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

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

A comparison of biomonitoring methodologies for surf zone fish communities.

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

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

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

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

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

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

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

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

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

The environmental cost of subsistence: Optimizing diets to minimize footprints.

The question of how to minimize monetary cost while meeting basic nutrient requirements (a subsistence diet) was posed by George Stigler in 1945. The problem, known as Stigler's diet problem, was famously solved using the simplex algorithm. Today, we are not only concerned with the monetary cost of food, but also the environmental cost. Efforts to quantify environmental impacts led to the development of footprint (FP) indicators. The environmental footprints of food production span multiple dimensions, including greenhouse gas emissions (carbon footprint), nitrogen release (nitrogen footprint), water use (blue and green water footprint) and land use (land footprint), and a diet minimizing one of these impacts could result in higher impacts in another dimension. In this study based on nutritional and population data for the United States, we identify diets that minimize each of these four footprints subject to nutrient constraints. We then calculate tradeoffs by taking the composition of each footprint's minimum diet and calculating the other three footprints. We find that diets for the minimized footprints tend to be similar for the four footprints, suggesting there are generally synergies, rather than tradeoffs, among low footprint diets. Plant-based food and seafood (fish and other aquatic foods) commonly appear in minimized diets and tend to most efficiently supply macronutrients and micronutrients, respectively. Livestock products rarely appear in minimized diets, suggesting these foods tend to be less efficient from an environmental perspective, even when nutrient content is considered. The results' emphasis on seafood is complicated by the environmental impacts of aquaculture versus capture fisheries, increasing in aquaculture, and shifting compositions of aquaculture feeds. While this analysis does not make specific diet recommendations, our approach demonstrates potential environmental synergies of plant- and seafood-based diets. As a result, this study provides a useful tool for decision-makers in linking human nutrition and environmental impacts.