Using patterns of variability to test for multiple community states on rocky intertidal shores

Predictions based on theory of multiple stable states suggest that larger perturbations should lead to more unpredictable patterns of succession. This prediction was tested in the Gulf of Maine using data from 60 intertidal plots of varying size that were experimentally cleared of the rockweed Ascophyllum nodosum and from 14 benchmark sites from throughout the Gulf. Rockweed was removed from the experimental clearings ranging from 1 to 8 m in diameter in 1996 and data collected in 2004 were used to test effects of clearing size and location on divergence and variability in species composition. Benchmark data were collected in 2005, and the 14 sites were from a dataset on 53 sites throughout the Gulf of Maine. The selected sites were randomly chosen from all sites with > 80% canopy cover by A. nodosum and were expected to be similar to uncleared control plots from the experiment. Experimental removal of A. nodosum resulted in clearings at 12 sites within 4 bays. Abundances of gastropods, barnacles, mussels, and fucoid algae and the percentage cover of barnacles, mussels, fucoid algae, bare space, and other species were sampled. CAP and PERMDISP analyses revealed significant differences in multivariate dispersion and variability with both clearing size and location. Variability generally increased with clearing size and location effects were related to the north-south positioning of the sites. Benchmark sites were similar to the experimental control plots but as variable as the largest clearings. Results suggest that succession in larger clearings has been more unpredictable than in small clearings. The pattern of variability in the experimental clearings is consistent with the predictions of multiple stable states. However, the large amount of variation among the benchmark sites was due to mussels and was unexpected. This unexpected variability underscores the importance of sampling benchmark sites as part of experiments.     

Field evidence of trait-mediated indirect interactions in a rocky intertidal food web

Studies on the implications of food web interactions to community structure have often focused on density-mediated interactions between predators and their prey. This approach emphasizes the importance of predator regulation of prey density via consumption (i.e. lethal effects), which, in turn, leads to cascading effects on the prey's resources. A more recent and contrasting view emphasizes the importance of non-lethal predator effects on prey traits (e.g. behaviour, morphology), or trait-mediated interactions. On rocky intertidal shores in New England, green crab ( Carcinus maenas ) predation is thought to be important to patterns of algal abundance and diversity by regulating the density of herbivorous snails ( Littorina littorea ). We found, however, that risk cues from green crabs can dramatically suppress snail grazing, with large effects on fucoid algal communities. Our results suggest that predator-induced changes in prey behaviour may be an important and under-appreciated component of food web interactions and community dynamics on rocky intertidal shores.     

Community stability: effects of limpet removal and reintroduction in a rocky intertidal community

Summary The stability of a high rocky intertidal community was assessed in a controlled field experiment in which the most common consumers, limpets, were temporarily removed. Compared to the unmanipulated plots, the exclusion plots developed greater algal abundance and altered species composition of both algae and barnacles. The community was not perturbed beyond its capacity to recover, since the community structure of the limpet-removal plots converged on the structure of the unmanipulated plots following limpet reintroduction. Different components of the community recovered at different rates, depending on whether or not the species had a size-related escape from the limpets. Algae had no size-related escape from limpets. The difference in algal abundance between manipulated and unmanipulated plots lasted less than six months after limpet reintroduction. Barnacles, however, had a size-related escape from limpets and therefore recovered more slowly. The difference in barnacle species composition between the perturbed and unperturbed plots lasted for 17 months after limpet reintroduction. The length of the limpet removal period (16 or 28 months) did not appear to affect the rate of community recovery.     

Divergent succession and implications for alternative states on rocky intertidal shores

It has been hypothesized that rockweed stands and mussel beds in sheltered bays in the Gulf of Maine, USA, are alternative community states. As a test of this hypothesis, experimental clearings of different sizes were established in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis to determine if successional changes in large clearings developed species assemblages distinctly different from the surrounding A. nodosum stands. Clearings ranging from 1 to 8 m in diameter were created at 12 sites in 4 bays on Swan's Island, Maine, in 1996 to mimic the effects of ice scour, and abundances of gastropods, barnacles, mussels and fucoid algae were monitored until 2002. ANOVAs and MDS showed strong effects of clearing size and divergent successional changes in large clearings. Large clearings were quickly filled in and remained dominated by the alga Fucus vesiculosus L. and the barnacle Semibalanus balanoides (L). There was no evidence for site-specific effects, and Mantel tests showed clearing size was a better predictor of species composition than geographic distances among sites. Results suggest that large pulse disturbances using clearings of 8 m in diameter can initiate divergent successional pathways and have a protracted effect on species composition. Results are also consistent with the hypothesis that mussel beds and rockweed stands in sheltered bays may be alternative community states.     

Generalizing from experiments: is predation strong or weak in the New England rocky intertidal?

Summary Petraitis (1990) recently critized previous generalizations regarding the effects of predation in the New England rocky intertidal region (e.g., Menge 1976; Lubchenco and Menge 1978). Contrary to Lubchenco's and my conclusions, Petraitis concluded that (1) barnacles and not mussels are the favored prey of dogwhelks and (2) barnacles and not dogwhelks control mussel abundances in the mid and low rocky intertidal region. I provide evidence that these criticisms are unwarranted. First, Lubchenco and I never claimed that diet composition reflected prey preference. Moreover, predators can influence prey abundance without preferring the prey. Hence, claims regarding preferences have no bearing on our conclusions. Second, Petraitis' experiments do not invalidate Lubchenco's and my experimental results. Reanalyses of our experimental data support the earlier conclusion that at wave-sheltered sites, whelks reduce the abundance of mussels independently of barnacle abundances. Further, at all but one of Lubchenco's and my study sites, predator densities were higher than at Petraitis' site. Thus, the absence of a predator effect in Petraitis's study was most likely due to low predator density rather than a lack of generality of our earlier results. This reevaluation therefore suggests that within a broader conceptual framework, Petraitis' apparently divergent results are actually consistent with ours.     

A test for top-down cascade in a detritus-based food web by litter-dwelling web spiders

A top-down cascade in detritus-based food webs has the potential to alter ecosystem functioning but its prevalence in terrestrial systems remains unclear. We manipulated the density of the dominant web spider Neriene brongersmai for 1.5 years to see whether the cascading effect would propagate down to the litter decomposition rate in the forest-floor of a Japanese cedar forest. Decreasing spider abundance increased collembolan density, but it did not change the biomass of microorganisms or the litter decomposition rate. We propose possible explanations for the absence of a top-down cascade and discuss the difficulty in generalizing the presence and absence of a cascading effect in soil ecosystems.     

Climate-driven interactions among rocky intertidal organisms caught between a rock and a hot place

To explore how climate may affect the structure of natural communities, we quantified the role of thermal stress in setting the high intertidal borders of the acorn barnacle, Semibalanus balanoides. At sites north and south of Cape Cod, a major faunal and thermal boundary on the east coast of North America, we examined the interacting effects of thermal stress and recruit density on individual survivorship. At hotter southern sites, particularly in bays, high intertidal barnacle survivorship was enhanced by experimental shading or by neighbors which ameliorate heat and desiccation stresses. In contrast, at cooler northern bay and coastal sites, neither shading nor group benefits increased barnacle survival, and mortality patterns were driven primarily by predators with largely boreal distributions. Our field results, like recent laboratory microcosm studies, suggest that predicting even simple community responses to climate change may be more complex than is currently appreciated. Received: 26 January 1999 / Accepted: 5 April 1999     

The structure and replenishment of rocky shore intertidal communities and biogeographic comparisons

Marine scientists have made many recent advances in understanding the connections between the structure of benthic communities, replenishment of populations through dispersal processes, and interactions with the nearshore water mass. In this review, some of the themes and models relating to these processes and interactions are discussed. Benthic–pelagic coupling models are in the early stages of development, but encompass oceanic processes such as upwelling and downwelling, the transport of larvae and their arrival back to shore to settle. Most current knowledge of these processes is based on a few taxa, especially barnacles and mussels. This is discussed with reference and comparison to macroalgae, which dominate much of the intertidal zone on temperate rocky shores, and have quite different life histories, transport and settlement processes. The role of key, habitat-dominating species is discussed, particularly their early life histories, with reference to the differences in community composition, nearshore dynamics and settlement processes in different countries. Finally, some suggestions are made for future work to fill gaps in understanding about rocky shore communities.     

An experimental test of the effects of variation in recruitment intensity on intertidal community composition

The rocky intertidal has been a model system for experimentally testing hypotheses regarding the factors that structure natural communities. Many ecologists have proposed that changes in the intensity of recruitment of individuals into a community should influence community structure. Past work investigating this hypothesis has primarily been surveys of recruitment and community structure across large spatial scales. Surprisingly, no researchers to date have manipulated recruitment into a rocky intertidal community to assess the response of interactions of the whole community to variation in recruitment intensity. We manipulated the densities of Balanus glandula and Chthamalus spp. recruits across a four-fold difference at two sites spanning the Monterey Bay, California, USA, to experimentally test if differences in recruitment intensity influenced initial changes in community composition and if these changes persisted through time. The results of this work indicate that differences in recruitment influence community composition initially, but that these changes can be short-lived.     

The structure of a plant-pollinator food web

The pollination biology literature is dominated by examples of specialization between plants and their pollinators. However, a recent review shows that it is generalization that prevails in the field, with most plants having a number of pollinators and most pollinators visiting a number of plants. Consequently, the vast majority of plant–pollinator interactions are embedded in a complex web of plant–pollinator interactions. These plant-pollinator webs can be studied in the manner of conventional food webs and the aim of this paper is to illustrate how contemporary methods of web construction and analysis can be applied to plant-pollinator communities.     

Diversity,heterogeneity and consumer pressure in a tropical rocky intertidal community

Summary Previous studies indicated that at Taboguilla Island (Gulf of Panama), persistence of many intertidal organisms depended on holes and crevices in the rock as refuges from both vertebrate (fishes) and invertebrate (crabs, gastropods, chitons) consumers. Here, we evaluate the influences of substratum heterogeneity and consumers on patterns of diversity of sessile organisms in this habitat. Local substratum topography is highly variable, ranging from smooth to irregular surfaces. Algal crusts typically dominate all low zone rock surfaces, and most other sessile spegies (invertebrates and foliose algae) occur in holes and crevices. Number (S) and diversity (H) of sessile species is lower on homogeneous surfaces than on heterogeneous surfaces. Rate of increase in S with area sampled is positively correlated with substratum heterogeneity; number of species sampled per transect at a homogeneous site would be about 10 vs 30 to 60 on a heterogeneous site. Large fishes and crabs forage intensively over both substratum types, but cannot enter holes and crevices to eat prey. Gastropods, chitons, limpets, and small crabs feed on both substrata but vary in abundance from hole to hole. Prey mortality is thus intense and constant on open surfaces, but variable in space and time in holes and crevices. When consumers are excluded from the general rock surface, algal crusts are settled upon and overgrown by foliose algae, hydrozoans, and sessile invertebrates, particularly bivalves. Both S and H first increase, as sessile species invade and become more abundant, and then decrease as the rock oyster Chama echinata begins to outcompete other species and dominate primary space. Hence, consumers normally keep local diversity low by removing most sessile prey from open surfaces.In these experiments, a consumer pressure gradient was established by removing 0, 1, 2, 3, and all of 4 distinct groups of consumers. As predicted by the intermediate disturbance hypothesis, lowest diversity occurred at lowest (total exclusion) and highest consumer pressure (normal condition). Highest diversity occurred at intermediate consumer pressure. Unexplained variation in this relationship is probably due to quantitative and qualitative differences in consumer regime, variation among plots in substratum heterogeneity, and insufficient time for competitive dominance by Chama to be fully expressed. On a small (0.25 m²) spatial scale, consumers maintain low diversity by keeping prey scarce and causing local extinctions. On larger spatial scales, they may maintain and even produce high diversity through their interaction with substratum heterogeneity and possibly low dispersal rates of sessile species.     

Epiphytes on the rocky intertidal red alga RhodomelaLarix (Turner) C. Agardh: Negative effects on the host and food for herbivores?

The diversity of epiflora and fauna associated with a dominant turf-forming alga was examined in intertidal communities on the central Oregon coast. Epiphytes associated with the red alga, Rhodomelalarix (Turner) C. Agardh, were examined by surveying intertidal areas for the presence of epiphytes, and by following changes in epiphyte cover in marked quadrats of R. larix. The alga is host for at least 17 species of sessile plants and animals. To determine the role of some of the larger epiphytes in the community, Rhodomela plants were marked and monitored and herbivore feeding was examined. Data suggest that epiphytes decrease the growth rate of their host, increase the probability of axis breakage and decrease reproductive output. Epiphytes provide food for littorine snails and gammarid amphipods that live in the beds of the plant. Amphipods were found to decrease epiphyte cover on R. larix in laboratory tanks, suggesting that these herbivores may have beneficial effects on the host plant.     

Linearity in the aggregate effects of multiple predators in a food web

Theory in community ecology often assumes that predator species have similar indirect effects and thus can be treated mathematically as a single functional unit (e.g. guild or trophic level). This assumption is questionable biologically because predator species typically differ in their effects, creating the potential for nonlinearities when they coexist. We evaluated the nature of indirect effects caused by three species of hunting spider predators, singly and in multiple species combinations, on grass and herb plants in experimental old-field food webs. Despite the potential for nonlinearity, indirect effects in different multiple predator combinations consistently did not differ significantly from the respective means of the single species effects. Thus, for this experimental system, the whole was simply the average of the parts. Consequently, models which abstract predator species as single trophic levels would successfully predict indirect effects in this system regardless of the composition of the predator fauna.     

Community convergence in a simple microbial food web

Previous studies of communities implicate many potential mechanisms that can create alternate stable states. These include density-dependent foraging behavior, size refuges reached by early colonists, environmental feedback following disturbance, and different initial densities of intraguild predators. Previous work shows that alternate states of varying stability can occur in food webs containing the intraguild predators Blepharisma americanum and Tetrahymena vorax. Differences in colonization history could create the alternate states, consisting of dominance by either Blepharisma or Tetrahymena, but it was unclear whether results depended on effects of initial density or only on changes in the resource base. We manipulated initial densities of both species to determine if density effects alone could create alternate stable states. Convergence of these communities over time indicated that differences in initial density did not create alternate stable states. By default, other factors influenced by colonization history, such as resource availability, may produce alternate states. Models of alternate stable-state phenomena should incorporate differences in resource availability in addition to direct competitive and predatory interactions to provide a more complete depiction of the causes of differences in community composition in otherwise similar habitats.     

Feeding guild structure of a rocky intertidal fish assemblage in central Chile

The identification and analysis of the guild structure of vertebrate assemblages has played a fundamental role in the understanding of the underlying mechanisms responsible for their community organization and structuring. This approach generally has not been undertaken for temperate water intertidal fish assemblages. In central Chile, fishes are important components of the intertidal community, but no studies attempting to understand their organization and structuring have been done. In the present study, the diets of 13 of the most abundant species which inhabit tidepools in the rocky intertidal zone of central Chile were determined. A total of 660 fishes was collected at 5 sites: Los Molles, Con-Cón, Quintay, El Tabo, and Las Cruces. Dietary overlap between all species pairs was calculated and a phenogram of dietary similarity was constructed and analyzed using a bootstrapping technique to objectively determine guild membership. The results showed that the intertidal fish assemblage of central Chile can be divided into three feeding guilds: two guilds consisting of carnivorous species and one guild of omnivorous and herbivorous species. The possible causes and implications of the resulting guild structure and the potential effects of predation by these fishes on other components of the intertidal community are discussed.     

Grazing dynamics in intertidal rockpools: Connectivity of microhabitats

Differences between rockpool and emergent rock communities are often attributed to their contrasting physical conditions. However, differences in grazing pressure between rockpools and open rock could also exert an important structuring role. Greater densities and/or the lack of tidal constraints on foraging may allow grazing intensity to be greater in rockpools. Here, wax discs were deployed to compare grazing intensity between rockpool and emergent rock habitats at each of three tidal heights on a moderately exposed shore in SW England. Grazing intensity was then examined in relation to herbivore density. Grazing intensity in pools was twice that on emergent rock, despite a lower density of herbivores in the rockpools. Of these herbivores, patellid limpets are the dominant grazers on rocky shores throughout the NE Atlantic and are recognised to have a major role in structuring intertidal communities. Thus, subsequent experiments focussed on the influence of limpets in determining the differences in consumer pressure between rockpools and emergent rock. Three alternative explanations were considered: (1) the effect of continuous immersion on grazing intensity in rockpools; (2) differences in limpet species abundance between the two habitats; (3) movement of limpets from emergent rock into pools to feed. The level of grazing pressure exerted by Patella ulyssiponensis (Gmelin), the predominant species living constantly immersed in rockpools, was similar to that of P. vulgata (Linnaeus) which is predominantly found on emergent rock. P. vulgata were observed moving from emergent rock into rockpools during high tide. Manipulative experiments confirmed that these foraging excursions resulted in a 2-fold increase in grazing intensity in the pools. Grazing activity of P. vulgata in rockpools was not consistent between sites and may be influenced by differences in wave exposure and/or the abundance of microbial resources. Elevated consumer pressure in rockpools may be an important factor influencing algal assemblages and probably explains the predominance of grazer resistant-species in these pools.     

The relative importance of herbivory and carnivory on the distribution of energy in a stochastic tri-trophic food web

A three-state, discrete-time Markov chain is used to model the dynamics of energy flow in a tri-trophic food web. The distribution of energy in the three trophic levels is related to the rates of flow between the trophic levels and calculated for the entire range of possible flow values. These distributions are then analysed for stability and used to test the idea that plants are resource-limited and herbivores are predation-limited. Low rates of death and decomposition, when coupled with low rates of herbivory and carnivory, tend to destabilize this food web. Food webs with higher rates of death and decomposition are relatively more stable regardless of rates of herbivory and carnivory. Plants are more prone to resource-limitation and herbivores are, in general, limited by their predators, which supports Hairston et al. (Am. Nat. 94 (1960) 421). The rate of decomposition often mediates the roles of top-down and bottom-up control of energy flow in the food web.     

Intertidal rocky shore biotopes of the Azores: a quantitative approach

Rocky littoral communities of the Azores are dominated by macroalgae where turf formations represent the main life form as on many subtropical and tropical shores. The present study aimed at describing and classifying benthic intertidal biotopes of the Azores using clear quantification and data analysis methodologies to be used in similar studies for spatial and/or temporal comparison. This numerical approach intends to have management and conservation applicability. Thirty study locations along the coast of São Miguel island were characterised by substratum type (cobbles, boulders or bedrock – different stability levels) and exposure to wave action (high, medium or low exposure), from June to August 2002. Algae, sessile and sedentary invertebrates were recorded along transects, and the more conspicuous taxa quantified. Community structure is mainly shaped by substratum stability – unstable cobble communities are separate from those of more stable boulders and bedrock. Boulders present an intermediate community composition between cobbles and bedrock. Exposure to wave action induces smaller variation in community composition. Sixteen biotopes have been recognised. Ultimately the present paper provides an objective set of biotope definition tools for a broad public whose interest focuses on spatial and temporal comparison of coastal communities and associated habitats.