Evaluating signatures of glacial refugia for North Atlantic benthic marine taxa

A goal of phylogeography is to relate patterns of genetic differentiation to potential historical geographic isolating events. Quaternary glaciations, particularly the one culminating in the Last Glacial Maximum ~21 ka (thousands of years ago), greatly affected the distributions and population sizes of temperate marine species as their ranges retreated southward to escape ice sheets. Traditional genetic models of glacial refugia and routes of recolonization include these predictions: low genetic diversity in formerly glaciated areas, with a small number of alleles/haplotypes dominating disproportionately large areas, and high diversity including "private" alleles in glacial refugia. In the Northern Hemisphere, low diversity in the north and high diversity in the south are expected. This simple model does not account for the possibility of populations surviving in relatively small northern periglacial refugia. If these periglacial populations experienced extreme bottlenecks, they could have the low genetic diversity expected in recolonized areas with no refugia, but should have more endemic diversity (private alleles) than recently recolonized areas. This review examines evidence of putative glacial refugia for eight benthic marine taxa in the temperate North Atlantic. All data sets were reanalyzed to allow direct comparisons between geographic patterns of genetic diversity and distribution of particular clades and haplotypes including private alleles. We contend that for marine organisms the genetic signatures of northern periglacial and southern refugia can be distinguished from one another. There is evidence for several periglacial refugia in northern latitudes, giving credence to recent climatic reconstructions with less extensive glaciation.

Manipulating Interaction Strengths and the Consequences for Trivariate Patterns in a Marine Food Web

We are experiencing a global extinction crisis as a result of climate change and human-induced alteration of natural habitats, with large predators at high trophic levels in food webs being particularly vulnerable. Unfortunately, there is a scarcity of food web data that can be used to assess how species extinctions alter the structure and stability of temporally and spatially replicated networks. We established a series of large experimental mesocosms in a shallow subtidal benthic marine system and constructed food webs for each replicate. After 6 months of community assembly, we removed large predators from the core communities of 20 experimental food webs, based on the strength of their trophic interactions, and monitored the changes in the networks' structure and stability over an 8-month period. Our analyses revealed the importance of allometric relationships and size-structuring in natural communities as a means of preserving food web structure and sustainability, despite significant changes in the diversity, stability and productivity of the system.

Comparison of granulometric methods and sampling strategies used in marine habitat classification and Ecological Status assessment

Sediment particle size analysis (PSA) is routinely used to support benthic macrofaunal community distribution data in habitat mapping and Ecological Status (ES) assessment. No optimal PSA Method to explain variability in multivariate macrofaunal distribution has been identified nor have the effects of changing sampling strategy been examined. Here, we use benthic macrofaunal and PSA grabs from two embayments in the south of Ireland. Four frequently used PSA Methods and two common sampling strategies are applied. A combination of laser particle sizing and wet/dry sieving without peroxide pre-treatment to remove organics was identified as the optimal Method for explaining macrofaunal distributions. ES classifications and EUNIS sediment classification were robust to changes in PSA Method. Fauna and PSA samples returned from the same grab sample significantly decreased macrofaunal variance explained by PSA and caused ES to be classified as lower. Employing the optimal PSA Method and sampling strategy will improve benthic monitoring. © 2012 Elsevier Ltd.

Predatory fish loss affects the structure and functioning of a model marine food web

The rate of species loss is increasing on a global scale and predators are most at risk from human-induced extinction. The effects of losing predators are difficult to predict, even with experimental single species removals, because different combinations of species interact in unpredictable ways. We tested the effects of the loss of groups of common predators on herbivore and algal assemblages in a model benthic marine system. The predator groups were fish, shrimp and crabs. Each group was represented by at least two characteristic species based on data collected at local field sites. We examined the effects of the loss of predators while controlling for the loss of predator biomass. The identity, not the number of predator groups, affected herbivore abundance and assemblage structure. Removing fish led to a large increase in the abundance of dominant herbivores, such as Ampithoids and Caprellids. Predator identity also affected algal assemblage structure. It did not, however, affect total algal mass. Removing fish led to an increase in the final biomass of the least common taxa (red algae) and reduced the mass of the dominant taxa (brown algae). This compensatory shift in the algal assemblage appeared to facilitate the maintenance of a constant total algal biomass. In the absence of fish, shrimp at higher than ambient densities had a similar effect on herbivore abundance, showing that other groups could partially compensate for the loss of dominant predators. Crabs had no effect on herbivore or algal populations, possibly because they were not at carrying capacity in our experimental system. These findings show that contrary to the assumptions of many food web models, predators cannot be classified into a single functional group and their role in food webs depends on their identity and density in 'real' systems and carrying capacities.

Why the size structure of marine communities can require decades to recover from fishing

A dynamic food-web model of more than 1000 species was used to quantify the recovery trajectory of marine community size-structure under different hypothetical fishing regimes, using the Northeast Atlantic as an example. Size-structure was summarised by four indicators: the Large Fish Indicator (LFI), the Large Species Indicator (LSI), the biomass-weighted mean maximum length of fish species (EMBED Equation.3) and the biomass-weighted mean maturation length of fish species (EMBED Equation.3). Time-series of these indicators recorded recovery following release from fishing with various size-selectivities, intensities and durations. In model simulations, fishing-induced trophic cascades were observed to distort fish community size-structure, but these did not have a large influence on recovery level or duration as measured by the four indicators. However, simulations showed that local extinctions of large fish species increased in number with both fishing intensity and duration, and could strongly limit the recovery level. Recovery of fish community size-structure to near equilibrium frequently took multiple decades in simulations; these long transient periods suggest that management interventions for size-structure recovery may require much longer than previously thought. Our results demonstrate the need for community-level modelling to set realistic targets for management of community size-structure.

Coastline configuration as a determinant of structure in larval assemblages

The richness and turnover of coastal larval pools set upper limits for biodiversity in coastal systems. For particular local systems, such as embayments, the characteristics of the local larval pool are determined by the relative contributions of locally produced and external larvae. The balance between these sources partially reflects the extent of tidal exchange and is hence related to system size and flushing time. Larvae of benthic marine invertebrates were sampled from 8 bays along the Irish coast to investigate the effect of coastline configuration on the characteristics of the larval pool. Flushing time explained 34.5% of the variability in species richness from a series of daily samples. Many of the potentially relevant environmental variables are correlated, limiting the potential for individual variables to be examined in isolation. We therefore used a principal components analysis to describe the major patterns in environmental variability across bays. The second principal component separated bays along a gradient of increasing depth, salinity, tidal range and flushing time. Scores along this component were generally better predictors of the larval pool than single variables, explaining as much as 61.2% of the variation in species richness, diversity and similarity between dates. Deeper bays, with more saline water and longer flushing times, tended to have richer and more diverse larval pools, with a greater consistency in species composition between sample dates. No relationship was found between environmental variables and larval abundance. Our results suggest that flushing time, particularly when in combination with topographic variables, chlorophyll, tidal range and salinity, may be a useful predictor for the richness and turnover of local larval pools.

Predator diversity enhances secondary production and decreases the likelihood of trophic cascades

We manipulated the diversity of top predators in a three trophic level marine food web. The food web included four top benthic marine fish predators (black goby, rock goby, sea scorpion and shore rockling), an intermediate trophic level of small fish, and a lower trophic level of benthic invertebrates. We kept predator density constant and monitored the response of the lower trophic levels. As top predator diversity increased, secondary production increased. We also observed that in the presence of the manipulated fish predators, the density of small gobiid fish (intermediate consumers) was suppressed, releasing certain groups of benthic invertebrates (caprellid amphipods, copepods, nematodes and spirorbid worms) from heavy intermediate predation pressure. We attribute the mechanism responsible for this trophic cascade to a trait-mediated indirect interaction, with the small gobiid fish changing their use of space in response to altered predator diversity. In the absence of top fish predators, a full-blown trophic cascade occurs. Therefore the diversity of predators reduces the likelihood of trophic cascades occurring and hence provides insurance against the loss of an important ecosystem function (i.e. secondary production).

Emerging asymmetric interactions between forage and predator fisheries impose management trade-offs

A size and trait-based marine community model was used to investigate interactions, with potential implications for yields, when a fishery targeting forage fish species (whose main adult diet is zooplankton) co-occurs with a fishery targeting larger-sized predator species. Predicted effects on the size structure of the fish community, growth and recruitment of fishes, and yield from the fisheries were used to identify management trade-offs among the different fisheries. Results showed that moderate fishing on forage fishes imposed only small effects on predator fisheries, whereas predator fisheries could enhance yield from forage fisheries under some circumstances.

Stressor intensity determines antagonistic interactions between species invasion and multiple stressor effects on ecosystem functioning

Biological invasions, nutrient enrichment and ocean warming are known to threaten biodiversity and ecosystem functioning. The independent effects of these ecological stressors are well studied, however, we lack understanding of their cumulative effects, which may be additive, antagonistic or synergistic. For example, the impacts of biological invasions are often determined by environmental context, which suggests that the effects of invasive species may vary with other stressors such as pollution or climate change. This study examined the effects of an invasive seaweed (Sargassum muticum) on the structure and functioning of a benthic marine assemblage and tested explicitly whether these effects varied with nutrient enrichment and ocean warming. Overall, the presence of Sargassum muticum increased assemblage productivity rates and warming altered algal assemblage structure, which was characterised by a decrease in kelp and an increase in ephemeral green algae. The effects of Sargassum muticum on total algal biomass accumulation, however, varied with nutrient enrichment and warming producing antagonistic cumulative effects on total algal biomass accumulation. These findings show that the nature of stressor interactions may vary with stressor intensity and among response variables, which leads to less predictable consequences for the structure and functioning of communities.

Using particle dispersal models to assist in the conservation and recovery of the overexploited native oyster (Ostrea edulis) in an enclosed sea lough

Oyster populations around the world have seen catastrophic decline which has been largely attributed to overexploitation, disease and pollution. While considerable effort and resources have been implemented into restoring these important environmental engineers, the success of oyster populations is often limited by poor understanding of site-specific dispersal patterns of propagules. Water-borne transport is a key factor controlling or regulating the dispersal of the larval stage of benthic marine invertebrates which have limited mobility. The distribution of the native oyster Ostrea edulis in Strangford Lough, Northern Ireland, together with their densities and population structure at subtidal and intertidal sites has been documented at irregular intervals between 1997 and 2013. This paper revisits this historical data and considers whether different prevailing environmental conditions can be used to explain the distribution, densities and population structure of O. edulis in Strangford Lough. The approach adopted involved comparing predictive 2D hydrodynamic models coupled with particle tracking to simulate the dispersal of oyster larvae with historical and recent field records of the distribution of both subtidal and intertidal, populations since 1995. Results from the models support the hypothesis that commercial stocks of O. edulis introduced into Strangford Lough in the 1990s resulted in the re-establishment of wild populations of oysters in the Northern Basin which in turn provided a potential source of propagules for subtidal populations. These results highlight that strategic site selection (while inadvertent in the case of the introduced population in 1995) for the re-introduction of important shellfish species can significantly accelerate their recovery and restoration.

Marine reserve designation, trophic cascades and altered community dynamics

Marine ecosystems and their associated populations are increasingly at risk from the cumulative impacts of many anthropogenic threats that increase the likelihood of species extinction and altered community dynamics. In response, marine reserves can be used to protect exploited species and conserve biodiversity. The increased abundance of predatory species in marine reserves may cause indirect effects along chains of multi-trophic interactions. These trophic cascades can arise through direct predation, density-mediated indirect interactions (DMIIs), or indirect behavioural effects, termed trait-mediated indirect interactions (TMIIs). The extent of algal cover and the abundance of 4 primary consumers were determined in Lough Hyne, which was designated Europe's first marine nature reserve in 1981. The primary consumers were the sea urchin Paracentrotus lividus, the topshell Gibbula cineraria, the oyster Anomia ephippium, and the scallop Chlamys varia. The abundances of 3 starfish species (Marthasterias glacialis, Asterias rubens, and Asterina gibbosa) were also determined, as were 2 potential crustacean predators, Necora puber and Carcinus maenas. These data were compared with historical data from a 1962 (prey) and a 1963 (predator) survey to determine the nature of community interactions over adjacent trophic levels. The present study reveals a breakdown in population structure of the 4 surveyed prey species. Marine reserve designation has led to an increase in predatory crabs and M. glacialis, a subsequent decrease in primary consumers, especially the herbivore P. lividus, and an increase in macroalgal cover which is indicative of a trophic cascade. The study shows that establishing a Marine Reserve does not guarantee that conservation benefits will be distributed equally.

Assessing the extent to which temporal changes in waterbird community composition are driven by either local, regional or global factors.

1. Lough Neagh and Lough Beg Special Protection Area (SPA, hereafter Lough Neagh) is an important non-estuarine site in Britain and Ireland for overwintering wildfowl. Multivariate analysis of the winter counts showed a state-shift in the waterbird community following winter 2000/2001, mostly due to rapid declines in abundance (46–57% declines in the mean mid-winter January counts between 1993–2000 and 2002–2009) of members of the diving duck guild (pochard Aythya ferina, tufted duck Aythya fuligula and goldeneye Bucephala clangula) and coot (Fulica atra), a submerged macrophyte feeder.
2. Only pochard showed correlations between declines at Lough Neagh and those of overall species flyway population indices to suggest that global changes could contribute to declines at the site. However, indices from the Republic of Ireland showed no overall decline in the rest of Ireland. Tufted duck indices at the site were inversely related to indices in Great Britain. Lough Neagh goldeneye indices were positively correlated with indices in the Republic of Ireland and Great Britain, suggesting that short-stopping could contribute to declines at the site. Coot declines at Lough Neagh did not correlate with trends elsewhere, suggesting local factors involved in the decline.
3. These analyses indicate that although there are potentially different explanations for the dramatic declines in these four waterbird species at this site, the simultaneous nature of the declines across two feeding guilds strongly
suggest that local factors (such as loss of submerged macrophytes and benthic invertebrates) were involved. An assessment of the food supply, local disturbance and other factors at Lough Neagh is required to find an explanation for the observed adverse trends in wintering numbers of the affected species.
4. This study highlights the potential of waterbird community structure to reflect the status of aquatic systems, but confirms the need to establish site-specific factors responsible for the observed changes in abundance of key waterbird species at a site.

Marine spatial planning from the perspective of a small seaside community in Ireland

Marine spatial planning (MSP) is advocated as a means of managing human uses of the sea in a manner that is consistent with the maintenance of the ecological goods and services of the marine environment. Support for the process is evident at international and national levels but the degree to which it is acceptable to local level stakeholders is not clear. An Daingean (formely Dingle) is a small sea-oriented town situated on the southwest coast of Ireland in which marine-based tourism and other relatively new uses of the sea are pursued along side traditional fishing activities. Stakeholders in An Daingean are found to be positively disposed to a local process of MSP that incorporates meaningful local involvement.