The biogeography of community assembly: latitude and predation drive variation in community trait distribution in a guild of epifaunal crustaceans.

While considerable evidence exists of biogeographic patterns in the intensity of species interactions, the influence of these patterns on variation in community structure is less clear. Studying how the distributions of traits in communities vary along global gradients can inform how variation in interactions and other factors contribute to the process of community assembly. Using a model selection approach on measures of trait dispersion in crustaceans associated with eelgrass (Zostera marina) spanning 30° of latitude in two oceans, we found that dispersion strongly increased with increasing predation and decreasing latitude. Ocean and epiphyte load appeared as secondary predictors; Pacific communities were more overdispersed while Atlantic communities were more clustered, and increasing epiphytes were associated with increased clustering. By examining how species interactions and environmental filters influence community structure across biogeographic regions, we demonstrate how both latitudinal variation in species interactions and historical contingency shape these responses. Community trait distributions have implications for ecosystem stability and functioning, and integrating large-scale observations of environmental filters, species interactions and traits can help us predict how communities may respond to environmental change.

The journey of hull-fouling mobile invaders: basibionts and boldness mediate dislodgement risk during transit.

Vessel hull-fouling is responsible for most bioinvasion events in the marine environment, yet it lacks regulation in most countries. Although experts advocate a preventative approach, research efforts on pre-arrival processes are limited. The performance of mobile epifauna during vessel transport was evaluated via laboratory simulations, using the well-known invasive Japanese skeleton shrimp (Caprella mutica), and its native congener C. laeviuscula as case study. The invader did not possess any advantage in terms of inherent resistance to drag. Instead, its performance was conditioned by the complexity of secondary substrate. Dislodgement risk was significantly reduced when sessile fouling basibionts were added, which provided refugia and boosted the probability of C. mutica remaining attached from 7 to 65% in flow exposure trials. Interestingly, the invader exhibited significantly higher exploratory tendency and motility than its native congener at zero-flow conditions. Implications in terms of en-route survivorship, invasion success and macrofouling management are discussed.

Reef communities show predictable undulations in linear abundance size spectra from copepods to sharks.

Among the more widely accepted general hypotheses in ecology is that community relationships between abundance and body size follow a log-linear size spectrum, from the smallest consumers to the largest predators (i.e. 'bacteria to whales'). Nevertheless, most studies only investigate small subsets of this spectrum, and note that extreme size classes in survey data deviate from linear expectations. In this study, we fit size spectra to field data from 45 rocky and coral reef sites along a 28° latitudinal gradient, comprising individuals from 0.125 mm to 2 m in body size. We found that 96% of the variation in abundance along this 'extended' size gradient was described by a single linear function across all sites. However, consistent 'wobbles' were also observed, with subtle peaks and troughs in abundance along the spectrum, which varied with sea temperature, as predicted by theory relating to trophic cascades.

Climate cascades affect coastal Antarctic seafloor ecosystem functioning.

Polar seafloor ecosystems are changing rapidly and dramatically, challenging previously held paradigms of extreme dynamical stability. Warming-related declines in polar sea ice are expected to alter fluxes of phytoplankton and under-ice algae to the seafloor. Yet, how changes in food flux cascade through to seafloor communities and functions remains unclear. We leveraged natural spatial and temporal gradients in summertime sea ice extent to better understand the trajectories and implications of climate-related change in McMurdo Sound, Antarctica. McMurdo Sound was expected to be one of the last coastal marine environments on Earth to be affected by planetary warming, but the situation may be changing. Comparing satellite observations of selected coastal sites in McMurdo Sound between 2010-2017 and 2002-2009 revealed more ice-free days per year, and shorter distances to open water during the warmest months each year, in the more recent period. Interdecadal Pacific Oscillation (IPO), Oceanic Niño Index (ONI) and Antarctic Oscillation (AAO) climate indices peaked concurrently between 2014 and 2017 when sea ice breakouts in McMurdo Sound were most spatially and temporally extensive. Increases in sediment chlorophyll a and phaeophytin content (indicating increased deposition of detrital algal food material) were recorded during 2014-2017 at three coastal study sites in McMurdo Sound following the major sea ice breakouts. Soft-sediment seafloor ecosystem metabolism (measured in benthic incubation chambers as dissolved oxygen and inorganic nutrient fluxes) was correlated with sediment algal pigment concentration. Epifaunal invertebrate density, particularly opportunistic sessile suspension feeders, and infaunal community composition also shifted with increased food supply. The ecological characteristics and functions measured at the food-poor sites shifted towards those observed at richer sites at a surprisingly fast pace. These results indicate the sensitivity of the benthos and shed light on Antarctic marine trophic cascades and trajectories of response of iconic high-latitude seafloor habitats to a warming climate.

Production of mobile invertebrate communities on shallow reefs from temperate to tropical seas.

Primary productivity of marine ecosystems is largely driven by broad gradients in environmental and ecological properties. By contrast, secondary productivity tends to be more variable, influenced by bottom-up (resource-driven) and top-down (predatory) processes, other environmental drivers, and mediation by the physical structure of habitats. Here, we use a continental-scale dataset on small mobile invertebrates (epifauna), common on surfaces in all marine ecosystems, to test influences of potential drivers of temperature-standardized secondary production across a large biogeographic range. We found epifaunal production to be remarkably consistent along a temperate to tropical Australian latitudinal gradient of 28.6°, spanning kelp forests to coral reefs (approx. 3500 km). Using a model selection procedure, epifaunal production was primarily related to biogenic habitat group, which explained up to 45% of total variability. Production was otherwise invariant to predictors capturing primary productivity, the local biomass of fishes (proxy for predation pressure), and environmental, geographical, and human impacts. Highly predictable levels of epifaunal productivity associated with distinct habitat groups across continental scales should allow accurate modelling of the contributions of these ubiquitous invertebrates to coastal food webs, thus improving understanding of likely changes to food web structure with ocean warming and other anthropogenic impacts on marine ecosystems.

Abiotic proxies for predictive mapping of nearshore benthic assemblages: implications for marine spatial planning.

Marine spatial planning (MSP) should assist managers in guiding human activities toward sustainable practices and in minimizing user conflicts in our oceans. A necessary first step is to quantify spatial patterns of marine assemblages in order to understand the ecosystem's structure, function, and services. However, the large spatial scale, high economic value, and density of human activities in nearshore habitats often makes quantifying this component of marine ecosystems especially daunting. To address this challenge, we developed an assessment method that employs abiotic proxies to rapidly characterize marine assemblages in nearshore benthic environments with relatively high resolution. We evaluated this assessment method along 300 km of the State of Maine's coastal shelf (<100 m depth), a zone where high densities of buoyed lobster traps typically preclude extensive surveys by towed sampling gear (i.e., otter trawls). During the summer months of 2010-2013, we implemented a stratified-random survey using a small remotely operated vehicle that allowed us to work around lobster buoys and to quantify all benthic megafauna to species. Stratifying by substrate, depth, and coastal water masses, we found that abiotic variables explained a significant portion of variance (37-59%) in benthic species composition, diversity, biomass, and economic value. Generally, the density, diversity, and biomass of assemblages significantly increased with the substrate complexity (i.e., from sand-mud to ledge). The diversity, biomass, and economic value of assemblages also decreased significantly with increasing depth. Last, demersal fish densities, sessile invertebrate densities, species diversity, and assemblage biomass increased from east to west, while the abundance of mobile invertebrates and economic value decreased, corresponding mainly to the contrasting water mass characteristics of the Maine Coastal Current system (i.e., summertime current direction, speed, and temperature). Integrating modeled predictions with existing GIS layers for abiotic conditions allowed us to scale up important assemblage attributes to define key foundational ecological principles of MSP and to find priority regions where some bottom-disturbing activities would have minimal impact to benthic assemblages. We conclude that abiotic proxies can be strong forcing functions for the assembly of marine communities and therefore useful tools for spatial extrapolations of marine assemblages in congested (heavily used) nearshore habitats.

Extending trait dispersion across trophic levels: Predator assemblages act as top-down filters on prey communities.

Studies of community assembly typically focus on the effects of abiotic environmental filters and stabilizing competition on functional trait dispersion within single trophic levels. Predation is a well-known driver of community diversity and composition, yet the role of functionally diverse predator communities in filtering prey community traits has received less attention. We examined functionally diverse communities of predators (fishes) and prey (epifaunal crustaceans) in eelgrass (Zostera marina) beds in two northern California estuaries to evaluate the filtering effects of predator traits on community assembly and how filters acting on predators influence their ability to mediate prey community assembly. Fish traits related to bottom orientation were correlated with more clustered epifauna communities, and epifauna were generally overdispersed while fishes were clustered, suggesting prey may be pushed to disparate areas of trait space to avoid capture by benthic sit-and-wait predators. We also found correlations between the trait dispersions of predator and prey communities that strengthened after accounting for the effects of habitat filters on predator dispersion, suggesting that habitat filtering effects on predator species pools may hinder their ability to affect prey community assembly. Our results present compelling observational evidence that specific predator traits have measurable impacts on the community assembly of prey, inviting experimental tests of predator trait means on community assembly and explicit comparisons of how the relative effects of habitat filters and intraguild competition on predators impact their ability to affect prey community assembly. Integrating our understanding of traits at multiple trophic levels can help us better predict the impacts of community composition on food web dynamics as regional species pools shift with climate change and anthropogenic introductions.

Facilitation of macrofaunal assemblages in marinas by the habitat-forming invader Amathia verticillata (Bryozoa: Gymnolaemata) across a spatiotemporal scale.

Widespread habitat-forming invaders inhabiting marinas, such as the spaghetti bryozoan Amathia verticillata, allow exploring facilitation processes across spatiotemporal contexts. Here we investigate the role of this bryozoan as habitat for native and exotic macrofaunal assemblages across different ecoregions of Western Mediterranean and East Atlantic coasts, and a monthly variation over a year. While only 7 (all peracarid crustaceans) of the 54 associated species were NIS, they dominated macrofaunal assemblages in terms of abundance, raising the potential for invasional meltdown. NIS richness and community structure differed among marinas but not among ecoregions, highlighting the importance of marina singularities in modulating facilitation at spatial scale. Despite facilitation did not depend on bryozoan abundance fluctuations, it was affected by its deciduous pattern, peaking in summer and disappearing in late winter. Monitoring A. verticillata in marinas, especially in summer periods, may improve the detection and management of multiple associated NIS.

Benthic taxonomic and functional diversity change across a depth gradient in the Southern Benguela Shelf ecoregion.

Functional trait-based approaches improve biodiversity assessment and have consequently been gaining traction in ecology, including marine benthic studies. However, taxonomic diversity is still the default biodiversity metric applied to monitor benthic community responses to environmental variation despite not always representing functional diversity change. Therefore, we used Biological Traits Analysis (BTA) to quantify functional diversity for infauna and epifauna communities collected from the same locations across a depth gradient in the Southern Benguela Shelf ecoregion. Infauna experienced an increase in functional uniqueness with depth, whereas epifauna experienced an increase in functional redundancy with depth. As a result, the epifauna species assemblage predicted 43 % of the epifauna trait assemblage, whereas the infauna species assemblage predicted only 8 % of the infauna trait assemblage. These findings suggest that taxonomic diversity and functional diversity changes are not congruent within and between marine benthic faunal groups. We emphasize the need to utilise both biodiversity metrics when quantifying marine biodiversity for conservation and management objectives.

Spatiotemporal stressors, not secondary structures or small temperature increases, control rapid facilitation of intertidal epifauna.

Small animals (epifauna) are ubiquitous in marine systems. Epifauna have high secondary production and provide trophic linkages between primary producers and higher-order consumers, like fish. Despite their importance, little is known about how these animals respond to warming or how their communities vary across spatiotemporal gradients. Here we use mimics of turf seaweed and invasive kelp holdfast to test, in a 5-factorial field experiment, whether intertidal epifauna are facilitated by different habitat structures, temperature conditions, and along cooccurring spatiotemporal gradients. We found that facilitation of epifauna by intertidal turf seaweed peaked in summer, at low elevation, in older habitats and at a less wave-exposed site. However, epifauna were not affected by the presence of a secondary structure like kelp holdfast mimics or small temperature increases from passive solar heating of black and white mimics. There were many significant two-way, but few higher order interactions, showing stronger facilitation under specific environmental conditions, like at low elevation in summer, or low elevation in old habitats. These results highlight that turf-associated epifauna are controlled by vertical elevation, season, hydrodynamics, and habitat age, and appear to be resilient to small temperature increases. Findings are important to better understand linkages between primary producers and higher order consumers and system-wide productivity, and because fast growing turf, facilitated by global warming and eutrophication, are increasingly outcompeting slower growing large perennial canopy forming seaweeds, like kelp and rockweeds.

Temporal variations of diversity and community structure of epifauna along the latitudinal and depth gradients in China seas over nearly 60 years.

To examine the diversity and community structure of epifauna along latitudinal and depth gradients in China seas and its temporal variations over nearly 60 years, a total of 61,424 records of epifauna were compiled. Latitudinal gradients of γ-diversity from 28 to 37°N showed two peaks of increase in 1958-1960; however, only one peak was observed near 28°N in 2000-2015. Along depth gradient, the peak of species richness shifted about 10 m deeper from about 50 m in 1958-1960 to about 60 m in 2000-2015. Values of α-diversity were higher in 2000-2015 than that in 1958-1960 in latitudes from 30 to 33°N and in depth from 50 to 75 m, respectively. Epifaunal communities were distributed basically along latitudinal and depth gradients. Spatial patterns of communities in the southern Yellow Sea and northern East China Sea were found relatively stable over nearly 60 years.

Drivers of variation in seagrass-associated amphipods across biogeographical areas.

Amphipods are one of the dominant epifaunal groups in seagrass meadows. However, our understanding of the biogeographical patterns in the distribution of these small crustaceans is limited. In this study, we investigated such patterns and the potential drivers in twelve Cymodocea nodosa meadows within four distinctive biogeographical areas across 2000 Km and 13° of latitude in two ocean basins (Mediterranean Sea and Atlantic Ocean). We found that species abundances in the assemblage of seagrass-associated amphipods differed among areas following a pattern largely explained by seagrass leaf area and epiphyte biomass, while the variation pattern in species presence/absence was determined by seagrass density and epiphyte biomass. Seagrass leaf area was also the most important determinant of greater amphipod total density and species richness, while amphipod density also increased with algal cover. Overall, our results evidenced that biogeographical patterns of variation in amphipod assemblages are mainly influenced by components of the habitat structure, which covary with environmental conditions, finding that structurally more complex meadows harboring higher abundance and richness of amphipods associated.

Facilitation of animals is stronger during summer marine heatwaves and around morphologically complex foundation species.

Foundation species create biogenic habitats, modify environmental conditions, augment biodiversity, and control animal community structures. In recent decades, marine heatwaves (MHWs) have affected the ecology of foundation species worldwide, and perhaps also their associated animal communities. However, no realistic field experiment has tested how MHWs affect animals that live in and around these foundation species. We therefore tested, in a four-factorial field experiment, if colonisation by small mobile marine animals (epifauna) onto plates with attached single versus co-occurring foundation species of different morphological complexities, were affected by 3-5°C heating (that mirrored a recent extreme MHW in the study area) and if the heating effect on the epifauna varied within and between seasons. For this experiment mimics of turf seaweed represented the single foundation species and holdfasts of seven common canopy-forming seaweed represented the co-occurring foundation species with different morphological complexities. We found that the taxonomic richness and total abundance of epifauna, dominated by copepods, generally were higher on heated plates with complex seaweed holdfasts in warmer summer trials. Furthermore, several interactions between test-factors were significant, e.g., epifaunal abundances, were, across taxonomic groups, generally higher in warmer than colder summer trials. These results suggest that, in temperate ecosystems, small, mobile, short-lived, and fast-growing marine epifauna can be facilitated by warmer oceans and morphologically complex foundation species, implying that future MHWs may increase secondary production and trophic transfers between primary producers and fish. Future studies should test whether these results can be scaled to other ecological species-interactions, across latitudes and biogeographical regions, and if similar results are found after longer MHWs or within live foundation species under real MHW conditions.

Sea spiders (Arthropoda, Pycnogonida) from ten recent research expeditions to the Antarctic Peninsula, Scotia Arc and Weddell Sea - data.

Background: This dataset contains information on specimens of Southern Ocean Pycnogonida (Arthropoda), that were collected from ten different research cruises, spanning 13 years. The individual aims and objectives of each cruise can be found in their cruise reports. The specimens have been collated into a single dataset, forming the basis of J. Maxwell's PhD. The dataset will be used to investigate the community structure of Antarctic pycnogonids and the factors which influence its composition. This dataset is published by SCAR-AntOBIS under the licence CC-BY 4.0. Please follow the guidelines from the SCAR and IPY Data Policies (https://www.scar.org/excom-meetings/xxxi-scar-delegates-2010-buenos-aires-argentina/4563-scar-xxxi-ip04b-scar-data-policy/file/) when using the data. If you have any questions regarding this dataset, please do not hesitate to contact us via the contact information provided in the metadata or via data-biodiversity-aq@naturalsciences.be. New information: This dataset adds vital occurrence and abundance data for pycnogonids from 10 previously unexamined research cruises from the Weddell Sea, Antarctic Penisula and the islands of the Scotia Arc. It includes the first pycnogonid data from the Prince Gustav Channel. The 197 sampling stations within this dataset represent an 11% increase in the number of stations where pycnogonids have been recorded in the Southern Ocean, southern South America and New Zealand waters and an 18% increase for above 60 degrees latitude. Presence data for any observed epifauna are also included.

Could the annual Saccorhiza polyschides replace a sympatric perennial kelp (Laminaria ochroleuca) when it comes to supporting the holdfast-associated fauna?

Kelp forests dominate temperate rocky shores worldwide but are declining globally with consequences for organisms that depend on them. In NW Iberia, the golden kelp (Laminaria ochroleuca) commonly occurs alongside a fast-growing annual that, unlike the golden kelp, does not seem to have receded in recent times (Saccorhiza polyschides). Here, we assessed whether the bulbous holdfast of S. polyschides could replace the intricate holdfast of L. ochroleuca as epifaunal habitat provider. Richness, diversity and total abundance of epifauna was similar in both seaweeds, while colonial/encrusting fauna was more abundant in L. ochroleuca. More importantly, each host supported a distinctive assemblage structure, indicating that S. polyschides seems an unsuitable replacement for L. ochroleuca as habitat provider for holdfast epifauna. Therefore, while S. polyschides may contribute substantially to the kelp forest canopy in some seasons, a regional decline of L. ochroleuca will likely alter the patterns of biodiversity within kelp stands.

The invasive seaweed Asparagopsis taxiformis erodes the habitat structure and biodiversity of native algal forests in the Mediterranean Sea.

Invasive seaweeds are listed among the most relevant threats to marine ecosystems worldwide. Biodiversity hotspots, such as the Mediterranean Sea, are facing multiple invasions and are expected to be severely affected by the introduction of new non-native seaweeds in the near future. In this study, we evaluated the consequences of the shift from the native Ericaria brachycarpa to the invasive Asparagopsis taxiformis habitat on the shallow rocky shores of Favignana Island (Egadi Islands, MPA, Sicily, Italy). We compared algal biomass and species composition and structure of the associated epifaunal assemblages in homogenous and mixed stands of E. brachycarpa and A. taxiformis. The results showed that the biomass of primary producers is reduced by 90% in the A. taxiformis invaded habitat compared to the E. brachycarpa native habitat. The structure of the epifaunal assemblages displayed significant variations among homogenous and mixed stands. The abundance, species richness and Shannon-Wiener diversity index of the epifaunal assemblages decreased by 89%, 78% and 40%, respectively, from homogenous stands of the native E. brachycarpa to the invasive A. taxiformis. Seaweed biomass was the structural attribute better explaining the variation in epifaunal abundance, species richness and diversity. Overall, our results suggest that the shift from E. brachycarpa to A. taxiformis habitat would drastically erode the biomass of primary producers and the associated biodiversity. We hypothesize that a complete shift from native to invasive seaweeds could ultimately lead to bottom-up effects on rocky shore habitats, with negative consequences for the ecosystem structure, functioning, and the services provided.

Impacts of habitat and predation on epifaunal communities from seagrass beds and artificial structures.

Within the coastal marine environment, the increased presence of artificial habitat can have negative impacts on the functioning of marine communities. Artificial structures provide a novel, hard surface for the colonization and growth of a variety of marine species and disproportionally favor introduced species. With the global rise in hardened shorelines, it is imperative to examine the ecological processes that occur within these habitats to those occurring in natural habitats. Here, we compared habitat differences in fouling community composition of different successional ages as well as the impact of predation on those communities. Specifically, we investigated how communities differed with respect to natural (seagrass beds) and artificial (docks) habitats and then exposed previously caged communities to predators to examine prey-specific effects within each habitat and on different aged communities. We found that habitat was a good predictor of community structure including both total species richness and introduced species richness higher in artificial habitats. We expected predators to increase available space allowing increased species co-existence, however, this was not the case. Predators in both habitats reduced richness despite having a strong impact on the percent cover of dominant groups. Predators also reduced introduced species richness, particularly in artificial habitats. Artificial structures are an important pathway of success for introduced species and results here show the importance of biotic resistance within these habitats, potentially limiting the spread of introduced species into natural habitat. Overall, species found within the different habitats could be predicted based on life history traits and predators did not increase the similarity of communities between habitats though still acted in a comparable way, reducing the dominant groups.

Fine-scale responses of mobile invertebrates and mesopredatory fish to habitat configuration.

As habitat-forming species continue to decline globally, it is important to understand how associated communities respond to habitat loss and fragmentation. Changes in the density and spatial configuration of habitat have important consequences for associated communities. However, tests of these factors are often confounded by morphological variation of habitat-formers, which can be resolved by using standardised habitat-mimics. Furthermore, few studies have incorporated the role of predators in mediating the observed effects. To test whether predators mediate the abundance of invertebrates among algal habitats of varying configuration (isolated vs patches, and positions within patches), we placed macroalgal mimics into subtidal estuarine habitats for one month to sample epifaunal communities. At the same time, we conducted underwater video surveys of fish communities to quantify fish communities and their feeding behaviour among the artificial habitats. Isolated habitats did not differ from patch habitats, however, patch edges had the highest epifaunal abundance, where fish were least commonly observed. Observed fish feeding was highest in the middle of patches and increased fish observations and feeding in habitats with reduced epifaunal communities suggest that mesopredatory fish are mediating epifauna in patches, with predation pressure altered by the spatial configuration of the habitat. This contrasts to previous studies that focus on predators that congregate outside patches and suggest that fragmentation leads to reduced invertebrate abundance at habitat edges in contrast to centres. However, this study highlights that in habitat patches housing small mesopredators that also benefit from the increased structure, the centre of the patch experiences higher predation and therefore fewer epifauna in contrast to patch edges and individual algal mimics.

Immediate effects of the 2019 oil spill on the macrobenthic fauna associated with macroalgae on the tropical coast of Brazil.

The 2019-2020 Brazil oil spill disaster affected several ecosystems and species-rich areas. The shallow-water reefs of Paiva beach are among the best-preserved coastal environments in the State of Pernambuco (NE Brazil), but were severely affected by oil arrival at the end of 2019. The reefs are densely covered by macroalgae as important biogenic substrates for benthic fauna. Based on that, herein, we provide a baseline assessment of the immediate impacts of the oil spill on the structure of epifaunal communities associated with the algae Jania capillacea and Penicillus capitatus. The benthic communities in both algae simplified (reduction of species richness and abundance) soon after oil arrival, while opportunistic taxa increased. After two months, the macrofaunal communities restructured to almost pre-disaster levels. However, polychaeta diversity remained low. Despite the apparently fast recovery of reef macrobenthos, a more detailed, long-term monitoring is necessary to evaluate the chronic effects.

Indirect effects shape macroalgal epifaunal communities.

We tested the response of algal epifauna to the direct effects of predation and the indirect consequences of habitat change due to grazing and nutrient supply through upwelling using an abundant intertidal rhodophyte, Gelidium pristoides. We ran a mid-shore field experiment at four sites (two upwelling sites interspersed with two non-upwelling sites) along 450 km of the south coast of South Africa. The experiment was started in June 2014 and ran until June 2015. Four treatments (predator exclusion, grazer exclusion, control, and procedural control) set out in a block design (n = 5) were monitored monthly for algal cover for the first 6 months and every 2 months for the last 6 months. Epifaunal abundance, species composition, algal cover, and algal architectural complexity (measured using fractal geometry) were assessed after 12 months. Predation had no significant effect on epifaunal abundances, while upwelling interacted with treatment. Grazing reduced the architectural complexity of algae, with increased fractal dimensions in the absence of grazers, and also reduced algal cover at all sites, though the latter effect was only significant for upwelling sites. Epifaunal community composition was not significantly affected by the presence of herbivores or predators but differed among sites independently of upwelling; sites were more similar to nearby sites than those farther away. In contrast, total epifaunal abundance was significantly affected by grazing, when normalized to algal cover. Grazing reduced the cover of algae; thus, epifaunal abundances were not affected by the direct top-down effects of predation but did respond to the indirect effects of grazing on habitat availability and quality. Our results indicate that epifaunal communities can be strongly influenced by the indirect consequences of biotic interactions.