate of environmental change that resident organisms will experience and because of the economic and societal value of this coastal region. Recent efforts by a research consortium – the Ocean Margin Ecosystems Group for Acidification Studies (OMEGAS) – has begun to characterize a portion of the CCLME; both describing the spatial mosaic of pH in coastal waters and examining the responses of key calcification-dependent benthic marine organisms to natural variation in pH and to changes in carbonate chemistry that are expected in the coming decades. In this review, we present the OMEGAS strategy of co-locating sensors and oceanographic observations with biological studies on benthic marine invertebrates, specifically measurements of functional traits such as calcification-related processes and genetic variation in populations that are locally adapted to conditions in a particular region of the coast. Highlighted in this contribution are (1) the OMEGAS sensor network that spans the west coast of the US from central Oregon to southern California, (2) initial findings of the carbonate chemistry amongst the OMEGAS study sites, and (3) an overview of the biological data that describes the acclimatization and the adaptation capacity of key benthic marine invertebrates within the CCLME.

Nearshore energy subsidies support Lake Michigan fishes and invertebrates following major changes in food web structure

Abstract: Aquatic food webs that incorporate multiple energy channels (e.g., nearshore benthic and pelagic) with varying productivity and turnover rates convey stability to biological communities by providing independent energy sources. Within the Lake Michigan food web, invasive dreissenid mussels have caused rapid changes to food web structure and potentially altered the channels through which consumers acquire energy. We used stable C and N isotopes to determine how Lake Michigan food web structure has changed in the past decade, coincident with the expansion of dreissenid mussels, decreased pelagic phytoplankton production, and increased nearshore benthic algal production. Fish and invertebrate samples collected from sites around Lake Michigan were analyzed to determine taxa-specific 13C:12C (δ13C) and 15N:14N (δ15N) ratios. Sampling took place during two distinct periods, 2002–2003 and 2010–2012, that spanned the period of dreissenid expansion, and included nearshore, pelagic and profundal fish and invertebrat...

Recovery of benthic marine communities from the end-Permian mass extinction at the low latitudes of eastern Panthalassa

Abstract: Based on the quantitative community analysis using species-level identifications, we track the restoration of benthic ecosystems after the end-Permian mass extinction throughout the Lower Triassic of the western USA. New data on the palaeoecology of the Thaynes Group and Sinbad Formation are provided, which fill a gap between the recently studied palaeoecology of the Griesbachian-Dienerian Dinwoody Formation and the Spathian Virgin Formation. In the Sinbad Formation and Thaynes Group, 17 species (12 genera) of bivalves, 7 species and genera of gastropods and 2 species and genera of brachiopods are recognized. The new bivalve genus Confusionella (Pteriidae) is described. A comprehensive review of the whole Lower Triassic succession of benthic ecosystems of the western USA indicates that midand inner shelf environments show incipient recovery signals around the Griesbachian-Dienerian transition, during the Smithian and, most profound, during the early Spathian. Ecological data from youngest strata of the Dinwoody Formation as well as stratigraphic ranges of species suggest that the late Dienerian was likely a time interval of environmental stress for benthic ecosystems. Despite some evidence for short-term environmental disturbances (e.g. shift of dominant taxa, transient drop in alpha-diversity) during the Smithian-Spathian transition, benthic ecosystems did not show any notable taxonomic turnover at that time, in contrast to the major crisis that affected ammonoids and conodonts. Whereas alpha-diversity of benthic communities generally increased throughout the Early Triassic, beta-diversity remained low, which reflects a persistently wide environmental range of benthic species. This observation is in accordance with a recently proposed model that predicts a time lag between increasing within-habitat diversity (alphadiversity) and the onset of taxonomic differentiation between habitats (beta-diversity) during biotic recoveries from mass extinction events. The observation that beta-diversity had not significantly increased during the Early Triassic might also provide an explanation for the comparably sluggish increase in benthic diversity during that time, which has previously been attributed to persistent environmental stress.

Effect of hypoxia and anoxia on invertebrate behaviour: Ecological perspectives from species to community level

Abstract: Coastal hypoxia and anoxia have become a global key stressor to marine ecosystems, with almost 500 dead zones recorded worldwide. By triggering cascading effects from the individual organism to the community-and ecosystem level, oxygen depletions threaten marine biodiversity and can alter ecosystem structure and function. By integrating both physiological function and ecological processes, animal behaviour is ideal for assessing the stress state of benthic macrofauna to low dissolved oxygen. The initial response of organisms can serve as an early warning signal, while the successive behavioural reactions of key species indicate hypoxia levels and help assess community degradation. Here we document the behavioural responses of a representative spectrum of benthic macrofauna in the natural setting in the Northern Adriatic Sea (Mediterranean). We experimentally induced small-scale anoxia with a benthic chamber in 24m depth to overcome the difficulties in predicting the onset of hypoxia, which often hinders full documentation in the field. The behavioural reactions were documented with a time-lapse camera. Oxygen depletion elicited significant and repeatable changes in general (visibility, locomotion, body movement and posture, location) and species-specific reactions in virtually all organisms (302 individuals from 32 species and 2 species groups). Most atypical (stress) behaviours were associated with specific oxygen thresholds: arm-tipping in the ophiuroid Ophiothrix quinquemaculata, for example, with the onset of mild hypoxia (< 2mLO(2) L-1), the emergence of polychaetes on the sediment surface with moderate hypoxia (< 1mLO(2) L-1), the emergence of the infaunal sea urchin Schizaster canaliferus on the sediment with severe hypoxia (< 0.5mLO(2) L-1) and heavy body rotations in sea anemones with anoxia. Other species changed their activity patterns, for example the circadian rhythm in the hermit crab Paguristes eremita or the bioherm-associated crab Pisidia longimana. Intra-and interspecific reactions were weakened or changed: decapods ceased defensive and territorial behaviour, and predator-prey interactions and relationships shifted. This nuanced scale of resolution is a useful tool to interpret present benthic community status (behaviour) and past mortalities (community composition, e.g. survival of tolerant species). This information on the sensitivity (onset of stress response), tolerance (mortality, survival), and characteristics (i. e. life habit, functional role) of key species also helps predict potential future changes in benthic structure and ecosystem functioning. This integrated approach can transport complex ecological processes to the public and decision-makers and help define specific monitoring, assessment and conservation plans