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.     

Distribution patterns of supralittoral arthropods: wrack deposits as a source of food and refuge on exposed sandy beaches (SW Spain)

Wrack deposits are a common feature of sandy beaches worldwide. Despite their relevance, the habitat features of wrack debris and their potential influence on the distribution of upper shore arthropods remain poorly understood. In this study, the distribution of supralittoral arthropods was analysed by sampling areas covered and not covered by wrack on two tidal levels during winter and summer from two sandy beaches of south-western Spain. Despite the differences between beaches and seasons, density, species richness and diversity were significantly higher in wrack patches than in bare sand. Moreover, community structure and composition varied between both microhabitats due to the influence of wrack deposits on the habitat selection by arthropod species. Food availability combined with temperature or moisture mostly explained the distribution pattern of assemblages. Moreover, these habitat features varied between wrack bands, the lower band being fresher and wetter than upper band. The density of the main species inhabiting wrack was correlated mainly with moisture. The present study provides evidences about the role of algal wrack structuring diversity and composition of supralittoral arthropods stressing the importance of the habitat heterogeneity to maintain abundant and diverse communities on exposed sandy beaches.     

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.     

Coupling between macroalgal inputs and nutrients outcrop in exposed sandy beaches

Fluxes of nutrients across habitats are of paramount relevance in ecological studies due to the implication in primary production, trophic structure and biodiversity. This study analyses the role of sandy beaches in the processing of organic matter. Three beaches with different macroalgal inputs were sampled throughout the annual cycle. The standing stock of wrack macroalgae on the beach surface and the nutrient concentration in the intertidal pore water (IPW) and in the surf zone water were measured monthly. Mean concentration of dissolved inorganic nitrogen and phosphate in the IPW increased from the low to the very high subsidized beach. Seasonal coupling was observed between the wrack biomass and the nutrient concentration throughout the year. Among the nutrient species, a variable relationship was found between the NH₄ ⁺/NO _x ^? ratio and the biomass of macroalgae deposited. These results provide evidences of the active role of sandy beaches in the processing of organic matter and in the nutrient cycling, remarking the feedback connectivity between sandy beaches and their neighbour ecosystems.     

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.     

Warming intensify CO2 flux and nutrient release from algal wrack subsidies on sandy beaches

Algal wrack subsidies underpin most of the food web structure of exposed sandy beaches and are responsible of important biogeochemical processes that link marine and terrestrial ecosystems. The response in decomposition of algal wrack deposits to global warming has not been studied in ocean‐exposed sandy beaches to date. With this aim, passive open top chambers (OTCs) were used to increase soil temperature within the range predicted by the IPCC for western Europe (between 0.5 and 1.5°C), following the hypothesis that the biogeochemical processing of macroalgal wrack subsidies would accelerate in response to temperature increase. The effect of temperature manipulation on three target substrates: fresh and aged macroalgae, and bare sand, was tested. Results indicated that a small warming (<0.5°C) affected the wrack decomposition process through traceable increases in soil respiration through CO₂ flux, inorganic nutrients within the interstitial environment (N and P), sediment organic contents measured through the amount of proteins and microbial pool through the total soil DNA. The different responses of soil variables in the studied substrates indicated that the decomposition stage of stranded macroalgae influences the biogeochemical processing of organic matter in sandy beaches. Thus, CO₂ fluxes, releases of organic and inorganic nutrients and microbial activity intensify in aged wrack deposits. Our results predict that expected global warming will increase the release of inorganic nutrients to the coastal ocean by 30% for the N (21 Gg/year) and 5.9% for P (14 Gg/year); that increase for the flow of C to the atmosphere as CO₂ was estimated in 8.2% (523 Gg/year). This study confirms the key role of sandy beaches in recycling ocean‐derived organic matter, highlighting their sensitivity to a changing scenario of global warming that predicts significant increases in temperature over the next few decades.     

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.     

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.     

DISPERSAL OF HORMOSIRA BANKSII (PHAEOPHYCEAE) VIA DETACHED FRAGMENTS: REPRODUCTIVE VIABILITY AND LONGEVITY1

Drifting, fertile thalli are well documented to be the primary long‐distance dispersal vector for many marine macroalgae, but little information about reproductive viability of drift is known. This study examined the reproductive viability and longevity of floating fragments of the intertidal Australasian fucoid Hormosira banksii (Turner) Decne. Beach wrack surveys and field experiments were conducted to test the model that long‐distance dispersal is achieved in H. banksii via floating, fertile fronds. High densities of beach wrack fragments were evident during summer compared to autumn. The majority of beach wrack occurred on sandy beaches rather than rocky shores. Both male and female fragments were present in the beach wrack. Detached fronds were capable of releasing gametes up to 8 weeks after detachment. Beach wrack produced high fertilization rates and recruited successfully onto artificial panels. Results suggest that detached fragments are reproductively viable and that floating, fertile fronds may be an important mechanism for facilitating long‐distance dispersal in this species. Nevertheless, the frequency of fronds reaching a suitable habitat and contributing to gene flow between populations, or colonizing new populations, may not be proportional to the total density of beach wrack.     

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.     

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.     

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.     

High Metabolic Rates in Beach Cast Communities

Metabolic hotspots at land–water interfaces are important in supporting biogeochemical processes. Here we confirm the generality of land–aquatic interfaces as biogeochemical hot spots by extending this concept to marine beach cast materials. In situ atmospheric pCO_2, from a respiration chamber (10 cm in diameter and 20 cm high) inserted into wrack deposits, was determined using a high-precision (±1 ppm) non-dispersive infrared gas analyzer (EGM-4, PP-systems) at 1 minute recording intervals. The wrack deposits supported high metabolic activities, with CO₂ fluxes averaging (±SE) 6.62 ± 0.88 μmol C m⁻² s⁻¹, compared to median value of 0.98 μmol C m⁻² s⁻¹ (mean 2.21 ± 1.25 μmol C m⁻² s⁻¹) for bare sand adjacent to deposits. Wrack metabolic rates ranged 40-fold across beaches, from a minimum of 0.57 ± 0.22 μmol C m⁻² s⁻¹ to a maximum of 20.8 ± 5.04 μmol C m⁻² s⁻¹, both derived from beaches with deposits dominated by Sargassum. Rates tended to increase significantly (F test, P < 0.05) from the shoreline to reach maximum rates at about 10 m from the shoreline, declining sharply further from the shoreline, and increased with increasing thickness of the deposits (maximum about 10 cm deep), declining for thicker deposits. Wrack differing in composition had similar metabolic rates, although deposits consisting of a mixture of seagrass and algae tended to show somewhat higher rates. Our results show a meter square of wrack deposit supports a metabolic rate equivalent to that supported by 3 m² of living seagrass or macroalgal habitat. In wrack, the marine environment provides organic material and moisture and the land environment provides oxygen to render wrack ecosystems an efficient metabolic reactor. Intense wrack metabolism should also be conducive to organismal growth by supporting the development of a cryptic, but diverse wrack-based food web.     

Impacts on biodiversity at Baltic Sea beaches

Sandy and shingle beaches were investigated in 2009 and 2010 along the coast of the Baltic Sea in northern Germany with the purpose of assessing biodiversity gradients from shingle to sandy beaches, from beach to primary dunes, and the impacts of tourism on biodiversity. On nine beach sites, ranging between 100% shingle and 99% sand without shingle, Carabidae, Staphylinidae, and Araneae were studied. Two of the six sandy beaches were open and four were closed to tourists. Additionally, trampling effects from tourists, species richness of plants, and plant cover were investigated on sixteen beaches. According to results, primary dunes showed higher species richness in carabids and spiders, but not in staphylinds. Shingle beaches exhibited lower species richness in Staphylinidae and Araneae, but not in Carabidae. As estimated by the Jackknife II method, shingle beaches were the lowest in total species richness. Trampling intensity ranged from 0 footprints m⁻² day⁻¹ on closed beaches, up to a maximum of 30 footprints m⁻² day⁻¹. On “intensively” used beaches (12 footprints m⁻² day⁻¹, on average), reduction of plant cover was more pronounced than on “extensively” used beaches (7 footprints m⁻² day⁻¹, on average). Both plant cover and plant species richness were lower on intensively and extensively used beaches than on closed beaches. In arthropods, only staphylind and spider species richness was significantly lower on open beaches than on closed beaches, but no differences were found in carabids. Referring to our results, trampling effects from tourists have high impact on species richness of sandy beaches, on both intensively and extensively used sites.     

The primary production of phytoplankton in the restored Maltañski Reservoir in Poland

Patterns in composition, abundance and diversity of the annelid fauna (Polychaeta and Oligochaeta) in 22 sandy beaches in Iceland were explored. The effect of exposure on annelid distribution was studied. A total of 5651 annelids were recorded from 160 core samples. Oligochaetes (chiefly Tubificidae) dominated the annelid assemblage whereas polychaetes represented a minor fraction. Polychaetes were relatively more abundant in exposed than in sheltered beaches, contrary to oligochaetes. Meiofaunal polychaete species were also more abundant in exposed than in sheltered beaches. Southwest beaches seemed more diverse in annelid species than northern ones. Annelid diversity did not differ between sheltered and exposed sites, but higher diversity was attained in fine sands at sheltered areas. Cluster analysis revealed large differences between beaches in the annelid community composition. The general patterns found suggest that beach exposure is a major factor conditioning macro- and meiofaunal polychaete and oligochaete distribution along the Icelandic coast.     

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     

Temporal patterns in the intertidal faunal community at the mouth of a tropical estuary

The use of intertidal sandy beaches by fish and macrocrustaceans was studied at different temporal scales at the mouth of a tropical estuary. Samples were taken along the lunar and diel cycles in the late dry and rainy seasons. Fish assemblage (number of species, density and biomass), crustaceans and wrack biomass, showed significant interactions among all studied factors, and the combination of moon phase and diel cycle, resulting in different patterns of environmental variables (depth, water temperature and dissolved oxygen), affected habitat use by the different species. Variances in faunal community were detected between seasons, stimulated by salinity fluctuations from freshwater input during the rainy season. These differences suggest an important cycling of habitats and an increase in connectivity between adjacent habitats (estuary and coastal waters). Moreover, the results showed that this intertidal sandy beach also provides an alternative nursery and protected shallow‐water area for the initial development phase of many marine and estuarine species. In addition, this intertidal habitat plays an important role in the maintenance of the ecological functioning of the estuarine–coastal ecosystem continuum.     

Recovery of sandy beach and maritime forest vegetation on Phuket Island (Thailand) after the major Indian Ocean tsunami of 2004

Question: How rapidly has the sandy beach and maritime forest vegetation on Phuket recovered and regenerated after the impact of the major Indian Ocean tsunami of 2004? What are the characteristics of sandy beach species for regenerating their populations and the invasion patterns of originally non-sandy beach species or other newcomers after the tsunami? Location: Phuket Island, southern Thailand. Methods: Species composition of beaches was studied on the same research plots 6 months before and 9 months after the tsunami. The changes in individual species cover before and after the tsunami were determined by χ² tests. Change in community composition was analysed by detrended correspondence analysis. The relationship between species and environmental factors was analysed by canonical correspondence analysis. Results: The sites disturbed by the tsunami were often invaded by annuals, especially grasses and asteraceous plants, rather than by perennials. In contrast, species with clonal growth by stolons decreased significantly. Factors determining the species habitat differences were soil hardness (penetration resistance of sandy soil), per cent silt content, soil water content and beach management. Habitat differences among originally non-sandy beach herbaceous species that expanded their population or moved to the coast after the disaster were defined by sand accretion or erosion caused by the tsunami. Many sandy beach herbaceous communities changed into Dactyloctenium aegyptium communities because of the tsunami were originally constituted by non-sandy beach D. aegyptium with Cenchrus echinatus. Although the forest floors of most maritime forests were invaded by originally non-sandy beach Tridax procumbens, Eleusine indica or D. aegyptium because of the tsunami, this did not result in a change in the vegetation unit, because species' loss was restricted to the understorey. In time, these forests will recover their previous community composition. Conclusions: Our results suggest that originally non-sandy beach native species invaded the disturbed beaches rapidly after the tsunami but their habitats differ. Sites where sand accumulated on a beach because of the tsunami were invaded by D. aegyptium and E. indica, whereas soil erosion permitted invasion by Digitania adscendens. Tridax procumbens establishes rapidly on wet sites with hard soil, high per cent silt content and low beach management pressure. Sandy beach species with subterranean long rhizomes are strongly tolerant of such disasters. We concluded that the species composition of the beaches disturbed by a temporary large disaster is determined by dormancy and growth forms, with radicoid form being influential.     

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.     

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.