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.     

Burrowing abilities and swash behavior of three crabs, Emerita analoga Stimpson, Blepharipoda occidentalis Randall, and Lepidopa californica Efford (Anomura, Hippoidea), of exposed sandy beaches

To investigate factors related to the distribution of intertidal species, and specific predictions of the swash exclusion hypothesis for exposed sandy beaches, we compared the burrowing abilities and swash behavior of three species of anomuran crabs in the superfamily Hippoidea (Emerita analoga, Blepharipoda occidentalis and Lepidopa californica) which commonly inhabit the intertidal and shallow subtidal zones of beaches along the California coast. Burrowing times in the laboratory increased significantly with crab size for all species in five sediment grain sizes ranging from fine sand to gravel (0.15 to 3.24 mm). For each species, burrowing times differed significantly among sand grain sizes, ranging from 0.3 to 21.5 s. Burrowing times for the hippid crab, E. analoga, were relatively constant across sediment types, while those of the albuneid crabs, B. occidentalis and L. californica, were rapid in fine to medium sands, and much slower in coarser sediments. Our results indicate that E. analoga is a substrate generalist while L. californica and B. occidentalis are substrate sensitive. Pre-burrowing times and behavior, distance moved, and burrowing times differed among the species in the swash zone. Combined times of preburrowing and burrowing were shorter than the swash period (6 s) for most E. analoga individuals. Fifty percent of the individuals of L. californica reached the substrate and burrowed in the swash period, while no individuals of B. occidentalis burrowed in that time. Pre-burrowing behavior and time may be valuable in explaining spatial and temporal patterns in the distribution of hippoid crabs on California beaches. Our results support predictions of the swash exclusion hypothesis concerning the burrowing and locomotory abilities of sandy beach macrofauna. The substrate generalist characteristics, and unique orientation and swimming abilities of the hippid crab, E. analoga, in intertidal swash may help explain the success of this species and its congeners, and have important implications for understanding patterns of macrofauna community structure on exposed sandy beaches in California and other regions.     

Swimming ability and burrowing time of two cirolanid isopods from different levels of exposed sandy beaches

Most of the macroinfauna from sandy beaches is highly mobile, emerging out of the sediment when the tide rises, and using the swash to migrate up and down the beach face or feed (searching for prey or carrion). After swash excursions, they usually burrow back into the sediment, maintaining zonation at low tide. Therefore, the different species abilities to emerge, move around and burrow under different swash climates and sediment conditions are expected to influence observed distribution patterns. Nonetheless, few attempts have been made to understand behavioral mechanisms of these organisms in moving fluids.In this study, we used a flume tunnel to investigate the orientation, swimming ability and burrowing time of two similar species of cirolanid isopods, Excirolana armata Dana and Excirolana braziliensis Richardson, under current velocities ranging from 5 to 30 cm·s⁻¹. E. armata inhabits middle levels of dissipative to intermediate beaches, while E. braziliensis is found towards the upper level of a wider range of beach states. Both species oriented countercurrent above a threshold velocity, which turned out to be significantly lower for E. armata than for E. braziliensis. E. armata proved to be a stronger swimmer as shown by the higher velocities surmounted, and the less drags experienced at the highest current velocity. E. armata also burrowed faster than E. braziliensis. Burrowing time was affected by sediment grain size and water content, but not by water flow. Once organisms managed to begin burrowing under different flow conditions, they were not affected by current velocity. Nonsaturated sand precluded burial, while coarse sand retarded it. Differences in the observed patterns of across-beach distribution may thus be the result of species-specific behavioral responses to swash climate, manifested in swimming ability, burying and orientation in directional flows.     

Non-Linear Interactions Determine the Impact of Sea-Level Rise on Estuarine Benthic Biodiversity and Ecosystem Processes

Sea-level rise induced by climate change may have significant impacts on the ecosystem functions and ecosystem services provided by intertidal sediment ecosystems. Accelerated sea-level rise is expected to lead to steeper beach slopes, coarser particle sizes and increased wave exposure, with consequent impacts on intertidal ecosystems. We examined the relationships between abundance, biomass, and community metabolism of benthic fauna with beach slope, particle size and exposure, using samples across a range of conditions from three different locations in the UK, to determine the significance of sediment particle size beach slope and wave exposure in affecting benthic fauna and ecosystem function in different ecological contexts. Our results show that abundance, biomass and oxygen consumption of intertidal macrofauna and meiofauna are affected significantly by interactions among sediment particle size, beach slope and wave exposure. For macrofauna on less sloping beaches, the effect of these physical constraints is mediated by the local context, although for meiofauna and for macrofauna on intermediate and steeper beaches, the effects of physical constraints dominate. Steeper beach slopes, coarser particle sizes and increased wave exposure generally result in decreases in abundance, biomass and oxygen consumption, but these relationships are complex and non-linear. Sea-level rise is likely to lead to changes in ecosystem structure with generally negative impacts on ecosystem functions and ecosystem services. However, the impacts of sea-level rise will also be affected by local ecological context, especially for less sloping beaches.     

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.     

Foraging activity of the sand beach isopod Scyphax ornatus Dana

Abstract

The terrestrial isopod Scyphax ornatus lives on exposed sand beaches. Adult isopods spent the daytime in burrows near the high water mark and made nightly foraging excursions over the uncovered middle beach. Hourly records of their night time distribution showed that Scyphax congregated near the water’s edge during the last 4 h of flood tide, where they fed on carrion. Insects (mostly drowned honey bees) were the main food item, but coelenterates, amphipods, crabs, and goose barnacles were also eaten. During ebb and early flood tides Scyphax occurred at low densities over the uncovered middle beach. Analysis of the distribution of large and small food items on the beach showed that by feeding during the flood tide, Scyphax exploited a rich concentration of carrion that accumulated in the swash zone and that was moved slowly up the beach by the incoming tide.     

Spatial and Temporal Variation in Enterococcal Abundance and Its Relationship to the Microbial Community in Hawaii Beach Sand and Water

Recent studies have reported high levels of fecal indicator enterococci in marine beach sand. This study aimed to determine the spatial and temporal variation of enterococcal abundance and to evaluate its relationships with microbial community parameters in Hawaii beach sand and water. Sampling at 23 beaches on the Island of Oahu detected higher levels of enterococci in beach foreshore sand than in beach water on a mass unit basis. Subsequent 8-week consecutive samplings at two selected beaches (Waialae and Kualoa) consistently detected significantly higher levels of enterococci in backshore sand than in foreshore/nearshore sand and beach water. Comparison between the abundance of enterococci and the microbial communities showed that enterococci correlated significantly with total Vibrio in all beach zones but less significantly with total bacterial density and Escherichia coli. Samples from the different zones of Waialae beach were sequenced by 16S rRNA gene pyrosequencing to determine the microbial community structure and diversity. The backshore sand had a significantly more diverse community and contained different major bacterial populations than the other beach zones, which corresponded to the spatial distribution pattern of enterococcal abundance. Taken together, multiple lines of evidence support the possibility of enterococci as autochthonous members of the microbial community in Hawaii beach sand.     

Simple estimate of filtration rates on a sandy beach

A new, simple and inexpensive method is proposed to determine the rate of water filtration in the swash zone of lakeshore beaches. Rates of water filtration on a sandy beach were determined by measuring the residue of a tracer injected into several cylinders buried vertically in the sandy layer at various locations from the shoreline to the tip of the swash zone. This methodology enabled the separate estimation of mean infiltration and exfiltration rates. The results obtained for seashore beaches were in accordance with previous data obtained by more conventional methods. The filtration rate results for lakeshore sandy beaches were almost equivalent to seashore sandy beaches, despite a shorter swash length. This paper indicates that this is caused by a larger grain size and shorter wave period. This method presents an easy approach to determine rates of water filtration in sandy beaches and is expected to further promote studies in this field.     

Fish community structure and food chain dynamics in the surf-zone of sandy beaches: The role of detached macrophyte detritus

The surf-zones of sandy beaches near Perth, Western Australia often harbour huge accumulations of detached macrophyte detritus. During 2.5 yr sampling, 29 species of fishes were captured over two sandy beaches in this region and the fish community was dominated by juveniles. There was a highly significant positive relationship between the number of fishes and the quantity of detached macrophytes taken in each surf-zone netting. Comparisons of total fish abundance on beaches with and without surf-zone accumulations of detached plants, showed that fishes were two to 10 times more abundant on the beach with weed accumulations, depending on the time of day, and date of sampling. However, despite the overall lower abundance of fishes on the open sandy beach, there was a significant increase in the number of fishes captured over the sandy beach at night. There were also two to five times the number of species over the beach with weed during the day, as opposed to equal numbers of species at night. Seven fish species made up >95% of the total catch and these species fell into two groups with regard to diurnal distribution patterns; those that were equally abundant in weed dominated or open surf-zones, and those that were weed-associated. Analyses of the diets of these fishes and the daytime distribution of an important avian piscivore in the surf-zone suggested that the large quantities of weed in the surf-zone of sandy beaches in this region provide both a rich feeding site for fishes, as well as a refuge from diurnal predators. At night, when visual feeding predators are absent, some fish species move to open sandy areas to feed. Because the majority of fishes in this surf-zone community feed on weed-associated prey, and the input of macrophyte detritus is the major source of primary production in the surf-zone, we argue that the food chain dynamics in the surf-zone in this region are fundamentally different to those of sandy beaches that have been studied previously.     

Habitat use by southern right whales,Eubalaena australis (Desmoulins, 1822), in their main northernmost calving area in the western South Atlantic

The subtropical and temperate coastal waters of the western South Atlantic are an important calving ground for southern right whales, Eubalaena australis. From 2002 to 2008, data on right whale distribution and habitat characteristics were collected in 14 bays along the coastline of Santa Catarina State, Brazil. Generalized linear models with a negative binomial error distribution were used to determine which environmental (beach morphotype, bay mouth width, bay inclination angle, north‐south and east‐west wind components), and temporal (month and year) variables best explained the aggregation pattern of individuals. Our results suggested that both cow‐calf pairs and adults unaccompanied by calves prefer bays with dissipative beaches, and that cow‐calf pairs apparently avoid bays facing southeast during days of strong east‐west winds. The number of sightings peaked in September and tended to increase over the study period. One particular embayment (Ribanceira beach) had considerably higher numbers of animals and may be considered a preferred spot in this calving ground. Our findings contribute to a better understanding of the species' habitat use and ecological requirements and should be taken into account if new management measures are implemented to further increase protection of southern right whales in the region.     

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.     

Horizontal and vertical distribution of meiofauna on sandy beaches of the North Sea (The Netherlands, Belgium, France)

Sandy intertidal zones were analysed for the presence of meiofauna. The material was collected on six macro-tidal sandy beaches along the North Sea (The Netherlands, France, Belgium), in order to analyse the vertical and horizontal meiofaunal distribution patterns. Eleven higher meiofauna taxa (one represented by larval stage—Copepoda nauplii) were recorded. The maximum total meiofauna abundance was observed on the Dutch beach (4,295±911 ind. 10 cm⁻²) in the Westerschelde estuary, while the lowest values (361±128 ind. 10 cm⁻²) were recorded in France at the Audresselles beach. Meiofauna of the different localities consisted mainly of nematodes, harpacticoids and turbellarians. Nematodes numerically dominated all sampled stations, comprising more than 45% of the total meiofauna density. Meiofauna was mainly concentrated at the sand surface with about 70% present in the uppermost 5 cm. Meiofauna occurred across the entire intertidal zone. A clear zonation pattern in the distribution of meiofauna taxa across the beaches was observed. The present work suggests that designation of exposed sandy beaches as physically controlled (McLachlan 1988) does not explain their biological variability.     

Crustaceans as ecological indicators of metropolitan sandy beaches health

We have investigated how indices of beach health perform in predicting the abundances of the crustaceans Emerita brasiliensis and Atlantorchestoidea brasiliensis from 22 metropolitan beaches in the cities of Rio de Janeiro and Niterói. Urbanization, Recreation and Conservation indices were used to assess sandy beaches health. Grain size and beach slope were used as morphodynamics indicators. Diagram from the principal component analysis clearly separated beaches with different urbanization and conservation levels. Generalized additive models (GAM's) were adjusted for species abundance using the indices and morphodynamic parameters as explanatory variables. Lower abundances were predicted for beaches with high levels of urbanization, whereas predictions of higher abundances occurred on beaches with high conservation levels. Using theoretic inference we showed that the urbanization index was the most important predictor for abundance of A. brasiliensis and the conservation index was the most important predictor for E. brasiliensis, reflecting different responses by upper tidal and intertidal species. A. brasiliensis occupies the intermediate and upper beach zones and E. brasiliensis is a swash zone filter-feeder that is more abundant in pristine beaches. Both species are highly subject to the impact of bathers and coastal modification. Unexpected, the recreation index did not show a negative effect on abundance predictions. Urbanization and conservation indices can be suitable metrics to measure anthropogenic effects on macrobenthic species. Moreover, mole crabs and sandhoppers species can be easily monitored. Coastal urbanization is a global phenomenon and we used the diagram of urbanization and conservation levels to expose possible directions for management strategies of metropolitan sandy beaches.     

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.     

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.     

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.     

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.     

Latitudinal patterns in abundance and life-history traits of the mole crab Emerita brasiliensis on South American sandy beaches

Demographic and life‐history attributes of the mole crab Emerita brasiliensis were analysed along 2700 km of the Atlantic coast of South America, including sandy beaches at the southernmost limit (Uruguay) and at the core of its geographical range (Brazil). Population features varied markedly within this range and exhibited systematic geographical patterns of variation. Abundance significantly increased from temperate to subtropical beaches, and the same held true for the asymptotic weight of males. Conversely, length at maturity and asymptotic weight of females increased from subtropical to temperate beaches, being inversely related to sea water temperature. Macroecological patterns in abundance and body weight showed the first large‐scale evidence of scaling of population density to body size for a sandy beach population. Mortality rates (both sexes) followed a nonlinear increase from low‐density temperate beaches to high‐density subtropical beaches. The effect of habitat quality and availability could explain discontinuities in the species distribution within its range, and also differential responses in life‐history attributes at a local scale. Asymmetries and converse latitudinal trends between sexes suggest that there is not a single general factor determining large‐scale patterns in life‐history traits of this species. Our results reinforce the view that density‐dependent and environmental factors operating together regulate sandy beach populations. The need to develop macroecological studies in sandy beach ecology is highlighted, as knowledge acquired from local to large spatial scales throws light on population structure and regulation mechanisms.     

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.