Recruitment of a threatened foundation oyster species varies with large and small spatial scales.

Understanding how habitat attributes (e.g., patch area and sizes, connectivity) control recruitment and how this is modified by processes operating at larger spatial scales is fundamental to understanding population sustainability and developing successful long-term restoration strategies for marine foundation species-including for globally threatened reef-forming oysters. In two experiments, we assessed the recruitment and energy reserves of oyster recruits onto remnant reefs of the oyster Saccostrea glomerata in estuaries spanning 550 km of coastline in southeastern Australia. In the first experiment, we determined whether recruitment of oysters to settlement plates in three estuaries was correlated with reef attributes within patches (distances to patch edges and surface elevation), whole-patch attributes (shape and size of patches), and landscape attributes (connectivity). We also determined whether environmental factors (e.g., sedimentation and water temperature) explained the differences among recruitment plates. We also tested whether differences in energy reserves of recruits could explain the differences between two of the estuaries (one high- and one low-sedimentation estuary). In the second experiment, across six estuaries (three with nominally high and three with nominally low sedimentation rates), we tested the hypothesis that, at the estuary scale, recruitment and survival were negatively correlated to sedimentation. Overall, total oyster recruitment varied mostly at the scale of estuaries rather than with reef attributes and was negatively correlated with sedimentation. Percentage recruit survival was, however, similar among estuaries, although energy reserves and condition of recruits were lower at a high- compared to a low-sediment estuary. Within each estuary, total oyster recruitment increased with patch area and decreased with increasing tidal height. Our results showed that differences among estuaries have the largest influence on oyster recruitment and recruit health and this may be explained by environmental processes operating at the same scale. While survival was high across all estuaries, growth and reproduction of oysters on remnant reefs may be affected by sublethal effects on the health of recruits in high-sediment estuaries. Thus, restoration programs should consider lethal and sublethal effects of whole-estuary environmental processes when selecting sites and include environmental mitigation actions to maximize recruitment success.

The influence of oyster reefs and surrounding sediments on nitrogen removal - An in-situ study along the East coast of Australia.

Oyster reefs play a crucial role in the removal of nitrogen (N) from aquatic systems by facilitating nutrient regeneration and denitrification, both in their tissues and shells and surrounding sediments. However, we still have a limited understanding about the contribution of each component of the reefs (e.g. oysters vs sediments) to N processes, and whether rates are dependent on site-specific characteristics. To address these knowledge gaps, we conducted an experiment across six oyster reefs along 1080 km of the Eastern Australian coast with different sediment characteristics. By using in-situ clear and dark incubation chambers, we assessed how benthic metabolism, nutrient and dinitrogen gas (N2) fluxes varied among the following treatments: 'oysters', 'sediments', and 'sediments + oysters' that were used to represent components of the whole reef habitat (i.e. reef matrix vs surrounding sediments vs the interaction among them, respectively), and sites. We found that during dark conditions and at siltier sites, N2 effluxes from oysters can be up to 23 times higher than sediments, while N2 effluxes from chambers with both sediments and oysters were similar to sediment treatments, and lower than oyster treatments. These results can be explained by sediment processes including nutrient assimilation by benthic microalgae and/or lower nutrient diffusion into interstitial space. Additionally, oyster treatments showed an uptake of nitrate (NO3-) that was likely converted into N2, whereas sediment treatments showed an overall release of NO3-. In dark conditions, ammonium (NH4+) fluxes remained consistent across treatments and sites, indicating that any exports from oyster excretion (in those treatments including oysters) were either counterbalanced by or comparable to exports from sediments. This study provides evidence that the crucial contribution of oyster reefs to N removal is dependent on interactions between reef components and environmental factors.

Propagules are not all equal: traits of vegetative fragments and disturbance regulate invasion success.

Invasion success is regulated by multiple factors. While the roles of disturbance and propagule pressure in regulating the establishment of non-native species are widely acknowledged, that of propagule morphology (a proxy for quality) is poorly known. By means of a multi-factorial field experiment, we tested how the number (5 vs. 10) and quality (intact, without fronds or without rhizoids) of fragments of the clonal invasive seaweed, Caulerpa cylindracea, influenced its ability to establish in patches of the native seagrass, Posidonia oceanica, exposed to different intensities of disturbance (0, 50, or 100% reduction in canopy cover). We hypothesized that the ability of fragments to establish would be greater for intact fragments (high quality) and reduced more by frond removal (low quality) than rhizoid removal (intermediate quality). At low propagule pressure or quality, fragment establishment was predicted to increase with increasing disturbance, whereas, at high propagule pressure or quality, it was predicted to be high regardless of disturbance intensity. Disturbance intensity, fragment number and quality had independent effects on C. cylindracea establishment success. Disturbance always facilitated fragment establishment. However, fragments retaining fronds, either intact or deprived of rhizoids, had higher establishment success than fragments deprived of fronds. Increasing propagule number had weak effects on the cover of C. cylindracea. Our results demonstrate that propagule traits enabling the acquisition of resources made available by disturbance can be more important than propagule number in determining the establishment and spread of clonal non-native plants. More generally, our study suggests that propagule quality is a key, yet underexplored, determinant of invasion success.

Relationships between the spread of Caulerpa filiformis and fish communities on temperate rocky reefs.

The previously sub-dominant native marine macrophyte Caulerpa filiformis is now dominant on many sub-tidal rocky reefs in New South Wales (NSW), Australia and is expanding its distribution. As C. filiformis is highly chemically defended and structurally different to co-occurring habitat-forming macrophytes, two key attributes that govern fish assemblages, we hypothesized that fish assemblages, particularly herbivorous fishes, would be different at sites where C. filiformis occurred from where it was previously absent and within sites, fish community structure would be correlated to the cover of C. filiformis. We investigated these hypotheses by determining reef-associated fish assemblage attributes (assemblage structure, species richness, total abundance, Shannon-Weiner diversity, abundance of herbivorous species) along transects within sites where C. filiformis was present and absent. Surprisingly, despite large patches and very high densities of C. filiformis on the reefs we sampled, at larger spatial scales (i.e., among sites) no fish assemblage metrics differed between sites with large stands of C. filiformis and sites without the alga. Moreover the abundance of one dominant herbivore, the rock cale Aplodactylus lophodon, was greater at sites within large beds of C. filiformis. At smaller spatial scales, however, i.e. within sites where C. filiformis was present, fish assemblages did vary as a function of C. filiformis cover along transects, although this was not consistent across sampling times. Overall, our results suggest that the potential effects of the spread of this alga on faunal communities warrants further investigation.

Disturbance-mediated facilitation by an intertidal ecosystem engineer.

Ecosystem engineers facilitate communities by providing a structural habitat that reduces abiotic stress or predation pressure for associated species. However, disturbance may damage or move the engineer to a more stressful environment, possibly increasing the importance of facilitation for associated communities. In this study, we determined how disturbance to intertidal boulders (i.e., flipping) and the subsequent movement of a structural ecosystem engineer, the tube-forming serpulid worm Galeolaria caespitosa, from the bottom (natural state, low abiotic stress) to the top (disturbed state, high abiotic stress) surface of boulders influenced the importance of facilitation for intertidal communities across two intertidal zones. Theory predicts stronger relative facilitation should occur in the harsher environments of the top of boulders and the high intertidal zone. To test this prediction, we experimentally positioned boulders with the serpulids either face up or face down for 12 months in low and high zones in an intertidal boulder field. There were very different communities associated with the different boulders and serpulids had the strongest facilitative effects on the more stressful top surface of boulders with approximately double the species richness compared to boulders lacking serpulids. Moreover, within the serpulid matrix itself there was also approximately double the species richness (both zones) and abundance (high zone only) of small invertebrates on the top of boulders compared to the bottom. The high relative facilitation on the top of boulders reflected a large reduction in temperature by the serpulid matrix on that surface (up to 10°C) highlighting a key role for modification of the abiotic environment in determining the community-wide facilitation. This study has demonstrated that disturbance and subsequent movement of an ecosystem engineer to a more stressful environment increased the importance of facilitation and allowed species to persist that would otherwise be unable to survive in that environment.

Habitat-former effects on prey behaviour increase predation and non-predation mortality.

Habitat-forming species can influence mortality on associated species via altering structural and non-structural abiotic conditions. Importantly, these effects can occur simultaneously and in opposite directions, although how they contribute to the net outcomes for predator-prey interactions remain unexplored. Seagrasses often have positive effects on associated fauna because their structure directly reduces predator encounter rates. However, we identified a 'risky' behaviour (shallower burial) in an infaunal bivalve at a high seagrass cover--likely induced by non-structural abiotic change--suggesting positive effects may be outweighed by risky behaviours. We determined whether the physical structure of the seagrass interacted with burial behaviour of clams to determine the predation and non-predation mortality and whether these interactions were mediated by the cover of the seagrass. Surveys on an intertidal sand flat in Tasmania, Australia showed that the highest densities of a dominant bivalve, Katelysia scalarina, occurred at low (33%) seagrass cover, but the lowest densities and the highest proportion of unburied clams occurred at high (100%) cover. A field experiment manipulating burial depth, seagrass cover and predator access demonstrated that unburied clams suffered very high predation and non-predation mortality compared to buried clams (~4x higher), which outweighed any positive effects of the seagrass structure in reducing predator access. Being unburied also had non-lethal consequences with surviving unburied clams having a reduced tissue biomass compared to buried clams. In this system, predation was driven by the availability of prey when they undertake a risky behaviour (shallow burial). However, significant changes in behaviour may only occur once a threshold of habitat-former density is reached. In this instance, changes in behaviour were likely due to seagrass effects on sediment redox potential, which decreased significantly above 33% seagrass cover. Our findings demonstrate that the negative effects of a habitat-former on the behaviour of associated species, via alteration of non-structural abiotic conditions, can outweigh any positive effects provided by increasing habitat structure as is commonly reported for habitat-formers.

The influence of bioturbator activity on sediment bacterial structure and function is moderated by environment.

Bioturbation in coastal sediments plays a crucial role in biogeochemical cycling. However, a key knowledge gap is the extent to which bioturbation influences bacterial community diversity and ecosystem processes, such as nitrogen cycling. This study paired bacterial diversity, bioturbation activity and in situ flux measurements of oxygen and nitrogen from bioturbated sediments at six estuaries along the East coast of Australia. Bacterial community diversity, composition and predicted functional profiles were similar across burrow and surface sediments but were significantly influenced by bioturbator activity (measured as number of burrows) at sites with higher fine grain content. Sediment oxygen demand increased with bioturbator activity but changes in nitrogen cycling (as measured by fluxes and predicted bacterial functional gene analysis) were more spatially variable and were unrelated to bioturbator activity and bacterial community shifts. This study highlights how bioturbator activity influences bacterial community structure and functioning and what implications this has for biogeochemical cycles in estuarine sediments.

Morphological traits and density of foundation species modulate a facilitation cascade in Australian mangroves.

Facilitation cascades are critical to the maintenance of biodiversity in a variety of habitats. Through a series of two experiments, we examined how the morphological traits and density of interacting foundation species influence the establishment and persistence of a facilitation cascade in temperate Australian mangrove forests. In this system, mangrove pneumatophores trap the free-living alga, Hormosira banksii, which, in turn, supports dense and diverse assemblages of epifaunal mollusks. The first experiment, which manipulated pneumatophore height and density, revealed that these two traits each had additive negative effects on the establishment, but additive positive effects on the persistence of the cascade. High densities of tall pneumatophores initially served as a physical barrier to algal colonization of pneumatophore plots, but over the longer-term enhanced the retention of algae. The increased algal biomass, in turn, facilitating epifaunal colonization. The second experiment demonstrated that the retention of algae by pneumatophores was influenced more by algal thallus length than vesicle diameter, and this effect occurred independent of pneumatophore height. Our study has extended facilitation theory by showing that the morphological traits and density of basal and intermediary facilitators influence both the establishment and persistence of facilitation cascades. Hence, attempts to use foundation species as a tool for restoration will require an understanding not only of the interactions among these, but also of the key traits that modify interrelationships.

Organic enrichment reduces sediment bacterial and archaeal diversity, composition, and functional profile independent of bioturbator activity.

Eutrophication is a worldwide issue that can disrupt ecosystem processes in sediments. Studies have shown that macrofauna influences sediment processes by engineering environments that constrain microbial communities. Here, we explored the effect of different sizes of the Sydney cockle (Anadara trapezia), on bacterial and archaeal communities in natural and experimentally enriched sediments. A mesocosm experiment was conducted with two enrichment conditions (natural or enriched) and 5 cockle treatments (small, medium, large, mixed sizes and a control). This study was unable to detect A. trapezia effects on microbial communities irrespective of body size. However, a substantial decrease of bacterial richness, diversity, and structural and functional shifts, were seen with organic enrichment of sediments. Archaea were similarly changed although the magnitude of effect was less than for bacteria. Overall, we found evidence to suggest that A. trapezia had limited capacity to affect sediment microbial communities and mitigate the effects of organic enrichment.

Exotic asphyxiation: interactions between invasive species and hypoxia.

Non-indigenous species (NIS) and hypoxia (<2 mg O2 l-1 ) can disturb and restructure aquatic communities. Both are heavily influenced by human activities and are intensifying with global change. As these disturbances increase, understanding how they interact to affect native species and systems is essential. To expose patterns, outcomes, and generalizations, we thoroughly reviewed the biological invasion literature and compiled 100 studies that examine the interaction of hypoxia and NIS. We found that 64% of studies showed that NIS are tolerant of hypoxia, and 62% showed that NIS perform better than native species under hypoxia. Only one-quarter of studies examined NIS as creators of hypoxia; thus, NIS are more often considered passengers associated with hypoxia, rather than drivers of it. Paradoxically, the NIS that most commonly create hypoxia are primary producers. Taxa like molluscs are typically more hypoxia tolerant than mobile taxa like fish and crustaceans. Most studies examine individual-level or localized responses to hypoxia; however, the most extensive impacts occur when hypoxia associated with NIS affects communities and ecosystems. We discuss how these influences of hypoxia at higher levels of organization better inform net outcomes of the biological invasion process, i.e. establishment, spread, and impact, and are thus most useful to management. Our review identifies wide variation in the way in which the interaction between hypoxia and NIS is studied in the literature, and suggests ways to address the number of variables that affect their interaction and refine insight gleaned from future studies. We also identify a clear need for resource management to consider the interactive effects of these two global stressors which are almost exclusively managed independently.

Habitat traits and predation interact to drive abundance and body size patterns in associated fauna.

Habitat-forming organisms provide three-dimensional structure that supports abundant and diverse communities. Variation in the morphological traits of habitat formers will therefore likely influence how they facilitate associated communities, either via food and habitat provisioning, or by altering predator-prey interactions. These mechanisms, however, are typically studied in isolation, and thus, we know little of how they interact to affect associated communities. In response to this, we used naturally occurring morphological variability in the alga Sargassum vestitum to create habitat units of distinct morphotypes to test whether variation in the morphological traits (frond size and thallus size) of S. vestitum or the interaction between these traits affects their value as habitat for associated communities in the presence and absence of predation. We found morphological traits did not interact, instead having independent effects on epifauna that were negligible in the absence of predation. However, when predators were present, habitat units with large fronds were found to host significantly lower epifaunal abundances than other morphotypes, suggesting that large frond alga provided low-value refuge from predators. The presence of predators also influenced the size structure of epifaunal communities from habitat units of differing frond size, suggesting that the refuge value of S. vestitum was also related to epifauna body size. This suggests that habitat formers may chiefly structure associated communities by mediating size-selective predation, and not through habitat provisioning. Furthermore, these results also highlight that habitat traits cannot be considered in isolation, for their interaction with biotic processes can have significant implications for associated communities.

Response of planktonic microbial assemblages to disturbance in an urban sub-tropical estuary.

Microbes are sensitive indicators of estuarine processes because they respond rapidly to dynamic disturbance events. As most of the world's population lives in urban areas and climate change-related disturbance events are becoming more frequent, estuaries bounded by cities are experiencing increasing stressors, at the same time that their ecosystem services are required more than ever. Here, using a multidisciplinary approach, we determined the response of planktonic microbial assemblages in response to seasonality and a rainfall disturbance in an urban estuary bounded by Australia's largest city, Sydney. We used molecular barcoding (16S, 18S V4 rRNA) and microscopy-based identification to compare microbial assemblages at locations with differing characteristics and urbanisation histories. Across 142 samples, we identified 8,496 unique free-living bacterial zOTUs, 8,175 unique particle associated bacterial zOTUs, and 1,920 unique microbial eukaryotic zOTUs. Using microscopy, we identified only the top <10% abundant, larger eukaryotic taxa (>10 µm), however quantification was possible. The site with the greater history of anthropogenic impact showed a more even community of associated bacteria and eukaryotes, and a significant increase in dissolved inorganic nitrogen following rainfall, when compared to the more buffered site. This coincided with a reduced proportional abundance of Actinomarina and Synechococcus spp., a change in SAR 11 clades, and an increase in the eukaryotic microbial groups Dinophyceae, Mediophyceae and Bathyoccocaceae, including a temporary dominance of the harmful algal bloom dinoflagellate Prorocentrum cordatum (syn. P. minimum). Finally, a validated hydrodynamic model of the estuary supported these results, showing that the more highly urbanised and upstream location consistently experienced a higher magnitude of salinity reduction in response to rainfall events during the study period. The best abiotic variables to explain community dissimilarities between locations were TDP, PN, modelled temperature and salinity (r = 0.73) for the free living bacteria, TP for the associated bacteria (r = 0.43), and modelled temperature (r = 0.28) for the microbial eukaryotic communities. Overall, these results show that a minor disturbance such as a brief rainfall event can significantly shift the microbial assemblage of an anthropogenically impacted area within an urban estuary to a greater degree than a seasonal change, but may result in a lesser response to the same disturbance at a buffered, more oceanic influenced location. Fine scale research into the factors driving the response of microbial communities in urban estuaries to climate related disturbances will be necessary to understand and implement changes to maintain future estuarine ecosystem services.

Invasive ecosystem engineer selects for different phenotypes of an associated native species.

Invasive habitat-forming ecosystem engineers modify the abiotic environment and thus represent a major perturbation to many ecosystems. Because native species often persist in these invaded habitats but have no shared history with the ecosystem engineer, the engineer may impose novel selective pressure on native species. In this study, we used a phenotypic selection framework to determine whether an invasive habitat-forming ecosystem engineer (the seaweed Caulerpa taxifolia) selects for different phenotypes of a common co-occurring native species (the bivalve Anadara trapezia). Compared to unvegetated habitat, Caulerpa habitat has lower water flow, lower dissolved oxygen, and sediments are more silty and anoxic. We determined the performance consequences of variation in key functional traits that may be affected by these abiotic changes (shell morphology, gill mass, and palp mass) for Anadara transplanted into Caulerpa and unvegetated habitat. Both linear and nonlinear performance gradients in Anadara differed between habitats, and these gradients were stronger in Caulerpa compared to unvegetated sediment. Moreover, in Caulerpa alternate phenotypes performed well, and these phenotypes were different from the dominant phenotype in unvegetated sediment. By demonstrating that phenotype-performance gradients differ between habitats, we have highlighted a role for Caulerpa as an agent of selection on native species.

Density-dependent facilitation cascades determine epifaunal community structure in temperate Australian mangroves.

Co-occurring foundation species can determine biological community structure via facilitation cascades. We examined the density dependencies of facilitation cascades, including how the density of a basal foundation species influences the density of secondary foundation species, and how the density of secondary foundation species influences community structure. The system in which we assessed density dependencies was a temperate mangrove forest in which pneumatophores trap the fucoid alga Hormosira banksii and provide substrate for the oyster, Saccostrea glomerata. The alga and oyster in turn determine benthic community structure. In the field, algal biomass was positively correlated with pneumatophore density. Oysters, by contrast, were highly over-dispersed and correlated with the presence/absence of pneumatophores. Epifaunal abundance and species richness were positively correlated with algal and oyster abundance, but their effects were independent. The positive effect of pneumatophore density on epifauna was primarily an indirect effect of trapping more algae. Pneumatophores did not directly influence invertebrate communities. Experiments revealed that, at very low pneumatophore densities, algal retention was insufficient to facilitate epifauna above that found on pneumatophores alone. At higher densities, however, increasing the density of pneumatophores increased algal retention, and the density and diversity of associated invertebrates. Shading by the mangrove canopy reduced algal biomass but did not modify the density-dependent nature of the cascade. Our results extend facilitation theory by showing that the density of both basal and secondary foundation species can be critical in triggering facilitation cascades. Our study also reveals that, where foundation species co-occur, multiple, independent cascades may arise from a single basal facilitator. These findings enhance our understanding of the role of density-dependent facilitation cascades in community assembly.

Heterogeneity within and among co-occurring foundation species increases biodiversity.

Habitat heterogeneity is considered a primary causal driver underpinning patterns of diversity, yet the universal role of heterogeneity in structuring biodiversity is unclear due to a lack of coordinated experiments testing its effects across geographic scales and habitat types. Furthermore, key species interactions that can enhance heterogeneity, such as facilitation cascades of foundation species, have been largely overlooked in general biodiversity models. Here, we performed 22 geographically distributed experiments in different ecosystems and biogeographical regions to assess the extent to which variation in biodiversity is explained by three axes of habitat heterogeneity: the amount of habitat, its morphological complexity, and capacity to provide ecological resources (e.g. food) within and between co-occurring foundation species. We show that positive and additive effects across the three axes of heterogeneity are common, providing a compelling mechanistic insight into the universal importance of habitat heterogeneity in promoting biodiversity via cascades of facilitative interactions. Because many aspects of habitat heterogeneity can be controlled through restoration and management interventions, our findings are directly relevant to biodiversity conservation.

Fine-scale responses of mobile invertebrates and mesopredatory fish to habitat configuration.

As habitat-forming species continue to decline globally, it is important to understand how associated communities respond to habitat loss and fragmentation. Changes in the density and spatial configuration of habitat have important consequences for associated communities. However, tests of these factors are often confounded by morphological variation of habitat-formers, which can be resolved by using standardised habitat-mimics. Furthermore, few studies have incorporated the role of predators in mediating the observed effects. To test whether predators mediate the abundance of invertebrates among algal habitats of varying configuration (isolated vs patches, and positions within patches), we placed macroalgal mimics into subtidal estuarine habitats for one month to sample epifaunal communities. At the same time, we conducted underwater video surveys of fish communities to quantify fish communities and their feeding behaviour among the artificial habitats. Isolated habitats did not differ from patch habitats, however, patch edges had the highest epifaunal abundance, where fish were least commonly observed. Observed fish feeding was highest in the middle of patches and increased fish observations and feeding in habitats with reduced epifaunal communities suggest that mesopredatory fish are mediating epifauna in patches, with predation pressure altered by the spatial configuration of the habitat. This contrasts to previous studies that focus on predators that congregate outside patches and suggest that fragmentation leads to reduced invertebrate abundance at habitat edges in contrast to centres. However, this study highlights that in habitat patches housing small mesopredators that also benefit from the increased structure, the centre of the patch experiences higher predation and therefore fewer epifauna in contrast to patch edges and individual algal mimics.

Shifts in biomass and structure of habitat-formers across a latitudinal gradient.

Global patterns of plant biomass drive the distribution of much of the marine and terrestrial life on Earth. This is because their biomass and physical structure have important consequences for the communities they support by providing food and habitat. In terrestrial ecosystems, temperature is one of the major determinants of plant biomass and can influence plant and leaf morphology. In temperate marine systems, macroalgae are major habitat-formers and commonly display highly variable morphology in response to local environmental conditions. Variation in their morphology, and thus habitat structure on temperate reefs, however, is poorly understood across large scales. In this study, we used a trait-based approach to quantify morphological variability in subtidal rocky reefs dominated by the algal genus Sargassum along a latitudinal gradient, in southeastern Australia (~900 km). We tested whether large-scale variation in sea surface temperature (SST), site exposure, and nutrient availability can predict algal biomass and individual morphology. We found Sargassum biomass declined with increasing maximum SST. We also found that individual morphology varied with abiotic ocean variables. Frond size and intraindividual variability in frond size decreased with increasing with distance from the equator, as SST decreased and nitrate concentration increased. The shape of fronds displayed no clear relationship with any of the abiotic variables measured. These results suggest climate change will cause significant changes to the structure of Sargassum habitats along the southeastern coast of Australia, resulting in an overall reduction in biomass and increase in the prevalence of thalli with large, highly variable fronds. Using a space-for-time approach means shifts in morphological trait values can be used as early warning signs of impending species declines and regime shifts. Consequently, by studying traits and how they change across large scales we can potentially predict and anticipate the impacts of environmental change on these communities.

Sublethal effects of a rapidly spreading native alga on a key herbivore.

Multiple anthropogenic stressors are causing a global decline in foundation species, including macrophytes, often resulting in the expansion of functionally different, more stressor-tolerant macrophytes. Previously subdominant species may experience further positive demographic feedback if they are exposed to weaker plant-herbivore interactions, possibly via decreased palatability or being structurally different from the species they are replacing. However, the consequences of the spread of opportunistic macrophytes for the local distribution and life history of herbivores are unknown.The green alga, Caulerpa filiformis, previously a subdominant macrophyte on low intertidal-shallow subtidal rock shores, is becoming locally more abundant and has spread into warmer waters across the coast of New South Wales, Australia.In this study, we measured (a) the distribution and abundance of a key consumer, the sea urchin Heliocidaris erythrogramma, across a seascape at sites where C. filiformis has become dominant, (b) performed behavioral field experiments to test the role of habitat selection in determining the local distribution of H. erythrogramma, and (c) consumer experiments to test differential palatability between previously dominant higher quality species like Ecklonia radiata and Sargassum sp. and C. filiformis and the physiological consequences of consuming it.At all sites, urchin densities were positively correlated with distance away from C. filiformis beds, and they actively moved away from beds. Feeding experiments showed that, while urchins consumed C. filiformis, sometimes in equal amounts to higher quality algae, there were strong sublethal consequences associated with C. filiformis consumption, mainly on reproductive potential (gonad size). Specifically, the gonad size of urchins that fed on C. filiformis was equivalent to that in starved urchins. There was also a tendency for urchin mortality to be greater when fed C. filiformis.Overall, strong negative effects on herbivore life-history traits and potentially their survivorship may establish further positive feedback on C. filiformis abundance that contributes to its spread and may mediate shifts from top-down to bottom-up control at locations where C. filiformis has become dominant.

Body size affects lethal and sublethal responses to organic enrichment: Evidence of associational susceptibility for an infaunal bivalve.

Eutrophication is an increasing problem worldwide and can disrupt ecosystem processes in which macrobenthic bioturbators play an essential role. This study explores how intraspecific variation in body size affects the survival, mobility and impact on sediment organic matter breakdown in enriched sediments of an infaunal bivalve. A mesocosm experiment was conducted in which monocultures and all size combinations of three body sizes (small, medium and large) of the Sydney cockle, Anadara trapezia, were exposed to natural or organically enriched sediments. Results demonstrate that larger body sizes have higher tolerance to enriched conditions and can reduce survival of smaller cockles when grown together. Also, large A. trapezia influenced sediment organic matter breakdown although a direct link to bioturbation activity was not clear. Overall, this study found that intraspecific variation in body size influences survival and performance of bioturbators in eutrophic scenarios.

Legacy Metal Contaminants and Excess Nutrients in Low Flow Estuarine Embayments Alter Composition and Function of Benthic Bacterial Communities.

Coastal systems such as estuaries are threatened by multiple anthropogenic stressors worldwide. However, how these stressors and estuarine hydrology shape benthic bacterial communities and their functions remains poorly known. Here, we surveyed sediment bacterial communities in poorly flushed embayments and well flushed channels in Sydney Harbour, Australia, using 16S rRNA gene sequencing. Sediment samples were collected monthly during the Austral summer-autumn 2014 at increasing distance from a large storm drain in each channel and embayment. Bacterial communities differed significantly between sites that varied in proximity to storm drains, with a gradient of change apparent for sites within embayments. We explored this pattern for embayment sites with analysis of RNA-Seq gene expression patterns and found higher expression of multiple genes involved in bacterial stress response far from storm drains, suggesting that bacterial communities close to storm drains may be more tolerant of localised anthropogenic stressors. Several bacterial groups also differed close to and far from storm drains, suggesting their potential utility as bioindicators to monitor contaminants in estuarine sediments. Overall, our study provides useful insights into changes in the composition and functioning of benthic bacterial communities as a result of multiple anthropogenic stressors in differing hydrological conditions.