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The article presents a meta-analysis of the restoration of oyster habitat worldwide to determine restoration-associated recovery of biodiversity and abundance of resident and transient fish and invertebrates in these habitats. Topics discussed include the effectiveness of the restoration at mitigating damage to oyster habitat ecosystems, the difference in diversity and abundance between unrestored and restored habitats, and the oyster habitat restoration method used.

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. Species interactions that can enhance habitat heterogeneity such as facilitation cascades of foundation species have been overlooked in biodiversity models. This study conducted 22 geographically distributed experiments in different ecosystems and biogeographical regions to assess the extent to which biodiversity is explained by three axes of habitat heterogeneity in facilitation cascades. [ABSTRACT FROM AUTHOR]

Urbanization is leading to biodiversity loss through habitat homogenization. The smooth, featureless surfaces of many marine urban structures support ecological communities, often of lower biodiversity, distinct from the complex natural habitats they replace. Eco-engineering (design for ecological co-benefits) seeks to enhance biodiversity and ecological functions on urban structures. We assessed the benefits to biodiversity of retrofitting four types of complex habitat panels to an intertidal seawall at patch (versus flat control panels) and site (versus unmodified control seawalls and reference rocky shores) scales. Two years after installation, patch-scale effects of complex panels on biodiversity ranged from neutral to positive, depending on the protective features they provided, though all but one design (honeycomb) supported unique species. Water-retaining features (rockpools) and crevices, which provided moisture retention and cooling, increased biodiversity and supported algae and invertebrates otherwise absent. At the site scale, biodiversity benefits ranged from neutral at the high- and mid-intertidal to positive at the low-intertidal elevation. The results highlight the importance of matching eco-engineering interventions to the niche of target species, and environmental conditions. While species richness was greatest on rockpool and crevice panels, the unique species supported by other panel designs highlights that to maximize biodiversity, habitat heterogeneity is essential. This article is part of the theme issue 'Ecological complexity and the biosphere: the next 30 years'. [ABSTRACT FROM AUTHOR]

Along urbanised coastlines, urban infrastructure is increasingly becoming the dominant habitat. These structures are often poor surrogates for natural habitats, and a diversity of eco-engineering approaches have been trialled to enhance their biodiversity, with varying success., We undertook a quantitative meta-analysis and qualitative review of 109 studies to compare the efficacy of common eco-engineering approaches (e.g. increasing texture, crevices, pits, holes, elevations and habitat-forming taxa) in enhancing the biodiversity of key functional groups of organisms, across a variety of habitat settings and spatial scales., All interventions, with one exception, increased the abundance or number of species of one or more of the functional groups considered. Nevertheless, the magnitude of effect varied markedly among groups and habitat settings. In the intertidal, interventions that provided moisture and shade had the greatest effect on the richness of sessile and mobile organisms, while water-retaining features had the greatest effect on the richness of fish. In contrast, in the subtidal, small-scale depressions which provide refuge to new recruits from predators and other environmental stressors such as waves, had higher abundances of sessile organisms while elevated structures had higher numbers and abundances of fish. The taxa that responded most positively to eco-engineering in the intertidal were those whose body size most closely matched the dimensions of the resulting intervention., Synthesis and applications. The efficacy of eco-engineering interventions varies among habitat settings and functional groups. This indicates the importance of developing site-specific approaches that match the target taxa and dominant stressors. Furthermore, because different types of intervention are effective at enhancing different groups of organisms, ideally a range of approaches should be applied simultaneously to maximise niche diversity. [ABSTRACT FROM AUTHOR]

Extensive development and construction in marine and coastal systems is driving a phenomenon known as “ocean sprawl”. Ocean sprawl removes or transforms marine habitats through the addition of artificial structures and some of the most significant impacts are occurring in sedimentary environments. Marine sediments have substantial social, ecological, and economic value, as they are rich in biodiversity, crucial to fisheries productivity, and major sites of nutrient transformation. Yet the impact of ocean sprawl on sedimentary environments has largely been ignored. Here we review current knowledge of the impacts to sedimentary ecosystems arising from artificial structures. Artificial structures alter the composition and abundance of a wide variety of sediment-dependent taxa, including microbes, invertebrates, and benthic-feeding fishes. The effects vary by structure design and configuration, as well as the physical, chemical, and biological characteristics of the environment in which structures are placed. The mechanisms driving effects from artificial structures include placement loss, habitat degradation, modification of sound and light conditions, hydrodynamic changes, organic enrichment and material fluxes, contamination, and altered biotic interactions. Most studies have inferred mechanism based on descriptive work, comparing biological and physical processes at various distances from structures. Further experimental studies are needed to identify the relative importance of multiple mechanisms and to demonstrate causal relationships. Additionally, past studies have focused on impacts at a relatively small scale, and independently of other development that is occurring. There is need to quantify large-scale and cumulative effects on sedimentary ecosystems as artificial structures proliferate. We highlight the importance for comprehensive monitoring using robust survey designs and outline research strategies needed to understand, value, and protect marine sedimentary ecosystems in the face of a rapidly changing environment. [ABSTRACT FROM AUTHOR]

The growing number of artificial structures in estuarine, coastal and marine environments is causing “ocean sprawl”. Artificial structures do not only modify marine and coastal ecosystems at the sites of their placement, but may also produce larger-scale impacts through their alteration of ecological connectivity - the movement of organisms, materials and energy between habitat units within seascapes. Despite the growing awareness of the capacity of ocean sprawl to influence ecological connectivity, we lack a comprehensive understanding of how artificial structures modify ecological connectivity in near- and off-shore environments, and when and where their effects on connectivity are greatest. We review the mechanisms by which ocean sprawl may modify ecological connectivity, including trophic connectivity associated with the flow of nutrients and resources. We also review demonstrated, inferred and likely ecological impacts of such changes to connectivity, at scales from genes to ecosystems, and potential strategies of management for mitigating these effects. Ocean sprawl may alter connectivity by: (1) creating barriers to the movement of some organisms and resources - by adding physical barriers or by modifying and fragmenting habitats; (2) introducing new structural material that acts as a conduit for the movement of other organisms or resources across the landscape; and (3) altering trophic connectivity. Changes to connectivity may, in turn, influence the genetic structure and size of populations, the distribution of species, and community structure and ecological functioning. Two main approaches to the assessment of ecological connectivity have been taken: (1) measurement of structural connectivity - the configuration of the landscape and habitat patches and their dynamics; and (2) measurement of functional connectivity - the response of organisms or particles to the landscape. Our review reveals the paucity of studies directly addressing the effects of artificial structures on ecological connectivity in the marine environment, particularly at large spatial and temporal scales. With the ongoing development of estuarine and marine environments, there is a pressing need for additional studies that quantify the effects of ocean sprawl on ecological connectivity. Understanding the mechanisms by which structures modify connectivity is essential if marine spatial planning and eco-engineering are to be effectively utilised to minimise impacts. [ABSTRACT FROM AUTHOR]

Ecological theory predicts that positive interactions among organisms will increase across gradients of increasing abiotic stress or consumer pressure. This theory has been supported by empirical studies examining the magnitude of ecosystem engineering across environmental gradients and between habitat settings at local scale. Predictions that habitat setting, by modifying both biotic and abiotic factors, will determine large-scale gradients in ecosystem engineering have not been tested, however. A combination of manipulative experiments and field surveys assessed whether along the east Australian coastline: (1) facilitation of invertebrates by the oyster Saccostrea glomerata increased across a latitudinal gradient in temperature; and (2) the magnitude of this effect varied between intertidal rocky shores and mangrove forests. It was expected that on rocky shores, where oysters are the primary ecosystem engineer, they would play a greater role in ameliorating latitudinal gradients in temperature than in mangroves, where they are a secondary ecosystem engineer living under the mangrove canopy. On rocky shores, the enhancement of invertebrate abundance in oysters as compared to bare microhabitat decreased with latitude, as the maximum temperatures experienced by intertidal organisms diminished. By contrast, in mangrove forests, where the mangrove canopy resulted in maximum temperatures that were cooler and of greater humidity than on rocky shores, we found no evidence of latitudinal gradients of oyster effects on invertebrate abundance. Contrary to predictions, the magnitude by which oysters enhanced biodiversity was in many instances similar between mangroves and rocky shores. Whether habitat-context modifies patterns of spatial variation in the effects of ecosystem engineers on community structure will depend, in part, on the extent to which the environmental amelioration provided by an ecosystem engineer replicates that of other co-occurring ecosystem engineers. [ABSTRACT FROM AUTHOR]

Predator-prey interactions are often highly co-evolved, with selection over time for prey with morphological and behavioral traits that minimize predation risk. Consequently, in many environments prey choose among potential habitats according to their refuge value. It is unclear, however, when presented with new habitats, if prey are able to evaluate the predation risk of these relative to familiar habitats and utilize these in accordance with their value. We tested whether, along the east coast of the USA, native mud crabs Panopeus herbstii utilize the non-native alga Gracilaria vermiculophylla according to its relative refuge value. Experiments examining predation by blue crabs Callinectes sapidus on mud crabs revealed that the non-native alga had an intermediate refuge value relative to native oysters, which were the most protective, and unvegetated sediment, which was the least. In subsequent choice experiments, mud crabs selected oysters over alga over unvegetated sediment, in accordance with habitat refuge values. Further, in field experiments, the use of Gracilaria by mud crabs was inversely related to the proximity of the alga to the preferred habitat type, oysters, and was reduced by the presence of a blue crab predator. Consequently, mud crabs are utilizing the non-native alga Gracilaria in accordance with its intermediate refuge value. The relative refuge value of non-native vs native habitat-forming species may provide a baseline expectation against which to measure the speed of learning and opportunism in the response of native prey to novel protective habitats. [ABSTRACT FROM AUTHOR]

Abstract: Beach nourishment is increasingly used to protect public beach amenity and coastal property from erosion and storm damage. Where beach nourishment uses fill sediments that differ in sedimentology from native beach sands, press disturbances to sandy beach invertebrates and their ecosystem services can occur. How long impacts persist is, however, unclear because monitoring after nourishment typically only extends for several months. Here, monitoring was extended for 3–4years following each of two spatially separated, replicate nourishment projects using unnaturally coarse sediments. Following both fill events, the contribution to beach sediments of gravel-sized particles and shell fragments was enhanced, and although diminishing through time, remained elevated as compared to control sites at the end of 3–4years of monitoring, including in the low intertidal and swash zones, where benthic macroinvertebrates concentrate. Consequently, two infaunal invertebrates, haustoriid amphipods and Donax spp., exhibited suppressed densities over the entire post-nourishment period of 3–4years. Emerita talpoida, by contrast, exhibited lower densities on nourished than control beaches only in the early summer of the first and second years and polychaetes exhibited little response to nourishment. The overall impact to invertebrates of nourishment was matched by multi-year reductions in abundances of their predators. Ghost crab abundances were suppressed on nourished beaches with impacts disappearing only by the fourth summer. Counts of foraging shorebirds were depressed for 4years after the first project and 2years after the second project. Our results challenge the view that beach nourishment is environmentally benign by demonstrating that application of unnaturally coarse and shelly sediments can serve as a press disturbance to degrade the beach habitat and its trophic services to shorebirds for 2–4years. Recognizing that recovery following nourishment can be slow, studies that monitor impacts for only several months are inadequate. [Copyright &y& Elsevier]

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. [ABSTRACT FROM AUTHOR]

Among the impacts of coastal settlements to estuaries, nutrient pollution is often singled out as a leading cause of modification to the ecological communities of soft sediments. Through sampling of 48 sites, distributed among 16 estuaries of New South Wales, Australia, we tested the hypotheses that (1) anthropogenic nutrient loads would be a better predictor of macrofaunal communities than estuarine geomorphology or local sediment characteristics; and (2) local environmental context, as determined largely by sediment characteristics, would modify the relationship between nutrient loading and community composition. Contrary to the hypothesis, multivariate multiple regression analyses revealed that sediment grain size was the best predictor of macrofaunal assemblage composition. When samples were stratified according to median grain size, relationships between faunal communities and nitrogen loading and latitude emerged, but only among estuaries with sandier sediments. In these estuaries, capitellid and nereid polychaetes and chironomid larvae were the taxa that showed the strongest correlations with nutrient loading. Overall, this study failed to provide evidence of a differential relationship between diffuse nutrient enrichment and benthic macrofauna across a gradient of 7° of latitude and 4°C temperature. Nevertheless, as human population growth continues to place increasing pressure on southeast Australian estuaries, manipulative field studies examining when and where nutrient loading will lead to significant changes in estuarine community structure are needed. [ABSTRACT FROM AUTHOR]

Abstract: Oyster mortality and subsequent degradation of shell matrices may influence associated epibiota by modifying processes of filtration and biodeposition and by changing habitat structure. In the Hawkesbury River, NSW, Australia, QX disease devastated aquaculture populations of the native Sydney rock oyster, Saccostrea glomerata, and threatened wild populations. To ascertain effects of this oyster mortality on associated epibiota, we compared epifaunal communities occupying 100% oyster shell cover among 5 sites along the estuary, ranging from 25ppt salinity and live oyster density of 96±13m−2, to 32ppt salinity and live oyster density of 3187±233m−2. Epifaunal richness was greatest closest to the estuarine mouth, where live oyster abundance was greatest. Epifaunal abundance, by contrast, generally increased with distance upstream, as oyster matrices were increasingly dominated by the shells of dead oysters. To ascertain the role of live and dead oysters as habitat providers, we carried out a manipulative experiment to test the hypothesis that the epifaunal assemblages that settle over a 4month period will significantly differ among constructed matrices comprised of live, whole dead and/or degraded (single valve) oysters. Treatments containing a component of single valves, which increase the availability of interstitial spaces and provide greater surface area for attachment, supported the greatest numbers of epibiota. Matrices comprised solely of live oysters supported fewest species and numbers of organisms. Results demonstrate that death and degradation of oysters alter the structure of associated communities, even where 100% cover of shell matrix is maintained. These results have important ramifications for management strategies and retaining estuarine biodiversity in the event that disease such as QX causes local oyster extinctions. [Copyright &y& Elsevier]

Intertidal rocky shores are considered among the most thermally stressful marine ecosystems, where many species live close to their upper thermal limit and depend on access to cool microclimates to persist through heat events. In such environments, the provision of cool microclimates by habitat-forming species enables persistence of associated species during high temperature events. We assessed whether, by maintaining cool microclimates through heat events, habitat formed by rock oysters (Saccostrea cucullata) provides temporal stability to associated invertebrate communities over periods of extreme temperatures. On three tropical rocky shores of Hong Kong, which experiences a monsoonal climate, we compared changes in microclimates and invertebrate communities associated with oyster and bare rock habitats between the cool and hot seasons. Oyster habitats were, across both seasons, consistently characterised by lower maximum temperatures and greater thermal stability than bare rock habitats. Invertebrate communities in the bare rock habitat were less diverse and abundant in the hot than the cool season, but communities in the cooler habitats provided by oysters did not display temporal change. These results demonstrate that microclimates formed by oysters provide temporal stability to associated communities across periods of temperature change and are key determinants of species distributions in thermally stressful environments. The conservation and restoration of oyster habitats may, therefore, build resilience in associated ecological communities subject to ongoing environmental change. • Oyster habitat reduces the impact of rising temperatures on intertidal organisms. • Microclimates within oyster habitat extend the distribution of associated animals. • Habitat-forming species can confer temporal stability for associated communities. [ABSTRACT FROM AUTHOR]

Abstract: Studies on rocky intertidal gastropods indicate habitat complexity and body size to be major determinants of dispersal patterns. Considerations of effects of habitat complexity and body size on soft sediment gastropods are, however, less common. In neither habitat has the interaction between habitat complexity and body size been considered despite the increasing recognition in the general ecological literature that complexity effects are body-size-dependent. We tested independent and interacting effects of habitat complexity and body size on movement of the mud-whelk, Pyrazus ebeninus, by marking large (61–85 mm) and small (31–55 mm) snails in sites with low and high densities of pneumatophores and determining the distance and direction of their dispersal over periods of 1 week, 2 weeks and 1 month. Contrary to our expectation, we found no effect of pneumatophore density on the distance of snail migration over each of the temporal scales; net distance travelled by snails was determined only by body size and idiosynchratic, site-specific factors. The direction of snail movement was, by contrast, influenced on some temporal scales by both pneumatophore density and snail size. Over 1 week, site effects dominated patterns of movement and neither size of snail nor density of pneumatophore produced statistically significant effects. As the temporal scale increased, effects of size and pneumatophore density became increasingly apparent. Over the 1-month period, large snails at all sites and small snails at sites with high pneumatophore density migrated down the shore, while small snails at sites with low pneumatophore displayed non-directional movement. Thus, overall this study provides only weak support for effects of pneumatophore density on snail movement. In combination with other studies, our results suggest that, in comparison to on rocky shores where habitat complexity has strong effects on the distribution, abundance and behaviour of gastropods in soft-sediment systems habitat complexity is a less important structuring agent. [Copyright &y& Elsevier]

Strategies for managing biological invasions are often based on the premise that characteristics of invading species and the invaded environment are key predictors of the invader's distribution. Yet, for either biological traits or environmental characteristics to explain distribution, adequate time must have elapsed for species to spread to all potential habitats. We compiled and analyzed a database of natural history and ecological traits of 138 coastal marine invertebrate species, the environmental conditions at sites to which they have been introduced, and their date of first introduction. We found that time since introduction explained the largest fraction (20%) of the variability in non-native range size, while traits of the species and environmental variables had significant, but minimal, influence on non-native range size. The positive relationship between time since introduction and range size indicates that non-native marine invertebrate species are not at equilibrium and are still spreading, posing a major challenge for management of coastal ecosystems. [ABSTRACT FROM AUTHOR]

Ecosystem engineers can influence biodiversity by enhancing complexity, and modifying the availability of resources. Understanding the mechanisms by which ecosystem engineers shape biodiversity is central to the concept of ‘ecological engineering’ of anthropogenic structures to enhance biodiversity. Here the presence and complexity of artificial turf was manipulated on an artificial structure to test the hypothesis that the colonisation of sessile invertebrates and mobile epibiota would vary with habitat complexity. Both sessile and mobile assemblage compositions differed according to the presence of artificial turf, and its complexity. Sessile invertebrates occupied greater proportions of available space on topographically simple ‘blank’ surfaces or low complexity artificial turf than those with high complexity turf, whereas mobile taxa were generally more abundant on the turf. However, the mobile assemblage was unrelated to the sessile assemblage when examined within each level of initial substrata complexity. Contrary to the increasing number of studies demonstrating nested hierarchical relationships between co-occurring ecosystem engineers, this study provides an example of an ecosystem engineer mimic (artificial turf) leading to the formation of habitat mosaics at small scales. The introduction of complex substrata to otherwise topographically simple artificial structures is a promising means of actively influencing assemblage composition. [ABSTRACT FROM AUTHOR]

Interest in oyster reef conservation and restoration is growing globally, but particularly in Australia, it is unclear the extent to which oyster reefs complement (versus replicate) habitat provisioning by other structured habitats in the seascape. Remote underwater video surveys of two east Australian estuaries revealed that at high tide, oyster reefs not only supported distinct fish communities to bare sediments but also to adjacent seagrass beds and mangrove forests. Fish observations in oyster reefs were close to double that of mangroves and seagrass, with species richness, abundance, feeding and wandering behaviours similar. Several species of blenny and goby were unique to oyster reefs and oyster-containing mangroves, whilst recreationally fished species such as bream and mullet were more abundant on oyster reefs than in other habitats. Resolving the association between oyster reefs and fish species within the broader seascape will assist in developing restoration and management strategies that maximise fisheries benefit. • The role of intertidal remnant oyster reefs as fish habitat was explored. • Oyster reefs provided habitat to a unique fish community within the seascape. • Fish observations were higher in oyster reefs, than in seagrass, mangroves and bare. • Fish sp. richness was higher in oyster reefs, than in seagrass, mangroves and bare. • Oyster reefs were utilised as refuge and foraging grounds by fish. [ABSTRACT FROM AUTHOR]