Predation intensity in a rocky intertidal community

Summary The predation intensity exerted by populations of the gastropod Thais lapillus at different study areas in the rocky intertidal community of New England is unrelated to predator density. Specifically, very similar intensities are exerted by populations differing in density by at least an order of magnitude. Predation intensity is, in part, a joint function of individual rates of prey consumption and various environmental characteristics. Major factors potentially affecting the individual feeding rates of Thais are (1) prey abundance and productivity, (2) other predators, (3) canopy-forming algae, (4) wave shock, (5) desiccation and (6) snail phenotype and/or history. The effects of the first two of these factors seem unimportant. The effects of the latter 4 on prey consumption rates were studied by estimating field feeding rates of snails held in cages with prey in microhabitats which were characterized by one of two alternative states of each factor. For example, microhabitats could be exposed or protected, at higher or lower levels in the mid intertidal, or under a canopy or not. In addition, exposed-phenotype or protected-phenotype snails were used in each experiment.All of factors (3) to (6) had statistically significant effects except wave shock. The latter would probably also have had a significant effect if the experiments had been performed in the stormier part of the year as well as late summer. The results indicate that sparse populations of Thais can exert intense predation pressure on their prey if they are in protected sites covered with a dense canopy (i.e. in cool, moist habitats in calm waters). Areas with sparser canopy (i.e. greater desiccation stress) and more severe wave shock or both apparently reduce average feeding rates of snails. This appears to explain the paradoxical lack of correlation between predation intensity and snail density.An unexpected result with potentially major implications is the nonlinear response of Thais feeding rates to combinations of factors (3) to (6). Four-way analyses of variance on experiments at exposed and protected sites indicate that 7 of 14 1st-order interactions, 2 of 8 2nd-order interactions, and even 1 of 2 3rd-order interactions are statistically significant. These results suggest that individual predators cannot be assumed to be identical, and that socalled higher order interactions cannot be safely ignored in models of interacting multi-species systems. Hence, it appears that to obtain a thorough understanding of the organization of natural communities, both field and theoretical ecologists alike should begin to grapple with such complexities of nature rather than ignore them.     

A test of the Menge-Sutherland model of community organization in a tropical rocky intertidal food web

Summary Menge and Sutherland (1976) predicted that in physically benign habitats: (1) community structure will be most strongly affected be predation, (2) the effect of predation will increase with a decrease in trophic position in the food web, (3) trophically intermediate species will be influenced by both predation and competition, and (4) competition will occur among prey species which successfully escape consumers. These predictions were tested in a tropical rocky intertidal community on the Pacific coast of Panama. The most abundant mobile species included fishes and crabs, which occupied the top trophic level, and predaceous gastropods and herbivorous molluscs, which occupied intermediate trophic levels. The most abundant sessile organisms were encrusting algae, foliose algae, barnacles, and bivalves. Diets were broad and overlapping, and 30.3% of the consumers were omnivorous. Each consumer group had strong effects on prey occurring at lower trophic levels: (1) Fishes and crabs reduced the abundance of predaceous snails, herbivorous molluscs, foliose algae, and sessile invertebrates. (2) Predaceous gastropods reduced the abundance of herbivorous molluscs and sessile invertebrates. (3) Herbivorous molluscs reduced the abundance of foliose algae and young stages of sessile invertebrates, and altered relative abundances of the encrusting algae. The encrusting algae, although normally the dominant space occupiers, proved to be inferior competitors for space with other sessile organisms when consumers were experimentally excluded. However, the crusts escaped consumers by virtue of superior anti-herbivore defenses and competed for space despite intense grazing. Observations do not support the hypothesis that the trophically intermediate species compete. Hence, with the exception of this last observation, the predictions of the Menge and Sutherland model were supported. Although further work is needed to evaluate other predictions of the model in this community, evidence from this study joins an increasing body of knowledge supporting the model. Contradictory evidence also exists, however, indicating that aspects of the model require revision.     

Top-down and bottom-up community regulation in marine rocky intertidal habitats

Strong top-down control by consumers has been demonstrated in rocky intertidal communities around the world. In contrast, the role of bottom-up effects (nutrients and productivity), known to have important influences in terrestrial and particularly freshwater ecosystems, is poorly known in marine hard-bottom communities. Recent studies in South Africa, New England, Oregon and New Zealand suggest that bottom-up processes can have important effects on rocky intertidal community structure. A significant aspect of all of these studies was the incorporation of processes varying on larger spatial scales than previously considered (10’s to 1000’s of km). In all four regions, variation in oceanographic factors (currents, upwelling, nutrients, rates of particle flux) was associated with different magnitudes of algal and/or phytoplankton abundance, availability of particulate food, and rates of recruitment. These processes led to differences in prey abundance and growth, secondary production, consumer growth, and consumer impact on prey resources. Oceanographic conditions therefore may vary on scales that generate ecologically significant variability in populations at the bottom of the food chain, and through upward-flowing food chain effects, lead to variation in top-down trophic effects. I conclude that top-down and bottom-up processes can be important joint determinants of community structure in rocky intertidal habitats, and predict that such effects will occur generally wherever oceanographic ‘discontinuities’ lie adjacent to rocky coastlines. I further argue that increased attention by researchers and of funding agencies to such benthic–pelagic coupling would dramatically enhance our understanding of the dynamics of marine ecosystems.     

Microhabitats, thermal heterogeneity, and patterns of physiological stress in the rocky intertidal zone.

Thermal stress has been considered to be among the most important determinants of organismal distribution in the rocky intertidal zone. Yet our understanding of how body temperatures experienced under field conditions vary in space and time, and of how these temperatures translate into physiological performance, is still rudimentary. We continuously monitored temperatures at a site in central California for a period of two years, using loggers designed to mimic the thermal characteristics of mussels, Mytilus californianus. Model mussel temperatures were recorded on both a horizontal and a vertical, north-facing microsite, and in an adjacent tidepool. We periodically measured levels of heat shock proteins (Hsp70), a measure of thermal stress, from mussels at each microsite. Mussel temperatures were consistently higher on the horizontal surface than on the vertical surface, and differences in body temperature between these sites were reflected in the amount of Hsp70. Seasonal peaks in extreme high temperatures ("acute" high temperatures) did not always coincide with peaks in average daily maxima ("chronic" high temperatures), suggesting that the time history of body temperature may be an important factor in determining levels of thermal stress. Temporal patterns in body temperature during low tide were decoupled from patterns in water temperature, suggesting that water temperature is an ineffective metric of thermal stress for intertidal organisms. This study demonstrates that spatial and temporal variability in thermal stress can be highly complex, and "snapshot" sampling of temperature and biochemical indices may not always be a reliable method for defining thermal stress at a site.     

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.     

Response of a rocky intertidal ecosystem engineer and community dominant to climate change

To evaluate how climate change might impact a competitively dominant ecological engineer, we analysed the growth response of the mussel Mytilus californianus to climate patterns [El Niño-Southern Oscillation, Pacific Decadal Oscillation (PDO)]. Mussels grew faster during warmer climatic events. Growth was initially faster on a more productive cape compared to a less productive cape. Growth rates at the two capes merged in 2002, coincidentally with a several year-long shift from warm to cool PDO conditions. To determine the mechanism underlying this response, we examined growth responses to intertidal sea and air temperatures, phytoplankton, sea level and tide height. Together, water temperature (32%) and food (12.5%) explained 44.5% of the variance in mussel growth; contributions of other factors were not significant. In turn, water temperature and food respond to climate-driven variation in upwelling and other, unknown factors. Understanding responses of ecosystem engineers to climate change will require knowing direct thermal effects and indirect effects of factors altered by temperature change.     

中街山列岛岩相潮间带大型底栖动物的群落格局

于2006年9月和2007年7月在中街山列岛4个有居民海岛岩相潮间带设立8个断面,对大型底栖动物的群落结构进行了调查。共获得大型底栖动物34种,其中腔肠动物1种,环节动物3种,甲壳动物10种,软体动物20种。优势种有日本笠藤壶(Tetraclita japonica)、紫贻贝(Mytilus edulis)和节蝾螺(Turb articulatus)等4种。以季节和站位为因子对多样性指数进行双因素方差分析,结果表明只有夏、秋季丰富度指数(P=0.028)存在显著差异,其他指数的方差分析没有显著差异(P>0.05)。3个潮带4个岛屿共12个站位的聚类分析和MDS标序分析表明,12个群落可分为3组(Ⅰ、Ⅱ和Ⅲ组)。大型底栖动物ABC曲线分析表明底栖动物群落尚未受到干扰。     

Anthropogenic harvesting pressure and changes in life history: insights from a rocky intertidal limpet

The importance of large breeding individuals for maintaining the health of marine fish and invertebrate populations has long been recognized. Unfortunately, decades of human harvesting that preferentially remove larger individuals have led to drastic reductions in body sizes of many of these species. Such size-selective harvesting is particularly worrisome for sequentially hermaphroditic species where the larger size classes are composed primarily of one sex. Whether these species can maintain stable sex ratios under sustained harvesting pressure depends on the level of plasticity of their life-history traits. Here, we show that populations of a marine limpet (Lottia gigantea) can adjust a fundamental aspect of their life history (the timing of sex change) when subjected to size-selective harvesting. As predicted by theoretical models, individuals from harvested populations change sex at smaller sizes and grow at slower rates compared to individuals from protected populations. In addition, the relative size at which the change from male to female occurs remains constant (~0.75; size at sex change/maximum size) across populations, regardless of harvesting pressure. Our results show that population-level demographic and life-history data, in conjunction with existing theory, can be sufficient to predict the responses of sequential hermaphrodites to harvesting pressure. Furthermore, they suggest such species can potentially adapt to size-selective harvesting.     

Effects of consumers and enrichment on abundance and diversity of benthic algae in a rocky intertidal community

Human alteration of nutrient cycling and the densities of important consumers have intensified the importance of understanding how nutrients and consumers influence the structure of ecological systems. We examined the effects of both grazing and nutrient enrichment on algal abundance and diversity in a high-intertidal limpet-macroalgal community on the South Island of New Zealand, a relatively nutrient-poor environment. We used a fully factorial design with three levels each of grazing (manipulations of limpet and snail densities) and nutrients (nutrient-diffusers attached to the rock). Top-down control by grazers appears to be the driving organizing mechanism for algal communities in this system, with strong negative effects of grazing on algal diversity and abundance across all levels of nutrient enrichment. However, in contrast to the conclusions drawn from the analysis of the whole algal community, there was an interactive effect of grazing and enrichment on foliose algae, an important component of the algal system. When herbivory was reduced to very low levels, enrichment generated increases in the abundance and biomass of foliose algae. As expected, top-down control was the primary determinant of algal community structure in this system, controlling abundance and diversity of macrophytes on the upper shore. Contrary to expectations, however, increased nutrients had no community-wide effects, although foliose algal abundance increases were greatest with high nutrients and reduced grazing. It seems likely that most of the corticated algal species have limited capacity to respond to nutrient pulses in this nutrient-poor environment.     

Scale, environment, and trophic status: the context dependency of community saturation in rocky intertidal communities

Our understanding of the relative influence of different ecological drivers on the number of species in a place remains limited. Assessing the relative influence of local ecological interactions versus regional species pools on local species richness should help bridge this conceptual gap. Plots of local species richness versus regional species pools have been used to address this question, yet after an active quarter-century of research on the relative influence of local interactions versus regional species pools, consensus remains elusive. We propose a conceptual framework that incorporates spatial scale and ecological interaction strength to reconcile current disparities. We then test this framework using a survey of marine rocky intertidal algal and invertebrate communities from the northeast Pacific. We reach two main conclusions. First, these data show that the power of regional species pools to predict local richness disintegrates at small spatial scales coincident with the scale of biological interactions, when studying ecologically interactive groups of species, and in generally more abiotically stressful habitats (e.g., the high intertidal). Second, conclusions of past studies asserting that the regional species pool is the primary driver of local species richness may be artifacts of large spatial scales or ecologically noninteractive groups of species.     

The grazing effect of limpets on the macroalgal community of a rocky intertidal shore

Abstract Abundance of macro-algae in the mid-littoral zone on a Victorian intertidal rocky shore varied seasonally, algae being rare in Summer and common in Winter and Spring. Field experiments demonstrated that, of two species of grazing limpets (Cellana tramoserica and Siphonana diemenensis), only Siphonaria had a major effect on the abundance of foliose algae (e.g. Scytopsiphon lomeniarid) and neither species had a great effect on the encrusting algae (Ralfsia spp.). There was no evidence of competition for food between the two species of limpets, in contrast with results found for similar species in New South Wales.     

Long term persistence in a rocky intertidal fish assemblage

Environmental Biology of Fishes - A rocky intertidal fish assemblage at Dillon Beach, CA was persistent over 42 months. This time period represents greater than one mean generation time for...     

Seasonal changes in community structure along a vertical gradient: patterns and processes in rocky intertidal sessile assemblages

Population Ecology - Here we considered two fundamental questions in community ecology regarding the relationship between seasonal changes in community structure and environmental gradients: (i)...     

Effects of Ulva spp. and seasonal disturbances on the recovery of a temperate rocky intertidal community

The inhibitory effects of ulvoid algae and their seasonality on the recovery and succession of an intertidal algal community were investigated under field conditions using a two‐factorial design. The experiment was conducted in the mid‐low intertidal zone on the Southern coast of Korea. In spring and fall, each of the ten 50 × 50 cm plots was disturbed artificially, and Ulva spp. were continuously excluded from five plots of the seasonal plots. The succession of spring plots was nearly twice as fast as that of the fall plots (spring, 4–6 months; fall, 10–15 months), indicating that the seasonal effects contributed significantly to the speed of the succession. The inhibition by Ulva species delayed the succession; however, the strength of this delay was limited largely by the duration and persistence of Ulva mass, which was also highly seasonal. The presence of Ulva spp. did not alter the final assemblage at the climax stage; instead, it caused a delay in community resilience. This study provides a few analytical factors for community‐wise comparison during the evaluation of the status of successional stages.     

Temporal patterns in the intertidal faunal community at the mouth of a tropical estuary

The use of intertidal sandy beaches by fish and macrocrustaceans was studied at different temporal scales at the mouth of a tropical estuary. Samples were taken along the lunar and diel cycles in the late dry and rainy seasons. Fish assemblage (number of species, density and biomass), crustaceans and wrack biomass, showed significant interactions among all studied factors, and the combination of moon phase and diel cycle, resulting in different patterns of environmental variables (depth, water temperature and dissolved oxygen), affected habitat use by the different species. Variances in faunal community were detected between seasons, stimulated by salinity fluctuations from freshwater input during the rainy season. These differences suggest an important cycling of habitats and an increase in connectivity between adjacent habitats (estuary and coastal waters). Moreover, the results showed that this intertidal sandy beach also provides an alternative nursery and protected shallow‐water area for the initial development phase of many marine and estuarine species. In addition, this intertidal habitat plays an important role in the maintenance of the ecological functioning of the estuarine–coastal ecosystem continuum.     

Spatio–temporal structure of earthworm community and soil heterogeneity in a tropical pasture

The spatio–temporal structure of an earthworm community was assessed in a tropical Colombian pasture and compared with the patterns of soil heterogeneity. Earth–worms were sampled at six dates during a period of one year, while soil properties were assessed at a single date at the end of this period. Sample design was spatially explicit and consisted in a regular grid of 5 m mesh. At each date, 120 samples were taken for earthworm evaluations, while the single soil sampling consisted in 60 points, Multivariate and geostatistical analyses were performed to describe the spatial structure of collected data. Mantel and partial ManteTs tests were used to assess the links between the spatio-temporal patterns of earthworm distribution and soil heterogeneity.
The results clearly demonstrated the existence of structured spatial patterns in the earthworm community which were stable in time, at least at the temporal scale concerned by the study period, it mostly corresponded to an alternation of patches where particular species assemblages dominated. The ecological significance of these assemblages is discussed. Soil properties also displayed a significant spatial organisation. Only a small part of the spatio–temporal structures of earthworm distribution was correlated with soil properties. Small species were more abundant in patches where soil was more compact, while large species were associated with high root biomass and total C levels. The question whether earthworm spatio–temporal distribution induces similar patterns in soil properties or the contrary occurs is discussed. Our results suggest that earthworm community organisation mostly corresponds to a non–equilibrium system where mobile earthworm populations influence soil properties. The overall effect of earthworm community on soil is difficult to assess due to the predominance of factors that preferentially express tbetnselves at large scales in the ecological hierarchy.     

Effects of avian grazing on the algal community and small invertebrates in the rocky intertidal zone

In this study of a rocky intertidal habitat in northern Japan, feeding by avian consumers had significant effects on algal assemblages and small herbivorous invertebrates. The effects of the birds on algae were different from those of invertebrate grazers such as urchins and gastropods. The abundance of the dominant algal species decreased during the grazing period, increased again after the grazing period, and indirectly affected algal species richness and evenness. Avian grazing also decreased the density of tube-dwelling amphipods on the dominant alga, but did not change the density of mobile and free-living isopods. These results suggest that avian grazers may act as habitat modifiers rather than exploitative competitors for the small herbivorous crustaceans. Avian herbivores consumed only the upper parts of large algal fronds, apparently reducing the amount of suitable microhabitat for the small herbivorous crustaceans, which are subject to a variety of physical or biological stress. Thus, avian herbivores function as ecosystem engineers, regulating community structure in a manner different to invertebrate herbivores in rocky intertidal habitats.