Top–down control by great blue herons Ardea herodias regulates seagrass‐associated epifauna

Top predators can influence the structure and function of plant and animal communities. In coastal marine systems, fish, shark and mammal population declines are major drivers of recent ecosystem‐level change. Cascading effects of predatory wading birds, however, are less understood, even though wading bird populations have declined in many regions. We quantified the effects of predation by the piscivorous great blue heron Ardea herodias fannini on fish, invertebrates and epiphytes living in eelgrass Zostera marina. We found that herons forage on benthic fish in seagrass meadows, and foraging intensity increased from late spring until midsummer. When we experimentally excluded herons, benthic fish abundance increased, and the invertebrate assemblage shifted to more shrimp‐dominated assemblages while grazing gammarid amphipod abundance declined. These shifts were associated with reduced epiphyte abundance when herons were excluded, reflecting a four‐level trophic cascade and mediated by shifts in the grazer assemblage. In summary, we found that a piscivorous wading bird species exerts top down control in a subtidal seagrass ecosystem. Losses and recovery of wading birds could have ecosystem‐level ecological consequences that may need to be considered in the context of concern for overfishing and predator recovery in marine coastal management.     

Possible effects of global environmental changes on Antarctic benthos: a synthesis across five major taxa

Because of the unique conditions that exist around the Antarctic continent, Southern Ocean (SO) ecosystems are very susceptible to the growing impact of global climate change and other anthropogenic influences. Consequently, there is an urgent need to understand how SO marine life will cope with expected future changes in the environment. Studies of Antarctic organisms have shown that individual species and higher taxa display different degrees of sensitivity to environmental shifts, making it difficult to predict overall community or ecosystem responses. This emphasizes the need for an improved understanding of the Antarctic benthic ecosystem response to global climate change using a multitaxon approach with consideration of different levels of biological organization. Here, we provide a synthesis of the ability of five important Antarctic benthic taxa (Foraminifera, Nematoda, Amphipoda, Isopoda, and Echinoidea) to cope with changes in the environment (temperature, pH, ice cover, ice scouring, food quantity, and quality) that are linked to climatic changes. Responses from individual to the taxon-specific community level to these drivers will vary with taxon but will include local species extinctions, invasions of warmer-water species, shifts in diversity, dominance, and trophic group composition, all with likely consequences for ecosystem functioning. Limitations in our current knowledge and understanding of climate change effects on the different levels are discussed.     

Biological monitoring of heavy-metal contamination in coastal waters off Casey Station, Windmill Islands, East Antarctica

Heavy-metal concentrations were determined in tissues of different species of benthic invertebrates collected in the Casey region (Australian Antarctic Territory) where an old waste-disposal tip site is a source of contamination. The species studied included the bivalve Laternula elliptica, starfish Notasterias armata, heart urchins Abatus nimrodi and A. ingens and gammaridean amphipod Paramoera walkeri. The specimens were collected at both reference and contaminated locations where lead was the priority element and copper was the next most important in terms of increased concentrations. The strong association between a gradient of contamination and concentrations in all species tested indicates that they are reflecting well the environmental changes, and that they appear as appropriate biological indicators of heavy-metal contamination. Aspects of the biology of species with different functional roles in the marine ecosystem are discussed in relation to their suitability for wider use in Antarctic monitoring programmes. For example, in terms of heavy-metal bioaccumulation, the bivalve appears as the most sensitive species to detect contamination; the starfish provides information on the transfer of metals through the food web while the heart urchin and gammarid give indications of the spatial and temporal patterns of the environmental contamination. The information gathered about processes of contaminant uptake and partitioning among different tissues and species could be used in later studies to investigate the behaviour and the source of contaminants.     

Iceberg Scour and Shell Damage in the Antarctic Bivalve Laternula elliptica

We document differences in shell damage and shell thickness in a bivalve mollusc (Laternula elliptica) from seven sites around Antarctica with differing exposures to ice movement. These range from 60% of the sea bed impacted by ice per year (Hangar Cove, Antarctic Peninsula) to those protected by virtually permanent sea ice cover (McMurdo Sound). Patterns of shell damage consistent with blunt force trauma were observed in populations where ice scour frequently occurs; damage repair frequencies and the thickness of shells correlated positively with the frequency of iceberg scour at the different sites with the highest repair rates and thicker shells at Hangar Cove (74.2% of animals damaged) compared to the other less impacted sites (less than 10% at McMurdo Sound). Genetic analysis of population structure using Amplified Fragment Length Polymorphisms (AFLPs) revealed no genetic differences between the two sites showing the greatest difference in shell morphology and repair rates. Taken together, our results suggest that L. elliptica exhibits considerable phenotypic plasticity in response to geographic variation in physical disturbance.     

Detecting growth under environmental extremes: Spatial and temporal patterns in nucleic acid ratios in two Antarctic bivalves

Growth of Antarctic benthic organisms is very slow due to temperature and food availability, and subtle differences in growth rate may be difficult to detect. Nucleic acid ratios (RNA/DNA, RNA/protein or total RNA concentration) are measures of protein synthesis potential and may be used to assess short-term growth rate in a range of marine organisms. We quantified nucleic acid ratios in the scallop Adamussium colbecki and the clam Laternula elliptica at five locations in the Ross Sea, Antarctica. We were able to detect species-specific, habitat-specific, and seasonal differences in nucleic acid ratios and related these to associated differences in primary productivity. By using nucleic acid ratios, future studies could relatively easily obtain a measure of growth rate from a multitude of locations with contrasting habitat characteristics, food availability and temperature regimes around the Antarctic continent. This would yield a unique understanding of spatial and temporal patterns in bivalve growth in this extreme environment.     

Population dynamics of the ubiquitous Antarctic benthic amphipod Orchomenella franklini and its vulnerability to environmental change

Comprehensive ecological research is still lacking for many of the species that dominate the Antarctic benthos, preventing an adequate understanding of their potential response to environmental change. Here, population dynamics were explored in one of the most ubiquitous nearshore Antarctic benthic amphipods, Orchomenella franklini. Sex, reproductive status and body length were recorded for over 6,000 individuals, sampled from a variety of locations and times at Casey station in East Antarctica. Several life history traits were revealed for O. franklini that exemplify adaptations predicted for a polar environment. These include delayed reproduction, extended brood incubation, low fecundity, longevity and seasonal breeding linked to the summer phytoplankton bloom. There was also preliminary evidence of inter-annual and spatial fluctuations in population structure, potentially reflecting local environmental heterogeneity such as sea-ice duration. The influence of both large scale and local environmental conditions on the ecology of O. franklini provides insight into the vulnerability of this species to environmental change.     

背角无齿蚌(Anodonta woodiana)对浅水系统底栖和浮游藻类竞争关系的影响

底栖藻类和浮游藻类之间的竞争关系对浅水生态系统的结构、功能具有重要的影响,双壳类可通过滤食控制浮游藻类,从而改变底栖藻类与浮游藻类之间的竞争结果。论文通过比较放养背角无齿蚌(Anodonta woodiana)(蚌处理组)与不放养背角无齿蚌(对照组)系统中底栖藻类、浮游藻类的生物量和优势种等的变化,研究了滤食性双壳类对底栖藻类和浮游藻类间竞争的影响。结果表明,背角无齿蚌可显著降低浮游藻类生物量,提高水体透明度和沉积物表面光照条件,从而显著提高底栖藻类的生物量;背角无齿蚌也改变了浮游藻类的优势种,使优势种由蓝藻转变成硅藻。因此,滤食性双壳类有利于促进浅水生态系统从混水态向清水态转变,本研究结果对富营养化浅水湖泊修复与管理具有一定的参考意义。     

Characteristics of shell microstructure and growth analysis of the Antarctic bivalve <Emphasis Type="Italic">Laternula elliptica</Emphasis> from Lützow-Holm Bay,Antarctica

Growth performance of the Antarctic bivalve Laternula elliptica was examined both by shell microstructural observation and by applying a fluorescent substance, tetracycline, as a shell growth marker. The shell was composed of two calcareous layers: the thick outer layer was homogeneous or granular in structure and the thin inner layer was nacreous. The architecture of Antarctic L. elliptica was different from that of temperate L. marilina, and the ratio of thickness between the outer and inner layers appeared to be different. The growth rate of the nacreous layer was analyzed to be very low. High correlations were found between the major axis of chondrophore and both shell length and shell dry weight, respectively. It is suggested that the chondrophore is an appropriate growth indicator, and combining the information of growth increments with the fluorescent method may be useful in estimating the bivalve growth performance in the Antarctic sea.     

Arginine metabolism of the Antarctic Bivalve <Emphasis Type="Italic">Laternula elliptica</Emphasis> (King &; Broderip, 1831): an ecophysiological approach

The potential aerobic ATP-generating pathway and the argininolytic capacity of the Antarctic bivalve Laternula elliptica in its main tissues were measured by the specific activity of the enzymes malate dehydrogenase (MDH), citrate synthase (CS) and arginase. The kidney showed the major potential for aerobic ATP-generating pathway and argininolytic capacity. High levels of CS and MDH activities indicated that renal tissue can be involved in activities that require a lot of energy such as excretion of metabolic end products, amino acids catabolism or even gluconeogenic activities related to inter-tissue metabolism. The fact that kidneys are the main site for arginase activity is very unusual for mollusks and could be related to the living habits of L. elliptica. Genetic expression of the L. elliptica renal arginase could be controlling the levels of l-arginine and forming urea in the excretory organ, which may not have its physiological functions directly affected by the seasonal retraction of its siphons. Compared to the bivalve Dreissena polymorpha, renal arginase of L. elliptica is more resistant to inhibition by copper and cadmium. This could be related to naturally high levels of these metals in the Antarctic marine environment and its bioaccumulation in the renal tissue of L. elliptica, as a probable advantage to its environmental adaptation. Different from other Antarctic animals that feed on Krill, the arginase of L. elliptica is much more sensitive to fluoride inhibition. However, diet composition of L. elliptica would be expected to be variable site to site and its high sensitivity to fluoride inhibition may be a matter of concern in areas near ornithogenic soils subjected to ice-melting processes.     

Consequences of Increasing Hypoxic Disturbance on Benthic Communities and Ecosystem Functioning

Disturbance-mediated species loss has prompted research considering how ecosystem functions are changed when biota is impaired. However, there is still limited empirical evidence from natural environments evaluating the direct and indirect (i.e. via biota) effects of disturbance on ecosystem functioning. Oxygen deficiency is a widespread threat to coastal and estuarine communities. While the negative impacts of hypoxia on benthic communities are well known, few studies have assessed in situ how benthic communities subjected to different degrees of hypoxic stress alter their contribution to ecosystem functioning. We studied changes in sediment ecosystem function (i.e. oxygen and nutrient fluxes across the sediment water-interface) by artificially inducing hypoxia of different durations (0, 3, 7 and 48 days) in a subtidal sandy habitat. Benthic chamber incubations were used for measuring responses in sediment oxygen and nutrient fluxes. Changes in benthic species richness, structure and traits were quantified, while stress-induced behavioral changes were documented by observing bivalve reburial rates. The initial change in faunal behavior was followed by non-linear degradation in benthic parameters (abundance, biomass, bioturbation potential), gradually impairing the structural and functional composition of the benthic community. In terms of ecosystem function, the increasing duration of hypoxia altered sediment oxygen consumption and enhanced sediment effluxes of NH₄ ⁺ and dissolved Si. Although effluxes of PO₄ ³⁻ were not altered significantly, changes were observed in sediment PO₄ ³⁻ sorption capability. The duration of hypoxia (i.e. number of days of stress) explained a minor part of the changes in ecosystem function. Instead, the benthic community and disturbance-driven changes within the benthos explained a larger proportion of the variability in sediment oxygen- and nutrient fluxes. Our results emphasize that the level of stress to the benthic habitat matters, and that the link between biodiversity and ecosystem function is likely to be affected by a range of factors in complex, natural environments.     

Assessing the sensitivity of bivalve populations to global warming using an individual‐based modelling approach

Climate change exposes benthic species populations in coastal ecosystems to a combination of different stressors (e.g., warming, acidification and eutrophication), threatening the sustainability of the ecological functions they provide. Thermal stress appears to be one of the strongest drivers impacting marine ecosystems, acting across a wide range of scales, from individual metabolic performances to geographic distribution of populations. Accounting for and integrating the response of species functional traits to thermal stress is therefore a necessary step in predicting how populations will respond to the warming expected in coming decades. Here, we developed an individual‐based population model using a mechanistic formulation of metabolic processes within the framework of the dynamic energy budget theory. Through a large number of simulations, we assessed the sensitivity of population growth potential to thermal stress and food conditions based on a climate projection scenario (Representative Concentration Pathway; RCP8.5: no reduction of greenhouse gas emissions). We focused on three bivalve species with contrasting thermal tolerance ranges and distinct distribution ranges along 5,000 km of coastline in the NE Atlantic: the Pacific oyster (Magallana gigas), and two mussel species: Mytilus edulis and Mytilus galloprovincialis. Our results suggest substantial and contrasting changes within species depending on local temperature and food concentration. Reproductive phenology appeared to be a core process driving the responses of the populations, and these patterns were closely related to species thermal tolerances. The nonlinear relationship we found between individual life‐history traits and response at the population level emphasizes the need to consider the interactions resulting from upscaling across different levels of biological organisation. These results underline the importance of a process‐based understanding of benthic population response to seawater warming, which will be necessary for forward planning of resource management and strategies for conservation and adaptation to environmental changes.     

Sublethal effects on reproduction in native fauna: are females more vulnerable to biological invasion?

Although invasive species are a major threat to survivorship of native species, we know little about their sublethal effects. In soft-sediment marine systems, mat-forming invasive species often have positive effects, facilitating recruitment and enhancing the diversity and abundance of native invertebrates. However, because mat-forming invasive species change the habitat in which they invade, and benthic invertebrates are sensitive to environmental disturbance, important sublethal effects on native species may exist. Using a model marine system we show that the widespread mat-forming invasive alga Caulerpa taxifolia (Vahl) C. Agardh has strong negative effects on the reproductive traits of a native bivalve Anadara trapezia (Deshayes, 1840) (e.g. timing of reproductive development and spawning, and follicle and gamete production) even though the invader has positive effects on recruitment. Moreover, gender specific responses occurred and indicated that females were more susceptible to invasion than males. Our results indicate that sublethal effects of an invasive species on reproductive traits will have severe consequences for fitness of the native species.     

Shallow-water benthic hydroids from Tethys Bay (Terra Nova Bay, Ross Sea, Antarctica)

The shallow-water hydrozoan Antarctic fauna is still poorly studied, and available knowledge mostly refers to samples gathered by traditional ship-operated gears. By scuba diving in the coastal areas off the Italian Antarctic station “Mario Zucchelli” (Ross Sea, Terra Nova Bay), in the austral summer 2002–2003, a total of 20 hydrozoan species were found, belonging to 10 families and 13 genera. As hypothesized, Anthoathecata (11 species), usually under-represented in collections from indirect sampling gears, are common as also are Leptothecata (9 species). Hydractiniidae and Hydractinia are the dominant family and genus, followed by Haleciidae and Halecium. A new species to science, Halecium exaggeratum sp. nov. is also described. Most species are either endemic to Antarctic waters or restricted to Antarctic/sub-Antarctic areas; only two species have a wider distribution. Material reared in aquaria at the Italian Antarctic Base Mario Zucchelli facilitated knowledge of the life cycle and reproductive biology of several species. In particular, Opercularella belgicae was found to liberate a medusa stage referable to Phialella, and the species is assigned here to that genus, as Phialella belgicae. Also, extraordinary is the complete absence or scant representation of the most typical Antarctic benthic hydroid genera (Antarctoscyphus, Oswaldella, Schizotricha, Staurotheca, and Symplectoscyphus), likely related to the shallow limits of sampling (down to 48 m).     

Extinction and recolonization of maritime Antarctica in the limpet Nacella concinna (Strebel, 1908) during the last glacial cycle: toward a model of Quaternary biogeography in shallow Antarctic invertebrates

Quaternary glaciations in Antarctica drastically modified geographical ranges and population sizes of marine benthic invertebrates and thus affected the amount and distribution of intraspecific genetic variation. Here, we present new genetic information in the Antarctic limpet Nacella concinna, a dominant Antarctic benthic species along shallow ice‐free rocky ecosystems. We examined the patterns of genetic diversity and structure in this broadcast spawner along maritime Antarctica and from the peri‐Antarctic island of South Georgia. Genetic analyses showed that N. concinna represents a single panmictic unit in maritime Antarctic. Low levels of genetic diversity characterized this population; its median‐joining haplotype network revealed a typical star‐like topology with a short genealogy and a dominant haplotype broadly distributed. As previously reported with nuclear markers, we detected significant genetic differentiation between South Georgia Island and maritime Antarctica populations. Higher levels of genetic diversity, a more expanded genealogy and the presence of more private haplotypes support the hypothesis of glacial persistence in this peri‐Antarctic island. Bayesian Skyline plot and mismatch distribution analyses recognized an older demographic history in South Georgia. Approximate Bayesian computations did not support the persistence of N. concinna along maritime Antarctica during the last glacial period, but indicated the resilience of the species in peri‐Antarctic refugia (South Georgia Island). We proposed a model of Quaternary Biogeography for Antarctic marine benthic invertebrates with shallow and narrow bathymetric ranges including (i) extinction of maritime Antarctic populations during glacial periods; (ii) persistence of populations in peri‐Antarctic refugia; and (iii) recolonization of maritime Antarctica following the deglaciation process.     

Shallow benthic fauna communities of South Georgia Island

Benthic communities in several fjords and sheltered bays of the north coast of South Georgia Island were examined using SCUBA and shore sampling in November 2004. It is one of the most northerly islands within the Polar Front and its well studied, terrestrial biota is described as sub Antarctic. The intertidal and subtidal zones and their fauna are, by comparison, little known. We describe the composition of the substratum and benthic communities of sites in several northern bays, including an exceptional community in the extremely sheltered Moraine Fjord with a 2 m sill at its entrance. In this, like those in some other fjordic systems, some taxa occur shallower than elsewhere, and in this instance, get large and are probably old. Elsewhere, we found the coastal fauna was fairly similar to Antarctic shallow communities in the southern Scotia Arc and Peninsula region. Of the taxa we found, we identified 53 to genus and 41 to species. Most of these were typical Antarctic shallow benthic taxa. Certain flatworms, nemerteans, bivalve and gastropod molluscs, amphipod and isopod crustaceans, asteroid echinoderms and stalked ascidians were very abundant, but some normally common Antarctic higher taxa and species were notably absent or rare.     

Respiratory responses of three Antarctic ascidians and a sea pen to increased sediment concentrations

Glacial retreat and subglacial bedrock erosion are consequences of rapid regional warming on the West Antarctic Peninsula. Sedimentation of fine-grained eroded particles can impact the physiology of filter-feeding benthic organisms. We investigated the effect of increasing concentrations of sediment on the oxygen consumption of suspension feeding species, the ascidians Molgula pedunculata, Cnemidocarpa verrucosa, Ascidia challengeri, and the pennatulid Malacobelemnon daytoni in Potter Cove (South Shetland Islands, Antarctica). In A. challengeri and C. verrucosa, oxygen consumption increased gradually up to a critical sediment concentration (C _crit) where species oxygen consumption was maximal (O _max in mg O₂?g^?1dm?day^?1) and further addition of sediments decreased respiration. C _crit was 200?mg?L^?1 for A. challengeri (O _max of 0.651?±?0.238) and between 100 and 200?mg?L^?1 for C. verrucosa (O _max of 0.898?±?0.582). Oxygen consumption of M. pedunculata increased significantly even at low sediment concentrations (15–50?mg sediment?L^?1). Contrary to the ascidians, sediment exposure did not affect oxygen consumption of the sea pen. The tiered response to sedimentation in the four species corroborates recent field observations that detected a reduction in the abundance of the sensitive ascidian M. pedunculata from areas strongly affected by glacial sediment discharge, whereas sea pens are increasing in abundance. Our investigation relates consequences (population shifts in filter-feeder communities) to causes (glacial retreat) and is of importance for modelling of climate change effects in Antarctic shallow coastal areas.     

Shallow gene pools in the high intertidal: extreme loss of genetic diversity in viviparous sea stars (Parvulastra)

We document an extreme example of reproductive trait evolution that affects population genetic structure in sister species of Parvulastra cushion stars from Australia. Self-fertilization by hermaphroditic adults and brood protection of benthic larvae causes strong inbreeding and range-wide genetic poverty. Most samples were fixed for a single allele at nearly all nuclear loci; heterozygotes were extremely rare (0.18%); mitochondrial DNA sequences were more variable, but few populations shared haplotypes in common. Isolation-with-migration models suggest that these patterns are caused by population bottlenecks (relative to ancestral population size) and low gene flow. Loss of genetic diversity and low potential for dispersal between high-intertidal habitats may have dire consequences for extinction risk and potential for future adaptive evolution in response to climate and other selective agents.     

Multiple stressors,nonlinear effects and the implications of climate change impacts on marine coastal ecosystems

Global climate change will undoubtedly be a pressure on coastal marine ecosystems, affecting not only species distributions and physiology but also ecosystem functioning. In the coastal zone, the environmental variables that may drive ecological responses to climate change include temperature, wave energy, upwelling events and freshwater inputs, and all act and interact at a variety of spatial and temporal scales. To date, we have a poor understanding of how climate‐related environmental changes may affect coastal marine ecosystems or which environmental variables are likely to produce priority effects. Here we use time series data (17 years) of coastal benthic macrofauna to investigate responses to a range of climate‐influenced variables including sea‐surface temperature, southern oscillation indices (SOI, Z4), wind‐wave exposure, freshwater inputs and rainfall. We investigate responses from the abundances of individual species to abundances of functional traits and test whether species that are near the edge of their tolerance to another stressor (in this case sedimentation) may exhibit stronger responses. The responses we observed were all nonlinear and some exhibited thresholds. While temperature was most frequently an important predictor, wave exposure and ENSO‐related variables were also frequently important and most ecological variables responded to interactions between environmental variables. There were also indications that species sensitive to another stressor responded more strongly to weaker climate‐related environmental change at the stressed site than the unstressed site. The observed interactions between climate variables, effects on key species or functional traits, and synergistic effects of additional anthropogenic stressors have important implications for understanding and predicting the ecological consequences of climate change to coastal ecosystems.