Weak Effects of Epibiota on the Abundances of Fishes Associated with Pier Pilings in Sydney Harbour

Biogenic habitats have profound effects on the distribution and abundances of many organisms. Epibiota are major biogenic components of hard substrata in marine habitats, particularly on artificial structures such as pier pilings, and have the potential to influence organisms associated with these structures. This study tested hypotheses about effects of epibiota on abundances of fishes associated with pilings in Middle Harbour, Sydney. Amount of epibiota had no effect on abundances of most species of fish, suggesting that epibiota are not important resources for these species. Abundances of the hulafish, Trachinops taeniatus, were, however, found to be greater around pilings with large than pilings with small amounts of epibiota at one site, at each of two times. Experimental removals of epibiota from pilings were done to test the hypothesis that epibiota are an important resource to T. taeniatus. Despite great temporal fluctuations in abundances over the duration of the experiment, the response of T. taeniatus to the removal of epibiota from pilings was as predicted, falling to zero following removal of epibiota. Although this suggests that epibiota are an important resource for this species, this pattern was spatially and temporally variable. We suggest that variation in the amount of epibiota has minor effects on the abundances of fishes around pilings.     

Surface composition and orientation interact to affect subtidal epibiota

Settlement panels were used to evaluate the effects of composition of the substratum (sandstone, concrete, wood) and orientation (vertical, horizontal undersides) on subtidal epibiota. It was predicted that both factors would influence the development of epibiotic assemblages, but that differences due to composition would be less marked on horizontal undersides compared to vertical panels. Differences in assemblages among sandstone, concrete and wooden panels orientated vertically were predicted to be similar to those described previously among vertical surfaces of sandstone rocky reefs and concrete and wooden urban structures (pilings and pontoons). Multivariate analyses indicated that assemblages were influenced greatly by orientation, whereas the effects of surface composition differed for the two orientations and among sites. Assemblages on wood were always significantly different from those on sandstone or concrete - patterns between the latter two surfaces depended on the orientation of the panels. The taxa that dominated these surfaces were not similar in identity nor abundance to those on urban structures of the same composition. The covers of most taxa were influenced by orientation alone or by surface composition for just one orientation. This study demonstrates the need for caution in generalizing about effects of orientation and surface composition because they may interact with each other and/or with other factors and they are certainly quite different for different taxa and among sites.     

Tadpole shrimp structure macroinvertebrate communities in playa lake microcosms

Laboratory microcosms were used to assess whether tadpole shrimp, Triops sp., affect community structure of other native macroinvertebrates in playa lakes of the Southern High Plains of Texas. Removal of tadpole shrimp shortly after hatching reduced abundances of many taxa, and decreased subsequent taxonomic richness and diversity. For many invertebrates, the presence of tadpole shrimp in low numbers had a positive effect on mean abundance. Direct effects of tadpole shrimp include the reduction of prey species abundance, which in turn may alter biotic interactions among other taxa. Indirect effects include physical modification of the environment during foraging through surface sediments. Results suggest that tadpole shrimp may be a key species controlling structure of macroinvertebrate communities in playa lakes.     

Epibiota communities of the introduced and indigenous macroalgal relatives<Emphasis Type="Italic"> Sargassum muticum</Emphasis> and<Emphasis Type="Italic"> Halidrys siliquosa</Emphasis> in Limfjorden (Denmark)

Sargassum muticum (Phaeophyceae, Fucales) has recently been introduced to Limfjorden (Denmark) where its closest relative is the indigenous Halidrys siliquosa. Previous studies have demonstrated large quantitative (canopy biomass) and qualitative (canopy persistence) differences in the habitat available to epibiota within the canopies of these two macroalgae. We therefore hypothesised that these algae would support different epibiota communities and tested this by sampling the epibiota of S. muticum and H. siliquosa on seven occasions throughout 1997 by enclosing entire thalli in mesh bags. We found 53 epibiota taxa and, with only one exception, they were all recorded on both host species. Species richness and abundance of epibiota exhibited clear seasonal variation on both host species, although epibiota biomass was seasonally constant on H. siliquosa but not on S. muticum. These patterns were consistent with the different life histories of the host species. There was a weakly negative correlation between thallus size and epibiota biomass for both host species. When taking species-specific seasonal variation in thallus size into consideration, S. muticum and H. siliquosa were found to support significantly different epibiota biomasses. Multivariate analyses showed that epibiota community structure was different, although highly overlapping, between the two species, whereas there was an almost parallel temporal development in epibiota community structure. We conclude that it is unlikely that the introduction of S. muticum to Limfjorden has caused major changes in local epibiota community structure. However, the standing stock of epibiota is likely to have increased.Communicated by H.-D. Franke     

Interactions between seagrasses and burrowing ghost shrimps and their influence on infaunal assemblages

The current study examined the direct interactions between intertidal seagrasses (Zosteraceae) and burrowing ghost shrimps (Callianassidae) and their influence on associated infaunal assemblages. Reciprocal transplant experiments conducted in two temperate regions revealed different interactions between both types of organism. In the U.S.A., seagrass prospered in all treatments, irrespective of the presence of ghost shrimp, whilst ghost shrimp declined in plots containing seagrass. In New Zealand, neither transplanted ghost shrimp nor seagrass became established in experimental plots, at the same time, neither type of organism appeared to be affected by the experimental addition of transplants. The differences in interactions between seagrass and ghost shrimp appeared to be related to seasonal differences in the timing of the transplant experiments and the pairing of particular ghost shrimp and seagrass species in each region. Infaunal assemblages showed distinct differences between seagrass and ghost shrimp treatments and reflected the dominant type of organism present. In treatments where transplanted seagrass or ghost shrimp became established, assemblage composition shifted in accordance with the type of transplanted organism. Differences in assemblage composition were characterised by higher relative abundances of discriminating taxa in treatments dominated by seagrass. The overall patterns of infaunal assemblage composition were correlated with a number of variables including the number of shoots, above-, below-ground seagrass biomass, % fines/sand, % total organic carbon, and sediment chlorophyll a. Findings from this study highlight the functional importance of intertidal seagrasses and burrowing ghost shrimps and reveal some of the ecological repercussions associated with changes in the distribution of these sympatric ecosystem engineers.     

Chironomid assemblages from seabird-affected High Arctic ponds

Seabirds can shunt nutrients and contaminants from marine to terrestrial ecosystems by forming dense breeding colonies and releasing wastes to these sites. A large colony of seabirds at Cape Vera (Devon Island, High Arctic Canada) has resulted in eutrophic conditions and potentially toxic concentrations of sedimentary metals in several freshwater ponds that drain their nesting sites. Here, we investigated the effects of elevated nutrient and sedimentary metal concentrations on the distribution of subfossil chironomids in surface sediments from 21 ponds that span a gradient of seabird influence. Although many ponds registered high nutrient concentrations (e.g., mean TP = 45 μg l ⁻¹), eutrophic taxa typical of temperate waters were not common, with most assemblages being dominated by morphotypes of Psectrocladius and Tanytarsina, as well as Corynoneura arctica-type, and Metriocnemus hygropetricus-type. Although the ponds within and outside the area influenced by seabirds contained largely similar taxa, variations did exist in the relative abundances of the different species. Lakewater pH was the only measured environmental variable that explained statistically significant amounts of variation in the chironomid assemblages. Although direct effects of pH on chironomids cannot be ruled out, pH is likely tracking production-related changes driven by limnetic dissolved inorganic carbon dynamics. Sediment cores collected from seabird-affected and seabird-free ponds showed a greater number of chironomid taxa and higher head capsule abundances in the pond receiving seabird inputs. Chironomid assemblages in both cores recorded increased abundances in recent decades, likely in response to warmer conditions and lengthened growing seasons.     

Tracking climate change in a dispersal-limited species: reduced spatial and genetic connectivity in a montane salamander

Tropical montane taxa are often locally adapted to very specific climatic conditions, contributing to their lower dispersal potential across complex landscapes. Climate and landscape features in montane regions affect population genetic structure in predictable ways, yet few empirical studies quantify the effects of both factors in shaping genetic structure of montane-adapted taxa. Here, we considered temporal and spatial variability in climate to explain contemporary genetic differentiation between populations of the montane salamander, Pseudoeurycea leprosa. Specifically, we used ecological niche modelling (ENM) and measured spatial connectivity and gene flow (using both mtDNA and microsatellite markers) across extant populations of P. leprosa in the Trans-Mexican Volcanic Belt (TVB). Our results indicate significant spatial and genetic isolation among populations, but we cannot distinguish between isolation by distance over time or current landscape barriers as mechanisms shaping population genetic divergences. Combining ecological niche modelling, spatial connectivity analyses, and historical and contemporary genetic signatures from different classes of genetic markers allows for inference of historical evolutionary processes and predictions of the impacts future climate change will have on the genetic diversity of montane taxa with low dispersal rates. Pseudoeurycea leprosa is one montane species among many endemic to this region and thus is a case study for the continued persistence of spatially and genetically isolated populations in the highly biodiverse TVB of central Mexico.     

Habitat segregation among freshwater shrimp species in an Amazonian rainforest stream system

1. Resource partitioning is a stabilising mechanism known to maintain species diversity in a variety of environments. Assemblages of stream shrimp species are structured by habitat features and predation. Therefore, segregation along habitat dimensions could facilitate coexistence among species in shrimp assemblages even when segregation is a result of predation pressure by fish species. These ecological interactions take place on a background modulated by biogeographic features, such as connectivity among drainages. However, these generalisations are mainly based on studies undertaken in temperate regions.
2. We investigated whether abundances of rainforest shrimp species are related to habitat dimensions, and whether habitat–abundance relationships might be mediated through fish-assemblage structure and the effect of drainages on connectivity.
3. We detected effects of habitat variation on densities of shrimp species, but the magnitudes of the effects were larger for some species than others. Fish-assemblage composition also affected shrimp densities. Two of the three species of shrimp showed some degree of habitat specialisation, but only along current-velocity, depth, and pH gradients. Habitat segregation among species occurred along the current-velocity and pH gradients. Relationships between density and environmental gradients differed between catchments for only one species and only along the pH gradient.
4. Our findings provide evidence that rainforest-stream shrimp species respond differently to environmental gradients and this could facilitate coexistence among species. However, interactions with fish seem to have a stronger effect on species densities, and consequently species segregation, than direct effects of the environmental gradients, resulting in apparent competition for these resources.

     

Floating pontoons create novel habitats for subtidal epibiota

Urban structures in the form of pontoons and pilings represent major coastal habitats for marine organisms and understanding the factors causing abundances of organisms to differ between these and natural habitat has been neglected in the study of coastal ecology. It has been proposed that composition of substrata explain differences previously described between subtidal assemblages of epibiota on rocky reef (sandstone) and pontoons (concrete) in Sydney Harbour, Australia. This study tested the hypothesis that differences in the composition of substratum (sandstone vs. concrete) independent of type of habitat (rocky reef vs. pontoon) affects the development of epibiotic assemblages. This was tested by experimentally providing substratum of the two types in both habitats. Epibiotic assemblages were unaffected by the composition of substratum but strongly affected by the type of habitat; demonstrating that pontoons constitute novel habitats for epibiota. This result highlights a need for determining how current ecological understanding of subtidal epibiota, which is heavily based on studies of urban structures (pilings and pontoons), relates to natural reef. Future tests of hypotheses about the nature of these differences will not only contribute to better ecological understanding of epibiota and their use of urban structures as habitats, but also to better predictions of future changes to the ecology of coastal habitats.     

Spatio-temporal changes of benthic macroinvertebrates in a cold Arkansas tailwater

This paper investigates spatial, seasonal and long-term changes in benthic macroinvertebrates in riffles of a cold tailwater. Cold tailwaters initially disrupt previously existing macroinvertebrate assemblages, but little is known about the long-term biological effects of a stable cold thermal regime. Assemblages at an upstream and downstream site of the Little Red River, Arkansas were investigated almost 30 years apart (1971 and 1999). Based upon published literature demonstrating the stability of benthic assemblages within unaltered environments, we predicted that the assemblages would be similar for each variable investigated. The benthic macroinvertebrate assemblages can be characterized as low diversity, with a total of 17 taxa identified. Isopods and Diptera comprised ~80% of all individuals. Other than chironomids, insects and particularly EPT taxa were poorly represented. Recent macroinvertebrate densities were significantly greater compared to the historical study period for the downstream site. Assemblage comparisons revealed moderate differences between study periods. Macroinvertebrate density was significantly greater upstream than downstream in the 1971 study period, yet taxa richness was significantly greater downstream for both study periods. Faunal composition was significantly different for upstream and downstream sites. Seasonal differences in numerical standing crop were identified for the 1971 upstream and 1999 downstream data sets. Low to moderate levels of seasonal, spatial and historical variation among benthic macroinvertebrate assemblages were attributed to environmental (temperature and flow) stability. The lack of aquatic insects other than chironomids over a 30-year period is indicative of the extreme constraints placed upon insect development within this cold regulated river.     

Microsatellite and mtDNA analysis of lake trout,Salvelinus namaycush,from Great Bear Lake,Northwest Territories: impacts of historical and contemporary evolutionary forces on Arctic ecosystems

Resolving the genetic population structure of species inhabiting pristine, high latitude ecosystems can provide novel insights into the post‐glacial, evolutionary processes shaping the distribution of contemporary genetic variation. In this study, we assayed genetic variation in lake trout (Salvelinus namaycush) from Great Bear Lake (GBL), NT and one population outside of this lake (Sandy Lake, NT) at 11 microsatellite loci and the mtDNA control region (d‐loop). Overall, population subdivision was low, but significant (global F_ST θ = 0.025), and pairwise comparisons indicated that significance was heavily influenced by comparisons between GBL localities and Sandy Lake. Our data indicate that there is no obvious genetic structure among the various basins within GBL (global F_ST = 0.002) despite the large geographic distances between sampling areas. We found evidence of low levels of contemporary gene flow among arms within GBL, but not between Sandy Lake and GBL. Coalescent analyses suggested that some historical gene flow occurred among arms within GBL and between GBL and Sandy Lake. It appears, therefore, that contemporary (ongoing dispersal and gene flow) and historical (historical gene flow and large founding and present‐day effective population sizes) factors contribute to the lack of neutral genetic structure in GBL. Overall, our results illustrate the importance of history (e.g., post‐glacial colonization) and contemporary dispersal ecology in shaping genetic population structure of Arctic faunas and provide a better understanding of the evolutionary ecology of long‐lived salmonids in pristine, interconnected habitats.     

Effects of early recruits on temperate sessile marine community composition depend on other species recruiting at the time

In many environments recruitment of dispersive propagules (e.g. seeds, spores and larvae) can vary from situations when particular taxa recruit in relative isolation to times when they recruit simultaneously with other, functionally quite different taxa. Differences in the identity and density of recruiting taxa can have important consequences on community structure, but it is still not clear how the effects of individual taxa on communities are modified when they recruit together with other species. Using an experimental approach we compared early development of a temperate marine sessile community after the recruitment of mixtures of botryllid ascidians and barnacles to that when barnacles or botryllid ascidians recruited alone. Communities exposed to recruitment of botryllid ascidians in isolation differed from those that received barnacles, a mixture of botryllids and barnacles or no recruitment in 2-week-old communities. These early differences were driven by higher abundances of the species that were present as initial recruits in experimental treatments. After 2 months communities also differed between barnacle and mixed recruitment treatments but not mixed and botryllid or botryllid and barnacle treatments. These differences were not directly due to differences in the abundances of our manipulated taxa but occurred because of two abundant arborescent bryozoans, Bugula dentata, which occupied more space in communities that initially received mixed recruitment than in those that received barnacle or no recruitment, and Zoobotryon verticillatum, which occupied more space in communities that initially received only barnacle recruitment than those that initially received botryllid or mixed recruitment. These effects did not persist, and communities did not differ after 6 months. These results suggest that, more generally, species may influence community dynamics differently when they recruit alongside other species than when they recruit in relative isolation.     

Impacts of past glaciation events on contemporary fish assemblages of the Ohio River basin

Aim We evaluated variation in fish assemblages on the basis of taxonomic composition and functional groups based on Pleistocene glacial boundaries in the Ohio River basin. We tested for the influence of habitat and hydrology on fish assemblage variation. Location Ohio River basin of North America, including the states of Ohio, Indiana and Illinois. Methods Fish collection sites were identified as Wisconsinan, pre‐Wisconsinan or unglaciated regions. Multivariate analyses, multi‐response permutation procedures, discriminant analysis and indicator species analyses were used to test whether taxonomic and functional assemblages were distinct among regions with varying glacial histories. Principal components analysis was used to identify habitat and water quality, as well as hydrological gradients that could be discerned by glacial region. Results We identified significant differences in both taxonomic and functional fish assemblage structure and habitat variation among regions that had different glaciation histories. The largest differences in taxonomic and functionally based fish communities were for unglaciated and pre‐Wisconsinan regions, while unglaciated and Wisconsinan regions were most similar. We correctly classified study regions in 71% and 60% of sites using taxonomy and functional analyses, respectively. Wisconsinan sites were characterized by Cyprinidae and Catostomidae assemblages with high abundances of tolerant fishes that tended to occur in habitats with reduced current velocity. Pre‐Wisconsinan sites were characterized by Cyprinidae, Catostomidae, Centrarchidae and Percidae families with increased abundances of intolerant fishes that tended to occur in habitats with coarser substrates and increased water velocity in streams of varying size. Unglaciated sites were characterized by Cyprinidae and Percidae families and were not closely associated with any habitat‐based functional group. Habitat in the unglaciated and pre‐Wisconsinan sites was significantly different from that in the Wisconsinan sites, which were characterized by increased channel structure and reduced stream size. Main conclusions Pleistocene glaciation events formed a lasting template of predictable regional differences in stream habitat and in the corresponding taxonomic and functional fish assemblage structures. While many factors impact the distribution of fishes, these results suggest that historical influences such as glaciation may be used to further explain the underlying mechanisms of spatial variation in fish assemblages.     

Geography and island geomorphology shape fish assemblage structure on isolated coral reef systems

We quantify the relative importance of multi‐scale drivers of reef fish assemblage structure on isolated coral reefs at the intersection of the Indian and Indo‐Pacific biogeographical provinces. Large (>30 cm), functionally‐important and commonly targeted species of fish, were surveyed on the outer reef crest/front at 38 coral reef sites spread across three oceanic coral reef systems (i.e. Christmas Island, Cocos (Keeling) Islands and the Rowley Shoals), in the tropical Indian Ocean (c. 1.126 x 106 km²). The effects of coral cover, exposure, fishing pressure, lagoon size and geographical context, on observed patterns of fish assemblage structure were modelled using Multivariate Regression Trees. Reef fish assemblages were clearly separated in space with geographical location explaining ~53 % of the observed variation. Lagoon size, within each isolated reef system was an equally effective proxy for explaining fish assemblage structure. Among local‐scale variables, ‘distance from port’, a proxy for the influence of fishing, explained 5.2% of total variation and separated the four most isolated reefs from Cocos (Keeling) Island, from reefs with closer boating access. Other factors were not significant. Major divisions in assemblage structure were driven by sister taxa that displayed little geographical overlap between reef systems and low abundances of several species on Christmas Island corresponding to small lagoon habitats. Exclusion of geographical context from the analysis resulted in local processes explaining 47.3% of the variation, highlighting the importance of controlling for spatial correlation to understand the drivers of fish assemblage structure. Our results suggest reef fish assemblage structure on remote coral reef systems in the tropical eastern Indian Ocean reflects a biogeographical legacy of isolation between Indian and Pacific fish faunas and geomorphological variation within the region, more than local fishing pressure or reef condition. Our findings re‐emphasise the importance that historical processes play in structuring contemporary biotic communities.     

Heterogeneous patterns of abundance of epigeic arthropod taxa along a major elevation gradient

Species diversity is the variable most commonly studied in recent ecological research. Ecological processes, however, are driven by individuals and affected by their abundances. Understanding the variation in animal abundances along climatic gradients is important for predicting changes in ecosystem processes under global warming. High abundances make arthropods, despite their small body sizes, important actors in food webs, yet abundance distributions of major arthropod taxa along climatic gradients remain poorly documented. We sampled arthropod assemblages in disturbed and undisturbed vegetation types along an elevational gradient of 860–4550 m asl on the southern slopes of Mt. Kilimanjaro, Tanzania. In our analysis, we focused on 13 taxa of arthropods that represented three major functional groups: predators, herbivores, and decomposers. Abundance patterns were unimodal for most of the taxa and functional groups, including decomposer arthropods, and most of them peaked at low elevations in lower montane forest. When we assigned beetles to functional groups, however, decomposer beetle abundances declined almost linearly, and abundances of predator beetles (ca. 2400 m asl) and herbivore beetles (ca. 3000 m asl, undisturbed vegetation) peaked at higher elevations and exhibited unimodal patterns. Temperature, not primary productivity, was the best predictor of abundance for most of the taxa and groups. Disturbance was only of minor importance. Our results revealed different trends in the response of arthropod abundance along the elevational gradient that depended on the level of taxonomic and functional resolution. This highlights the need for more comparisons of different taxa along the same climatic gradients.     

Seawater-atmosphere O(2) exchange rates in open-top laboratory microcosms: application for continuous estimates of planktonic primary production and respiration

Foraging by predatory fish is thought to be one of the primary ecological processes affecting the abundances of plants and animals in subtidal habitats. The importance of this process was assessed on the subtidal surfaces of urban structures (pontoons and pilings) that represent major coastal habitats for marine organisms. Fish feed with greater intensity on epibiota attached to pilings than pontoons and it was hypothesised that greater predation on pilings explained why the structure of epibiotic assemblages differs between these habitats. I predicted that the structure of epibiotic assemblages would develop differently between pilings and pontoons in the presence of fish (plates open to predation) but not in the absence of fish (plates inside exclusion cages). Results revealed large differences in abundance between pilings and pontoons that were largely independent of the caged and uncaged plates. Predation may be intense (as it appeared on pilings) but unimportant because it does not explain observed abundances of prey (epibiota between pilings and pontoons).     

Protist assemblages in winter sea ice: setting the stage for the spring ice algal bloom

This study documents, for the first time, the abundance and species composition of protist assemblages in Arctic sea ice during the dark winter period. Lack of knowledge of sea-ice assemblages during the dark period has left questions about the retention and survival of protist species that initiate the ice algal bloom. Sea-ice and surface water samples were collected between December 27, 2007 and January 31, 2008 within the Cape Bathurst flaw lead, Canadian Beaufort Sea. Samples were analyzed for protist identification and counts, chlorophyll (chl) a, and total particulate carbon and nitrogen concentrations. Sea-ice chl a concentrations (max. 0.27 μg l⁻¹) and total protist abundances (max. 4 × 10³ cells l⁻¹) were very low, indicating minimal retention of protists in the ice during winter. The diversity of winter ice protists (134 taxa) was comparable to spring ice assemblages. Pennate diatoms dominated the winter protist assemblage numerically (averaging 77% of total protist abundances), with Nitzschia frigida being the most abundant species. Only 56 taxa were identified in surface waters, where dinoflagellates were the dominant group. Our results indicate that differences in the timing of ice formation may have a greater impact on the abundance than structure of protist assemblages present in winter sea ice and at the onset of the spring ice algal bloom.     

The quest for absolute abundance: The use of internal standards for DNA‐based community ecology

To characterize microbiomes and other ecological assemblages, ecologists routinely sequence and compare loci that differ among focal taxa. Counts of these sequences convey information regarding the occurrence and relative abundances of taxa, but provide no direct measure of their absolute abundances, due to the technical limitations of the sequencing process. The relative abundances in compositional data are inherently constrained and difficult to interpret. The incorporation of internal standards (ISDs; colloquially referred to as ‘spike‐ins’) into DNA pools can ameliorate the problems posed by relative abundance data and allow absolute abundances to be approximated. Unfortunately, many laboratory and sampling biases cause ISDs to underperform or fail. Here, we discuss how careful deployment of ISDs can avoid these complications and be an integral component of well‐designed studies seeking to characterize ecological assemblages via sequencing of DNA.     

Foraging by the mud snail, Ilyanassa obsoleta (Say), modulates spatial variation in benthic community structure

We investigated the foraging behavior of the mud snail, Ilyanassa obsoleta, and its consequences for macrobenthic community structure on mud flats on Long Island, NY, USA. Field sampling demonstrated strong spatial heterogeneity in the population densities of I. obsoleta. We experimentally tested three hypotheses: (i) I. obsoleta are strongly attracted to areas with high levels of detritus; (ii) local abundances of deposit-feeding annelids are limited by detritus; and (iii) the foraging activities of I. obsoleta negatively affects annelid assemblages. We manipulated the density of mud snails using inclusion fences and the levels of detritus using dried Ulva. Results showed that high densities of I. obsoleta were attracted to areas enriched with Ulva detritus. In addition, high densities of snails negatively affected abundances of annelids, with the opportunistic species, Capitella spp. and Paranais litoralis, being most affected. The addition of Ulva detritus had more specific effects on annelid assemblages. Only Capitella spp. showed a significant positive response, although previous evidence has demonstrated that higher experimental detrital inputs stimulated growth of other species of annelids and microphytobenthos. In an experimental treatment with enhanced detritus and low densities of snails, we found population abundances of opportunistic annelids (up to 200,000 m⁻²) substantially larger than has ever been recorded in 5 years of sampling. Because mud snails in natural areas actively search, locate and exploit areas with enhanced detritus and their foraging negatively affects abundances of opportunistic worms, I. obsoleta probably controls the upper limits of annelid abundance in the field. Foraging behavior of I. obsoleta therefore modulates spatial variation in benthic community structure in an environment where limiting resources are patchily distributed.     

Response of deposit-feeders to exclusion of epibenthic predators in a Mediterranean intertidal flat

Deposit-feeders are common components of macrofaunal assemblages in intertidal soft sediments. Predation has been considered to have a central role in affecting their distribution and population dynamics. This study investigates the effect of epibenthic predators on deposit-feeders, inhabiting the shallow layers of the sediment (surface and subsurface) and also the deepest layers (e.g., burrowing shrimp Upogebia pusilla; Petagna). The experiment was conducted in summer 2000 (August-September) at three different sites on an intertidal flat in Mediterranean Sea. In the field, predators were excluded using cages, placed on the surface of the sediment. It was predicted that under the cages, (i) abundances of animals would increase and (ii) species composition of assemblages would change as an effect of the exclusion of predators. Potential artefacts caused by the cages were controlled using partial cages (PC). Composition of organic matter and porosity were also analysed under PC and in natural controls to test whether the presence of cages would change sediment characteristics on the surface. At the end of the experiment, there were no significant differences in the composition of organic matter between PC and the undisturbed sediment (UC). After 8 weeks from the beginning of the experiment, exclusion of predators enhanced the abundance of U. pusilla. There were, however, no clear-cut changes in the species composition of macrofaunal assemblages and densities of animals did not increase under the cages. Indeed, some animals (Oligochaeta, Brania oculata, and Tanais dulongii) were less abundant under the cages (EC) than outside (PC and UC). We propose that predation might play a role in regulating interspecific relationships among some surface deposit-feeders and the burrowing shrimp U. pusilla.