A seasonal stable isotope survey of the food web associated to a peri-urban rocky shore

We characterized the structure and functioning of the food web associated to an intertidal rocky shore in the harbour of Brest (Western Brittany, France), through a seasonal δ¹³C and δ¹⁵N survey. The dominance of benthic-derived organic matter, and particularly resuspended sediment, in the local pool of suspended organic matter, is highlighted. This benthic-derived organic matter dominates the diet of filter-feeders, including a certain degree of selectivity displayed by various taxa belonging to this trophic group. The food web structure appeared very stable temporally, which suggested the dominance of mixed diets, rather than the consumption of pure sources. It was constituted of 4 trophic levels, including a large diversity of predators, likely favoured by the diversity of microhabitats that characterize this intertidal habitat. From our results, the trophic functioning of this peri-urban rocky shore community does not appear functionally simplified as previously reported for other anthropized and/or artificial rocky ecosystems. We suggest that the topological complexity of rocky habitats, including a diversity of microhabitats, might be responsible of the important animal diversity, in spite of the anthropogenic disturbances characterizing this peri-urban area.     

Effect of canopy removal on a rocky shore community metabolism and structure

The effects of the permanent removal of the canopy-forming alga Fucus serratus was studied in terms of both functioning and diversity on a mid–low rocky shore ecosystem in the south-west English Channel (48°N 43.686′, 3°W 59.282′). Ecosystem functioning was examined as net or gross primary productivity (NPP or GPP) and respiration (Resp) measured through CO₂ fluxes. Diversity was examined as number and composition of species. Measurements were performed in situ, during emersion times, without altering target assemblages. The experiment was designed with two treatments [control (C) or canopy removed (CR)] and five replicates, and was conducted over an 18-month period (from February 2006 to August 2007) to integrate the seasonal variability. The mean GPP and Resp were severely reduced in CR treatment compared to control throughout the survey. The mean NPP was not affected at first, due to the development of opportunistic green algae, but was drastically reduced after 9 months of experiment. The canopy removal affected neither the number of species nor their distribution among trophic groups, and the algal community was only slightly affected. The abundance and biomass of mobile invertebrates, however, were greatly reduced in the absence of canopy. This indicates an important effect of the dominant alga on the higher trophic levels of the community. At this tidal level, the canopy did not seem to affect the community by dampening the environmental stress but by providing food, habitat or both.     

Description of a Tetraclita stalactifera panamensis community on a rocky intertidal Pacific shore of Panama

The midlittoral zone at Paitilla Beach, Panama, Is occupied by a group of organisms closely associated with Tetraclita stalactifera panamensis. Upon death, this cirripede provides housing for a number of species bound by common substrata and interacting with each other due to their close physical proximity; the faunistic group associated with each barnacle qualifies then, as a small, well-defined community which represents a subsample of the more comprehensive midlittoral community dominated by T. stalactifera panamensis. In patches where it is abundant, T. stalactifera panamensis occupies 28% of the surface area available. It has a relative abundance of only 1% of the specimens found in the zone, but all the other 95 species (with an average of 26, 810 individuals/m²) identified from the area are associated either with the parietal canals, or the external or internal surfaces of T. stalactifera panamensis tests. Each one of these structures represents a habitat which provides surface for attachment, shelter, or both, to 37 species of polychaetes, 32 of mollusks, 20 of crustaceans, 3 of actiniarians, 2 of pycogonids, 2 of sipunculans and an unknown number of tanaid, nemertean and turbellarian species. The single most diverse space-niche is represented by the parietal canals (H=0.783), occupied mostly by polychaetes. dead barnacles harbor an average of 131 individuals per test, which compose a fauna of higher diversity (H=0.678) than that associated with live barnacles (H=0.163). The latter contain 53 specimens per test on the average. Although there is considerable overlapping of species associated with the different T. stalactifera panamensis space-niches, mollusks are clearly more abundant on internal surfaces, polychaetes in parietal, canals; Balanus spp. on external surfaces, and anemones at the base of the parapet on the external surfaces of the tests. Over 220 species of macro-invertebrates were identified from the rocky intertidal at Paitilla. Most of the species are concentrated in the lower (157 species, 1824 specimens/m²) and midlittoral (96 species, 26, 810 specimens/m²) intertidal zones. Splash zone is depauperate (6 species, 1578 specimens/m²). Of the entire fauna identified from the intertidal, 73 species (33%) occur in more than one zone and 147 (67%) are restricted to a particular belt. This last category of zone-restricted species breaks down into 42% for the lower intertidal, 17% for the midlittoral, 6% for the intermediate zone, and 2% for the splash zone.     

Immunological methods for food web analysis in a soft-bottom benthic community

Basic to nearly any prediction concerning the behavior and structure of entire communities or of their components is knowledge of trophic connections among species. A major impediment to such understanding of soft-bottom benthos is methodological. Because none of the routinely available methods of food web analysis (e.g. visual gut content analysis, direct observation of feeding, tracer techniques) is generally suitable for examining all trophic interactions of benthic infauna, we rought to evaluate the potential of immunological methods for identifying predatorprey relationships in one typical, estuarine, intertidal sand flat. Whole-organism extracts of individual macro- and meiofaunal taxa were injected into rabbits to produce antisera of varying specificity. Double immunodiffusion precipitin tests of antiserum specificity revealed both phyletic and trophic relationships among 20 taxa. Using relatively unspecific antisera, preliminary analysis of the stomach contents of a few surface deposit-feeders and particle browsers was successful, giving positive identification of several trophic links which would otherwise have gone undetected. The production of taxon-specific antisera is expected to provide the methodological tool necessary to document the breadth of trophic connections in a marine benthic food web.     

The minimal area problem in invertebrate communities of Mediterranean rocky substrata

Different ways of obtaining information about minimal area are studied. The classical species-area curve lacks objectivity. Curves based on similarity indices are objective, in so far that a critical threshold can be defined, where one admits that the minimal area is reached. Comparison is made between Sørensen's qualitative similarity index (based on presence-absence) and Kulczynski's quantitative similarity index (based on numbers of individuals and/or percent cover). In the shallowwater rocky habitats studied, two types of community were distinguished. The first, miniaturized communities are found in dark caves. Their minimal area is about 4,000 cm². The second, coarser, communities occur on open rocks, either horizontal, sloping or vertical. Their minimal area is 20,000 cm². Calculations were based mainly on Octocorallia (Coelenterata) which are assumed to be characteristic of the community as a whole.     

Community structure of the rocky shore macrobenthos in southern Hokkaido,Japan

Relationships between community structure and environmental factors were sought through multivariate analysis (quantification analysis and correspondence analysis) of data obtained in a benthic macrofauna survey of a rocky low-tide platform located at Usujiri, southern Hokkaido, Japan. A total of 21 species were found. However, their annual mean abundance was dominated by only a few species. Three dominant species of the platform (Chthamalus challengeri, Littorina brevicula andSeptifer virgatus) accounted for an average of 92% of total abundance. Three environmental factors with associated species groups were examined. Among them, micro-topographic characteristics and wave action defined the main correspondence variables.L. brevicula was a characteristic member of the boulder and sheltered field.S. virgatus was the dominant species of sloped surfaces.C. challengeri, S. virgatus, andMytilus edulis were characteristic of ledge or bench environments. Finally,C. challengeri was characteristic of nip and high wave-exposure environments, given its exclusive appearences in these locations. However, as the degree of exposure depends on the scale of wash through the platform, it is directly limited by the microtopographic properties of low-tide platforms. Specific environmental influences such as exposure and height above the datum-plane were rather disapparent. Any overall effects on the community structure were synthesized by those of microtopography. After complete consideration, it was concluded that the community structure of the macrobenthos in this low-tide platform was found to be primarily under the influence of microtopographic characteristics.     

Microscale genetic differentiation in a sessile invertebrate with cloned larvae: investigating the role of polyembryony

Microscale genetic differentiation of sessile organisms can arise from restricted dispersal of sexual propagules, leading to isolation by distance, or from localised cloning. Cyclostome bryozoans offer a possible combination of both: the localised transfer of spermatozoa between mates with limited dispersal of the resulting larvae, in association with the splitting of each sexually produced embryo into many clonal copies (polyembryony). We spatially sampled 157 colonies of Crisia denticulata from subtidal rock overhangs from one shore in Devon, England at a geographic scale of ca. 0.05 to 130 m plus a further 21 colonies from Pembrokeshire, Wales as an outgroup. Analysis of molecular variance (AMOVA) revealed that the majority (67%) of genetic variation was distributed among individuals within single rock overhangs, with only 16% of variation among different overhangs within each shore and 17% of variation between the ingroup and outgroup shores. Despite local genetic variation, pairwise genetic similarity analysed by spatial autocorrelation was greatest at the smallest inter-individual distance we tested (5 cm) and remained significant and positive across generally within-overhang comparisons (<4 m). Spatial autocorrelation and AMOVA analyses both indicated that patches of C. denticulata located on different rock overhangs tended to be genetically distinct, with the switch from positive to negative autocorrelation, which is often considered to be the distance within which individuals reproduce with their close relatives or the radius of a patch, occurring at the 4–8 m distance class. Rerunning analyses with twenty data sets that only included one individual of each multilocus genotype (n = 97) or the single data set that contained just the unique genotypes (n = 67) revealed that the presence of repeat genotypes had an impact on genetic structuring (PhiPT values were reduced when shared genotypes were removed from the dataset) but that it was not great and only statistically evident at distances between individuals of 1–2 m. Comparisons to a further 20 randomisations of the data set that were performed irrespective of genotype (n = 97) suggested that this conclusion is not an artefact of reduced sample size. A resampling procedure using kinship coefficients, implemented by the software package GENCLONE gave broadly similar results but the greater statistical power allowed small but significant impacts of repeat genotypes on genetic structure to be also detected at 0.125–0.5 and 4–16 m. Although we predict that a proportion of the repeat multilocus genotypes are shared by chance, such generally within-overhang distances may represent a common distance of cloned larval dispersal. These results suggests that closely situated potential mates include a significant proportion of the available genetic diversity within a population, making it unlikely that, as previously hypothesised, the potential disadvantage of producing clonal broods through polyembryony is offset by genetic uniformity within the mating neighbourhood. We also report an error in the published primer note of Craig et al. (Mol Ecol Notes 1:281–282, 2001): loci Cd5 and Cd6 appear to be the same microsatellite. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Shading facilitates sessile invertebrate dominance in the rocky subtidal Gulf of Maine

Dramatic shifts in community composition occur between vertical and horizontal rocky surfaces in subtidal environments worldwide, yet the forces mediating this transition are poorly understood. Vertical rock walls are often covered by lush, diverse communities of sessile suspension-feeding invertebrates, while adjacent horizontal substrates are dominated by algae, or corals in the tropics. Multiple factors, including light, sedimentation, water flow, and predation have been proposed to explain this pattern, but experimental tests of these hypotheses are lacking. We manipulated light level and predation to test whether variation in these mechanisms could be responsible for the shift in composition of sessile communities between vertical and horizontal surfaces in the rocky subtidal Gulf of Maine. Shaded horizontally oriented granite plots were dominated by invertebrates (e.g., ascidians, barnacles, bryozoans) after 25 months. Unshaded plots were dominated by macroalgae, which was virtually absent in shaded plots. Exclusion of grazers with cages had no effect on percent cover of invertebrates or algae. Preferential settlement of invertebrate larvae to shaded plots, due to larval behaviors such as negative phototaxis, did not seem to play a role. Shading likely affects post-settlement mortality of invertebrates by alleviating competition for space with algae, although greater abundance of micropredators in algal-dominated communities could also be important. Communities on shaded plots lacked many taxa present on natural wall communities, likely due to greater disturbance on horizontal substrates and/or lack of sufficient time for colonization of these taxa. These results suggest that light plays a key role in controlling the structure, composition, and function of shallow subtidal communities.     

The primary productivity of marine macrophytes from a rocky intertidal community

This study represents the first report of primary production rates for Southern California intertidal producers. The production rates of 18 marine macrophytes near Wilson Cove, San Clemente Island, are close to those for other marine algal communities. No apparent relationship was revealed between the productivity of an alga and the division to which it belongs; however, productivity was associated with growth form. Encrusting prostrate forms were the lowest producers in terms of g C/m²/h and g C/g dry weight/h; sheet-like and finely-branched forms showed a greater productivity than coarsely-branched forms. Gelidium pusillum and Ulva californica had considerably greater production rates than the other algae measured.     

Patterns in species composition of rocky shore communities in sub-arctic fjords of eastern Iceland

Quantitative data on plants and animals from 20 transects of the eulittoral zone in 8 fjords of sub-arctic east Iceland were analysed by a number of classification and ordination techniques. Field work was carried out during two periods of spring tides in June and July 1987. The study was restricted to gently sloping stable bedrock, but transects were otherwise randomly chosen. Species composition was strongly influenced by the relative position of the transects along the length of the fjords. Species richness increased from heads towards mouths. Increased temperature fluctuations towards inner parts of fjords are suggested as an explanation for these patterns, while salinity is unlikely to be a critical factor. The analyses also indicate that degree of exposure, as measured both by a physically derived index and by a simple biological index, is an important factor in structuring these eulittoral communities, although affecting species richness only slightly, if at all. Classification of transects results in a number of community types which, however, are not well defined. A classification based on the presence/absence of three canopy-forming fucoids yields groups of transects differing in species composition and in degree of exposure. The rocky eulittoral communities of the fjords of eastern Iceland resemble those of northern Norway, but differ in that several key species of the latter communities are missing, probably largely due to the isolation of Iceland.     

Hydrology and grazing jointly control a large-river food web

Inputs of fresh water and grazing both can control aquatic food webs, but little is known about the relative strengths of and interactions between these controls. We use long-term data on the food web of the freshwater Hudson River estuary to investigate the importance of, and interactions between, inputs of fresh water and grazing by the invasive zebra mussel (Dreissena polymorpha). Both freshwater inputs and zebra mussel grazing have strong, pervasive effects on the Hudson River food web. High flow tended to reduce population size in most parts of the food web. High grazing also reduced populations in the planktonic food web, but increased populations in the littoral food web, probably as a result of increases in water clarity. The influences of flow and zebra mussel grazing were roughly equal (i.e., within a factor of 2) for many variables over the period of our study. Zebra mussel grazing made phytoplankton less sensitive to freshwater inputs, but water clarity and the littoral food web more sensitive to freshwater inputs, showing that interactions between these two controlling factors can be strong and varied.     

Adaptations in a hierarchical food web of southeastern Lake Michigan

Two issues in ecological network theory are: (1) how to construct an ecological network model and (2) how do entire networks (as opposed to individual species) adapt to changing conditions? We present a novel method for constructing an ecological network model for the food web of southeastern Lake Michigan (USA) and we identify changes in key system properties that are large relative to their uncertainty as this ecological network adapts from one time point to a second time point in response to multiple perturbations. To construct our food web for southeastern Lake Michigan, we followed the list of seven recommendations outlined in Cohen et al. [Cohen, J.E., et al., 1993. Improving food webs. Ecology 74, 252–258] for improving food webs. We explored two inter-related extensions of hierarchical system theory with our food web; the first one was that subsystems react to perturbations independently in the short-term and the second one was that a system's properties change at a slower rate than its subsystems’ properties. We used Shannon's equations to provide quantitative versions of the basic food web properties: number of prey, number of predators, number of feeding links, and connectance (or density). We then compared these properties between the two time-periods by developing distributions of each property for each time period that took uncertainty about the property into account. We compared these distributions, and concluded that non-overlapping distributions indicated changes in these properties that were large relative to their uncertainty. Two subsystems were identified within our food web system structure (p < 0.001). One subsystem had more non-overlapping distributions in food web properties between Time 1 and Time 2 than the other subsystem. The overall system had all overlapping distributions in food web properties between Time 1 and Time 2. These results supported both extensions of hierarchical systems theory. Interestingly, the subsystem with more non-overlapping distributions in food web properties was the subsystem that contained primarily benthic taxa, contrary to expectations that the identified major perturbations (lower phosphorous inputs and invasive species) would more greatly affect the subsystem containing primarily pelagic taxa. Future food-web research should employ rigorous statistical analysis and incorporate uncertainty in food web properties for a better understanding of how ecological networks adapt.     

High intertidal community organization on a rocky headland in Maine,USA

A mosaic patchwork of the barnacle Balanus balanoides L., the mussel Mytilus edulis L., and the alga Fucus vesiculosus L. was found in the transitional region between the mid and high intertidal zones on a rocky headland on Mount Desert Island, Maine, USA. The development of the mosaic was observed by following recruitment and survival of B. balanoides in denuded patches located at the same tidal level along a 60 m stretch of shore. Barnacle recruitment was least under canopies of F. vesiculosus and greatest in open areas kept moist at low tide by surf. Barnacle survival after settlement was least under the F. vesiculosus canopy due to the whiplash effect of the algal fronds in the surf and greatest in open areas free from competition from mussels. In open areas, early mortality was correlated with settlement density. In areas of dense settlement (60 spat cm^-2) up to 90% mortality resulted within 5 months from crowding associated with growth. In older individuals crowding produced hummocks of elongated, weakly attached barnacles which were more prone to removal by surf than uncrowded barnacles. Mussels exerted competitive dominance over barnacles for space and the presence of mussel beds prevented further barnacle recruitment. Mussels suffered extensive mortality during winter storms when surf removed dense mats of weakly attached mussels. The patchy distribution of mussels and barnacles results from irregular rock substrata producing numerous environmental patches with respect to wave exposure and drainage at low tide, and from densitydependent mortality of both mussels and barnacles which creates patches of new colonizable space within each environmental patch.     

Direct and indirect effects of ants on a forest-floor food web

Interactions among predators that prey on each other and are potential competitors for shared prey (intraguild [IG] predators) are widespread in terrestrial ecosystems and have the potential to strongly influence the dynamics of terrestrial food webs. Ants and spiders are abundant and ubiquitous terrestrial IG predators, yet the strength and consequences of interactions between them are largely unknown. In the leaf-litter food web of a deciduous forest in Kentucky (USA), we tested the direct and indirect effects of ants on spiders and a category of shared prey (Collembola) by experimentally subsidizing ants in open plots in two field experiments. In the first experiment, ant activity was increased, and the density of ants in the litter was doubled, by placing carbohydrate and protein baits in the center of each plot. Gnaphosa spiders were almost twice as abundant and Schizocosa spiders were half as abundant in baited plots relative to controls. There were more tomocerid Collembola in baited plots, suggesting possible indirect effects on Collembola caused by ant-spider interactions. The second experiment, in which screening of two mesh sizes selectively excluded large and small worker ants from a sugar bait, revealed that the large ants, primarily Camponotus, could alone induce similar effects on spiders. Gnaphosa biomass density was almost twice as high in the plots where large ants were more active, whereas Schizocosa biomass density was reduced by half in these plots. Although tomocerid densities did not differ between treatments, tomocerid numbers were negatively correlated with the activity of Formica, another large ant species. Path analysis failed to support the hypothesis that the ant Camponotus indirectly affected tomocerid Collembola through effects on densities of spiders. However, path analysis also revealed other indirect effects of Camponotus affecting tomocerids. These results illustrate the complexity of interactions between and within two major IG predator groups with disparate predatory behaviors, complexities that will have consequences for functioning of the forest-floor food web.     

Tagging studies on the growth of the sea hare Aplysia kurodai on an intertidal rocky shore

A total of 1,427 juvenile Aplysia kurodai were tagged and released during spring tides in January and February, 1973, on the rocky shore of Nabeta Bay, Japan. Recapture, weighing and release were repeated within a defined observation area during 10 spring tides from late January to early June. Recapture rates decreased with time and became significantly low after the end of the spawning season (March and April in this area). Of 1,427 individuals released, 550 (ca. 39%) were never recaptured. Only 68 individuals (ca. 5%) were recaptured 5 to 10 times during the observation period. The growth curves of these 68 individuals displayed a common growth pattern of body weight increasing almost linearly, attaining a maximum during the spawning season and then decreasing gradually. The maximum weight varied fairly widely from one individual to another. Laboratory measurements indicated a growth pattern similar to that observed in the field, and a fairly remarkable daily fluctuation in body weight was observed under laboratory conditions. The described tagging-recapture method, with slight modifications, should prove useful for future studies on the life history and ecology of A. kurodai.     

Temperature-dependent ranges of coexistence in a model of a two-prey-one-predator microbial food web

The objective of our study was to analyze the effects of temperature on the population dynamics of a three-species food web consisting of two prey bacteria (Pedobacter sp. and Acinetobacter johnsonii) and a protozoan predator (Tetrahymena pyriformis) as model organisms. We assessed the effects of temperature on the growth rates of all three species with the objective of developing a model with four differential equations based on the experimental data. The following hypotheses were tested at a theoretical level: Firstly, temperature changes can affect the dynamic behavior of a system by temperature-dependent parameters and interactions and secondly, food web response to temperature cannot be derived from the single species temperature response. The main outcome of the study is that temperature changes affect the parameter range where coexistence is possible within all three species. This has significant consequences on our ideas regarding the evaluation of effects of global warming.     

Polychaete assemblages along a rocky shore on the South Adriatic coast (Mediterranean Sea): patterns of spatial distribution

This study reports on the distribution of polychaete assemblages along a depth gradient from 5 to 25 m, along a stretch of rocky coast near Otranto (South Adriatic sea, Italy). Three depths were sampled in May and November 2000. At each depth three sites (about 100 m apart) were sampled by scraping off three replicate quadrats of 20×20 cm. The experimental design enabled identification of across- and along-shore spatial patterns of variation of polychaete assemblages. A total of 4,168 specimens, belonging to 152 taxa were collected. Multivariate analysis showed that the polychaete assemblages differed significantly among depths with the clearest differences between the shallowest sites (5 m) and the deeper ones (15–25 m). A considerable source of variation among sites at each level of the shore was also exhibited by the analyses, with the greatest differences among sites within depths recorded at 5 m. The species most contributing to the differentiation of assemblages among depths and sites within each depth were identified. Some potential causes of the observed differences are hypothesized and discussed. The importance of quantitative observation to provide the context for studying the underlying ecological processes is also stressed.Communicated by R. Cattaneo-Vietti, Genova