Predation in the marine intertidal amphipod Echinogammarus marinus Leach: implications of inter- and intra-individual variation

Studies of competition, predator–prey dynamics and food webs typically consider conspecifics as equal, however, individuals from the same population that are seemingly identical can show considerable variation with regards to a number of processes. Such phenomena may be demonstrated in terms of diet, and the quantities and types of resources that are consumed are commonly considered. The marine amphipod Echinogammarus marinus, a recently demonstrated predator on intertidal rocky shores, has been shown to consume a wide range of food types but it is unknown how this may vary between individuals. Here, we investigated the variation that occurs both among and within individuals of a population of E. marinus with respect to the mean numbers consumed of a common prey item, the isopod Jaera nordmanni. First, by comparing the length of starvation times, used as a proxy for hunger level, individuals maintained without food for up to 24 h consumed significantly less prey during feeding trials than those starved for 48 h and longer. The degree of inter-individual variation within each starvation period was also found to differ, with greater variation among individuals starved for shorter periods of time than those starved for longer time periods. Secondly, we tested whether individual amphipods tracked over time consumed consistently similar numbers of prey or whether they showed intra-individual variation, and if so, to what degree. We found that the numbers of prey consumed per individual could be predicted in the short-term between consecutive feeding trials, however over the long-term this relationship broke down. These results are discussed with respect to potential physiological and behavioural mechanisms, as well as the implications that such variation may have for stability of prey populations in the field.

Coastal upwelling drives intertidal assemblage structure and trophic ecology

ecosystems. Coastal oceanic upwelling, for example, has been associated with elevatedbiomass and abundance patterns of certain functional groups, e.g., corticated macroalgae.In the upwelling system of Northern Chile, we examined measures of intertidal macrobenthiccomposition, structure and trophic ecology across eighteen shores varying in theirproximity to two coastal upwelling centres, in a hierarchical sampling design (spatial scalesof >1 and >10 km). The influence of coastal upwelling on intertidal communities was confirmedby the stable isotope values (δ13C and δ15N) of consumers, including a dominantsuspension feeder, grazers, and their putative resources of POM, epilithic biofilm, andmacroalgae. We highlight the utility of muscle δ15N from the suspension feeding mussel,Perumytilus purpuratus, as a proxy for upwelling, supported by satellite data and previousstudies. Where possible, we used corrections for broader-scale trends, spatial autocorrelation,ontogenetic dietary shifts and spatial baseline isotopic variation prior to analysis. Ourresults showed macroalgal assemblage composition, and benthic consumer assemblagestructure, varied significantly with the intertidal influence of coastal upwelling, especiallycontrasting bays and coastal headlands. Coastal topography also separated differences inconsumer resource use. This suggested that coastal upwelling, itself driven by coastlinetopography, influences intertidal communities by advecting nearshore phytoplankton populationsoffshore and cooling coastal water temperatures. We recommend the isotopic valuesof benthic organisms, specifically long-lived suspension feeders, as in situ alternativesto offshore measurements of upwelling influence

Nitrate and phosphate uptake characteristics of three species of brown algae cultured at low salinity.

Nitrate and phosphate uptake mechanisms have been characterised under conditions of 100 and 50% seawater in 3 common brown algae of NW Europe: Fucus vesiculosus, F. serratus and Laminaria digitata. Under low salinity, the growth rate and internal nitrate accumulation of F. serratus significantly increased (20 and 48%, respectively), but no significant changes were observed for F. vesiculosus and L. digitata. However, nitrate uptake rates were reduced in L. digitata, so that this species was less adaptable to low salinity than the Fucus species. Both F. vesiculosus and F. serratus reached a steady-state uptake rate after acclimation regardless of the salinity treatment. All 3 species had a high capacity for storing inorganic N and P intracellularly. The results for F. serratus pointed to a dual mechanism of adaptation to the special characteristics of the intertidal environment where it grows. Non-saturating (low affinity) nitrate uptake and biphasic (double Michaelis-Menten curve) phosphate uptake are adaptations to high nutrient concentrations. Temporal partition of cellular energy for carbon metabolism and nutrient uptake is also suggested as an adaptation to the transient nutrient inputs occurring in these environments.

Dispersal mode and assessments of recovery on the shores of Gruinard, the 'anthrax island'

Formaldehyde run-off was an unintended impact of the anthrax decontamination procedure on the island of Gruinard. The death of intertidal organisms was observed where formaldehyde reached the shore during 1986. The extent to which shores on Gruinard have recovered was assessed with survey work in 2000. Recovery estimates were based on the hypothesis that the process of recolonization is partly dependent on species' dispersal capability. Underdevelopment of the assemblage of species lacking planktonic dispersal stages (direct developers) is therefore evidence that the process of recolonization is ongoing, rather than complete. A novel multivariate test showed that, when comparing quadrats from Gruinard and nearby mainland shores, assemblages of direct developing molluscs were significantly more distinct than assemblages of molluscs with planktonic dispersal stages. The average densities of species with direct development were generally lower on Gruinard than on mainland shores. While some species with direct development have similar densities on Gruinard and on the surrounding shores, the recovery of the overall assemblage was still incomplete after 14 years. In contrast, the harvested species, Littorina littorea, appeared to benefit from the absence of humans visiting Gruinard's shores. Populations of L. littorea on Gruinard contained significantly higher proportions of large individuals. Depending on the dispersal capabilities of different species, Gruinard is either still in recovery or acts as a reserve.

Distinct patterns of nitrate reductase activity in brown algae: Light and ammonium sensitivity in Laminaria digitata is absent in Fucus species

Fucus and Laminaria species, dominant seaweeds in the intertidal and subtidal zones of the temperate North Atlantic, experience tidal cycles that are not synchronized with light:dark (L:D) cycles. To investigate how nutrient assimilation is affected by light cycles, the activity of nitrate reductase (NR) was examined in thalli incubated in outdoor tanks with flowing seawater and natural L:D cycles. NR activity in Laminaria digitata (Huds.) Lamour. showed strong diel patterns with low activities in darkness and peak activities near midday. This diel pattern was controlled by light but not by a circadian rhythm. In contrast, there was no diel variation in NR activity in Fucus serratus L., F. vesiculosus (L.) Lamour., and F. spiralis L. either collected directly from the shore or maintained in the outdoor tanks. In laboratory cultures, transfer to continuous darkness suppressed NR activity in L. digitata, but not in F. vesiculosus; continuous light increased NR activity in L. digitata but decreased activity in F. vesiculosus. Furthermore, 4 d enrichment with ammonium (50 mu mol . L-1 pulses), resulted in NR activity declining by > 80% in L. digitata, but no significant changes in F. serratus. Seasonal differences in maximum NR activity were present in both genera with activities highest in late winter and lowest in summer. This is the first report of NR activity in any alga that is not strongly regulated by light and ammonium. Because light and tidal emersion do not always coincide, Fucus species may have lost the regulation of NR by light that has been observed in other algae and higher plants.

Limpet grazing and loss of Ascophyllum nodosum canopies on decadal time scales

The role of limpet grazing in preventing the development of algal canopies is a recurrent theme in intertidal ecology. Less is known about interactions of limpets with the long-term dynamics of established canopies. Aerial photographs indicate that intertidal canopy cover has declined over the past 44 yr in Strangford Lough, Northern Ireland. There has been a loss of the previously continuous cover of Ascophyllum nodosum (L.) Le Jolis in the mid-shore. A barnacles dominated assemblage now fills gaps in the A. nodosum canopy. The rates at which barnacle patches become established and grow have increased since 1990. Changes in canopy cover have been accompanied by increases in limpet densities since the 1980s. Measurements between 2003 and 2004 showed no increase in length of A. nodosum fronds when limpets Patella vulgata had access to the algal holdfasts. In contrast, when limpets were experimentally excluded from the holdfasts, there was net frond growth. In the Isle of Man, which is climatically similar to Strangford Lough but has fewer limpets, growth occurred regardless of limpet grazing. The breaking force for A. nodosum declined with increasing local densities of limpets. A. nodosum is a sheltered shore species, potentially vulnerable to changes in wave exposure. There is no evidence, however, that Strangford Lough has become windier over the past 3 decades. Variation in wave exposure among locations within the lough was not related to rates of barnacle patch creation or expansion, Limpet population density has increased following a series of mild winters. Climate change may have a role in causing canopy loss, not by direct effects on the limpet populations.

Variation in shell morphology of Calliostoma zizyphinum (Gastropoda : Trochidae)

Differences in shell morphology in the intertidal prosobranch mollusc Calliostoma zizyphinum were studied from a number of sites within four geographical reo ions of the British Isles with varying exposures to wave action and crab predation. Mean values of damage scarring were highest in shells sampled from Strangford Lough, Northern Ireland, and lowe;t in individuals from the Atlantic Coast. Shells collected front the Isle of Man were smaller than those from either the Atlantic coast of Ireland or Strangford Lough. Shells front Strangford Lough had taller shells (higher aspect ratio) than shells from the County Down and Atlantic Coast and shells from the County Down coast were more squat (lower aspect ratio) than those from all other areas. Shells from Strangford Lough were significantly thinner than shells from all other geographical areas. The relationships between shell damage scarring and shell size and tallness were not consistent among areas. Shells from Strangford Lough are notable in showing a steep, positive relationship between shell size and scarring and a steep, negative relationship between scarring and tallness. The unusual shell phenotypes observed in shells from Strangford 1,Lough may be explained by rapid shell growth, which would not only allow Calliostoma to attain a size refuge from crab predators but also to recover successfully from repeated crab attacks on the shell lip. Such a strategy would result in larger, thinner shells with a high number of damage scars.

Foraging ecology, fluctuating food availability and energetics of wintering brent geese

Changing energy requirements and dramatic shifts in food availability are major factors driving behaviour and distribution of herbivores. We investigate this in wintering East Canadian High Arctic light-bellied brent geese Branta bernicla hrota in Northern Ireland. They followed a sequential pattern of habitat use, feeding on intertidal Zostera spp. in autumn and early winter before moving to predominantly saltmarsh and farmland in late winter and early spring. Night-time feeding occurred throughout and made a considerable contribution to the birds' daily energy budget, at times accounting for > 50% of energy intake. Nocturnal feeding, however, is limited to the intertidal, possibly because of predation risk on terrestrial habitat, and increases with moonlight. The amount of Zostera spp., declined dramatically after the arrival of birds, predominantly, but not entirely, due to consumption by the birds. Birds gained fat reserves in the first 2 months but then this was dramatically lost as their major food source collapsed and their daily energy intake declined. Single birds consistently fared worse than paired birds and pairs with juveniles fared better than those without suggesting a benefit of having a family to compete for food. Many birds leave the Lough at this time of reduced Zostera spp. for other sea inlets in Ireland but some remain. Body condition of the latter gradually improved in early spring and reflected a heavy reliance on terrestrial habitats, particularly farmland, to meet the birds' daily energy requirements. However, even in the period immediately before migration to the breeding ground, the birds did not regain the amount of abdominal fatness observed in November. The dramatic changes in available food and requirements of the birds drive the major changes seen in foraging behaviour as the birds evade starvation in the wintering period.

Age and palaeoenvironmental significance of an inter-tidal peat bed at Ballywoolen, Bann estuary, Co. Londonderry.

A recently exposed inter-tidal peat bed at Ballywoolen, Bann estuary, Co. Londonderry, has yielded new information about mid-Holocene coastal environmental change in the northeast of Ireland. Pollen analytical data and wood detritus demonstrate that peat accumulation occurred in a terrestrial environment that was free from marine influence. Radiocarbon dates suggest that the peat accumulated rapidly during a period of low relative sea level subsequent to the maximum of Holocene relative sea-level rise along the north coast of Northern Ireland. The absence of marine/brackish indicator taxa at the site suggests that the tidal range was somewhat less than that at present and/or that the channel of the river was located some distance east of its present alignment. The dates indicate that the low stand lasted for at least ~0.2 ka and possibly for ~1.1 ka. Stable, woodland-dominated landscapes are indicated at both this site and neighbouring ones around ~6.4-5.3 cal ka BP. There is no evidence for large-scale aeolian sand movement or human impact on the landscape during the period of peat accumulation.

delta C-13 and delta N-15 reveal significant differences in the coastal foodwebs of the seas surrounding Trinidad and Tobago

This study assessed nearshore, marine ecosystem function around Trinidad and Tobago (TT). The coastline of TT is highly complex, bordered by the Atlantic Ocean, the Caribbean Sea, the Gulf of Paria and the Columbus Channel, and subject to local terrestrial runoff and regional riverine inputs (e.g. the Orinoco and Amazon rivers). Coastal organisms can assimilate energy from allochthonous and autochthonous Sources, We assessed whether stable isotopes delta C-13 and delta N-15 Could be used to provide a rapid assessment of trophic interactions in primary consumers around the islands. Filter-feeding (bivalves and barnacles) and grazing organisms (gastropods and chitons) were collected from 40 marine sites during the wet season. The flesh of organisms was analysed for delta C-13 and delta N-15. Results indicate significant variation in primary consumers (by feeding guild and sampling zone). This variation was linked to different energy Sources being assimilated by consumers. Results suggest that offshore production is fuelling intertidal foodwebs; for example, a depleted delta C-13 signature in grazers from the Gulf of Paria, Columbus Channel and the Caribbean and Atlantic coastline of 9 Tobago indicates that carbon with an offshore origin (e.g. phytoplankton and dissolved organic matter) is more important than benthic or littoral algae (luring the wet season. Results also confirm findings from other studies indicating that much of the coastline is subject to Cultural eutrophication. This Study revealed that ecosystem function is spatially variable around the coastline of TT, This has clear implications for marine resource management, as a single management approach is unlikely to be successful at a national level.

Do mussel patches provide a refuge for algae from grazing gastropods?

On rocky shores, cover of macroalgae is often greater growing epibiotically on mussels compared to algae growing directly attached to rock. A survey of two shores on the east coast of Ireland confirmed that mussel beds contained greater percentage algal cover and more diverse algal assemblages compared to those on rock. The reasons for this difference are not clear. It has been suggested that mussel beds provide a refuge for algae from grazing gastropods. Surprisingly, we found no evidence to support this. Using wax discs, gastropod grazing patterns were found to be similar within the mussel beds as on rock. The mussel beds do not appear to provide a refuge for algae from grazing activity at this scale and we suggest other possible mechanisms for the prevalence of epibiotic algal cover on mussels. Intertidal grazers may in fact affect the epibiotic algae on mussels and thereby affect indirectly the persistence of mussel beds.

The European oyster (Ostrea edulis) and its epibiotic succession

This study aimed to investigate intra- and inter-site differences in the epibiota on the European oyster Ostrea edulis of different ages and on hard substrata to assess their contribution to biodiversity. This research has shown that shells of O. edulis do show higher species diversity than non-living hard substrata and as oysters grow larger epibiotic diversity will increase. This investigation has revealed that O. edulis possesses biogenic engineering qualities and the value of O. edulis as a vehicle for increasing biodiversity should not be underestimated. Relatively few studies have examined the shell surfaces of Ostrea edulis, which, together with their associated epifauna provide a large portion of the natural hard substrata for sessile filter-feeding organisms, including oysters, in coastal and estuarine ecosystems. It can be considered a keystone species which has an influence on intertidal and subtidal community structures.

Deposit-feeding mechanisms and resource partitioning in tropical holothurians

Aspidochirote holothurians found on tropical reef flats feed on particulate deposits which form a variety of substrata. The synaptid holothurian Opheodesoma grisea (Semper) feeds in a similar manner by scraping deposits from the surfaces of sea grasses. Distributional and gut content analyses showed that species partitioning is on the basis of substratum and particle size preference. Scanning electron microscopy revealed that the tentacles of aspidochirotes have a nodular surface while those of O. grisea have a tessellated surface structure. The twelve different species examined were shown to have different tentacular surface textures which bore an apparent relationship with the mean particle sizes selected by the different species. Light microscope studies of tentacle sections confirmed earlier observations on the extent of the water vascular system in aspidochirote and pinnate tentacles. From these observations a functional interpretation is proposed for tentacular operation and the means of particle selection in such holothurians.