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Communities are shaped by a variety of ecological and environmental processes, each acting at different spatial scales. Seminal research on rocky shores highlighted the effects of consumers as local determinants of primary productivity and community assembly. However, it is now clear that the species interactions shaping communities at local scales are themselves regulated by large-scale oceanographic processes that generate regional variation in resource availability. Upwelling events deliver nutrient-rich water to coastal ecosystems, influencing primary productivity and algae-herbivore interactions. Despite the potential for upwelling to alter top-down control by herbivores, we know relatively little about the coupling between oceanographic processes and herbivory on tropical rocky shores, where herbivore effects on producers are considered to be strong and nutrient levels are considered to be limiting. By replicating seasonal molluscan herbivore exclusion experiments across three regions exposed to varying intensity of seasonal upwelling, separated by hundreds of kilometers along Panama's Pacific coast, we examine large-scale environmental determinants of consumer effects and community structure on tropical rocky shores. At sites experiencing seasonal upwelling, grazers strongly limited macroalgal cover when upwelling was absent, leading to dominance by crustose algae. As nutrients increased and surface water cooled during upwelling events, increases in primary productivity temporarily weakened herbivory, allowing foliose, turf and filamentous algae to replace crusts. Meanwhile, grazer effects were persistently strong at sites without seasonal upwelling. Our results confirm that herbivores are key determinants of tropical algal cover, and that the mollusk grazing guild can control initial stages of macroalgal succession. However, our focus on regional oceanographic conditions revealed that bottom-up processes regulate top-down control on tropical shorelines. This study expands on the extensive body of work highlighting the influence of upwelling on local ecological processes by demonstrating that nutrient subsidies delivered by upwelling events can weaken herbivory in tropical rocky shores.

International audience; Understanding the drivers of geographical variation in species distributions, and the resulting community structure, constitutes one of the grandest challenges in ecology. Geographical patterns of species richness and composition have been relatively well studied. Less is known about how the entire set of trophic and non-trophic ecological interactions, and the complex networks that they create by gluing species together in complex communities, change across geographical extents. Here, we compiled data of species composition and three types of ecological interactions occurring between species in rocky intertidal communities across a large spatial extent (~970 km of shoreline) of central Chile, and analyzed the geographical variability in these multiplex networks (i.e., comprising several interaction types) of ecological interactions. We calculated nine network summary statistics common across interaction types, and additional network attributes specific to each of the different types of interactions. We theninvestigated potential environmental drivers of this multivariate network organization. These included variation in sea surface temperature and coastal upwelling, the main drivers of productivity in nearshore waters. Our results suggest that structural properties of multiplex ecological networks are affected by local species richness and modulated by factors influencing productivity and environmental predictability. Our results show that non-trophic negative interactions are more sensitive to spatially structured temporal environmental variation than feeding relationships, with non-trophic positive interactions being the least labile to it. We also show that environmental effects are partly mediated through changes in species richness and partly through direct influences on species interactions, probably associated to changes in environmental predictability and to bottom-up nutrient availability. Our findings highlight the need for a comprehensive picture of ecological interactions and their geographical variability if weare to predict potential effects of environmental changes on ecological communities

[Excerpt] Best known as a "love them or hate them" luxury food, or for their pearls, oysters are also ecosystem engineers, forming vast oyster reefs. Oyster reefs provide habitat for a myriad of species, and support fisheries, improve water quality and provide coastal protection. These services are estimated to be worth US$5,500–$99,000 per hectare per year (Grabowski et al. 2012). Globally, oyster reefs have declined by 85% through destructive overfishing, coastal development, pollution, and introduced competitors, predators and diseases (Beck et al. 2011). Active restoration is becoming an increasingly popular tool to bring back lost oyster reefs and the ecosystem services they provide (Fitzsimons et al. 2019). However, restoration is not always successful, and knowledge about how reefs naturally form and function is vital to improve restoration success. Oyster larvae only settle on hard substrates. Reefs proliferate because oyster shells provide a settlement surface, and oysters provide chemical and sound cues that facilitate larval settlement (Lillis et al. 2013). However, these reefs often form on intertidal sand and mud banks. This raises the question, how do oyster reefs form on mud banks in the absence of hard surfaces?

Surf zones, classified from reflective to dissipative, separate the ocean from shore and subsidies from the coastal ocean must pass through surf zones to reach the shore. We have observed that variations in phytoplankton concentrations in the water over the intertidal zone varied with surf-zone hydrodynamics and we hypothesized that this variation would alter growth rates, population structure, and reproductive output of Mytilus californianus and Balanus glandula. From May 2016 to April 2017, along 7 km of Cape Arago, Oregon, USA surf-zone phytoplankton concentrations were determined weekly at nine sites with varying surf-zone hydrodynamics as indicated by surf-zone widths. Throughout the year, concentrations of phytoplankton in wider, more dissipative surf zones were, on average, 16× higher than in narrow, more reflective surf zones. Similar to previous observations, surf-zone width explained >90% of the variability in phytoplankton concentrations in the surf-zone. On average, ~83% of B. glandula had egg lamellae at more dissipative shores compared to only 8% at more reflective. An index of potential reproductive output by barnacle populations as measured by ash free dry mass (AFDM) of egg lamellae was ~243× larger at more dissipative than at reflective shores and surf-zone phytoplankton concentrations and surf-zone width explained ~96 and 92% of the variability in this index. On average, density of M. californianus was ~2× higher at more reflective shores, but 60% of these individuals were smaller and non-reproductive compared to only 24% at the more dissipative sites. The gonad tissue mass/m2 of medium sized mussels was ~5× greater at more dissipative than reflective shores. Surf-zone phytoplankton concentrations and surf-zone width explained ~80% and 65% of the variability in individual mussel gonad mass and ~69% and 56% of the variability in mussel population gonad mass, respectively. M. californianus were out-planted to assess growth rates and, after 5 months, average body mass was ~3× greater at more dissipative than reflective shores. Surf-zone phytoplankton concentrations and width explained ~85% and 92% of the mass increase, respectively. Phytoplankton subsidies varied with surf-zone hydrodynamics altering the growth and reproductive output of two ecologically important intertidal filter feeders.

Ecosystem engineers facilitate communities by providing a structural habitat that reduces abiotic stress or predation pressure for associated species. However, disturbance may damage or move the engineer to a more stressful environment, possibly increasing the importance of facilitation for associated communities. In this study, we determined how disturbance to intertidal boulders (i.e., flipping) and the subsequent movement of a structural ecosystem engineer, the tube-forming serpulid worm Galeolaria caespitosa, from the bottom (natural state, low abiotic stress) to the top (disturbed state, high abiotic stress) surface of boulders influenced the importance of facilitation for intertidal communities across two intertidal zones. Theory predicts stronger relative facilitation should occur in the harsher environments of the top of boulders and the high intertidal zone. To test this prediction, we experimentally positioned boulders with the serpulids either face up or face down for 12 months in low and high zones in an intertidal boulder field. There were very different communities associated with the different boulders and serpulids had the strongest facilitative effects on the more stressful top surface of boulders with approximately double the species richness compared to boulders lacking serpulids. Moreover, within the serpulid matrix itself there was also approximately double the species richness (both zones) and abundance (high zone only) of small invertebrates on the top of boulders compared to the bottom. The high relative facilitation on the top of boulders reflected a large reduction in temperature by the serpulid matrix on that surface (up to 10°C) highlighting a key role for modification of the abiotic environment in determining the community-wide facilitation. This study has demonstrated that disturbance and subsequent movement of an ecosystem engineer to a more stressful environment increased the importance of facilitation and allowed species to persist that would otherwise be unable to survive in that environment.

The 18.6-yr lunar-nodal cycle drives changes in tidal amplitude globally, affecting coastal habitat formation, species and communities inhabiting rocky shores, and salt marsh vegetation. However, the cycle's influence on salt marsh fauna lacked sufficient long-term data for testing its effect. We circumvented this problem by using salt marsh mosquito records obtained over a period of over four decades in two estuaries in the northeastern USA. Salt marsh mosquito habitat is near the highest tide level where the impact of the nodal cycle on flood frequency is greatest. Wavelet spectral and cross-correlation analyses revealed periodicity in salt marsh mosquito abundance that was negatively correlated with tidal amplitude. Tidal amplitude was a significant predictor of salt marsh mosquito abundance with the cycle maxima coinciding with lower mosquito populations, possibly due to access by predatory fish. However, these effects were detected only at the location with extensive salt marsh habitat and astronomical tides and were weakened or lacked significance at the location with small microtidal salt marshes and wind-driven tides. Mosquitoes can serve as proxy indicators for numerous invertebrate species on the salt marsh. These predictable cycles and their effects need to be taken into consideration when investigating, restoring, or managing intertidal communities that are also facing sea-level rise.

© 2017 by the Ecological Society of America Ecological invasions are facilitated by pre-adaptation and phenotypic plasticity, upon which evolution can act. The rapid invasion of the intertidal grass Spartina alterniflora in China during the last 36 yr is a test case for the roles of these mechanisms. A previous study of S. alterniflora in China found strong latitudinal clines in vegetative and sexual traits and concluded that most of this variation was due to phenotypic plasticity. Recent observations suggested provenance-by-environment interactions, and we employed common gardens at multiple latitudes as a test of this idea. Phenotypically, field plant height, which correlates strongly with biomass and other indices of vegetative performance in this species, showed a hump-shaped relationship across 10 sites, covering 19° of latitude; field seed set increased linearly with latitude. To assess the role of plasticity vs. genetic differentiation in these patterns, we grew plants from the ten field sites in three common gardens at low (20.9° N), mid (28.3° N), and high (38.0° N) latitudes to maturity, at 18 months. Plant height varied among common gardens, with the tallest plants at mid latitude, mirroring the field pattern, consistent with the previous study. Within the gardens, latitude of origin also affected plant height. Seed set varied among the gardens, with the greatest values at high latitudes, again mirroring the field pattern and indicating substantial plasticity. Evidence of evolution was found as increasing seed set with latitude among provenances within common gardens. However, the effect differed among common gardens, with the greatest slope in the high-latitude garden, lower slope in the mid-latitude garden, and no relationship in the low-latitude garden, indicating a provenance-by-environment interaction. The number of surviving plants also suggested a provenance-by-environment interaction; no relationship with latitude among provenances in the two southern gardens and increasing survival with latitude in the northern garden. Field seedling density was ~200-fold greater at high than at low latitude sites. The profuse seed germination and recruitment in the north could have created high selection intensity resulting in evolution of reproductive traits at high latitudes with the result that the mechanisms of the invasion differ with latitude.

Prey often retreat into the safety of refuges for protection from predators. This shift into refuge can reduce foraging opportunities, escalating the costs of risk and the strength of nonconsumptive effects. Such costs, however, may be shaped by the variation in resources that refuges harbor for prey foraging (i.e., refuge quality), and change dynamically via impacts on prey state. Despite its potential importance, we lack an explicit understanding of how refuge quality impacts prey performance under risk. Using a rocky intertidal food chain, we examined the interaction between predation risk and the amount of resources available for prey in refuge. We found that refuges with more resources greatly reduce the costs of refuge use, and that nonconsumptive effects are thereby weakened by as much as one-half, with especially strong impacts on prey growth and growth efficiency. These results suggest that failure to consider refuge quality could result in overestimation of the negative effects associated with prey refuge use.

Understanding the relative roles of species interactions and environmental factors in structuring communities has historically focused on local scales where manipulative experiments are possible. However, recent interest in predicting the effects of climate change and species invasions has spurred increasing attention to processes occurring at larger spatial and temporal scales. The "meta-ecosystem" approach is an ideal framework for integrating processes operating at multiple scales as it explicitly considers the influence of local biotic interactions and regional flows of energy, materials, and organisms on community structure. Using a comparative-experimental design, we asked (1) what is the relative importance of local biotic interactions and oceanic processes in determining rocky intertidal community structure in the low zone within the Northern California Current System, and (2) what factors are most important in regulating this structure and why? We focused on functional group interactions between macrophytes and sessile invertebrates and their consumers (grazers, predators), how these varied across spatial scales, and with ocean-driven conditions (upwelling, temperature) and ecological subsidies (nutrients, phytoplankton, sessile invertebrate recruits). Experiments were conducted at 13 sites divided across four capes in Oregon and northern California. Results showed that biotic interactions were variable in space and time but overall, sessile invertebrates had no effect on macrophytes while macrophytes had weakly negative effects on sessile invertebrates. Consumers, particularly predators, also had weakly negative effects on both functional groups. Overall, we found that 40-49% of the variance in community structure at the local scale was explained by external factors (e.g., spatial scale, time, upwelling, temperature, ecological subsidies) vs. 19-39% explained by functional group interactions. When individual functional group interaction strengths were used, only 2-3% of the variation was explained by any one functional group while 28-54% of the variation was explained by external factors. We conclude that community structure in the low intertidal zone is driven primarily by external factors at the regional scale with local biotic interactions playing a secondary role.

Most sedentary marine animals disperse from their place of origin during their initial life stages as larvae. The delivery of planktonic larvae back to coastal adult habitats after weeks or months of offshore development is commonly thought to be stochastic, resulting in large recruitment fluctuations and making predictive understanding of population dynamics difficult. Time series of invertebrate settlement on intertidal shores have been used to infer how various oceanographic processes deliver planktonic larvae ashore. However, the possibility that successful settlement may involve a series of different transport mechanisms, which are sequentially utilized by late-stage larvae, has received little attention. To address this, we monitored both the delivery of mussel and barnacle larvae to inner-shelf moorings positioned 200-1400 m from the shore, and larval settlement in the intertidal adult habitat, at two contrasting sites: a headland forming an upwelling center and a downstream bay. Model selection was employed to determine the most likely scenario(s) of larval onshore transport from four a priori transport mechanisms individually and in combination: (1) upwelling or relaxation/downwelling, (2) tidal motions, (3) diurnal sea breezes, and (4) surface waves. Mussel larvae were delivered to the inner shelf during upwelling in the bay, but during downwelling at the headland, and were further transported to the shore by surface waves at both locales. In contrast, the delivery of barnacle larvae to the inner shelf occurred during relaxation/downwelling events at both sites, and intertidal settlement coincided with spring tides, suggesting a role for internal tides in their onshore transport. Thus, sequential mechanisms appear to be utilized by larvae to get to the shore, involving interactions of regional-scale upwelling/downwelling processes and local-scale tidal and surface-wave processes, which differ among taxa and among sites with different topography. A bottleneck for larval delivery across the surf zone may be a result of out-of-phase steps in sequential transport mechanisms leaving larvae lost "in transit."

I investigated predation on mussels occurring at upper levels of a sheltered rocky shore on Santa Catalina Island, USA. Despite continuous recruitment, the mussels Mytilus californianus, M. edulis, and Septifer bifurcatus seldom survived to lengths >1.5 cm., and they remained inconspicuous beneath a canopy of perennial algae. Surveys using scuba at high tide and gut—content analyses showed that intertidal mussels were preyed upon by whelks (Ceratostoma nuttali and Maxwellia gemma), diurnal fishes (Halichoeres semisinctus and Semicossyphus pulcher), and nocturnal spiny lobsters (Panulirus interruptus). Whelks and fishes foraged primarily on mid to low shore levels, the lower half of the mussels' vertical range. However, the lobsters foraged over the entire intertidal zone and were the only predator to specialize on mussels. Exclusion of lobsters from upper shore levels caused significant increases in the densities and sizes of the mussels and herbivorous molluscs. Increases in the percent cover of mussels coincided with decreases in the percent cover understory algae. Previous studies open temperate shores indicate that mussel beds form in upper shore refugia where brief submergence intervals preclude predators that require long foraging times and are intolerant of exposure. On some sheltered temperate shores, predators that move quickly with the tides may influence upper shore zonation. Pronounced variation in prey population structure can result from shifts in the relative effectiveness of different foraging characteristics within diverse predator faunas.

Understanding the role of larger-scale processes in modulating the assembly, structure, and dynamics of communities is critical for forecasting the effects of climate-change and managing ecosystems. Developing this comprehensive perspective is difficult though, because species interactions are complex, interdependent, and dynamic through space and time. Typically, experiments focus on tractable subsets of interactions that will be most critical to investigate and explain shifts in communities, but qualitatively base these choices on experience, natural history, and theory. One quantitative approach to identify the putative forces regulating communities, without reducing system complexity, is estimating transition probabilities among species occupying space (i.e., multispecies Markov chain models). Although not mechanistic, these models estimate the relative frequency and importance of ecological pathways in community assembly and dynamics, and can serve as a framework to identify how pathways change across large scales and which are most important to investigate further. Here, we demonstrate this method in the Gulf of Maine (GOM) intertidal zone, where research has largely focused on the local-scale processes that influence communities, while the mechanisms responsible for more regional shifts in communities are less clear. Transition probabilities of faunal elements were quantified bimonthly for ~2.5 yr in local intertidal communities at three replicate sites in the southern, mid-coast, and northern GOM. Transitions related to mortality, colonization, and replacement by mussels, barnacles, red algae, and encrusting corallines differed regionally, suggesting specific pathways related to consumer pressure and recruitment vary across the GOM with shifting intertidal community structure. Combined with species abundance data and insights from previous research, we develop and evaluate the pathways by which communities likely change in the GOM. Species interactions in local communities can be complex, and this complexity should be incorporated into hypothesis building, experiments, theory, interpretations, and forecasts in ecology. Such a comprehensive approach will be critical to understand how regional shifts in local interactions can drive large-scale community change.

Nonnative species that form novel habitats strongly affect ecosystem processes. The effects of these ecosystem engineers can be both positive and negative but the mechanisms behind their effects are not well described. In this study we determined the relative importance of three main mechanisms by which invasive ecosystem engineers can facilitate native fauna. The engineer may provide new physical structure that reduces harsh abiotic conditions or gives refuge from predation (both engineering mechanisms), or provide a new profitable food resource (a trophic mechanism). The invasive seaweed Gracilaria vermiculophylla is a novel addition to estuarine intertidal mudflats of the southeastern United States. The epifaunal amphipod Gammarus mucronatus is up to 100 times more abundant on Gracilaria-invaded mudflats compared to uninvaded mudflats. Feeding assays, a survivorship experiment and stable isotope analysis demonstrated that Gammarus consumes little Gracilaria and cannot survive on Gracilaria alone. However, the structural engineering effects of Gracilaria greatly enhanced the survivorship of Gammarus in the presence of predators during high tide and when exposed to harsh abiotic conditions during low tide. Our results demonstrate that invasive ecosystem engineers can dramatically affect the distribution and abundance of native species by providing a novel protective structure.

Predicting the effects of climate change on ecological communities requires an understanding of how environmental factors influence both physiological processes and species interactions. Specifically, the net impact of temperature on community structure depends on the relative response of physiological energetic costs (metabolism) and energetic gains (ingestion of resources) that mediate the flow of energy throughout a food web. However, the relative temperature scaling of metabolic and ingestion rates have rarely been measured for multiple species within an ecological assemblage and it is not known how, and to what extent, these relative scaling differences vary among species. To investigate the relative influence of these processes, I measured the temperature scaling of metabolic and ingestion rates for a suite of rocky intertidal species using a multiple regression experimental design. I compared oxygen consumption rates (as a proxy for metabolic rate) and ingestion rates by estimating the temperature scaling parameter of the universal temperature dependence (UTD) model, a theoretical model derived from first principles of biochemical kinetics and allometry. The results show that consumer metabolic rates (energetic costs) tend to be more sensitive to temperature than ingestion rates (energetic gains). Thus, as temperature increases, metabolic rates tend to increase faster relative to ingestion rates, causing the overall energetic efficiencies of these rocky intertidal invertebrates to decline. Metabolic and ingestion rates largely scaled in accordance with the UTD model; however, nonlinearity was evident in several cases, particularly at higher temperatures, in which alternative models were more appropriate. There are few studies where multiple rate dependencies are measured on multiple species from the same ecological community. These results indicate that there may be general patterns across species in the temperature scaling of biological rates, with important implications for forecasting temperature effects on ecological communities.

Patterns in rocky intertidal seaweed biodiversity influence the resilience and functioning of these important primary producer communities. In turn, seaweed biodiversity patterns are the result of many ecological factors. We determined the influences of thermal and desiccation stress, herbivory, and nutrients on seaweed biodiversity on a northern California rocky shoreline. In a fully crossed design at two tidal heights at wave-protected and exposed sites, we deployed screens to reduce stress, removed herbivores, and added nutrients for 18 months. The treatments reduced temperature, increased relative humidity, decreased herbivore abundances, and increased nitrogen in both seawater and seaweeds. Seaweed abundance and biodiversity (cover, biomass, species richness, diversity, evenness, and community composition) were influenced by tidal height, physical stress, and herbivores. Wave exposure affected all response variables except biomass and evenness. Stress and herbivores had independent additive effects on seaweed abundance and diversity. Physical stress did not make the community as a whole more susceptible to herbivores, and screens had overarching positive effects on seaweed biodiversity even though they also had positive effects on herbivore abundance. Nutrients had virtually no effect on seaweed biodiversity, and we observed no bottom-up effects of nutrient addition on herbivore density or biomass. Small green algae and diatoms were important contributors to overall algal cover and to changes in composition across treatments, but larger macroalgae dominated the species richness response. The striking absence of interactions between stress and herbivory highlights how seaweed communities can respond independently to important drivers of biodiversity. Thus, nonadditive, potentially synergistic effects do not necessarily complicate the understanding of how seaweed biodiversity responds to environmental change.

Increasing evidence shows that spatial interactions between sedentary organisms can structure communities and promote landscape complexity in many ecosystems. Here we tested the hypothesis that reef-forming mussels (Mytilus edulis L.), a dominant intertidal ecosystem engineer in the Wadden Sea, promote abundances of the burrowing bivalve Cerastoderma edule L. (cockle) in neighboring habitats at relatively long distances coastward from mussel beds. Field surveys within and around three mussel beds showed a peak in cockle densities at 50-100 m toward the coast from the mussel bed, while cockle abundances elsewhere in the study area were very low. Field transplantation of cockles showed higher survival of young cockles (2-3 years old) and increased spat fall coastward of the mussel bed compared to within the bed and to areas without mussels, whereas growth decreased within and coastward of the mussel bed. Our measurements suggest that the observed spatial patterns in cockle numbers resulted from (1) inhibition effects by the mussels close to the beds due to preemptive algal depletion and deteriorated sediment conditions and (2) facilitation effects by the mussels farther away from the beds due to reduction of wave energy. Our results imply that these spatial, scale-dependent interactions between reef-forming ecosystem engineers and surrounding communities of sedentary benthic organisms can be an important determinant of the large-scale community structure in intertidal ecosystems. Understanding this interplay between neighboring communities of sedentary species is therefore essential for effective conservation and restoration of soft-bottom intertidal communities.

Knowledge of dispersal vectors used by organisms is essential to the understanding of population and community dynamics. We report on ice rafting, a vector by which intertidal benthic invertebrates can be transported well outside their normal dispersal range during winter in temperate climates. We found multiple invertebrate taxa in sediment-laden ice blocks sampled in the intertidal zone. A large proportion of individuals were alive and active when freed from the ice. Using radio tracking, we found that ice blocks can travel over 20 km within a few days. Given the abundance of highly mobile ice blocks carrying viable invertebrates, we conclude that ice-rafting is likely an important dispersal vector, contributing to spatial community dynamics in intertidal systems. This mechanism helps explain observed genetic structure of populations, but it also raises concerns about potential negative impacts of climate change on connectivity between populations.

Biogeochemical functioning of ecosystems is central to nutrient cycling, carbon balance, and several ecosystem services, yet it is not always clear why levels of function might vary among systems. Wetlands are widely recognized for their ability to alter concentrations of solutes and particles as water moves through them, but we have only general expectations for what attributes of wetlands are linked to variability in these processes. We examined changes in several water quality variables (dissolved oxygen, dissolved organic carbon, nutrients, and suspended particles) to ascertain which constituents are influenced during tidal exchange with a range of 17 tidal freshwater wetlands along the Hudson River, New York, USA. Many of the constituents showed significant differences among wetlands or between flooding and ebbing tidal concentrations, indicating wetland-mediated effects. For dissolved oxygen, the presence of even small proportional cover by submerged aquatic vegetation increased the concentration of dissolved oxygen in water returned to the main channel following a daytime tidal exchange. Nitrate concentrations showed consistent declines during ebbing tides, but the magnitude of decline varied greatly among sites. The proportional cover by graminoid-dominated high intertidal vegetation accounted for over 40% of the variation in nitrate decline. Knowing which water-quality alterations are associated with which attributes helps suggest underlying mechanisms and identifies what functions might be susceptible to change as sea level rise or salinity intrusion drives shifts in wetland vegetation cover.

Accurately evaluating the strengths of direct (i.e., consumptive and non-consumptive) effects and indirect (density- and trait-mediated) interactions is crucial for understanding the mechanisms of the maintenance and dynamics of an ecosystem. However, an in situ evaluation has not been conducted for a long enough period of time to fully consider the seasonality and life histories of the community components. We conducted a 9-month (from summer to spring) field experiment in an intertidal rocky shore ecosystem involving the carnivorous snail, Thais clavigera, its prey, the limpet Siphonaria sirius, and their resources, the cyanobacterium (blue-green alga) Lithoderma sp. and the green algae Ulva spp. From summer to autumn, the predation pressure was high, and the consumptive and non-consumptive effects of the predator had opposite (positive and negative, respectively) effects on the prey. Both the density- and trait-mediated indirect interactions decreased the coverage of Lithoderma and increased the coverage of Ulva. As the predation pressure decreased in autumn, the predator affected both the adults and the new recruits of the prey. The trait-mediated interactions still existed, but the density-mediated interactions were not detected. From winter to spring, no direct effects or indirect interactions were detected because of the low predation pressure. Our investigation highlights previously unnoticed processes-showing that the strengths of the direct effects and indirect interactions fluctuate greatly with the seasonality of the ecosystem components.

Ecological theory predicts that positive interactions among organisms will increase across gradients of increasing abiotic stress or consumer pressure. This theory has been supported by empirical studies examining the magnitude of ecosystem engineering across environmental gradients and between habitat settings at local scale. Predictions that habitat setting, by modifying both biotic and abiotic factors, will determine large-scale gradients in ecosystem engineering have not been tested, however. A combination of manipulative experiments and field surveys assessed whether along the east Australian coastline: (1) facilitation of invertebrates by the oyster Saccostrea glomerata increased across a latitudinal gradient in temperature; and (2) the magnitude of this effect varied between intertidal rocky shores and mangrove forests. It was expected that on rocky shores, where oysters are the primary ecosystem engineer, they would play a greater role in ameliorating latitudinal gradients in temperature than in mangroves, where they are a secondary ecosystem engineer living under the mangrove canopy. On rocky shores, the enhancement of invertebrate abundance in oysters as compared to bare microhabitat decreased with latitude, as the maximum temperatures experienced by intertidal organisms diminished. By contrast, in mangrove forests, where the mangrove canopy resulted in maximum temperatures that were cooler and of greater humidity than on rocky shores, we found no evidence of latitudinal gradients of oyster effects on invertebrate abundance. Contrary to predictions, the magnitude by which oysters enhanced biodiversity was in many instances similar between mangroves and rocky shores. Whether habitat-context modifies patterns of spatial variation in the effects of ecosystem engineers on community structure will depend, in part, on the extent to which the environmental amelioration provided by an ecosystem engineer replicates that of other co-occurring ecosystem engineers.

Complexity is well accepted as one of the primary drivers of biodiversity, however, empirical support for such positive associations is often confounded with surface area and undermined by other potential explanatory factors, especially the type of structural component (e.g., pits, crevices, overhangs, etc.). In the present study, sample units (artificial substrates) of equal surface area (± 0.2%) were used to simultaneously examine the independent effects of complexity and different structural component types on species richness (S), abundance (N), and community composition. We created simple and complex concrete substrates of four different geometric designs using novel software. The substrates (n = 8) were mounted onto granite seawalls (at two tidal heights) on two islands south of Singapore Island. After 13 months of colonization, all 384 tiles were collected and their assemblages compared. A total of 53 744 individuals of 70 species/morphospecies were collected and identified. Our results show that greater complexity can support greater species richness and different communities that are independent of surface area. Furthermore, the type of structure (e.g., "pits," "grooves," "towers") can have an effect on richness and community composition that is independent of complexity.

Mussel beds and rockweed stands (fucoid algae) have been shown to be ilternative states on rocky intertidal shores in New England, and here the hypothesis that variation in recruitment provides opportunity for the development of alternative community states was tested. Disturbance by ice scour opens patches for development of alternative states, and in winter 1996-1997, 60 experimental clearings of differing sizes were established on Swan's Island, Maine, USA. Half of the plots were re-cleared during the winter of 2010-2011. Recruitment data for barnacles, mussels, and fucoid algae collected from 1997 to 2012 were used to (1) test for persistence of scale-dependent thresholds, (2) estimate the magnitudes and sources of variation, (3) fit a surface of alternative states as defined by the cusp catastrophe, and (4) test if 1997 recruitment would predict 2010-2011 recruitment in re-scraped plots (i.e., a test of divergence, which is expected in systems with alternative states). For barnacles and mussels, recruitment varied enormously year to year and among sites, but showed consistent patterns over the long-term with respect to clearing size. Average recruitment prior to re-clearing was a good predictor of recruitment afterwards. In contrast, over 50% of the variance in fucoid recruitment was unexplained with weak effects among years and locations. Past fucoid recruitment was a poor predictor of subsequent recruitment. The cusp analysis indicated that fucoid recruitment defines the alternative states. Fucoid recruitment was largely unpredictable and suggests long-term, small-scale priority effects drive the development of alternative states. These observations strongly reinforce the notion that long-term and well- replicated experiments are necessary to develop robust tests of ecological theory.

Trait-mediated indirect interactions (TMII) play an important role in structuring natural communities, and numerous studies have experimentally demonstrated their presence in a variety of systems. However, these studies have largely examined the presence or absence of traits that are responsible for these interactions, without considering natural variation between individuals in the extent to which these traits are manifested. We used a well-documented TMII to investigate the importance of individual behavior type for determining the strength of the TMII. The toadfish Opsanus tau has an indirect positive influence on bivalve survival because the mud crab Panopeus herbstii, a consumer of bivalves, reduces foraging effort in the presence of toadfish. We quantified variation in the strength of persistent individual mud crab responses to toadfish and resulting variation in the strength of TMII. We demonstrate that the strength of this TMII is strongly influenced by mud crab size and behavior type, strengthening with the intensity of response of individual mud crabs to toadfish predator cues. Further, we demonstrate that the spatial distribution within intertidal oyster reefs of crabs with different behavior types is not random; mud crabs inhabiting subtidal areas, where predator cues are more persistent, are significantly less responsive to toadfish cues than mud crabs from intertidal areas. This spatial behavioral structure should lead to spatial variation in the strength of TMII. Given the widespread importance of TMII and the broad occurrence of individual personality or behavior types across numerous taxa, these results should be generally applicable. The distribution of behavior types within a population may therefore be a useful metric for improving our ability to predict the strength of TMII.

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.

Ongoing statistical sophistication allows a shift from describing species' spatial distributions toward statistically disentangling the possible roles of environmental variables in shaping species distributions. Based on a landscape-scale benthic survey in the Dutch Wadden Sea, we show the merits of spatially explicit generalized estimating equations (GEE). The intertidal macrozoobenthic species, Macoma balthica, Cerastoderma edule, Marenzelleria viridis, Scoloplos armiger, Corophium volutator, and Urothoe poseidonis served as test cases, with median grain-size and inundation time as typical environmental explanatory variables. GEEs outperformed spatially naive generalized linear models (GLMs), and removed much residual spatial structure, indicating the importance of median grain-size and inundation time in shaping landscape-scale species distributions in the intertidal. GEE regression coefficients were smaller than those attained with GLM, and GEE standard errors were larger. The best fitting GEE for each species was used to predict species' density in relation to median grain-size and inundation time. Although no drastic changes were noted compared to previous work that described habitat suitability for benthic fauna in the Wadden Sea, our predictions provided more detailed and unbiased estimates of the determinants of species-environment relationships. We conclude that spatial GEEs offer the necessary methodological advances to further steps toward linking pattern to process.

The spatial structure of seagrass landscapes is typically ascribed to the direct influence of physical factors such as hydrodynamics, light, and sediment transport. We studied regularly interspaced banded patterns, formed by elongated patches of seagrass, in a small-scale intertidal ecosystem. We investigated (1) whether the observed spatial patterns may arise from feedback interactions between seagrass and its abiotic environment and (2) whether changes in abiotic conditions may lead to predictable changes in these spatial patterns. Field measurements, experiments, and a spatially explicit computer model identified a ‘‘scale-dependent feedback’’ (a mechanism for spatial self-organization) as a possible cause for the banded patterns. Increased protection from uprooting by improved anchoring with increasing seagrass density caused a local positive feedback. Sediment erosion around seagrass shoots increased with distance through the seagrass bands, hence causing a long-range negative feedback. Measurements across the depth gradient of the intertidal, together with model simulations, demonstrated that seagrass cover and mean patch size were predictably influenced by additional external stress caused by light limitation and desiccation. Thus, our study provides direct empirical evidence for a consistent response of spatial self-organized patterns to changing abiotic conditions, suggesting a potential use for self-organized spatial patterns as stress indicators in ecosystems.

Green macroalgal blooms have substantially altered marine community structure and function, specifically by smothering seagrasses and other primary producers that are critical to commercial fisheries and by creating anoxic conditions in enclosed embayments. Bottom-up factors are viewed as the primary drivers of these blooms, but increasing attention has been paid to biotic controls of species composition. In Washington State, USA, blooms are often dominated by Ulva spp. intertidally and Ulvaria obscura subtidally. Factors that could cause this spatial difference were examined, including competition, grazer preferences, salinity, photoacclimation, nutrient requirements, and responses to nutrient enrichment. Ulva specimens grew faster than Ulvaria in intertidal chambers but not significantly faster in subtidal chambers. Ulva was better able to acclimate to a high-light environment and was more tolerant of low salinity than Ulvaria. Ulvaria had higher tissue N content, chlorophyll, chlorophyll b: chlorophyll a, and protein content than Ulva. These differences suggest that nitrogen availability could affect species composition. A suite of five grazers preferred Ulva to Ulvaria in choice experiments. Thus, bottom-up factors allow Ulva to dominate the intertidal zone while resistance to grazers appears to allow Ulvaria to dominate the subtidal zone. While ulvoid algae are in the same functional-form group, they are not functionally redundant.

We show that a higher vertebrate can graze surficial intertidal biofilm, previously only considered a food source for rasping invertebrates and a few specialized fish. Using evidence from video recordings, stomach contents, and stable isotopes, we describe for the first time the grazing behavior of Western Sandpipers (Calidris mauri) and estimate that biofilm accounts for 45-59% of their total diet or 50% of their daily energy budget. Our finding of shorebirds as herbivores extends the trophic range of shorebirds to primary consumers and potential competitors with grazing invertebrates. Also, given individual grazing rates estimated at seven times body mass per day and flock sizes into the tens of thousands, biofilm-feeding shorebirds could have major impacts on sediment dynamics. We stress the importance of the physical and biological processes maintaining biofilm to shorebird and intertidal conservation.

Reproductive output is a central attribute of life history, providing a measure of individual and population performance. The fields of ecology, biogeography, and evolutionary biology take disparate approaches in addressing spatial variation in reproduction, and thus we lack clear predictions for how reproductive output should vary geographically. We empirically investigate these contrasting theoretical approaches by determining geographic patterns in reproductive output for intertidal populations of the purple sea urchin, Strongylocentrotus purpuratus, at 15 sites spanning a large geographic distance (9 degrees span of latitude) from central California, USA, to Baja California, Mexico. Contrary to predictions from biogeography, some of the highest values of reproductive output are at sites near the species' southern range boundary. Additionally, sea urchins do not exhibit a latitudinal gradient in reproduction, but rather show considerable mesoscale variation in reproductive output. Spatial analyses reveal that this variation is correlated with coastal topographic features that are known to influence the pattern of nearshore nutrient flux and circulation. We hypothesize that urchins' reproductive output may be driven by the spatial distribution of their food supply, drift macroalgae, the abundance of which is influenced by both nutrient supply and alongshore transport processes that are coupled to topographic features. Large-scale studies such as this provide valuable insight into the causes of species' range limits, population connectivity, habitat reserve design, and forecasting the effects of climate change on species' distributions.

Populations of intertidal species span a steep environmental gradient driven by differences in emersion time. In spite of strong differential selection on traits related to this gradient, the small spatial scale over which differences occur may prevent local adaptation, and instead may favor a single intermediate phenotype, or nongenetic mechanisms of differentiation. Here I examine whether a common macroalga, Silvetia compressa, exhibits phenotypic differentiation across the intertidal gradient and evaluate how local adaptation, developmental plasticity, and maternal effects may interact to shape individual phenotypes. Reciprocal transplants of both adults and embryos showed a "home-height advantage" in two of the three populations tested. In laboratory trials, the progeny of upper-limit individuals survived exposure to air significantly better than lower-limit progeny from the same population. I compared the emersion tolerance of full-sib families generated from gametes produced in the field to those produced under common garden conditions. The relative advantage of upper-limit lineages was robust to maternal environment during gametogenesis; this pattern is consistent with genetic differentiation. The possible role of local adaptation has historically been ignored in studies of intertidal zonation. In S. compressa, phenotypic differentiation may have important consequences for vertical range, both within and among sites.

We investigated the nonconsumptive effects (NCEs) of predatory dogwhelks (Nucella lapillus) on intertidal barnacle (Semibalanus balanoides) recruitment through field experiments on the Gulf of St. Lawrence coast and the Atlantic coast of Nova Scotia, Canada. We studied the recruitment seasons (May-June) of 2011 and 2013. In 2011, the Gulf coast had five times more nearshore phytoplankton (food for barnacle larvae and recruits) during the recruitment season and yielded a 58% higher barnacle recruit density than the Atlantic coast at the end of the recruitment season. In 2013, phytoplankton levels and barnacle recruit density were similar on both coasts and also lower than for the Gulf coast in 2011. Using the comparative-experimental method, the manipulation of dogwhelk presence (without allowing physical contact with prey) revealed that dogwhelk cues limited barnacle recruitment under moderate recruit densities (Atlantic 2011/2013 and Gulf 2013) but had no effect under a high recruit density (Gulf 2011). Barnacle recruits attract settling larvae through chemical cues. Thus, the highest recruit density appears to have neutralized dogwhelk effects. This study suggests that the predation risk perceived by settling larvae may decrease with increasing recruit density and that prey food supply may indirectly influence predator NCEs on prey recruitment.

Ecological studies of plant distributions along environmental gradients, such as plant zonation in salt marshes, have primarily focused on abiotic stress and plant interactions (competition and facilitation). A decades-old paradigm is that the stressful and benign boundaries of salt marsh plants are determined by abiotic stress and competition, respectively. Although consumers have long been recognized as mediating algal and sessile animal zonation in the rocky intertidal, their role in generating plant zonation in salt marshes remains largely unexplored. We examined the zonation of two annual succulents, Salicornia europaea and Suaeda salsa, along an elevation gradient in a northern Chinese salt marsh, with and without manipulating the common herbivorous crab Helice tientsinensis. Salicornia occupies waterlogged, low-salinity habitats, whereas Suaeda dominates non-waterlogged, hypersaline habitats at higher elevations. We first conducted a pot experiment crossing salinity, waterlogging, and competition, followed by a field experiment with removal of competitors, and found that neither waterlogging nor salinity stress explained the absence of either species from the other's zone, while Suaeda competitively excluded Salicornia from the upper non-waterlogged zone. We then conducted field and lab herbivory experiments, which showed that Helice preferentially grazed Suaeda at waterlogged low elevations and that Helice grazing on Suaeda increased with waterlogging. These results reveal that while competition plays a role in the zonation by excluding Salicornia from the upper Suaeda zone, crab grazing limits the success of Suaeda in the lower Salicornia zone. These findings challenge the idea that plant interactions and abiotic stress are sufficient to explain marsh zonation in all cases, and highlight an overlooked role of consumers, a role potentially general across diverse intertidal ecosystems. Future models of plant distributions should consider how consumer pressure interacts with plant interactions and abiotic stress across environmental gradients.

Extreme heat events cause patchy mortality in many habitats. We examine biophysical mechanisms responsible for patchy mortality in beds of the competitively dominant ecosystem engineer, the marine mussel Mytilus californianus, on the west coast of the United States. We used a biophysical model to predict daily fluctuations in body temperature at sites from southern California to Washington and used results of laboratory experiments on thermal tolerance to determine mortality rates from body temperature. In our model, we varied the rate of thermal conduction within mussel beds and found that this factor can account for large differences in body temperature and consequent mortality during heat waves. Mussel beds provide structural habitat for other species and increase local biodiversity, but, as sessile organisms, they are particularly vulnerable to extreme weather conditions. Identifying critical biophysical mechanisms related to mortality and ecological performance will improve our ability to predict the effects of climate change on these vulnerable ecosystems.

Canopy-forming macroalgae are key species on temperate rocky shores. However, there is a lack of understanding of how the relative balance of physical and biological factors controls the establishment and persistence of intertidal macroalgae. Here we present an integrated study of the relative importance of wave-induced forces and grazing for the recruitment and survival of the canopy-forming intertidal macroalgae Fucus vesiculosus and F. spiralis. A set of overtopped breakwaters provided a nearly unconfounded gradient in wave exposure between seaward and landward sides. A biomechanical analysis was performed based on empirical measurements of maximum drag forces in breaking waves, a model of long-term maximum wave height, and the breaking stress of Fucus spp. The estimated maximum flow speed (7-8 m/s) on the seaward side of the breakwaters was predicted to completely dislodge or prune Fucus spp. larger than approximately 10 cm, while dislodgment was highly unlikely on the landward side for all sizes. Experimental transplantation of Fucus spp. supported the biomechanical analysis but also suggested that mechanical abrasion may further limit survival in wave-exposed locations. Experimental removal of the limpet Patella vulgata, which was the principal grazer at this site, resulted in recruitment of Fucus spp. on the seaward side. We present a model of limpet grazing that indicates that limpet densities5-20 individuals/m2 provide a proximate mechanism preventing establishment of Fucus spp., whereas wave action2 m/s reduces persistence through dislodgment and battering. In a conceptual model we further propose that recruitment and survival of juvenile Fucus spp. are controlled indirectly by wave exposure through higher limpet densities at exposed locations. This model predicts that climate change, and in particular an increased frequency of storm events in the northeast Atlantic, will restrict fucoids to more sheltered locations.

Besides the “normal” challenge of obtaining adequate intake rates in a patchy and dangerous world, shorebirds foraging in intertidal habitats face additional environmental hurdles. The tide forces them to commute between a roosting site and feeding grounds, twice a day. Moreover, because intertidal food patches are not all available at the same time, shorebirds should follow itineraries along the best patches available at a given time. Finally, shorebirds need additional energy stores in order to survive unpredictable periods of bad weather, during which food patches are covered by extreme tides. In order to model such tide-specific decisions, we applied stochastic dynamic programming in a spatially explicit context. Two assumptions were varied, leading to four models. First, birds had either perfect (ideal) or no (non-ideal) information about the intake rate at each site. Second, traveling between sites was either for free or incurred time and energy costs (non-free). Predictions were generated for three aspects of foraging: area use, foraging routines, and energy stores. In general, non-ideal foragers should feed most intensely and should maintain low energy stores. If traveling for such birds is free, they should feed at a random site; otherwise, they should feed close to their roost. Ideal foragers should concentrate their feeding around low tide (especially when free) and should maintain larger energy stores (especially when non-free). If traveling for such birds is free, they should feed at the site offering the highest intake rate; otherwise, they should trade off travel costs and intake rate. Models were parameterized for Red Knots (Calidris canutus) living in the Dutch Wadden Sea in late summer, an area for which detailed, spatially explicit data on prey densities and tidal heights are available. Observations of radio-marked knots (area use) and unmarked knots (foraging routines, energy stores) showed the closest match with the ideal/non-free model. We conclude that knots make state-dependent decisions by trading off starvation against foraging-associated risks, including predation. Presumably, knots share public information about resource quality that enables them to behave in a more or less ideal manner. We suggest that our modeling approach may be applicable in other systems where resources fluctuate in space and time. [KEYWORDS: Calidris canutus ; daily routine ; foraging behavior; patch use ; Red Knot ; state-dependent decisions ; stochastic dynamic programming ; tidal regime ; Wadden Sea.]

Marine mesograzers use individual seaweeds as both habitat and food. Previous studies by M. E. Hay, J. E. Duffy, and others have shown that mesograzers can acquire enemy-free space by inhabiting and consuming seaweeds that are chemically defended against larger, more mobile consumers. However, their studies, carried out in North Carolina and two tropical regions, consider only a small number of mesograzer species and focus primarily on fishes, even though some other large consumers can destroy mesograzers' host seaweeds. We asked whether the previous findings could be generalized to (1) mesograzers at the community level, (2) large consumers other than fishes, and (3) a different ecosystem in another region, namely, diverse temperate rocky reef communities in Australia and New Zealand. Results from both Australasian regions were similar. In multiple-choice feeding assays larger grazers (three fishes, four gastropods, and three sea urchins) varied in breadth of preferences, but most often preferred to eat green algae of the family Ulvales and the red alga Pterocladiella capillacea, all of which were restricted to the intertidal zone. Most of the mesograzers surveyed (seven amphipods, three isopods, a gastropod, and a sea urchin) preferred to eat the seaweed species from which they were collected, many of which were either inaccessible or less palatable to many larger grazers. This difference in mesograzer vs. larger grazer feeding preferences was consistent with the findings of Hay and Duffy et al. However, community-level patterns of mesograzer distribution and abundance were not consistent with the extensions of the theory developed here. Seaweeds that were less palatable to the larger grazers did not tend to support more mesograzer individuals, mesograzer species, or specialist mesograzers (for either the entire mesograzer assemblage or just for sedentary mesograzers), but the evenness of entire mesograzer assemblages was significantly higher on seaweeds that were less palatable to larger grazers. We argue that spatial refuges on Australasian rocky reefs and differences in the biology of both the seaweed and animal species in Australasia vs. previously studied locations enable palatable seaweeds to persist in the face of herbivory by larger grazers, which reduces the selective pressure on mesograzers to inhabit unpalatable species.

The paradigm that net seaweed photosynthesis in the stressful intertidal zone decreases predictably with increasing tidal elevation has not been tested widely. We measured the variation in net photosynthesis rates of a common intertidal brown seaweed (Fucus gardneri) collected from 14 sites and two tidal heights on San Juan Island, Washington (USA) in water and air, in summer and winter. We also measured temperature, relative humidity, and thallus hydration in situ. Mean net photosynthesis in air was two orders of magnitude lower than in water, a greater mean difference than found among sites and between tidal elevations and seasons. Thalli never fully desiccated even in summer, and net photosynthesis was always positive. Rates in air and water varied little and nonsignificantly between tidal elevations and were not consistently lower at upper elevations, where in situ conditions were not always drier and hotter. Although photosynthetic rates per se did not vary predictably across the tidal gradient, mid-elevation thalli gained 25 times more carbon because they were exposed to air for shorter periods. Thalli experienced stressful conditions that reduce net photosynthesis during exposure, but the length of time exposed to air, rather than desiccation, determined daily photosynthetic carbon acquisition.

The onshore deposition of macroalgal and macrophyte wrack provides a potentially significant marine subsidy to intertidal and supratidal herbivore and decomposer communities. Based on the study of daily input loads to beaches, we estimated summer wrack deposition of up to 140 Mg (dry mass)/km shoreline in Barkley Sound, British Columbia. However, input rates were highly variable depending on beach type, nearshore hydrodynamics, and buoyancy characteristics of the wrack. Cobble beaches retained ;10 times and 30 times more wrack than did gravel and sand beaches, respectively. Cobble and gravel beaches also differed in species composition of new (fresh) wrack input, with Ma- crocystis integrifolia being characteristic for the former and Nereocystis luetkeana for the latter, which we attribute to buoyancy characteristics of the floating debris. On sand beaches, Phyllospadix spp. and Enteromorpha spp. were the dominant wrack species. Species com- position of freshly deposited wrack also depended on wave exposure, but predictability based on the species pool within a beach's catchment was restricted. Drift lines of aging wrack differed from freshly deposited wrack in species composition, probably due to wrack decomposition that results in fluxes of nutrients and energy between the adjacent marine and terrestrial habitats. We hold that the characteristics of a given beach, e.g., substratum and wave exposure, and their effects on wrack input, will have important ecological and biogeochemical implications for the marine-terrestrial ecotone.

Biological rhythms with lunar components are common in nature. In the sea, the moon's gravitational pull on earth is the principal cause of the tides, which normally reach maximum amplitudes every new and full moon. Many populations synchronize spawning to this time. Some choose either the new or the full moon, implying that moonlight is important; but one lunar phase usually has higher tides than the other, and many species select the phase with the higher tide to improve the offshore transport of their progeny. However, tidal dominance by one lunar phase is not constant; it switches between new and full moon every seven months. We tested the influence of this 14-month “syzygy inequality cycle” (SIC) on lunar synchrony by sampling 11 populations of intertidal crabs at two locations in East Africa for 21 months. Eight populations synchronized larval release with the SIC. Tidal cues were more important than moonlight in entraining the reproductive rhythm, although two populations synchronized spawning to t...

Biologically mediated modifications of the abiotic environment, also called ecosystem engineering, can significantly affect a broad range of ecosystems. Nevertheless, remarkably little work has focused on the costs and benefits that ecosystem engineers obtain from traits that underlie their ecosystem engineering capacity. We addressed this topic by comparing two autogenic engineers, which vary in the degree in which they affect their abiotic environment via their physical structure. That is, we compared two plant species from the intertidal coastal zone (Spartina anglica and Zostera noltii), whose shoots are exposed to similar currents and waves, but differ in the extent that they modify their environment via reduction of hydrodynamic energy. Our results indicate that there can be trade-offs related to the traits that underlies autogenic ecosystem engineering capacity. Dissipation of hydrodynamic forces from waves was roughly a factor of three higher in vegetation with stiff leaves compared to those with flexible leaves. Drag was highest and most sensitive to hydrodynamic forces in stiff vegetation that does not bend with the flow. Thus, shoot stiffness determines both the capacity to reduce hydrodynamic energy (i.e., proxy for ecosystem engineering capacity) and the drag that needs to be resisted (i.e., proxy for associated costs). Our study underlines the importance of insight in the trade-offs involved in ecosystem engineering as a first step toward understanding the adaptive nature of ecosystem engineering. [KEYWORDS: autogenic ecosystem engineers ; current velocity ; drag ; extended phenotype ; niche construction ; sediment accretion ; Spartina anglica ; trade-offs ; waves; Zostera noltii.]

Many ecological phenomena are characterized by context dependency, and the relationship between diversity and productivity is no exception. We examined the re- lationship between macroalgal diversity and nutrient availability by evaluating the effects of reduced nutrients and their subsequent replacement via local-scale nutrient loading in tide pools. Macroalgae in Oregon coast high-intertidal pools have evolved in a nitrate-rich upwelling ecosystem, but instead of settling on low-intertidal reefs (where algae are often immersed in nutrient-rich nearshore waters) these individuals have colonized high-zone pools, where they are isolated from the ocean for extended periods of time and are subjected to extended periods of nitrate depletion. In some pools, this nutrient stress was ameliorated by a positive interaction: the excretion of ammonium by invertebrates. We conducted ex- perimental manipulations to quantify invertebrate-mediated ammonium loading and ma- croalgal ammonium uptake in high-intertidal pools. Variation in tide pool volume and invertebrate biomass created a gradient of local-scale nutrient inputs, allowing us to address the relationship between nitrogen loading and algal diversity. Slow-growing species tolerant of low nitrogen availability were joined by fast-growing species with higher nitrogen re- quirements in pools with higher ammonium loading rates. A fourfold increase in the am- monium loading rate was associated with a doubling in the number of macroalgal species, and macroalgal assemblages in more species-rich pools were characterized by higher rates of biomass-specific ammonium uptake. These patterns contrast with productivity-diversity relationships in terrestrial systems, where local-scale nutrient enrichments generally result in reduced producer diversity due to displacement of subordinate species by aggressive competitors. Our data suggest that the effect of enrichment on diversity is context-depen- dent. Each ecosystem has a critical level of nutrient availability, determined by the level of nutrients typically available in that system. Below this critical level, local-scale nutrient additions increase diversity, but above it, diversity declines with enrichment.

Canopy-forming plants and algae, in addition to their role as food, can positively affect higher trophic levels by providing protection from predation or stressful abiotic conditions. In the low rocky intertidal zone of the Pacific Northwest, the chiton Katharina tunicata is closely associated with the canopy-forming alga Hedophyllum sessile. In a 27-month field experiment, I used experimental devices to mimic characteristics of the Hedophyllum canopy and monitored the abundance and physiology of Katharina to determine the mechanism behind this association. The data show that this association is due to selection of shaded microhabitats by Katharina, and that neither predation nor Hedophyllum per se influenced the distribution of the chiton. Exclusion of visual predators had no effect on chiton abundance, and tests of the effects of shade independent of Hedophyllum presence indicated that the shade produced by the alga was critical to the chiton. Seasonal fluctuations in levels of 70 kDa (1.16 × 10−19 g) heat shock proteins (Hsp70) indicate that Katharina experience sublethal physiological stress during the spring and summer. The corresponding use of shaded areas during spring and summer low tides suggests that the chiton's distribution is limited during these seasons by the availability of refugia from sublethal abiotic stresses. Subtle variations in environmental conditions are also important to this association; the strength of the positive interaction varied annually, becoming more positive in warm summers (1998 and 2000) and neutral in a cooler summer (1999). On a finer scale, physiological measurements indicate that the response of chitons to summertime temperatures depends on both absolute temperature change and the animal's recent thermal history. In particular, the seasonal onset of daytime low tides in the spring is associated with increased Hsp70 levels, indicating that this is a significant physiological event for Katharina. The importance of facultative positive interactions that allow consumers to escape from sublethal abiotic stresses will likely be variable in time but can have large impacts on communities both by affecting the small-scale distribution of consumers over the short term and potentially by affecting the persistence of organisms over the long term.

One of the most commonly predicted effects of global ocean warming on marine communities is a poleward shift in the distributional boundaries of species with an associated replacement of cold-water species by warm-water species. However, these types of predictions are imprecise and based largely on broad correlations in uncontrolled studies that examine changes in the distribution or abundances of species in relation to seawater temperature. Our study used an 18-year sampling program in intertidal and subtidal habitats and before-after, control-impact analyses. We show that a 3.58C rise in seawater temper- ature, induced by the thermal outfall of a power-generating station, over 10 years along 2 km of rocky coastline in California resulted in significant community-wide changes in 150 species of algae and invertebrates relative to adjacent control areas experiencing natural temperatures. Contrary to predictions based on current biogeographic models, there was no trend toward warmer-water species with southern geographic affinities replacing colder- water species with northern affinities. Instead, the communities were greatly altered in apparently cascading responses to changes in abundance of several key taxa, particularly habitat-forming subtidal kelps and intertidal foliose red algae. Many temperature-sensitive algae decreased greatly in abundance, whereas many invertebrate grazers increased. The responses of these benthic communities to ocean warming were mostly unpredicted and strongly coupled to direct effects of temperature on key taxa and indirect effects operating through ecological interactions.

Many organisms have evolved morphological and behavioral traits that reduce their susceptibility to predation. However, few studies have explicitly investigated the relationships between defensive traits and susceptibility. Here we demonstrate a negative correlation between morphological defenses and behavioral avoidance across several species of marine gastropod that is linked to vulnerability to crab predation. Snails that had relatively taller shell spires (high aspect ratio) showed greater responsiveness when exposed to predation cues than did species with disc-like shells (low aspect ratio). Our results suggest that the snail species most vulnerable to predation compensated by showing the highest levels of behavioral avoidance, and hence may be at a disadvantage in competition with less vulnerable species. This has important implications because the behavioral response of herbivorous gastropods to predation cues may play a central role in structuring rocky intertidal communities through trait-mediated indirect effects.

Epilithic biofilms play a key role in marine ecosystems. They also provide a tractable system to investigate the relative roles of environmental stressors, bottom-up physicochemical factors and top-down biological control in regulating communities. Patterns of photosynthetic microbial biomass were recorded over a four-year period at several tidal levels on shores in the Isle of Man. Photosynthetic biomass and the abundance of diatoms were consistently greater during winter than summer. Biomass was negatively correlated with insolation stress and air temperature, but was not correlated with grazing intensity, dissolved nutrients, sea temperature, or planktonic chlorophyll. Field experiments confirmed that reducing insolation stress led to substantial increases in photosynthetic biomass, predominantly of diatoms and macroalgal germlings. Reducing grazing intensity also led to considerable increases in photosynthetic biomass, but reducing desiccation stress or increasing nutrient availability had no effect. Although grazing can regulate microalgal biomass, seasonal patterns of grazing activity were driven by temperature and were decoupled from photosynthetic biomass. Our study demonstrates the importance of physiological stresses for the direct and indirect regulation of the balance between primary producers and consumers. Based on these findings, we present a model that combines the roles of stressors together with bottom-up forcing and top-down regulation in controlling communities on wave-exposed shores.

Mussel beds and algal canopies on Gulf of Maine (USA) rocky shores have been hypothesized to represent stochastically determined community states, but we recently found that they were highly deterministic, consumer-controlled states in a tidal river. Mussel beds dominated high-flow habitats with high larval delivery and low consumer pressure, whereas algal canopies dominated low-flow habitats with high consumer pressure. In this paper we examine the alternative-community-state hypothesis on rocky shores on the open coast of Maine. We hypothesized that community states would be less deterministic due to reduced consumer pressure and more stochastic flow patterns. At 10 mussel-bed and 10 seaweed-canopy sites on the open coast of central Maine, we created 9-m2 and 1-m2 clearings and an unmanipulated control area, each containing control (no caging), caged, and cage-control quadrats. Independent of disturbance patch size, mussel beds and seaweed canopies rapidly began to return to their original community type in the absence of consumers (crabs and snails). With consumers present, recovery of the original communities did occur, but at a much slower pace. Our results suggest that mussel beds and algal canopies in the Gulf of Maine are deterministic consumer-controlled states, rather than being stochastically determined. If stochastic alternative community states exist in this system, they occur in a very restricted subset of habitats where the strong consumer control and determinism we have found in both tidal river and open coast habitats are relaxed.

The influence of predation on rocky intertidal community structure has long emphasized the importance of indirect interactions. Most efforts in this area have focused on the density-mediated, or lethal effects, of predators on prey density. Recently, there has been growing interest in trait-mediated indirect interactions (TMIIs): the presence of a predator in the environment influences the interaction between two other species (prey and their resource) by altering a trait of the prey species. For example, waterborne cues released by predators can cause changes in prey species behavior, such as feeding rates, thereby altering the impact of the prey species on their resources. Thus, TMIIs represent the nonlethal effects of predators that contrast with the more traditional emphasis on lethal indirect effects. Marine ecologists are just beginning to explore the role of TMIIs in their systems. We examined whether risk cues released by a ubiquitous crab predator (Carcinus maenas) influence the abundance of two dominant species in the rocky intertidal zone (barnacles [Semibalanus balanoides] and fucoid algae [Ascophyllum nodosum]) by altering the behavior of two of its snail prey (Nucella lapillus and Littorina littorea). We found that the presence of green crab risk cues can have strong cascading indirect effects on the abundance of barnacles and fucoid algae. N. lapillus exposed to risk cues consumed up to 29% fewer barnacles compared to conspecifics feeding in the absence of risk cues, whereas L. littorea exposed to risk cues consumed 459% fewer fucoids compared to conspecifics feeding in the absence of risk cues. These cascading interactions appear to reflect suppression of snail feeding by predator risk cues. In both food chains, snails exhibited more refuge-seeking behavior and grew less in the presence of risk cues. Our experiments suggest that TMIIs may have an important and underappreciated influence on species interactions that shape community dynamics on rocky intertidal shores. Corresponding Editor: P. T. Raimondi

How and why populations fluctuate and what drives the magnitude of fluctuations are questions that have long intrigued ecologists. Dispersal may dampen population fluctuations through the effect of spreading offspring over heterogeneous habitats. The planktonic period common in many marine organisms, therefore, could dampen population fluctuations through larval dispersal. However, emphasis on the hazards of planktonic life predicts that species that have a longer planktonic period have greater fluctuations in adult populations than species with shorter or no planktonic period. I analyzed the population variation of 570 time series from the literature for intertidal and benthic subtidal marine species and found that time series of adult populations for species with no planktonic period had greater fluctuations than time series for species with a planktonic period (both short and long planktonic period). In addition, there was no difference in fluctuation of adult and recruit time series between species with long and short planktonic periods. The planktonic period did not appear to result in increased population fluctuations, as was widely assumed. Rather, the planktonic period may be acting to decrease population fluctuations, potentially by dispersal. Corresponding Editor: J. E. Duffy.