Artificial light at night and warming impact grazing rates and gonad index of the sea urchin Centrostephanus rodgersii.

Artificial light at night (ALAN) is a growing threat to coastal habitats, and is likely to exacerbate the impacts of other stressors. Kelp forests are dominant habitats on temperate reefs but are declining due to ocean warming and overgrazing. We tested the independent and interactive effects of ALAN (dark versus ALAN) and warming (ambient versus warm) on grazing rates and gonad index of the sea urchin Centrostephanus rodgersii. Within these treatments, urchins were fed either 'fresh' kelp or 'treated' kelp. Treated kelp (Ecklonia radiata) was exposed to the same light and temperature combinations as urchins. We assessed photosynthetic yield, carbon and nitrogen content and C : N ratio of treated kelp to help identify potential drivers behind any effects on urchins. Grazing increased with warming and ALAN for urchins fed fresh kelp, and increased with warming for urchins fed treated kelp. Gonad index was higher in ALAN/ambient and dark/warm treatments compared to dark/ambient treatments for urchins fed fresh kelp. Kelp carbon content was higher in ALAN/ambient treatments than ALAN/warm treatments at one time point. This indicates ocean warming and ALAN may increase urchin grazing pressure on rocky reefs, an important finding for management strategies.

Investigating the interactive effects of habitat type and light intensity on rocky shores.

Light availability and habitat complexity are two key drivers of community assembly. Urbanisation has been shown to affect both, with important consequences to ecological communities. On the intertidal, for instance, studies have shown that light intensity is greater on natural rocky shores than on less complex artificial habitats (seawalls), though different habitats can also experience similar light intensities, for example when shaded by urban structures. Understanding therefore how these factors individually, and combined, affect communities is important to understand the mechanisms driving changes in community structure, and consequently provide solutions to tackle the increasing homogenisation of habitats and lightscapes in urbanised spaces through smart infrastructure designs. Here, we assessed how different light levels affect the recruitment of communities in rock pools and on emergent rock on an intertidal rocky shore. We cleared 30 patches of emergent rock and 30 rock pools and manipulated light using shades with different light transmissions (full light, procedural control, 75%, 35%, and 15% light transmission, full shade) and assessed mobile and sessile communities monthly for 6 months. Effects of reducing light levels were generally stronger on rock than in pools. Fully shaded plots supported double the amount of mobile organisms than plots in full sunlight, in both habitats. Algal cover was higher in pools compared to rock, and at intermediate light levels, but effects varied with site. This study highlights the importance of variable light conditions and different habitats for rocky shore communities, which should be considered in future coastal developments to retain natural biodiversity.

Variable effects of substrate colour and microtexture on sessile marine taxa in Australian estuaries.

Concrete infrastructure in coastal waters is increasing. While adding complex habitat and manipulating concrete mixtures to enhance biodiversity have been studied, field investigations of sub-millimetre-scale complexity and substrate colour are lacking. Here, the interacting effects of 'colour' (white, grey, black) and 'microtexture' (smooth, 0.5 mm texture) on colonisation were assessed at three sites in Australia. In Townsville, no effects of colour or microtexture were observed. In Sydney, spirorbid polychaetes occupied more space on smooth than textured tiles, but there was no effect of microtexture on serpulid polychaetes, bryozoans and algae. In Melbourne, barnacles were more abundant on black than white tiles, while serpulid polychaetes showed opposite patterns and ascidians did not vary with treatments. These results suggest that microtexture and colour can facilitate colonisation of some taxa. The context-dependency of the results shows that inclusion of these factors into marine infrastructure designs needs to be carefully considered.

Complexity-functioning relationships differ across different environmental conditions.

Habitat complexity is widely considered an important determinant of biodiversity, and enhancing complexity can play a key role in restoring degraded habitats. However, the effects of habitat complexity on ecosystem functioning - as opposed to biodiversity and community structure - are relatively poorly understood for artificial habitats, which dominate many coastlines. With Greening of Grey Infrastructure (GGI) approaches, or eco-engineering, increasingly being applied around the globe, it is important to understand the effects that modifying habitat complexity has on both biodiversity and ecological functioning in these highly modified habitats. We assessed how manipulating physical (primary substrate) and/or biogenic habitat (bivalves) complexity on intertidal artificial substrata affected filtration rates, net and gross primary productivity (NPP and GPP, respectively) and community respiration (CR) - as well as abundance of filter feeders and macro-algae and habitat use by cryptobenthic fish across six locations in three continents. We manipulated both physical and biogenic complexity using 1) flat or ridged (2.5 cm or 5 cm) settlement tiles that were either 2) unseeded or seeded with oysters or mussels. Across all locations, increasing physical and biogenic complexity (5 cm seeded tiles) had a significant effect on most ecological functioning variables, increasing overall filtration rates and community respiration of the assemblages on tiles but decreasing productivity (both GPP and NPP) across all locations. There were no overall effects of increasing either type of habitat complexity on cryptobenthic fish MaxN, total time in frame or macro-algal cover. Within each location, there were marked differences in the effects of habitat complexity. In Hobart, we found higher filtration, filter feeder biomass and community respiration on 5 cm tiles compared to flat tiles. However, at this location, both macro-algae cover and GPP decreased with increasing physical complexity. Similarly in Dublin, filtration, filter feeder biomass and community respiration were higher on 5 cm tiles compared to less complex tiles. In Sydney, filtration and filter feeder biomass were higher on seeded than unseeded tiles, and fish MaxN was higher on 5 cm tiles compared to flat tiles. On unseeded tiles in Sydney, filter feeder biomass also increased with increasing physical complexity. Our findings suggest that GGI solutions via increased habitat complexity are likely to have trade-offs among potentially desired functions, such as productivity and filtration rates, and variable effects on cryptobenthic fish communities. Importantly, our results show that the effects of GGI practices can vary markedly according to the environmental context and therefore should not be blindly and uniformly applied across the globe.

Predicting the impact of sea-level rise on intertidal rocky shores with remote sensing.

Sea-level rise is an inevitable consequence of climate change and threatens coastal ecosystems, particularly intertidal habitats that are constrained by landward development. Intertidal habitats support significant biodiversity, but also provide natural buffers from climate-threats such as increased storm events. Predicting the effects of climate scenarios on coastal ecosystems is important for understanding both the degree of habitat loss for associated ecological communities and the risk of the loss of coastal buffer zones. We take a novel approach by combining remote sensing with the IUCN Red List of Ecosystem criteria to assess this impact. We quantified the extent of horizontal intertidal rocky shores along ~200 km of coastline in Eastern Australia using GIS and remote-sensing (LiDAR) and used this information to predict changes in extent under four different climate change driven sea-level rise scenarios. We then applied the IUCN Red List of Ecosystems Criterion C2 (habitat degradation over the next 50 years based on change in an abiotic variable) to estimate the status of this ecosystem using the Hawkesbury Shelf Marine Bioregion as a test coastline. We also used four individual rocky shores as case studies to investigate the role of local topography in determining the severity of sea-level rise impacts. We found that, if the habitat loss within the study area is representative of the entire bioregion, the IUCN status of this ecosystem is 'near threatened', assuming that an assessment of the other criteria would return lower categories of risk. There was, however, high spatial variability in this effect. Rocky shores with gentle slopes had the highest projected losses of area whereas rocky shores expanding above the current intertidal range were less affected. Among the sites surveyed in detail, the ecosystem status ranged from 'least concern' to 'vulnerable', but reached 'endangered' under upper estimates of the most severe scenario. Our results have important implications for conservation management, highlighting a new link between remote sensing and the IUCN Red List of Ecosystem criteria that can be applied worldwide to assess ecosystem risk to sea-level rise.

The matrix influences direct and indirect effects of an anthropogenic disturbance on marine organisms.

The magnitude and direction of direct and indirect effects of disturbances can be context-dependent, with the matrix (surrounding habitat) in which populations are embedded either mitigating or worsening the impacts of disturbances. Chemical disturbances are particularly harmful and can affect organisms directly or indirectly. We used bleach, a common stressor in marine systems, to test hypotheses about direct and indirect effects of anthropogenic disturbances on intertidal grazers and the influence of the surrounding macro-algal matrix on such effects. We manipulated the contaminant, food (biofilm) and surrounding macro-algal matrix. Fewer limpets were found in contaminated areas. Bleach had a strong direct negative effect on limpets and caused a reduction in biofilm food, indirectly affecting limpets. This effect was strongest in the presence of macro-algal matrix. Anthropogenic disturbances can have major consequences via direct and indirect effects on key interacting species. We showed that such effects are, however, context-dependent. Capsule: Pollution is a major driver of biodiversity declines. We show that direct and indirect effects of contaminants on organisms depend on the context in which they occur.

Application of management tools to integrate ecological principles with the design of marine infrastructure.

Globally the coastal zone is suffering the collateral damage from continuing urban development and construction, expanding resource sectors, increasing population, regulation to river flow, and on-going land change and degradation. While protection of natural coastal habitat is recommended, balancing conservation with human services is now the challenge for managers. Marine infrastructure such as seawalls, marinas and offshore platforms is increasingly used to support and provide services, but has primarily been designed for engineering purposes without consideration of the ecological consequences. Increasingly developments are seeking alternatives to hard engineering and a range of ecological solutions has begun to replace or be incorporated into marine and coastal infrastructure. But too often, hard engineering remains the primary strategy because the tools for managers to implement ecological solutions are either lacking or not supported by policy and stakeholders. Here we outline critical research needs for marine urban development and emerging strategies that seek to mitigate the impacts of marine infrastructure. We present case studies to highlight the strategic direction necessary to support management decisions internationally.

Antagonistic Effects of Light Pollution and Warming on Habitat-Forming Seaweeds.

Artificial Light at Night (ALAN) is an emerging global stressor that is likely to interact with other stressors such as warming, affecting habitat-forming species and ecological functions. Seaweeds are dominant habitat-forming species in temperate marine ecosystems, where they support primary productivity and diverse ecological communities. Warming is a major stressor affecting seaweed forests, but effects of ALAN on seaweeds are largely unknown. We manipulated ALAN (0 lx vs. 25 lx at night) and temperature (ambient vs. +1.54°C warming) to test their independent and interactive effects on the survival, growth (biomass, total-, blade- and stipe-length) and function (photosynthesis, primary productivity and respiration) on the juveniles of two habitat-forming seaweeds, the kelp Ecklonia radiata and the fucoid Sargassum sp. Warming significantly increased Ecklonia mortality; however, ALAN did not affect mortality. ALAN had positive effects on Ecklonia biomass, total and blade growth rates and gross primary productivity; however, warming largely counterbalanced these effects. We found no significant effects of warming or ALAN on Ecklonia photosynthetic yield, stipe length, net primary productivity or respiration rates. We found no effects of ALAN or warming on Sargassum for any of the measured variables. Synthesis. Our findings indicate that ALAN can have positive effects on seaweed growth and functioning, but such effects are likely species-specific and can be counterbalanced by warming, suggesting an antagonistic interaction between these global stressors. These findings can help us to predict and manage the effects of these stressors on seaweeds, which underpin coastal biodiversity.

Morpho-physiological traits and tissue burdens of Ecklonia radiata linked to environmental variation in an urban estuary.

Organisms respond to their environment in various ways, including moving, adapting, acclimatising or a combination of responses. Within estuarine habitats, organisms are exposed to naturally variable environmental conditions. In urbanised estuaries, these natural variations can interact with human stressors such as habitat modification and pollution. Here, we investigated trait variation in the golden kelp Ecklonia radiata across an urban estuary - Sydney Harbour, Australia. We found that kelp morphology differed significantly between the more human-modified inner and the less modified outer harbour. Kelp individuals were smaller, had fewer laminae, and lacked spines in the inner harbour where it was warmer, more contaminated and less light was available. Inner harbour populations were characterised by lower tissue nitrogen and higher lead concentrations. These findings provide insights into how environmental variation could affect kelp morphology and physiology, and the high trait variation suggests adaptive capacity in E. radiata.

Influence of habitat features on the colonisation of native and non-indigenous species.

Marine artificial structures provide substrates on which organisms can settle and grow. These structures facilitate establishment and spread of non-indigenous species, in part due to their distinct physical features (substrate material, movement, orientation) compared to natural habitat analogues such as rocky shores, and because following construction, they have abundant resources (space) for species to colonise. Despite the perceived importance of these habitat features, few studies have directly compared distributions of native and non-indigenous species or considered how functional identity and associated environmental preferences drive associations. We undertook a meta-analysis to investigate whether colonisation of native and non-indigenous species varies between artificial structures with features most closely resembling natural habitats (natural substrates, fixed structures, surfaces oriented upwards) and those least resembling natural habitats (artificial materials, floating structures, downfacing or vertical surfaces), or whether functional identity is the primary driver of differences. Analyses were done at global and more local (SE Australia) scales to investigate if patterns held regardless of scale. Our results suggest that functional group (i.e., algae, ascidians. barnacles, bryozoans, polychaetes) rather than species classification (i.e., native or non-indigenous) are the main drivers of differences in communities between different types of artificial structures. Specifically, there were differences in the abundance of ascidians, barnacles, and polychaetes between (1) upfacing and downfacing/vertical surfaces, and (2) floating and fixed substrates. When differences were detected, taxa were most abundant on features least resembling natural habitats. Results varied between global and SE Australian analyses, potentially due to reduced variability across studies in the SE Australian dataset. Thus, the functional group and associated preferences of the highest threat NIS in the area should be considered in design strategies (e.g., ecological engineering) to limit their establishment on newly built infrastructure.

Current extent and future opportunities for living shorelines in Australia.

Living shorelines aim to enhance the resilience of coastlines to hazards while simultaneously delivering co-benefits such as carbon sequestration. Despite the potential ecological and socio-economic benefits of living shorelines over conventional engineered coastal protection structures, application is limited globally. Australia has a long and diverse coastline that provides prime opportunities for living shorelines using beaches and dunes, vegetation, and biogenic reefs, which may be either natural ('soft' approach) or with an engineered structural component ('hybrid' approach). Published scientific studies, however, have indicated limited use of living shorelines for coastal protection in Australia. In response, we combined a national survey and interviews of coastal practitioners and a grey and peer-reviewed literature search to (1) identify barriers to living shoreline implementation; and (2) create a database of living shoreline projects in Australia based on sources other than scientific literature. Projects included were those that had either a primary or secondary goal of protection of coastal assets from erosion and/or flooding. We identified 138 living shoreline projects in Australia through the means sampled starting in 1970; with the number of projects increasing through time particularly since 2000. Over half of the total projects (59 %) were considered to be successful according to their initial stated objective (i.e., reducing hazard risk) and 18 % of projects could not be assessed for their success based on the information available. Seventy percent of projects received formal or informal monitoring. Even in the absence of peer-reviewed support for living shoreline construction in Australia, we discovered local and regional increases in their use. This suggests that coastal practitioners are learning on-the-ground, however more generally it was stated that few examples of living shorelines are being made available, suggesting a barrier in information sharing among agencies at a broader scale. A database of living shoreline projects can increase knowledge among practitioners globally to develop best practice that informs technical guidelines for different approaches and helps focus attention on areas for further research.

Urban shading and artificial light at night alter natural light regimes and affect marine intertidal assemblages.

Urban development in many coastal cities has resulted in altered natural light regimes, with many coastal habitats being artificially shaded during the daytime by built structures such as seawalls and piers, while artificial light emitted from buildings and associated infrastructure creates pollution at night. As a result, these habitats may experience changes to community structure and impacts on key ecological processes such as grazing. This study investigated how changes to light regimes affect the abundance of grazers on natural and artificial intertidal habitats in Sydney Harbour, Australia. We also examined whether differences in patterns of responses to shading or artificial light at night (ALAN) varied across different areas within the Harbour, characterised by different overall levels of urbanisation. As predicted, light intensity was greater during the daytime on rocky shores than seawalls at the more urbanised sites of the harbour. We found a negative relationship between the abundance of grazers and increasing light during the daytime on rocky shores (inner harbour) and seawalls (outer harbour). We found similar patterns at night on rocky shores, with a negative relationship between the abundance of grazers and light. However, on seawalls, grazer abundances increased with increasing night-time lux levels, but this was mainly driven by one site. Overall, we found the opposite patterns for algal cover. Our findings corroborate those of previous studies that found that urbanisation can significantly affect natural light cycles, with consequences to ecological communities.

The impacts of artificial light at night on the ecology of temperate and tropical reefs.

Despite 22% of the world's coastal regions experiencing some degree of light pollution, and biologically important artificial light at night (ALAN) reaching large portions of the seafloor (greater than 75%) near coastal developments, the impacts of ALAN on temperate and tropical reefs are still relatively unknown. Because many reef species have evolved in response to low-light nocturnal environments, consistent daily, lunar, and seasonal light cycles, and distinct light spectra, these impacts are likely to be profound. Recent studies have found ALAN can decrease reproductive success of fishes, alter predation rates of invertebrates and fishes, and impact the physiology and biochemistry of reef-building corals. In this paper, we integrate knowledge of the role of natural light in temperate and tropical reefs with a synthesis of the current literature on the impacts of ALAN on reef organisms to explore potential changes at the system level in reef communities exposed to ALAN. Specifically, we identify the direct impacts of ALAN on individual organisms and flow on effects for reef communities, and present potential scenarios where ALAN could significantly alter system-level dynamics, possibly even creating novel ecosystems. Lastly, we highlight large knowledge gaps in our understanding of the overall impact of ALAN on reef systems. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Contaminant pulse following wildfire is associated with shifts in estuarine benthic communities.

Novel combinations of climatic conditions due to climate change and prolonged fire seasons have contributed to an increased occurrence of "megafires". Such large-scale fires pose an unknown threat to biodiversity due to the increased extent and severity of burn. Assessments of wildfires often focus on terrestrial ecosystems and effects on aquatic habitats are less documented, particularly in coastal environments. In a novel application of eDNA techniques, we assessed the impacts of the 2019-2020 Australian wildfires on the diversity of estuarine benthic sediment communities in six estuaries in NSW, Australia, before and after the fires. Estuaries differed in area of catchment burnt (0-92%) and amount of vegetative buffer that remained post-fire between burnt areas and waterways. We found greater dissimilarities in the composition and abundance of eukaryotic and bacterial sediment communities in estuaries from burnt catchments with no buffer compared to those with an intact buffer or from unburnt catchments. Shifts in composition in highly burnt catchments were associated with increased concentrations of nutrients, carbon, including fire-derived pyrogenic carbon, and copper, which was representative of multiple highly correlated trace metals. Changes in the relative abundances of certain taxonomic groups, such as sulfate-reducing and nitrifying bacterial groups, in the most impacted estuaries indicate potential consequences for the functioning of sediment communities. These results provide a unique demonstration of the use of eDNA to identify wildfire impacts on ecological communities and emphasize the importance of vegetative buffers in limiting wildfire-associated impacts.

Complexity-biodiversity relationships on marine urban structures: reintroducing habitat heterogeneity through eco-engineering.

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'.

Variations in benthic fluxes of sediments near pier pilings and natural rocky reefs.

Marine artificial structures such as pilings are replacing natural habitats, and modifying surrounding areas, often resulting in local decreases in species diversity and facilitation of bioinvasion. Most research on the impacts of artificial structures in marine ecosystems has primarily focused on rocky bottom habitats and biodiversity, overlooking the effects of these structures on the functioning of nearby sedimentary habitats. Here we compared, for the first time, benthic metabolism (O2 fluxes) and sediment-water nutrient (inorganic nitrogen, phosphate, and dissolved organic nitrogen) fluxes in shallow water sediments adjacent to pilings and natural reefs. We also measured sediment properties (grain size, total organic carbon, total nitrogen, C:N ratio and chlorophyll-a content). We found that sediments near pilings were generally finer with greater C:N ratios than those near reefs, while differences in other sediment properties between types of habitats were dependent on the site. We found significant differences in the oxygen consumption, primary productivity, and net ecosystem metabolism in sediments around pilings compared to sediments near natural reefs, but these patterns differed by site. Net nutrient fluxes were similar in sediments near pilings and reefs at both sites. This study showed that although pilings can be associated with changes in the functioning of sedimentary habitats, patterns and the direction of change seem to vary depending on local conditions.

Wildfires cause rapid changes to estuarine benthic habitat.

Estuaries are one of the most valuable biomes on earth. Although humans are highly dependent on these ecosystems, anthropogenic activities have impacted estuaries worldwide, altering their ecological functions and ability to provide a variety of important ecosystem services. Many anthropogenic stressors combine to affect the soft sedimentary habitats that dominate estuarine ecosystems. Now, due to climate change, estuaries and other marine areas might be increasingly exposed to the emerging threat of megafires. Here, by sampling estuaries before and after a megafire, we describe impacts of wildfires on estuarine benthic habitats and justify why megafires are a new and concerning threat to coastal ecosystems. We (1) show that wildfires change the fundamental characteristics of estuarine benthic habitat, (2) identify the factors (burnt intensity and proximity to water's edge) that influence the consequences of fires on estuaries, and (3) identify relevant indicators of wildfire impact: metals, nutrients, and pyrogenic carbon. We then discuss how fires can impact estuaries globally, regardless of local variability and differences in catchment. In the first empirical assessment of the impact of wildfires on estuarine condition, our results highlight indicators that may assist waterway managers to empirically detect wildfire impacts in estuaries and identify catchment factors that should be included in fire risk assessments for estuaries. Overall, this study highlights the importance of considering fire threats in current and future estuarine and coastal management.

Light pollution: a landscape-scale issue requiring cross-realm consideration.

Terrestrial, marine and freshwater realms are inherently linked through ecological, biogeochemical and/or physical processes. An understanding of these connections is critical to optimise management strategies and ensure the ongoing resilience of ecosystems. Artificial light at night (ALAN) is a global stressor that can profoundly affect a wide range of organisms and habitats and impact multiple realms. Despite this, current management practices for light pollution rarely consider connectivity between realms. Here we discuss the ways in which ALAN can have cross-realm impacts and provide case studies for each example discussed. We identified three main ways in which ALAN can affect two or more realms: 1) impacts on species that have life cycles and/or stages in two or more realms, such as diadromous fish that cross realms during ontogenetic migrations and many terrestrial insects that have juvenile phases of the life cycle in aquatic realms; 2) impacts on species interactions that occur across realm boundaries, and 3) impacts on transition zones or ecosystems such as mangroves and estuaries. We then propose a framework for cross-realm management of light pollution and discuss current challenges and potential solutions to increase the uptake of a cross-realm approach for ALAN management. We argue that the strengthening and formalisation of professional networks that involve academics, lighting practitioners, environmental managers and regulators that work in multiple realms is essential to provide an integrated approach to light pollution. Networks that have a strong multi-realm and multi-disciplinary focus are important as they enable a holistic understanding of issues related to ALAN.

Supporting urban ecosystem services across terrestrial, marine and freshwater realms.

The terrestrial, freshwater and marine realms all provide essential ecosystem services in urban environments. However, the services provided by each realm are often considered independently, which ignores the synergies between them and risks underestimating the benefits derived collectively. Greater research collaboration across these realms, and an integrated approach to management decisions can help to support urban developments and restoration projects in maintaining or enhancing ecosystem services. The aim of this paper is to highlight the synergies and trade-offs among ecosystem services provided by each realm and to offer suggestions on how to improve current practice. We use case studies to illustrate the flow of services across realms. In our call to better integrate research and management across realms, we present a framework that provides a 6-step process for conducting collaborative research and management with an Australian perspective. Our framework considers unifying language, sharing, and understanding of desired outcomes, conducting cost-benefit analyses to minimise trade-offs, using multiple modes of communication for stakeholders, and applying research outcomes to inform regulation. It can be applied to improve collaboration among researchers, managers and planners from all realms, leading to strategic allocation of resources, increased protection of urban natural resources and improved environmental regulation with broad public support.

Sublethal effects of contaminants on marine habitat-forming species: a review and meta-analysis.

Contaminants may affect ecosystem functioning by reducing the fitness of organisms and these impacts may cascade through ecosystems, particularly if the sensitive organisms are also habitat-forming species. Understanding how sub-lethal effects of toxicants can affect the quality and functions of biogenic habitats is critical if we are to establish effective guidelines for protecting ecosystems. We carried out a global systematic review and meta-analysis critically evaluating contaminant effects on properties of habitat-formers linked to ecosystem functioning. We reviewed a total of 95 publications. However, 40% of publications initially captured by the literature search were identified as having flaws in experimental design and ~11% did not present results in an appropriate way and thus were excluded from the quantitative meta-analysis. We quantitatively reviewed 410 studies from 46 publications, of which 313 (~76%) were on plants and seaweeds, that is macro-algae, saltmarsh plants and seagrasses, 58 (~14%) studied corals and 39 (~10%) looked at toxicant impacts on bivalves, with 70% of those on mussels and the remaining studies on oysters. Response variables analysed were photosynthetic efficiency, amount of chlorophyll a (as a proxy for primary production) and growth of plants, seaweeds and corals as well as leaf area of plants. We also analysed filtration, growth and respiration rates of bivalves. Our meta-analysis found that chemical contaminants have a significant negative impact on most of the analysed functional variables, with the exception of the amount of chlorophyll a. Metals were the most widely harmful type of contaminant, significantly decreasing photosynthetic efficiency of kelps, leaf area of saltmarsh plants, growth of fucoids, corals and saltmarsh plants and the filtration rates of bivalves. Organic contaminants decreased the photosynthetic efficiency of seagrass, but had no significant effects on bivalve filtration. We did not find significant effects of polycyclic aromatic hydrocarbons on any of the analysed functional variables or habitat-forming taxa, but this could be due to the low number of studies available. A meta-regression revealed that relationships between concentrations of metal contaminants and the magnitude of functional responses varied with the type of metal and habitat-former. Increasing concentrations of contaminants significantly increased the negative effects on the photosynthetic efficiency of habitat-formers. There was, however, no apparent relationship between ecologically relevant concentrations of metals and effect sizes of photosynthetic efficiency of corals and seaweeds. A qualitative analysis of all relevant studies found slightly different patterns when compared to our quantitative analysis, emphasising the need for studies to meet critical inclusion criteria for meta-analyses. Our study highlights links between effects of contaminants at lower levels of organisation (i.e. at the biochemical and/or physiological level of individuals) and ecological, large-scale impacts, through effects on habitat-forming species. Contaminants can clearly reduce the functioning of many habitat-forming marine species. We therefore recommend the adoption of routine measures of functional endpoints in monitoring and conservation programs to complement structural measures.