Sub-lethal effects of off-road vehicles (ORVs) on surf clams on sandy beaches

Ecological impacts of vehicle traffic are a significant environmental management issue on many sandy shores. Impacts usually focus on lethal effects of vehicles to organisms, but sub-lethal effects which could reduce the fitness of macrofauna populations are equally possible but unknown. Consequently, we measured changes in body condition and burrowing performance of the beach clam Donax deltoides subjected to vehicle traffic on sandy shores in eastern Australia. Body mass index of clams on beaches open to traffic was 16% lower, but gonadosomatic index and relative valve thickness were not consistently linked to vehicle access to beaches. By contrast, off-road vehicles significantly impaired the burrowing performance of clams. After experimental exposure to ORV traffic (30 passes) and dislodgement from the sediment, the time taken for clams to re-bury into the sand doubled irrespective of the vehicle weight used. Because burrowing is such a critical function in the behavioural repertoire of sandy beach animals, the traffic-induced changes to organisms' performance found in this study may increase mortality by causing displacement to less favourable habitats by swash, and by intensifying the risk of predation and desiccation. When assessing the ecological impacts of vehicles on beach fauna, it is thus important to consider both lethal and sub-lethal effects.     

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.     

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.     

Effects of Fishing and Fishing Closures on Beach Clams: Experimental Evaluation across Commercially Fished and Non-Fished Beaches before and during Harvesting

Management responses to reconcile declining fisheries typically include closed areas and times to fishing. This study evaluated this strategy for a beach clam fishery by testing the hypothesis that changes in the densities and size compositions of clams from before to during harvesting would differ between commercially fished and non-fished beaches. Sampling was spatially stratified across the swash and dry sand habitats on each of two commercially fished and two non-fished beaches, and temporally stratified across three six-week blocks: before, early and late harvesting. Small-scale spatio-temporal variability in the densities and sizes of clams was prevalent across both habitats and the components of variation were generally greatest at the lowest levels examined. Despite this, differences in the densities and sizes of clams among individual beaches were evident, but there were few significant differences across the commercially fished versus non-fished beaches from before to during harvesting. There was no evidence of reduced densities or truncated size compositions of clams on fished compared to non-fished beaches, contrasting reports of some other organisms in protected areas. This was probably due to a combination of factors, including the current levels of commercial harvests, the movements and other local-scale responses of clams to ecological processes acting independently across individual beaches. The results identify the difficulties in detecting fishing-related impacts against inherent levels of variability in clam populations. Nevertheless, continued experimental studies that test alternate management arrangements may help refine and determine the most suitable strategies for the sustainable harvesting of beach clams, ultimately enhancing the management of sandy beaches.     

Human-induced changes in the trophic functioning of sandy beaches

The increasing anthropogenic disturbance on coastal ecosystems has threatened ecological interactions and ecosystems functioning. To investigate if human pressure affects the trophic structure of sandy beaches, mass-balanced models were applied on two Brazilian sandy beaches with distinct human impact degree. The food web models included detritus, phytoplankton, macroinvertebrates, fish and seabirds. Macroinvertebrates in non-urbanized sectors represented the highest production fraction consumed by predators. The energy transfer and the cycling indicator were more efficient in the non-urbanized sectors than urbanized ones. The results indicate that macroinvertebrates sensitive to direct human impact such as trampling are important to the trophic functioning of sandy beaches. Establishing a threshold for the number of beachgoers or dispersing recreational activities to avoid crowds may be tangible ways to mitigate the trampling impact on macroinvertebrates.     

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 (~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.     

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.     

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.     

NOTICES

Ward, D. 1990. The demography, diet and reproductive success of African Black Oystercatchers on a sandy beach. Ostrich 61:125-133.

The biology of African Black Oystercatchers Haematopus moquini on a sandy beach is described and compared with earlier studies of this species on mixed and rocky shores. Adult oystercatchers specialized on the wedge clam Donax serra. There was no difference in the size of clams taken by adult and immature oystercatchers, although chicks were fed smaller clams. The abundance of oystercatchers and the biomass of Donax serra was positively correlated. There was no significant relationship between clutch size, egg volume or clutch volume and prey biomass.     

Talitrid and Tylid crustaceans bioecology as a tool to monitor and assess sandy beaches’ ecological quality condition

In the last decades, the sharp increase of human activities on sandy beaches has introduced several detrimental impacts on these ecosystems, highlighting the importance of developing environmental quality assessments and sustainable management and protection plans for these environments. The study of key species at the population level represents an adequate and important approach to the ecosystem's ecological quality if the impacts of environmental disturbances, namely derived from human activities, are to be assessed. Talitrid and Tylid crustaceans often form abundant populations in temperate exposed sandy beaches, and might represent the bulk of the macrofaunal communities. In the present case study, the comparison of distinct Talitrid and Tylid populations on Atlantic (Portugal) and Mediterranean (Italy and Tunisia) beaches allowed to assess the adaptation of these animals on local and macro scale scenarios and evaluate the importance of bioecological studies as auxiliary tools in environmental monitoring and in the ecological quality assessment of sandy beaches. The results revealed that Talitrid and Tylid populations have a strong plasticity over geographic gradients and to local environments, presenting a high degree of variation on population ecology, namely latitudinal clines on several life history traits. Also, since the studies were performed on sandy beaches with similar degrees of exposure to wave action and low degrees of human disturbance, the present work may play a relevant role as reference knowledge in environmental monitoring studies and as an ecological quality assessment tool that might be used to evaluate the impacts of environmental disturbances on sandy beach ecosystems, namely from human origin.     

Differential effects of native and invasive algal wrack on macrofaunal assemblages inhabiting exposed sandy beaches

Many sandy beaches worldwide receive large amounts of drift seaweed, known as wrack, from offshore algal beds and closer rocky intertidal shores. Despite the important influence of algal wrack on macrofaunal assemblages from different coastal systems, relatively little attention has been paid to the macrofaunal responses in sandy beaches to macrophyte wrack supplies. Algal wrack is a key resource, i.e. for food and/or refuge, for beach invertebrates while its availability can affect diversity and abundance of intertidal animals including shorebirds, but the role of certain types of wrack and its location on the shore has not been examined experimentally to date. In this paper, we use experimental manipulation of two species of brown seaweeds, i.e. artificial wrack patches made up of the native macroalgae Saccorhiza polyschides and the invasive species Sargassum muticum, to test hypotheses about influences on macrofaunal assemblages inhabiting the drift line and supratidal levels of exposed beaches. Results pointed out that different types of wrack deposits were not used uniformly by invertebrates. Nutritional value differed between the two species of wrack. In most cases, the carbohydrates, lipids and organic carbon content were greater in patches of S. muticum than in patches of S. polyschides. Data also provided evidences that nutritional content and microclimatic conditions of wrack deposits, i.e. temperature and humidity, might affect macrofaunal assemblages.     

Helpful habitant or pernicious passenger: interactions between an infaunal bivalve,an epifaunal hydroid and three potential predators

The importance of positive interspecific interactions within physically stressful habitats has received increased attention from community ecologists. The exposed sandy beach is an example of a physically rigorous environment where biological interactions have long been considered insignificant. We examined the interaction between the infaunal clam, Donax variabilis, and the hydroid, Lovenella gracilis, on exposed sandy beaches in North Carolina. Epibiotic occupation of Donax by hydroids has been repeatedly observed on ocean beaches but rarely investigated. By providing a stable substrate for attachment, the clam facilitates the persistence of the hydroid in the intertidal beach; however, benefits or costs experienced by the host as a result of this association are unknown. By exposing clams with and without hydroid colonies to multiple types of clam predators, we tested the effectiveness of the hydroid, which possesses stinging nematocysts, in defending its host. The hydroid defended the clam against one common predator, the Florida pompano ( Trachinotus carolinus). Against speckled crabs ( Arenaeus cribrarius) and ghost crabs ( Ocypode quadrata), however, the hydroid offered no protection for its host and instead facilitated predation. The epibiotic hydroid, which projects above the surface of the sand, allowed the crabs to more readily detect clams below the surface. In the field, we evaluated the effect of the hydroid on the tidally synchronized migrations and burrowing speed of the clam. The hydroid, which can form large colonies on the posterior end of the clam, had no effect on Donax burrowing speed, but did reduce the speed of transport of clams by wave swash. Depending on relative predation pressure, the occupation of D. variabilis by L. gracilis can alternately be characterized as beneficial or detrimental to the host.     

Genetic structure as a response to anthropogenic and extreme weather disturbances of a coastal dune dwelling spider,Arctosa sanctaerosae

The continued increase in the number of tourists visiting the Northern Gulf Coast (NGC), USA, in the last century, and the resulting sprawl of large cities along the coast, has degraded and fragmented the available habitat of Arctosa sanctaerosae, a wolf spider endemic to the secondary dunes of the white sandy beaches of the NGC. In addition to anthropogenic disturbance to this coastal region, hurricanes are an additional and natural perturbation to the ecosystem. The data presented here explore the status of populations of this species spanning the entire known range and the factors influencing population demography including anthropogenic disturbance and severe tropical storms. Using microsatellite markers, we were able to document the genetic structure of A. sanctaerosae, including current and historic patterns of migration. These results combined with ecological and census data reveal the characteristics that have influenced population persistence: ecological variables affecting the recovery of the population clusters after severe tropical storms, genetic fragmentation due to anthropogenic disturbance, and their interaction. These findings demonstrate the significance that the high traffic beach communities of the NGC and their impact on the once intact contiguous dune ecosystem have on recovery after severe tropical storms. Contemporary modeling methods that compare current and historic levels of gene flow suggest A. sanctaerosae has experienced a single, contiguous population subdivision, and the isolates reduced in size since the onset of commercial development of the NGC. These results point to the need for monitoring of the species and increased protection for this endangered habitat.     

Response of macroalgae and macroinvertebrates to anthropogenic disturbance gradients in rocky shores

The compliance of macroalgal and macroinvertebrate assemblages to anthropogenic disturbance gradients (e.g., nutrient enrichment) was investigated at intertidal rocky shores. Macroalgae and macroinvertebrates presented parallel behavior, both showing shifts in the communities’ structural variation along the gradients, in which an higher number of opportunistic species (and higher abundances) were found in more stressful sites (close to the disturbance source), in contrast to less disturbed sites (far from the disturbance source), which showed higher presence of more sensitive species (and higher abundance of several of them).The macroinvertebrate abundance and taxonomic composition, which are parameters required by the Water Framework Directive (WFD) to be included in tools for the ecological quality status assessment, responded to the disturbance gradient. Results suggest that the macroinvertebrate biological element might be considered an indicator of disturbance in intertidal rocky shores as good as the macroalgae, and therefore the development of a specific methodology based solely on benthic macroinvertebrates of rocky shores, presently a gap in the ecological quality status assessment for the WFD, seems feasible.     

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.     

Complex, dynamic combination of physical, chemical and nutritional variables controls spatio-temporal variation of sandy beach community structure

Sandy beach ecological theory states that physical features of the beach control macrobenthic community structure on all but the most dissipative beaches. However, few studies have simultaneously evaluated the relative importance of physical, chemical and biological factors as potential explanatory variables for meso-scale spatio-temporal patterns of intertidal community structure in these systems. Here, we investigate macroinfaunal community structure of a micro-tidal sandy beach that is located on an oligotrophic subtropical coast and is influenced by seasonal estuarine input. We repeatedly sampled biological and environmental variables at a series of beach transects arranged at increasing distances from the estuary mouth. Sampling took place over a period of five months, corresponding with the transition between the dry and wet season. This allowed assessment of biological-physical relationships across chemical and nutritional gradients associated with a range of estuarine inputs. Physical, chemical, and biological response variables, as well as measures of community structure, showed significant spatio-temporal patterns. In general, bivariate relationships between biological and environmental variables were rare and weak. However, multivariate correlation approaches identified a variety of environmental variables (i.e., sampling session, the C∶N ratio of particulate organic matter, dissolved inorganic nutrient concentrations, various size fractions of photopigment concentrations, salinity and, to a lesser extent, beach width and sediment kurtosis) that either alone or combined provided significant explanatory power for spatio-temporal patterns of macroinfaunal community structure. Overall, these results showed that the macrobenthic community on Mtunzini Beach was not structured primarily by physical factors, but instead by a complex and dynamic blend of nutritional, chemical and physical drivers. This emphasises the need to recognise ocean-exposed sandy beaches as functional ecosystems in their own right.     

Humans alter habitat selection of birds on ocean‐exposed sandy beaches

Aim Resource‐selection functions (RSFs) can quantify and predict the density of animal populations across heterogeneous landscapes and are important conservation tools in areas subject to human disturbance. Sandy beach ecosystems have comparatively low habitat heterogeneity and structural relief in the intertidal zone, but intense human use. We aimed to develop predictive RSFs for birds on ocean‐exposed sandy beaches at two spatial scales, 25 ha (local scale) and 250 ha (landscape scale), and to test whether habitat selection of birds that commonly use the surf–beach–dune interface is influenced by the rates of human activities. Location Moreton and North Stradbroke Island, eastern Australia. Methods Avifauna and human activities were mapped on three sandy beaches covering 79 km of coastline for 15 months. Habitat characteristics of the surf–beach–dune interface were derived from remote sensing and ground surveys. RSFs were developed for 12 species of birds at two spatial scales: 25 ha (local scale) and 250 ha (landscape scale). Results  At local (25 ha) and landscape scales (250 ha), dune dimensions and the extent and type of vegetation structure were important predictors of bird density. Adding the frequency of human activities improved the predictive power of RSFs, suggesting that habitat selection of birds on beaches is modified by human use of these environments. Human activities occurred mostly in the mid‐ to lower intertidal zone of the beach, overlapping closely with the preferred habitats of Silver Gulls (Larus novaehollandiae), Pied Oystercatchers (Haematopus longirostris), Red‐capped Plovers (Charadrius ruficapillus) and endangered Little Terns (Sternula albifrons). Main conclusions In addition to demonstrating the appropriateness of RSFs to the surf–beach–dune interface, our results stress the need for systematic conservation planning for these ecosystems, where ecological values have traditionally been subsidiary to the maintenance of sand budgets and erosion control.     

Bottom‐up control in the benthos of ocean‐exposed sandy beaches?

Bottom‐up control is a fundamental structuring force in food webs. Food webs of ocean‐exposed sandy beaches are predicted to be bottom‐up controlled systems, underpinned by imported organic matter rather than in situ primary production. This ecological model of resource‐based regulation of biological assemblages is juxtaposed against a prevailing paradigm built around a dominance of physical drivers in sandy beach ecosystems. Surprisingly, given the apparently ubiquitous energetic subsidies of beach food webs, the central premise of bottom‐up control has not been tested. Here we experimentally manipulated in situ nutrient levels on a sandy beach to test food web responses at the levels of primary producers (benthic microalgae) and their grazers (meiofauna). The meiofauna community as a whole appeared most strongly influenced by the local physical environment, particularly changes in sediment grain size – this supports the traditional ‘environmental control paradigm’. We also detected a significant, positive response of two consumer groups of the meiofauna (nematodes, ostracods) to nutrient enrichment that supports a model of biological, bottom‐up control. Although the predicted response of elevated producer biomass following nutrient enrichment was not detected, intense grazing pressure on new, stimulated production may have masked positive responses by the primary producers. Multichannel regulation of food webs is likely for many exposed sandy beaches, albeit an often lower importance of in situ bottom‐up forces compared with stronger environmental control.     

Enhancement of habitat heterogeneity and species richness on rocky shores inundated by sand

Summary Many rocky shores are subject to periodic inundation by sand, which is often thought to reduce species richness by eliminating organisms intolerant of sand scour or sand smothering. However, regular disturbance (e.g. inundation) should promote richness by preventing the development of low diversity climax communities. A study of faunal richness on 10 regularly inundated shores showed that inundation does promote richness, but by increasing habitat heterogeneity. Some species are excluded from parts of the shore by sand, but because of the patchiness of sand deposits they are rarely excluded from the entire shore. Other species are found only on rocks associated with sand, while typically sandy shore animals occur in the sand deposits themselves. Total richness (281 species) was greater than for local noninundated shores and sandy beaches combined.     

In depth review of the ecology of arenicolous marine fungi

As an ecotone, sandy beaches exist within a multi-dimensional mesh of environmental gradients, shaped by numerous parameters (e.g. temperature, humidity, wave action, sand particle size and salinity). These limit the proliferation of a narrow group of fungal species. Obligate arenicolous marine fungi are an ecological assemblage of sand-associated heterotrophs that inhabit sandy beaches. These organisms have evolved to cope with dynamic beach conditions, having a cosmopolitan distribution across tropical, subtropical and temperate regions. Herein we provide an overview of published works relating to the fungi of sandy beaches, focusing on the past half-century. We outline a broad range of topics in ecology including fungal adaptive traits to intertidal conditions at the morphological and genetic levels, temporal and spatial patterns in community structure, and species variations in substrate preference. Collectively, these concepts should encourage marine mycologists to embrace a holistic set of perspectives to shape the outlook for beach ecology.