Effects of sandy beach cusps on wrack accumulation,sediment characteristics and macrofaunal assemblages

Beach‐cast marine macroalgae and seagrasses, collectively termed wrack, provide shelter and habitat for beach fauna and can often provide a large input of energy and nutrients to sandy beaches. Wrack deposition on sandy beaches varies spatially and could be affected by morphological features on the beach face such as cusps. This study tested a series of hypotheses regarding the differences in wrack deposits, sediments and macrofaunal assemblages between cusp bays and horns on two beaches in South Australia. Bays had greater cover and larger pieces of wrack than horns. Sediment organic matter content was greater on horns than in bays but mean particle size did not differ consistently between bays and horns. Macrofaunal diversity was higher in bays and this pattern was probably driven by differences in the cover of wrack between bays and horns. Cusp morphology thus influences the distribution of wrack on the beach face, which in turn influences the distribution of macrofauna. Studies of sandy beaches with cusps should therefore be explicitly designed to sample cusp features and associated wrack deposits.     

Cusps and pipis on a sandy ocean beach in New South Wales

The effect of small-scale variation in beach morphology has been largely ignored in studies of beach macrofauna. This study examined the distribution and abundance of the large bivalve, Donax deltoides (commonly known as the ‘pipi’) in relation to beach cusps (rhythmically spaced undulations of the beachface consisting of scalloped bays and flanking horns on a scale of tens of metres). No differences were found in the abundance of pipis downshore of cusp horns versus cusp bays at any of three sites. In contrast, previous work on a relatively low-energy (reflective) beach reported greater abundances of bivalves below cusp bays than below cusp horns. Results suggest that the swash climate on the higher-energy (more dissipative) beach studied here produces weaker cusping and weaker patterns in the active or passive redistribution of beach fauna by swash action. Wider more dissipative beaches also allow beach macrofauna to remain on flatter parts of the beach, further below cusps than on reflective beaches. The more rigorous study design used here may also contribute to the difference in results between this study and previous studies. However, populations of pipis below cusp bays did contain a greater proportion of small individuals (≤20 mm length) than populations on cusp horns, suggesting some effect related to beach cusps. Small pipis live closer to the beach surface as they have shorter siphons and are subject to larger rates of dislodgement and movement from cusp horns to cusp bays by swash action than are large pipis. Future ecological studies of beaches with cusps should take beach cusps and the processes producing them into account as they may affect the distribution, abundance and the size distribution of beach invertebrates.     

Sandy beaches at the brink

Sandy beaches line most of the world's oceans and are highly valued by society: more people use sandy beaches than any other type of shore. While the economic and social values of beaches are generally regarded as paramount, sandy shores also have special ecological features and contain a distinctive biodiversity that is generally not recognized. These unique ecosystems are facing escalating anthropogenic pressures, chiefly from rapacious coastal development, direct human uses — mainly associated with recreation — and rising sea levels. Beaches are increasingly becoming trapped in a 'coastal squeeze' between burgeoning human populations from the land and the effects of global climate change from the sea. Society's interventions (e.g. shoreline armouring, beach nourishment) to combat changes in beach environments, such as erosion and shoreline retreat, can result in severe ecological impacts and loss of biodiversity at local scales, but are predicted also to have cumulative large-scale consequences worldwide. Because of the scale of this problem, the continued existence of beaches as functional ecosystems is likely to depend on direct conservation efforts. Conservation, in turn, will have to increasingly draw on a consolidated body of ecological theory for these ecosystems. Although this body of theory has yet to be fully developed, we identify here a number of critical research directions that are required to progress coastal management and conservation of sandy beach ecosystems.     

Diversity and Transport of Microorganisms in Intertidal Sands of the California Coast

Forced by tides and waves, large volumes of seawater are flushed through the beach daily. Organic material and nutrients in seawater are remineralized and cycled as they pass through the beach. Microorganisms are responsible for most of the biogeochemical cycling in the beach; however, few studies have characterized their diversity in intertidal sands, and little work has characterized the extent to which microbes are transported between different compartments of the beach. The present study uses next-generation massively parallel sequencing to characterize the microbial community present at 49 beaches along the coast of California. In addition, we characterize the transport of microorganisms within intertidal sands using laboratory column experiments. We identified extensive diversity in the beach sands. Nearly 1,000 unique taxa were identified in sands from 10 or more unique beaches, suggesting the existence of a group of “cosmopolitan” sand microorganisms. A biogeographical analysis identified a taxon-distance relationship among the beaches. In addition, sands with similar grain size, organic carbon content, exposed to a similar wave climate, and having the same degree of anthropogenic influence tended to have similar microbial communities. Column experiments identified microbes readily mobilized by seawater infiltrating through unsaturated intertidal sands. The ease with which microbes were mobilized suggests that intertidal sands may represent a reservoir of bacteria that seed the beach aquifer where they may partake in biogeochemical cycling.     

Influence of Land Use,Nutrients, and Geography on Microbial Communities and Fecal Indicator Abundance at Lake Michigan Beaches

Microbial communities within beach sand play a key role in nutrient cycling and are important to the nearshore ecosystem function. Escherichia coli and enterococci, two common indicators of fecal pollution, have been shown to persist in the beach sand, but little is known about how microbial community assemblages are related to these fecal indicator bacteria (FIB) reservoirs. We examined eight beaches across a geographic gradient and range of land use types and characterized the indigenous community structure in the water and the backshore, berm, and submerged sands. FIB were found at similar levels in sand at beaches adjacent to urban, forested, and agricultural land and in both the berm and backshore. However, there were striking differences in the berm and backshore microbial communities, even within the same beach, reflecting the very different environmental conditions in these beach zones in which FIB can survive. In contrast, the microbial communities in a particular beach zone were similar among beaches, including at beaches on opposite shores of Lake Michigan. The differences in the microbial communities that did exist within a beach zone correlated to nutrient levels, which varied among geographic locations. Total organic carbon and total phosphorus were higher in Wisconsin beach sand than in beach sand from Michigan. Within predominate genera, fine-scale sequence differences could be found that distinguished the populations from the two states, suggesting a biogeographic effect. This work demonstrates that microbial communities are reflective of environmental conditions at freshwater beaches and are able to provide useful information regarding long-term anthropogenic stress.     

The relationship between heritage,recreational quality and geomorphological vulnerability in the coastal zone: A case study of beach systems in the Canary Islands

The relationships between geomorphological vulnerability, recreational quality and heritage at beaches in the Canary Islands (Spain) are studied using three sets of indicators. Processes and their interactions are analysed at urban, semi-urban and natural beaches. Natural, cultural and landscape heritage acts as an attractor of recreational activity, which has intensified significantly on Canarian beaches in recent decades. Overcrowding has negatively impacted conservation of the natural and cultural heritage of the beaches, and has increased human pressure, leading to an increase in geomorphological susceptibility. The resulting reduction in geomorphological resilience has had a negative impact on beach facilities. Despite this common pattern, the relationships between these processes have particularities associated with the different types of beach (urban, semi-urban and natural). In order to establish new priorities in coastal public policy, these beach diagnoses and indices should be discussed and debated by the different actors involved in beach management.     

Some physical,chemical, and microbiological characteristics of two beaches of anglesey

Observations during 1971 and 1972 of some of the physical, chemical, and microbiological characteristics of contrasting Anglesey beaches, Newborough and Llanddona, are reported. The fine sandy beach at Newborough was observed to be very unstable and topographical changes were recorded. In particular, the movement of a sand wave across the intertidal zone from low water to extinction at the foot of the dune system was observed. The more extensive fine sandy beach at Llanddona had greater stability.Chemically, each beach was variable both spatially and temporally, with ill-defined patterns of concentration changes. Sand from Newborough beach was low in organic carbon (0.07–0.40 mg C/g dry sand) and well aerated, and the soluble inorganic nitrogen in the ground water (up to 30 μg-at. N/l) was dominated by nitrate form (up to 22 μg NO₃-N/l). By contrast, Llanddona sand had a more variable organic carbon content (0.22–2.25 mg C/g dry sand), was wetter, and poorly aerated with consequent sulphide lenses; its dissolved inorganic nitrogen (over 70 μg-at. N/l) was completely dominated by the ammonium form.Microbiologically, the beaches possessed dissimilar bacterial floras, and sediment from Llanddona gave higher bacterial counts than that from Newborough. For both beaches it is shown that estimated bacterial numbers decreased with depth as well as down the intertidal zone.     

Multi-Dimensional assessment of beach systems on the Catalan coast from a pragmatic and epistemological perspective

In recent decades, much of the progress observed in the study of beaches to aid policy has focused on producing sound substantive and/or contextual knowledge. However, attention has not usually been paid to the procedural manner in which it has been produced. Guided by an open learning process, this study aims to develop and present an expanded reviewed assessment of the Catalan coast operationalized by means of the Double-Looped Beach Quality Index (DL-BQI). Such openness helps us to understand which narratives of beach management in Catalonia are operating (and which are excluded), the extent to which they are unfolding, and the capacity of the DL-BQI to provide information for policymaking. Data collection enabled assessment of 96 representative beaches on the Catalan coast, that may find correspondence with other geographic locations. Both the instrument used for the assessment – the DL-BQI – and its results were co-evaluated at a multi-stakeholder meeting. In general, beaches have mainly been conditioned to supply elements for tourism and leisure. Different pressures have arisen from the orientation towards such activities, usually according to beach type, most of them negatively impacting the ecological assets and natural heritage of beaches. The economic growth narrative, albeit predominant, was shown to be deficient for tackling complex coastal problems, i.e. intensifying ecological deterioration. Alternatively, the study shows the importance for regional coastal policy of considering the value of beaches as natural systems, as advocated, to a greater or lesser extent, by the narratives of sustainability, environmental protection and integrated coastal zone management. This research innovates on considering substantive, contextual and procedural dimensions of knowledge quality for the governance of beach systems, by relating dimensions, narratives and outcomes.     

Exploring macroinvertebrate species distributions at regional and local scales across a sandy beach geographic continuum

Exposed sandy beaches are highly dynamic ecosystems where macroinvertebrate species cope with extremely variable environmental conditions. The majority of the beach ecology studies present exposed beaches as physically dominated ecosystems where abiotic factors largely determine the structure and distribution of macrobenthic communities. However, beach species patterns at different scales can be modified by the interaction between different environmental variables, including biotic interactions. In this study, we examined the role of different environmental variables for describing the regional and local scale distributions of common macrobenthic species across 39 beaches along the North coast of Spain. The analyses were carried out using boosted regression trees, a relatively new technique from the field of machine learning. Our study showed that the macroinvertebrate community on exposed beaches is not structured by a single physical factor, but instead by a complex set of drivers including the biotic compound. Thus, at a regional scale the macrobenthic community, in terms of number of species and abundance, was mainly explained by surrogates of food availability, such as chlorophyll a. The results also revealed that the local scale is a feasible way to construct general predictive species-environmental models, since relationships derived from different beaches showed similar responses for most of the species. However, additional information on aspects of beach species distribution can be obtained with large scale models. This study showed that species-environmental models should be validated against changes in spatial extent, and also illustrates the utility of BRTs as a powerful analysis tool for ecology data insight.     

Wrack promotes the persistence of fecal indicator bacteria in marine sands and seawater

Algae on freshwater beaches can serve as reservoirs for fecal indicator bacteria (FIB). Wrack (especially kelp) at marine beaches might sustain FIB as well. This study examines the relationship between beach wrack, FIB, and surrounding water and sediment at marine beaches along the California coast. Surveys of southern and central California beaches were conducted to observe environmental wrack-associated FIB concentrations. FIB concentrations normalized to dry weight were the highest in stranded dry wrack, followed by stranded wet and suspended 'surf' wrack. Laboratory microcosms were conducted to examine the effect of wrack on FIB persistence in seawater and sediment. Indigenous enterococci and Escherichia coli incubated in a seawater microcosm containing wrack showed increased persistence relative to those incubated in a microcosm without wrack. FIB concentrations in microcosms containing wrack-covered sand were significantly higher than those in uncovered sand after several days. These findings implicate beach wrack as an important FIB reservoir. The presence of wrack may increase water and sediment FIB levels, altering the relationship between FIB levels and actual health risk while possibly leading to beach closures. Further work will need to investigate the possibility of FIB growth on wrack and the potential for pathogen presence.     

Major food web properties of two sandy beaches with contrasting morphodynamics, and effects on the stability

We determined major structural properties influencing the food webs of two sandy beaches with contrasting morphodynamics in the Atlantic coast of Uruguay: reflective (narrow and steep) and dissipative beaches (wide and flat). Furthermore, we evaluated how these characteristics could influence the stability of the local food webs. To this end, we examined the correlation of several food web properties with different ecosystem types (including freshwater habitats, estuary, marine, and terrestrial environments) using a principal components analysis. Sandy beach food web components included detritus, phytoplankton, zooplankton, benthic invertebrates, fishes, and seabirds. Our results revealed that the dissipative beach presented higher trophic levels, a higher number of trophic species, more links per species, as well as a higher proportion of intermediate trophic species, but lower connectance and proportion of omnivorous species than the reflective beach. The variation in the food web properties was explained by two principal components. Sandy beach food webs contribute mainly to one dimension of the principal components analysis that was determined by the number of trophic species, links per species, the trophic similarity, and the characteristic path length. We suggest that species and link characteristics, such as predominance of scavengers and detritivorous, the relatively high connectance and the short path length are drivers in the food web structure and may play a role in the community dynamic.     

Evaluation of beach vegetation for conservation/rehabilitation in Tottori Prefecture,Southwest Japan

We examined 17 beaches in Tottori Prefecture to evaluate the current status of beach vegetation for conservation/rehabilitation purposes. We conducted a floristic survey and a beach vegetation status survey, and analyzed the beach environment using aerial photographs and a geographic information system. The health of the beach vegetation was evaluated by principal component analysis (PCA) of beach vegetation and environmental variables. PCA of beach vegetation variables, where axis 1 represented the scale of beach vegetation and axis 2 represented the quality of beach vegetation, resulted in four categories. PCA of beach environment variables, with axis 1 representing the synthetic component of grain/area/extension and axis 2 representing beach widths, also resulted in four categories. A beach vegetation list was determined from a floristic survey, and three conclusions were made based on this list. We first determined the priority of the beach plant species with regard to conservation. Based on their occurrence, the beach plants were divided into four types that were ordered with regard to their priority for beach plant conservation. We next focused on the occurrence of beach plants listed in the Red data book for Tottori Prefecture and reexamined the validity of the extinction risk category. Our conclusion was that this category needs to be updated. Finally, we compared the number of beach plant species of the survey site beaches with that of the Tottori Sand Dunes, where more natural conditions prevail. We found that only seven beaches equaled or surpassed the Tottori Sand Dunes in terms of the number of plant species. Based on these findings, we concluded that beaches other than the Tottori Sand Dunes are in need of conservation/rehabilitation.     

Spatial variation in population dynamics of the sand-burrowing amphipod Haustorioides japonicus

Densities of sandy beach organisms along coastlines often vary considerably even within small local scales. In order to understand the demographic basis of density variations in shore animals, we examined the population dynamics of the amphipod Haustorioides japonicus Kamihira, which is an ideal subject because it lacks a planktonic dispersal stage in its life history. Quantitative sampling at three adjacent sandy beaches along the Sea of Japan coast revealed that the spring density, i.e., the density of overwintered individuals, was similar among the three beaches. However, the amount of recruitment of newly hatched juveniles from late spring to summer greatly differed among the beaches, resulting in large spatial variations in the summer density. Cohort analyses revealed that at all beaches, young-of-the-year (YOY) individuals reproduced and then died, and that individuals which recruited in late summer overwintered. Moreover, growth, fecundity, and survival schedules of overwintered and YOY individuals were very similar among beaches. These observations suggest that the difference in recruitment success, i.e., the survival of newly hatched juveniles, is the primary cause of the inter-population variation in the summer density. It is likely that variations in salinity and sandy sediment affect the recruitment success of newly hatched individuals. This study indicates that biological productivity may differ substantially even among adjacent sandy beaches, with important implications for beach management.     

Indicator microbes correlate with pathogenic bacteria, yeasts and helminthes in sand at a subtropical recreational beach site

Aims: Research into the relationship between pathogens, faecal indicator microbes and environmental factors in beach sand has been limited, yet vital to the understanding of the microbial relationship between sand and the water column and to the improvement of criteria for better human health protection at beaches. The objectives of this study were to evaluate the presence and distribution of pathogens in various zones of beach sand (subtidal, intertidal and supratidal) and to assess their relationship with environmental parameters and indicator microbes at a non‐point source subtropical marine beach. Methods and Results: In this exploratory study in subtropical Miami (Florida, USA), beach sand samples were collected and analysed over the course of 6 days for several pathogens, microbial source tracking markers and indicator microbes. An inverse correlation between moisture content and most indicator microbes was found. Significant associations were identified between some indicator microbes and pathogens (such as nematode larvae and yeasts in the genus Candida), which are from classes of microbes that are rarely evaluated in the context of recreational beach use. Conclusions: Results indicate that indicator microbes may predict the presence of some of the pathogens, in particular helminthes, yeasts and the bacterial pathogen Staphylococcus aureus including methicillin‐resistant forms. Indicator microbes may thus be useful for monitoring beach sand and water quality at non‐point source beaches. Significance and Impact of the Study: The presence of both indicator microbes and pathogens in beach sand provides one possible explanation for human health effects reported at non‐point sources beaches.     

Issues to Be Considered in Counting Burrows as a Measure of Atlantic Ghost Crab Populations,an Important Bioindicator of Sandy Beaches

The use of indirect estimates of ghost-crab populations to assess beach disturbance has several advantages, including non-destructiveness, ease and low cost, although this strategy may add some degree of noise to estimates of population parameters. Resolution of these shortcomings may allow wider use of these populations as an indicator of differences in quality among beaches. This study analyzed to what extent the number of crab burrows may diverge from the number of animals, considering beach morphology, burrow depth and signs of occupation as contributing factors or indicators of a higher or lower occupation rate. We estimated the occupation rate of crabs in burrows on nine low-use beaches, which were previously categorized as dissipative, intermediate or reflexive. Three random 2-m-wide transects were laid perpendicular to the shoreline, where burrows were counted and excavated to search for crabs. The depth and signs of recent activity around the burrows were also recorded. The occupation rate differed on the different beaches, but morphodynamics was not identified as a grouping factor. A considerable number of burrows that lacked signs of recent activity proved to be occupied, and the proportions of these burrows also differed among beaches. Virtually all burrows less than 10 cm deep were unoccupied; the occupation rate tended to increase gradually to a burrow depth of 20–35 cm. Other methods (water, smoke, and traps) were applied to measure the effectiveness of excavating as a method for burrow counts. Traps and excavation proved to be the best methods. These observations illustrate the possible degree of unreliability of comparisons of beaches based on indirect measures. Combining burrow depth assessment with surrounding signs of occupation proved to be a useful tool to minimize biases.     

Fine-scale differentiation between sockeye salmon ecotypes and the effect of phenotype on straying

A long-standing goal of evolutionary biology is to understand the factors that drive population divergence, local adaptation and speciation. In particular, the effect of selection against dispersers on gene flow and local adaptation has attracted interest, although empirical data on phenotypic characters of dispersers are scarce. Here, we used genetic and phenotypic data from beach and creek ecotypes of sockeye salmon (Oncorhynchus nerka) in Little Togiak Lake, Alaska, to examine the relationship between gene flow and phenotypic and genetic differentiation. Despite close geographic proximity, both genetic and phenotypic differentiation between beach and creek fish was high and significant in all sampling years, with beach males having deeper bodies than creek males. Strays, or fish that did not return to their natal sites to spawn as determined by genetic assignment, tended to morphologically resemble the fish in the population that they joined. Male strays from beaches to creeks were shallower bodied than other beach fish, and male strays from creeks to beaches were deeper bodied than other creek males. Our results indicated that selection against strays may be moderated by the strays' phenotypic similarity to individuals in the recipient populations, but comparison of assignment results with long-term estimates of gene flow from F(ST) still suggested that strays had low reproductive success.     

Climate,landscape and microenvironment interact to determine plant composition in naturally discrete gravel beach communities

Question: What is the relative importance of national‐, regional‐ and within‐beach‐scale influences on vegetation composition and floristic affinities of New Zealand gravel beaches? Location: Coastal New Zealand. Methods: We sampled vegetation composition at 61 gravel beaches, quantifying site factors and adjacent landscape characteristics. Site, climate and geographic relationships between gravel beaches and related ecosystems were inferred using GIS data layers. To simultaneously investigate influences at different spatial scales, we used ordination and variation partitioning to examine relationships between composition and environment, and hierarchical models to understand floristic affinities with related ecosystems. Results: At a national scale, compositional variation among beaches reflects mean annual temperature and spring vapour pressure deficit; within regions, proximity of native woody vegetation and coastal turfs are important; within‐beach variation is related to substrate stability and particle size distribution. The gravel beach flora is 50% exotic, reflecting the highly modified nearby landscapes; 30% of species are characteristic of coastal sands, 20% of braided riverbeds and 8% of coastal turfs. Affinities with coastal sand communities are unrelated to microsite sandiness or area of sand dunes within 50 km. Affinities with braided riverbeds are related to the bed area of those rivers draining within 200 km and proportion of gravel in the substrate. Affinities with coastal turfs are related to proximity to the nearest turf and the proportion of humus in the substrate. Conclusions: Examining multiple scales of influence in a landscape context is essential to understand composition of naturally discrete ecosystems that span wide geographic ranges and to underpin their conservation management.     

Talitrid orientation as bioindicator of shoreline stability: Protected headland-bays versus exposed extended beaches

The behaviour of talitrids, being a local adaptation to beaches, is known to be related to environmental stability. The use of behavioural responses of resident populations as bioindicator of shoreline stability has been tested under various conditions, including after soft and hard engineering actions to stabilise eroded beaches. Port structures likely have impact on sediment longshore transportation and shoreline stability. The question was whether talitrid orientation behaviour could be proposed as bioindicator of impacts also for sandy bays of limited extension and highly used for recreation, such as those in the vicinity of touristic port structures. Orientation experiments were carried out on a set of sandy beaches of different extension and morphology, each of them in the vicinity of a touristic port, across the Mediterranean coasts. The protocol included field orientation tests of populations of talitrids, then analysed in terms of orientation precision seawards (considering sun compass orientation as the most locally adapted behavioural mechanism) in different seasons (before and after the touristic season) and times of day. The populations from more protected (either naturally or artificially) headland-bays showed a higher precision of orientation with respect to the shoreline direction than those from extended beaches, more subject to changes in longshore sedimentary transport as consequence of natural and human activities. The distance from the port and touristic pressure had no influence on talitrid orientation. An important stabilising factor for the sandy beach ecosystems, including talitrid populations and their behavioural adaptation, appeared to be the presence of seagrass banquette. The behavioural data point out that biotic information proceeding from local animal populations linked to beach sediments may complement sedimentology data and allow scaling the impacts occurring on a developed coastline. This becomes particularly relevant when considering interdisciplinary approaches to monitoring strategies.     

Climate‐change impacts on sandy‐beach biota: crossing a line in the sand

Sandy ocean beaches are iconic assets that provide irreplaceable ecosystem services to society. Despite their great socioeconomic importance, beaches as ecosystems are severely under‐represented in the literature on climate‐change ecology. Here, we redress this imbalance by examining whether beach biota have been observed to respond to recent climate change in ways that are consistent with expectations under climate change. We base our assessments on evidence coming from case studies on beach invertebrates in South America and on sea turtles globally. Surprisingly, we find that observational evidence for climate‐change responses in beach biota is more convincing for invertebrates than for highly charismatic turtles. This asymmetry is paradoxical given the better theoretical understanding of the mechanisms by which turtles are likely to respond to changes in climate. Regardless of this disparity, knowledge of the unique attributes of beach systems can complement our detection of climate‐change impacts on sandy‐shore invertebrates to add rigor to studies of climate‐change ecology for sandy beaches. To this end, we combine theory from beach ecology and climate‐change ecology to put forward a suite of predictive hypotheses regarding climate impacts on beaches and to suggest ways that these can be tested. Addressing these hypotheses could significantly advance both beach and climate‐change ecology, thereby progressing understanding of how future climate change will impact coastal ecosystems more generally.     

Mortalities caused by off-road vehicles (ORVs) to a key member of sandy beach assemblages,the surf clam <Emphasis Type="Italic">Donax deltoides</Emphasis>

Sandy beaches are prime recreational areas, but human use of beaches is not without ecological consequences. Driving of off-road vehicles on beaches for recreational pursuits is perhaps the physically most severe form of direct anthropogenic disturbance on sandy shores. Potential management and conservation interventions lack, however, data on how sensitive beach species are to vehicle impacts. We therefore experimentally quantified the link between beach traffic and lethal damages caused by vehicles to sandy shore invertebrates, using surf clams (Donax deltoides) as the biological response variable. Although clams had some tolerance against vehicles at low traffic volumes (5 vehicle passes), more than half of them were killed at higher traffic volumes (75 passes) in situations where cars traversed soft sand and turned across the beach face. Overall, both traffic volume and driver behaviour (i.e. straight vs. turning vehicle tracks) determined the incidence of direct crushing of clams under vehicles. Our data demonstrate that recreational use of ORVs is a source of mortality for beach invertebrates, but equally caution against extrapolating impact data from hard-shelled clams to potentially more sensitive soft-bodied species. Robust management interventions that seek to mitigate ecological damage from beach traffic will therefore require information on the functional relationship between the form, intensity and frequency of human disturbance and the biological responses for entire faunal assemblages on sandy shores. Handling editor: T. P. Crowe