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.

The seagrass holobiont: understanding seagrass-bacteria interactions and their role in seagrass ecosystem functioning.

This review shows that the presence of seagrass microbial community is critical for the development of seagrasses; from seed germination, through to phytohormone production and enhanced nutrient availability, and defence against pathogens and saprophytes. The tight seagrass-bacterial relationship highlighted in this review supports the existence of a seagrass holobiont and adds to the growing evidence for the importance of marine eukaryotic microorganisms in sustaining vital ecosystems. Incorporating a micro-scale view on seagrass ecosystems substantially expands our understanding of ecosystem functioning and may have significant implications for future seagrass management and mitigation against human disturbance.

Microorganisms facilitate uptake of dissolved organic nitrogen by seagrass leaves.

Microorganisms play a critical role in nitrogen cycling by mineralising dissolved organic nitrogen (DON) compounds into bioavailable inorganic forms (DIN). Although DIN is crucial for seagrass growth, the hypothesis that seagrass leaf associated-microorganisms could convert DON to forms available for plant uptake has never been tested. We conducted a laboratory-based experiment in which seagrass (Posidonia sinuosa) leaves were incubated with 15N-amino acids (aa), with and without associated microorganisms. Samples were collected after 0.5, 2, 6 and 12 h. Both bulk stable isotope and nanoscale secondary ion mass spectrometry (NanoSIMS) analysis showed high accumulation of 15N within seagrass leaf tissues with an associated microbiota, but not in plants devoid of microorganisms. These results significantly change our understanding of the mechanisms of seagrass nitrogen use and provide evidence that seagrass microbiota increase nitrogen availability for uptake by seagrass leaves by mineralising aa, thus enhancing growth and productivity of these important coastal ecosystems.

Coastal connectivity and spatial subsidy from a microbial perspective.

The transfer of organic material from one coastal environment to another can increase production in recipient habitats in a process known as spatial subsidy. Microorganisms drive the generation, transformation, and uptake of organic material in shallow coastal environments, but their significance in connecting coastal habitats through spatial subsidies has received limited attention. We address this by presenting a conceptual model of coastal connectivity that focuses on the flow of microbially mediated organic material in key coastal habitats. Our model suggests that it is not the difference in generation rates of organic material between coastal habitats but the amount of organic material assimilated into microbial biomass and respiration that determines the amount of material that can be exported from one coastal environment to another. Further, the flow of organic material across coastal habitats is sensitive to environmental change as this can alter microbial remineralization and respiration rates. Our model highlights microorganisms as an integral part of coastal connectivity and emphasizes the importance of including a microbial perspective in coastal connectivity studies.

Accelerating Tropicalization and the Transformation of Temperate Seagrass Meadows.

Climate-driven changes are altering production and functioning of biotic assemblages in terrestrial and aquatic environments. In temperate coastal waters, rising sea temperatures, warm water anomalies and poleward shifts in the distribution of tropical herbivores have had a detrimental effect on algal forests. We develop generalized scenarios of this form of tropicalization and its potential effects on the structure and functioning of globally significant and threatened seagrass ecosystems, through poleward shifts in tropical seagrasses and herbivores. Initially, we expect tropical herbivorous fishes to establish in temperate seagrass meadows, followed later by megafauna. Tropical seagrasses are likely to establish later, delayed by more limited dispersal abilities. Ultimately, food webs are likely to shift from primarily seagrass-detritus to more direct-consumption-based systems, thereby affecting a range of important ecosystem services that seagrasses provide, including their nursery habitat role for fishery species, carbon sequestration, and the provision of organic matter to other ecosystems in temperate regions.

Evaluation of geostatistical estimators and their applicability to characterise the spatial patterns of recreational fishing catch rates.

Western Australians are heavily engaged in recreational fishing activities with a participation rate of approximately 30%. An accurate estimation of the spatial distribution of recreational catch per unit effort (catch rates) is an integral component for monitoring fish population changes and to develop strategies for ecosystem-based marine management. Geostatistical techniques such as kriging can provide useful tools for characterising the spatial distributions of recreational catch rates. However, most recreational fishery data are highly skewed, zero-inflated and when expressed as ratios are impacted by the small number problem which can influence the estimates obtained from the traditional kriging. The applicability of ordinary, indicator and Poisson kriging to recreational catch rate data was evaluated for three aquatic species with different behaviours and distribution patterns. The prediction performance of each estimator was assessed based on cross-validation. For all three species, the accuracy plot of the indicator kriging (IK) showed a better agreement between expected and empirical proportions of catch rate data falling within probability intervals of increasing size, as measured by the goodness statistic. Also, indicator kriging was found to be better in predicting the latent catch rate for the three species compared to ordinary and Poisson kriging. For each species, the spatial maps from the three estimators displayed similar patterns but Poisson kriging produced smoother spatial distributions. We show that the IK estimator may be preferable for the spatial modelling of catch rate data exhibiting these characteristics, and has the best prediction performance regardless of the life history and distribution patterns of those three species.

Mechanisms and ecological role of carbon transfer within coastal seascapes.

Worldwide, coastal systems provide some of the most productive habitats, which potentially influence a range of marine and terrestrial ecosystems through the transfer of nutrients and energy. Several reviews have examined aspects of connectivity within coastal seascapes, but the scope of those reviews has been limited to single systems or single vectors. We use the transfer of carbon to examine the processes of connectivity through multiple vectors in multiple ecosystems using four coastal seascapes as case studies. We discuss and compare the main vectors of carbon connecting different ecosystems, and then the natural and human-induced factors that influence the magnitude of effect for those vectors on recipient systems. Vectors of carbon transfer can be grouped into two main categories: detrital particulate organic carbon (POC) and its associated dissolved organic and inorganic carbon (DOC/DIC) that are transported passively; and mobile consumers that transport carbon actively. High proportions of net primary production can be exported over meters to hundreds of kilometers from seagrass beds, algal reefs and mangroves as POC, with its export dependent on wind-generated currents in the first two of these systems and tidal currents for the last. By contrast, saltmarshes export large quantities of DOC through tidal movement, while land run-off plays a critical role in the transport of terrestrial POC and DOC into temperate fjords. Nekton actively transfers carbon across ecosystem boundaries through foraging movements, ontogenetic migrations, or 'trophic relays', into and out of seagrass beds, mangroves or saltmarshes. The magnitude of these vectors is influenced by: the hydrodynamics and geomorphology of the region; the characteristics of the carbon vector, such as their particle size and buoyancy; and for nekton, the extent and frequency of migrations between ecosystems. Through a risk-assessment process, we have identified the most significant human disturbances that affect the integrity of connectivity among ecosystems. Loss of habitat, net primary production (NPP) and overfishing pose the greatest risks to carbon transfer in temperate saltmarsh and tropical estuaries, particularly through their effects on nekton abundance and movement. In comparison, habitat/NPP loss and climate change are likely to be the major risks to carbon transfer in temperate fjords and temperate open coasts through alteration in the amount of POC and/or DOC/DIC being transported. While we have highlighted the importance of these vectors in coastal seascapes, there is limited quantitative data on the effects of these vectors on recipient systems. It is only through quantifying those subsidies that we can effectively incorporate complex interactions into the management of the marine environment and its resources.

Spatial patterns in herbivory on a coral reef are influenced by structural complexity but not by algal traits.

BACKGROUND: Patterns of herbivory can alter the spatial structure of ecosystems, with important consequences for ecosystem functions and biodiversity. While the factors that drive spatial patterns in herbivory in terrestrial systems are well established, comparatively less is known about what influences the distribution of herbivory in coral reefs. METHODOLOGY AND PRINCIPAL FINDINGS: We quantified spatial patterns of macroalgal consumption in a cross-section of Ningaloo Reef (Western Australia). We used a combination of descriptive and experimental approaches to assess the influence of multiple macroalgal traits and structural complexity in establishing the observed spatial patterns in macroalgal herbivory, and to identify potential feedback mechanisms between herbivory and macroalgal nutritional quality. Spatial patterns in macroalgal consumption were best explained by differences in structural complexity among habitats. The biomass of herbivorous fish, and rates of herbivory were always greater in the structurally-complex coral-dominated outer reef and reef flat habitats, which were also characterised by high biomass of herbivorous fish, low cover and biomass of macroalgae and the presence of unpalatable algae species. Macroalgal consumption decreased to undetectable levels within 75 m of structurally-complex reef habitat, and algae were most abundant in the structurally-simple lagoon habitats, which were also characterised by the presence of the most palatable algae species. In contrast to terrestrial ecosystems, herbivory patterns were not influenced by the distribution, productivity or nutritional quality of resources (macroalgae), and we found no evidence of a positive feedback between macroalgal consumption and the nitrogen content of algae. SIGNIFICANCE: This study highlights the importance of seascape-scale patterns in structural complexity in determining spatial patterns of macroalgal consumption by fish. Given the importance of herbivory in maintaining the ability of coral reefs to reorganise and retain ecosystem functions following disturbance, structural complexity emerges as a critical feature that is essential for the healthy functioning of these ecosystems.

Food web analysis in two permanently open temperate estuaries: consequences of saltmarsh loss?

Saltmarsh vegetation, seston and microphytobenthos are all conspicuous components of most temperate estuaries and they potentially contribute to the estuarine food chain. Yet their relative contributions are unclear, as is the significance of saltmarsh losses through natural and human-induced impacts. This study aimed to quantitatively determine the contribution of various types of primary producers to detritus in the Walpole-Nornalup Estuary and Leschenault Inlet, two permanently open estuaries in SW Australia, and, estimate the flow of different types of detritus to higher trophic levels, using carbon ((13)C) and nitrogen ((15)N) stable isotopes as tracers. Results of the mixing model indicated that seston, microphytobenthos and to some extent seagrass and fringing saltmarsh were the main contributors to the detrital pool in both estuaries. However, the relative contribution of different primary producers varied both within and between estuaries. The contribution of saltmarsh was higher at sites close to rivers and dense fringing vegetation, while seston, microphytobenthos and seagrass dominated the detrital material at other sites. Benthic harpacticoid copepods were shown to feed on detritus though they appeared to actively select for different components of the detritus depending on site and estuary. Isotopic signatures of other consumers indicated that fish and invertebrates derived nutrients from MPB and detritus, either directly as food or indirectly through feeding on invertebrates. The overall contribution of saltmarsh to detritus was lower in Leschenault Inlet than in Walpole-Nornalup Estuary, possibly as a result of increased clearing of fringing vegetation around Leschenault Inlet. This pattern was however not reflected in harpacticoid food. Therefore, although losses of fringing saltmarsh around estuaries have the potential to significantly affect estuarine food webs, the significance of such losses will be site- and estuary-dependent.