Spatial synchrony cascades across ecosystem boundaries and up food webs via resource subsidies.

Cross-ecosystem subsidies are critical to ecosystem structure and function, especially in recipient ecosystems where they are the primary source of organic matter to the food web. Subsidies are indicative of processes connecting ecosystems and can couple ecological dynamics across system boundaries. However, the degree to which such flows can induce cross-ecosystem cascades of spatial synchrony, the tendency for system fluctuations to be correlated across locations, is not well understood. Synchrony has destabilizing effects on ecosystems, adding to the importance of understanding spatiotemporal patterns of synchrony transmission. In order to understand whether and how spatial synchrony cascades across the marine-terrestrial boundary via resource subsidies, we studied the relationship between giant kelp forests on rocky nearshore reefs and sandy beach ecosystems that receive resource subsidies in the form of kelp wrack (detritus). We found that synchrony cascades from rocky reefs to sandy beaches, with spatiotemporal patterns mediated by fluctuations in live kelp biomass, wave action, and beach width. Moreover, wrack deposition synchronized local abundances of shorebirds that move among beaches seeking to forage on wrack-associated invertebrates, demonstrating that synchrony due to subsidies propagates across trophic levels in the recipient ecosystem. Synchronizing resource subsidies likely play an underappreciated role in the spatiotemporal structure, functioning, and stability of ecosystems.

A marine protected area network does not confer community structure resilience to a marine heatwave across coastal ecosystems.

Marine protected areas (MPAs) have gained attention as a conservation tool for enhancing ecosystem resilience to climate change. However, empirical evidence explicitly linking MPAs to enhanced ecological resilience is limited and mixed. To better understand whether MPAs can buffer climate impacts, we tested the resistance and recovery of marine communities to the 2014-2016 Northeast Pacific heatwave in the largest scientifically designed MPA network in the world off the coast of California, United States. The network consists of 124 MPAs (48 no-take state marine reserves, and 76 partial-take or special regulation conservation areas) implemented at different times, with full implementation completed in 2012. We compared fish, benthic invertebrate, and macroalgal community structure inside and outside of 13 no-take MPAs across rocky intertidal, kelp forest, shallow reef, and deep reef nearshore habitats in California's Central Coast region from 2007 to 2020. We also explored whether MPA features, including age, size, depth, proportion rock, historic fishing pressure, habitat diversity and richness, connectivity, and fish biomass response ratios (proxy for ecological performance), conferred climate resilience for kelp forest and rocky intertidal habitats spanning 28 MPAs across the full network. Ecological communities dramatically shifted due to the marine heatwave across all four nearshore habitats, and MPAs did not facilitate habitat-wide resistance or recovery. Only in protected rocky intertidal habitats did community structure significantly resist marine heatwave impacts. Community shifts were associated with a pronounced decline in the relative proportion of cold water species and an increase in warm water species. MPA features did not explain resistance or recovery to the marine heatwave. Collectively, our findings suggest that MPAs have limited ability to mitigate the impacts of marine heatwaves on community structure. Given that mechanisms of resilience to climate perturbations are complex, there is a clear need to expand assessments of ecosystem-wide consequences resulting from acute climate-driven perturbations, and the potential role of regulatory protection in mitigating community structure changes.

A comparison of biomonitoring methodologies for surf zone fish communities.

Surf zones are highly dynamic marine ecosystems that are subject to increasing anthropogenic and climatic pressures, posing multiple challenges for biomonitoring. Traditional methods such as seines and hook and line surveys are often labor intensive, taxonomically biased, and can be physically hazardous. Emerging techniques, such as baited remote underwater video (BRUV) and environmental DNA (eDNA) are promising nondestructive tools for assessing marine biodiversity in surf zones of sandy beaches. Here we compare the relative performance of beach seines, BRUV, and eDNA in characterizing community composition of bony (teleost) and cartilaginous (elasmobranch) fishes of surf zones at 18 open coast sandy beaches in southern California. Seine and BRUV surveys captured overlapping, but distinct fish communities with 50% (18/36) of detected species shared. BRUV surveys more frequently detected larger species (e.g. sharks and rays) while seines more frequently detected one of the most abundant species, barred surfperch (Amphistichus argenteus). In contrast, eDNA metabarcoding captured 88.9% (32/36) of all fishes observed in seine and BRUV surveys plus 57 additional species, including 15 that frequent surf zone habitats. On average, eDNA detected over 5 times more species than BRUVs and 8 times more species than seine surveys at a given site. eDNA approaches also showed significantly higher sensitivity than seine and BRUV methods and more consistently detected 31 of the 32 (96.9%) jointly observed species across beaches. The four species detected by BRUV/seines, but not eDNA were only resolved at higher taxonomic ranks (e.g. Embiotocidae surfperches and Sygnathidae pipefishes). In frequent co-detection of species between methods limited comparisons of richness and abundance estimates, highlighting the challenge of comparing biomonitoring approaches. Despite potential for improvement, results overall demonstrate that eDNA can provide a cost-effective tool for long-term surf zone monitoring that complements data from seine and BRUV surveys, allowing more comprehensive surveys of vertebrate diversity in surf zone habitats.

Uptake of polycyclic aromatic hydrocarbons via high-energy water accommodated fraction (HEWAF) by beach hoppers (Amphipoda, Talitridae) using different sandy beach exposure pathways.

Sandy beach ecosystems are highly dynamic coastal environments subject to a variety of anthropogenic pressures and impacts. Pollution from oil spills can damage beach ecosystems through the toxic effects of hydrocarbons on organisms and the disruptive nature of large-scale clean-up practices. On temperate sandy beaches, intertidal talitrid amphipods are primary consumers of macrophyte wrack subsidies and serve as prey for higher trophic level consumers, such as birds and fish. These integral organisms of the beach food web can be exposed to hydrocarbons by direct contact with oiled sand through burrowing and by the consumption of oiled wrack. We experimentally evaluated the primary polycyclic aromatic hydrocarbon (PAH) exposure pathway via high-energy water accommodated fraction (HEWAF) for a species of talitrid amphipod (Megalorchestia pugettensis). Our results indicated that tissue PAH concentrations in talitrids were six-fold higher in treatments that included oiled sand compared to those with only oiled kelp and the controls.

The role of inputs of marine wrack and carrion in sandy-beach ecosystems: a global review.

Sandy beaches are iconic interfaces that functionally link the ocean with the land via the flow of organic matter from the sea. These cross-ecosystem fluxes often comprise uprooted seagrass and dislodged macroalgae that can form substantial accumulations of detritus, termed 'wrack', on sandy beaches. In addition, the tissue of the carcasses of marine animals that regularly wash up on beaches form a rich food source ('carrion') for a diversity of scavenging animals. Here, we provide a global review of how wrack and carrion provide spatial subsidies that shape the structure and functioning of sandy-beach ecosystems (sandy beaches and adjacent surf zones), which typically have little in situ primary production. We also examine the spatial scaling of the influence of these processes across the broader land- and seascape, and identify key gaps in our knowledge to guide future research directions and priorities. Large quantities of detrital kelp and seagrass can flow into sandy-beach ecosystems, where microbial decomposers and animals process it. The rates of wrack supply and its retention are influenced by the oceanographic processes that transport it, the geomorphology and landscape context of the recipient beaches, and the condition, life history and morphological characteristics of the macrophyte taxa that are the ultimate source of wrack. When retained in beach ecosystems, wrack often creates hotspots of microbial metabolism, secondary productivity, biodiversity, and nutrient remineralization. Nutrients are produced during wrack breakdown, and these can return to coastal waters in surface flows (swash) and aquifers discharging into the subtidal surf. Beach-cast kelp often plays a key trophic role, being an abundant and preferred food source for mobile, semi-aquatic invertebrates that channel imported algal matter to predatory invertebrates, fish, and birds. The role of beach-cast marine carrion is likely to be underestimated, as it can be consumed rapidly by highly mobile scavengers (e.g. foxes, coyotes, raptors, vultures). These consumers become important vectors in transferring marine productivity inland, thereby linking marine and terrestrial ecosystems. Whilst deposits of organic matter on sandy-beach ecosystems underpin a range of ecosystem functions and services, they can be at variance with aesthetic perceptions resulting in widespread activities, such as 'beach cleaning and grooming'. This practice diminishes the energetic base of food webs, intertidal fauna, and biodiversity. Global declines in seagrass beds and kelp forests (linked to global warming) are predicted to cause substantial reductions in the amounts of marine organic matter reaching many beach ecosystems, likely causing flow-on effects for food webs and biodiversity. Similarly, future sea-level rise and increased storm frequency are likely to alter profoundly the physical attributes of beaches, which in turn can change the rates at which beaches retain and process the influxes of wrack and animal carcasses. Conservation of the multi-faceted ecosystem services that sandy beaches provide will increasingly need to encompass a greater societal appreciation and the safeguarding of ecological functions reliant on beach-cast organic matter on innumerable ocean shores worldwide.

Diet of a threatened endemic fox reveals variation in sandy beach resource use on California Channel Islands.

The coastal zone provides foraging opportunities for insular populations of terrestrial mammals, allowing for expanded habitat use, increased dietary breadth, and locally higher population densities. We examined the use of sandy beach resources by the threatened island fox (Urocyon littoralis) on the California Channel Islands using scat analysis, surveys of potential prey, beach habitat attributes, and stable isotope analysis. Consumption of beach invertebrates, primarily intertidal talitrid amphipods (Megalorchestia spp.) by island fox varied with abundance of these prey across sites. Distance-based linear modeling revealed that abundance of giant kelp (Macrocystis pyrifera) wrack, rather than beach physical attributes, explained the largest amount of variation in talitrid amphipod abundance and biomass across beaches. δ13C and δ15N values of fox whisker (vibrissae) segments suggested individualism in diet, with generally low δ13C and δ15N values of some foxes consistent with specializing on primarily terrestrial foods, contrasting with the higher isotope values of other individuals that suggested a sustained use of sandy beach resources, the importance of which varied over time. Abundant allochthonous marine resources on beaches, including inputs of giant kelp, may expand habitat use and diet breadth of the island fox, increasing population resilience during declines in terrestrial resources associated with climate variability and long-term climate change.

Species identity drives ecosystem function in a subsidy-dependent coastal ecosystem.

Declines in species diversity carry profound implications for ecosystem functioning. Communities of primary producers and consumers interact on evolutionary as well as ecological time scales, shaping complex relationships between biodiversity and ecosystem functioning. In subsidized ecosystems, resource inputs are independent of consumer actions, offering a simplified view of the relationship between species diversity and function for higher trophic levels. With food webs supported by substantial but variable inputs of detritus from adjacent marine ecosystems, sandy beaches are classic examples of subsidized ecosystems. We investigated effects of consumer species diversity and identity on a key ecological function, consumption of kelp wrack from nearshore giant kelp (Macrocystis pyrifera) forests. We assessed effects of species richness on kelp consumption by experimentally manipulating richness of six common species of invertebrate detritivores in laboratory mesocosms and conducting field assays of kelp consumption on beaches. Consumer richness had no effect on kelp consumption in the field and a slight negative effect in laboratory experiments. Kelp consumption was most strongly affected by the species composition of the detritivore community. Species identity and body size of intertidal detritivores drove variation in kelp consumption rates in both experiments and field assays. Our results provide further evidence that species traits, rather than richness per se, influence ecosystem function most, particularly in detrital-based food webs with high functional redundancy across species. On sandy beaches, where biodiversity is threatened by rising sea levels and expanding development, our findings suggest that loss of large-bodied consumer species could disproportionally impact ecosystem function.

Multiple climate change-driven tipping points for coastal systems.

As the climate evolves over the next century, the interaction of accelerating sea level rise (SLR) and storms, combined with confining development and infrastructure, will place greater stresses on physical, ecological, and human systems along the ocean-land margin. Many of these valued coastal systems could reach "tipping points," at which hazard exposure substantially increases and threatens the present-day form, function, and viability of communities, infrastructure, and ecosystems. Determining the timing and nature of these tipping points is essential for effective climate adaptation planning. Here we present a multidisciplinary case study from Santa Barbara, California (USA), to identify potential climate change-related tipping points for various coastal systems. This study integrates numerical and statistical models of the climate, ocean water levels, beach and cliff evolution, and two soft sediment ecosystems, sandy beaches and tidal wetlands. We find that tipping points for beaches and wetlands could be reached with just 0.25 m or less of SLR (~ 2050), with > 50% subsequent habitat loss that would degrade overall biodiversity and ecosystem function. In contrast, the largest projected changes in socioeconomic exposure to flooding for five communities in this region are not anticipated until SLR exceeds 0.75 m for daily flooding and 1.5 m for storm-driven flooding (~ 2100 or later). These changes are less acute relative to community totals and do not qualify as tipping points given the adaptive capacity of communities. Nonetheless, the natural and human built systems are interconnected such that the loss of natural system function could negatively impact the quality of life of residents and disrupt the local economy, resulting in indirect socioeconomic impacts long before built infrastructure is directly impacted by flooding.

Toxicity of Refugio Beach Oil to Sand Crabs (Emerita analoga), Blue Mussels (Mytilus sp.), and Inland Silversides (Menidia beryllina).

Monterey formation crude oil spilled from an onshore pipeline and entered the surf zone near Refugio State Beach, Santa Barbara County, California (USA) on 19 May 2015. During this season, early life stages of many marine fish and invertebrates were present. Surf zone water and beach porewater samples were collected during the 4 mo after the spill and 2 yr later for chemical analyses. Elevated polycyclic aromatic hydrocarbon (PAH) and total petroleum hydrocarbon concentrations were observed in surf zone water and porewater near the release point, declining with distance and time. Early life stage toxicity was investigated by conducting 6- and 7-d static renewal bioassays with sand crab (Emerita analoga) post larvae (megalopae) and inland silverside larvae (Menidia beryllina), respectively, and a 48-h blue mussel (Mytilus sp.) embryo development bioassay. Dilutions of a high-energy water accommodated fraction of the Refugio Beach oil and a seawater control were prepared to simulate surf zone PAH concentrations (nominal PAH45 ; 0, 0.5, 1, 5, 10, 50, 100, and 500 µg/L). The PAH45 median lethal concentrations (LC50s), based on measured concentrations, were 381 µg/L for Mytilus sp., 75.6 µg/L for Menidia, and 40.9 µg/L for Emerita. Our results suggest that PAH concentrations in coastal waters of the spill-affected area were potentially lethal to early life stages of fish and invertebrates. Environ Toxicol Chem 2021;40:2578-2586. © 2021 SETAC.

Local scale processes drive long-term change in biodiversity of sandy beach ecosystems.

Evaluating impacts to biodiversity requires ecologically informed comparisons over sufficient time spans. The vulnerability of coastal ecosystems to anthropogenic and climate change-related impacts makes them potentially valuable indicators of biodiversity change. To evaluate multidecadal change in biodiversity, we compared results from intertidal surveys of 13 sandy beaches conducted in the 1970s and 2009-11 along 500 km of coast (California, USA). Using a novel extrapolation approach to adjust species richness for sampling effort allowed us to address data gaps and has promise for application to other data-limited biodiversity comparisons. Long-term changes in species richness varied in direction and magnitude among beaches and with human impacts but showed no regional patterns. Observed long-term changes in richness differed markedly among functional groups of intertidal invertebrates. At the majority (77%) of beaches, changes in richness were most evident for wrack-associated invertebrates suggesting they have disproportionate vulnerability to impacts. Reduced diversity of this group was consistent with long-term habitat loss from erosion and sea level rise at one beach. Wrack-associated species richness declined over time at impacted beaches (beach fill and grooming), despite observed increases in overall intertidal richness. In contrast richness of these taxa increased at more than half (53%) of the beaches including two beaches recovering from decades of off-road vehicle impacts. Over more than three decades, our results suggest that local scale processes exerted a stronger influence on intertidal biodiversity on beaches than regional processes and highlight the role of human impacts for local spatial scales. Our results illustrate how comparisons of overall biodiversity may mask ecologically important changes and stress the value of evaluating biodiversity change in the context of functional groups. The long-term loss of wrack-associated species, a key component of sandy beach ecosystems, documented here represents a significant threat to the biodiversity and function of coastal ecosystems.

Macroscale patterns in body size of intertidal crustaceans provide insights on climate change effects.

Predicting responses of coastal ecosystems to altered sea surface temperatures (SST) associated with global climate change, requires knowledge of demographic responses of individual species. Body size is an excellent metric because it scales strongly with growth and fecundity for many ectotherms. These attributes can underpin demographic as well as community and ecosystem level processes, providing valuable insights for responses of vulnerable coastal ecosystems to changing climate. We investigated contemporary macroscale patterns in body size among widely distributed crustaceans that comprise the majority of intertidal abundance and biomass of sandy beach ecosystems of the eastern Pacific coasts of Chile and California, USA. We focused on ecologically important species representing different tidal zones, trophic guilds and developmental modes, including a high-shore macroalga-consuming talitrid amphipod (Orchestoidea tuberculata), two mid-shore scavenging cirolanid isopods (Excirolana braziliensis and E. hirsuticauda), and a low-shore suspension-feeding hippid crab (Emerita analoga) with an amphitropical distribution. Significant latitudinal patterns in body sizes were observed for all species in Chile (21° - 42°S), with similar but steeper patterns in Emerita analoga, in California (32°- 41°N). Sea surface temperature was a strong predictor of body size (-4% to -35% °C-1) in all species. Beach characteristics were subsidiary predictors of body size. Alterations in ocean temperatures of even a few degrees associated with global climate change are likely to affect body sizes of important intertidal ectotherms, with consequences for population demography, life history, community structure, trophic interactions, food-webs, and indirect effects such as ecosystem function. The consistency of results for body size and temperature across species with different life histories, feeding modes, ecological roles, and microhabitats inhabiting a single widespread coastal ecosystem, and for one species, across hemispheres in this space-for-time substitution, suggests predictions of ecosystem responses to thermal effects of climate change may potentially be generalised, with important implications for coastal conservation.

The Effects of Anthropogenic Structures on Habitat Connectivity and the Potential Spread of Non-Native Invertebrate Species in the Offshore Environment.

Offshore structures provide habitat that could facilitate species range expansions and the introduction of non-native species into new geographic areas. Surveys of assemblages of seven offshore oil and gas platforms in the Santa Barbara Channel revealed a change in distribution of the non-native sessile invertebrate Watersipora subtorquata, a bryozoan with a planktonic larval duration (PLD) of 24 hours or less, from one platform in 2001 to four platforms in 2013. We use a three-dimensional biophysical model to assess whether larval dispersal via currents from harbors to platforms and among platforms is a plausible mechanism to explain the change in distribution of Watersipora and to predict potential spread to other platforms in the future. Hull fouling is another possible mechanism to explain the change in distribution of Watersipora. We find that larval dispersal via currents could account for the increase in distribution of Watersipora from one to four platforms and that Watersipora is unlikely to spread from these four platforms to additional platforms through larval dispersal. Our results also suggest that larvae with PLDs of 24 hours or less released from offshore platforms can attain much greater dispersal distances than larvae with PLDs of 24 hours or less released from nearshore habitat. We hypothesize that the enhanced dispersal distance of larvae released from offshore platforms is driven by a combination of the offshore hydrodynamic environment, larval behavior, and larval release above the seafloor.

Responses of dune plant communities to continental uplift from a major earthquake: sudden releases from coastal squeeze.

Vegetated dunes are recognized as important natural barriers that shelter inland ecosystems and coastlines suffering daily erosive impacts of the sea and extreme events, such as tsunamis. However, societal responses to erosion and shoreline retreat often result in man-made coastal defence structures that cover part of the intertidal and upper shore zones causing coastal squeeze and habitat loss, especially for upper shore biota, such as dune plants. Coseismic uplift of up to 2.0 m on the Peninsula de Arauco (South central Chile, ca. 37.5º S) caused by the 2010 Maule earthquake drastically modified the coastal landscape, including major increases in the width of uplifted beaches and the immediate conversion of mid to low sandy intertidal habitat to supralittoral sandy habitat above the reach of average tides and waves. To investigate the early stage responses in species richness, cover and across-shore distribution of the hitherto absent dune plants, we surveyed two formerly intertidal armoured sites and a nearby intertidal unarmoured site on a sandy beach located on the uplifted coast of Llico (Peninsula de Arauco) over two years. Almost 2 years after the 2010 earthquake, dune plants began to recruit, then rapidly grew and produced dune hummocks in the new upper beach habitats created by uplift at the three sites. Initial vegetation responses were very similar among sites. However, over the course of the study, the emerging vegetated dunes of the armoured sites suffered a slowdown in the development of the spatial distribution process, and remained impoverished in species richness and cover compared to the unarmoured site. Our results suggest that when released from the effects of coastal squeeze, vegetated dunes can recover without restoration actions. However, subsequent human activities and management of newly created beach and dune habitats can significantly alter the trajectory of vegetated dune development. Management that integrates the effects of natural and human induced disturbances, and promotes the development of dune vegetation as natural barriers can provide societal and conservation benefits in coastal ecosystems.

Metrics to assess ecological condition, change, and impacts in sandy beach ecosystems.

Complexity is increasingly the hallmark in environmental management practices of sandy shorelines. This arises primarily from meeting growing public demands (e.g., real estate, recreation) whilst reconciling economic demands with expectations of coastal users who have modern conservation ethics. Ideally, shoreline management is underpinned by empirical data, but selecting ecologically-meaningful metrics to accurately measure the condition of systems, and the ecological effects of human activities, is a complex task. Here we construct a framework for metric selection, considering six categories of issues that authorities commonly address: erosion; habitat loss; recreation; fishing; pollution (litter and chemical contaminants); and wildlife conservation. Possible metrics were scored in terms of their ability to reflect environmental change, and against criteria that are widely used for judging the performance of ecological indicators (i.e., sensitivity, practicability, costs, and public appeal). From this analysis, four types of broadly applicable metrics that also performed very well against the indicator criteria emerged: 1.) traits of bird populations and assemblages (e.g., abundance, diversity, distributions, habitat use); 2.) breeding/reproductive performance sensu lato (especially relevant for birds and turtles nesting on beaches and in dunes, but equally applicable to invertebrates and plants); 3.) population parameters and distributions of vertebrates associated primarily with dunes and the supralittoral beach zone (traditionally focused on birds and turtles, but expandable to mammals); 4.) compound measurements of the abundance/cover/biomass of biota (plants, invertebrates, vertebrates) at both the population and assemblage level. Local constraints (i.e., the absence of birds in highly degraded urban settings or lack of dunes on bluff-backed beaches) and particular issues may require alternatives. Metrics - if selected and applied correctly - provide empirical evidence of environmental condition and change, but often do not reflect deeper environmental values per se. Yet, values remain poorly articulated for many beach systems; this calls for a comprehensive identification of environmental values and the development of targeted programs to conserve these values on sandy shorelines globally.

Ecological implications of extreme events: footprints of the 2010 earthquake along the Chilean coast.

Deciphering ecological effects of major catastrophic events such as earthquakes, tsunamis, volcanic eruptions, storms and fires, requires rapid interdisciplinary efforts often hampered by a lack of pre-event data. Using results of intertidal surveys conducted shortly before and immediately after Chile's 2010 M(w) 8.8 earthquake along the entire rupture zone (ca. 34-38°S), we provide the first quantification of earthquake and tsunami effects on sandy beach ecosystems. Our study incorporated anthropogenic coastal development as a key design factor. Ecological responses of beach ecosystems were strongly affected by the magnitude of land-level change. Subsidence along the northern rupture segment combined with tsunami-associated disturbance and drowned beaches. In contrast, along the co-seismically uplifted southern rupture, beaches widened and flattened increasing habitat availability. Post-event changes in abundance and distribution of mobile intertidal invertebrates were not uniform, varying with land-level change, tsunami height and coastal development. On beaches where subsidence occurred, intertidal zones and their associated species disappeared. On some beaches, uplift of rocky sub-tidal substrate eliminated low intertidal sand beach habitat for ecologically important species. On others, unexpected interactions of uplift with man-made coastal armouring included restoration of upper and mid-intertidal habitat seaward of armouring followed by rapid colonization of mobile crustaceans typical of these zones formerly excluded by constraints imposed by the armouring structures. Responses of coastal ecosystems to major earthquakes appear to vary strongly with land-level change, the mobility of the biota and shore type. Our results show that interactions of extreme events with human-altered shorelines can produce surprising ecological outcomes, and suggest these complex responses to landscape alteration can leave lasting footprints in coastal ecosystems.

Competitive interactions in macroinfaunal animals of exposed sandy beaches.

The influence of biotic interactions in structuring macroinfaunal communities of exposed sandy beaches, an unstable habitat characterized by strong physical forces, is generally considered negligible. We investigated the hypothesis that competitive interactions during burrowing could potentially affect the intertidal distribution and abundance of macroinfaunal animals of sandy beaches using two species of invertebrates, a hippid crab, Emerita analoga, and a bivalve, Mesodesma donacium, common along the coast of Chile. Spatial overlap in the intertidal distributions of these species was dynamic, varying with abundance, location, time of year and tide. Highest density zones of each species were often distinctly separated at low tide and spatial overlap in their distributions decreased significantly with increasing density, suggesting density dependence of the interactions. Negative relationships between densities of the two species at the smallest spatial scale examined also suggested active interactions among individuals. Over a tidal cycle, peak densities of the two species overlapped suggesting that interactions could occur frequently. Burrowing performance of E. analoga varied between size classes in three experimental densities of clams (5, 10 and 15 clams 0.008 m(-2)) and in controls with no clams. Burrowing times of large crabs were significantly longer (approximately twofold) in all densities of clams than in controls, while those of small crabs did not differ significantly among treatments and controls. Large crabs also displaced clams from the sand while burrowing suggesting that two mechanisms of direct interference can occur, both of which could increase exposure of individuals involved to active swash and transport across or along the beach with potentially negative consequences. Our results suggest that competitive interactions capable of affecting zonation and population and community biology on a number of scales can occur among burrowing macroinfauna on exposed sandy beaches. Those interactions could be more ecologically significant than previously appreciated and may contribute to patterns observed in community structure and zonation on sandy beaches. Our results illustrate the potential importance of negative biological interactions in a physically stressful environment.