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Over the past few decades, tool use has been described in a variety of animals across taxa. However, there are relatively limited reports and studies of tool use in marine environments. Among marine fish though, labrids have been reported to be proficient at using tools. Here, we describe the first observations of tool use in two species of tropical wrasses—Halichoeres hortulanus (checkerboard wrasse) and Thalassoma jansenii (Jansen's wrasse)—and the first report of tool use from the wild for the wrasse Thalassoma lunare (Moon wrasse). These observations were made incidentally as part of size-specific predation assays for a small burrow-dwelling sea urchin (Echinostrephus molaris) in the Lakshadweep Archipelago, Indian Ocean. We documented multiple instances of anvil use by the wrasses H. hortulanus, and T. jansenii, and one instance of anvil use by T. lunare. All three species were recorded carrying the assay urchins to a nearby coral or benthic substrate and striking them repeatedly to break off their spines and crack their tests before consuming them. The behaviour was observed multiple times in different wrasse individuals. Also, sea urchin test size was a good predictor of the size of the fish that preyed on them. As naturalists spend more time observing fish in their natural environment, records of tool use in fish are increasing. Often these observations are incidental, yet documenting them carefully is critical, so we can ask larger questions about the evolution and development of animal intelligence. This study adds to observations of predation-associated tool use by fishes expanding reports of this behaviour taxonomically as well as geographically. [ABSTRACT FROM AUTHOR]

Introduction: Light gradients are ubiquitous in marine systems as light reduces exponentially with depth. Seagrasses have a set of mechanisms that help them to cope with light stress gradients. Physiological photoacclimation and clonal integration help to maximize light capture and minimize carbon losses. These mechanisms can shape plants minimum light requirements (MLR), which establish critical thresholds for seagrass survival and help us predict ecosystem responses to the alarming reduction in light availability. Methods: Using the seagrass Cymodocea nodosa as a case study, we compare the MLR under di fferent carbon model scenarios, which include photoacclimation and/or self-facilitation (based on clonal integration) and that where parameterized with values from field experiments. Results: Physiological photoacclimation conferred plants with increased tolerance to reducing light, approximately halving their MLR from 5-6% surface irradiance (SI) to ? 3% SI. In oligotrophic waters, this change in MLR could translate to an increase of several meters in their depth colonization limit. In addition, we show that reduced mortality rates derived from self-facilitation mechanisms (promoted by high biomass) induce bistability of seagrass meadows along the light stress gradient, leading to abrupt shifts and hysteretic behaviors at their deep limit. Discussion: The results from our models point to (i) the critical role of physiological photoacclimation in conferring greater resistance and ability to recover (i.e., resilience), to seagrasses facing light deprivation and (ii) the importance of self-facilitating reinforcing mechanisms in driving the resilience and recovery of seagrass systems exposed to severe light reduction events. [ABSTRACT FROM AUTHOR]

Aim: The global vertical depth distribution of seagrass species remains poorly understood. Locally, the abundance and distribution of seagrasses is determined by light penetration, but at global levels each seagrass species has very distinct maximum distributional depth ranges, indicating that plant‐associated traits must also influence their specific depth ranges. Seagrass‐specific attributes, such as plant size or architecture, growth or reproductive strategy and their physiological and/or morphological acclimatization potential, have been suggested to be responsible for this variety of vertical distributions. We investigate here whether these species‐specific traits drive differences in the global maximum vertical distribution of seagrasses. Location: Global. Time period: Publications between 1982 and 2020. Major taxa studied: Seagrasses (order Alismatales). Methods: We tested whether the species‐specific maximum vertical distribution of seagrasses can be predicted by (1) their rhizome diameter (a proxy for plant size); (2) their functional resilience (growth/reproductive strategy); or (3) their acclimatization capacity. For the last aspect, we used a systematic review followed by meta‐analytical approaches to select key seagrass traits that could potentially acclimatize to extreme light ranges across different seagrasses. Results: We found that vertical distribution is best explained by the species‐specific acclimatization capacity of various seagrass traits, including saturation irradiance (physiological trait), leaves per shoot (morphological trait) and above‐ground biomass (structural trait). In contrast, our results indicate no predictive power of seagrass size or growth/reproductive strategy on the vertical distribution of seagrasses. Main conclusions: Across the globe, the ability of seagrass species to thrive at a wide range of depths is strongly linked to the species‐specific acclimatization capacity of key traits at different organizational levels. [ABSTRACT FROM AUTHOR]

Predictable aggregations of large marine mammals are valuable conservation targets, but aggregated populations can also be exposed to site‐level threats.The globally vulnerable Dugong dugon has a wide distribution but is found in large numbers mainly in Australia and the Arabian Gulf. Though Australian dugong populations are well studied, much less is known of the dugongs in the Arabian Gulf.The spatial and temporal persistence of dugongs around Bahrain, with a focus on large dugong groups (>50 dugongs), was determined using an occupancy modelling framework supported by historical records, structured interviews, citizen science network reports, and small‐scale boat and unmanned aerial vehicle surveys.Historical records and current distributional studies confirmed that large dugong groups have been reliably sighted around Hawar Island (Bahrain) since at least 1986, forming large, clumped groups that persist almost year round. The largest recorded so far in the world, these fluid groups (maximum of ~700 dugongs) account for ~60% of the dugongs found in Bahrain and ~12% of all dugongs in the Arabian Gulf.The delineated occupancy core area of large dugong groups (~145 km2) straddles the Bahrain–Qatar border, reflecting the transboundary nature of these groups.Careful management of human‐induced stressors (in particular, fishing, boating, and coastal development) combined with regular monitoring of Hawar Island's large dugong groups and their seagrass habitat is critical to safeguard this globally important population.The effectiveness of any conservation management is predicated on strengthening cooperation among all range states in the Arabian Gulf. A key recommendation of this study is to establish a regional network of marine protected areas encompassing core aggregation sites for dugongs, particularly the Hawar Islands in Bahrain, the north‐western waters of Qatar, Marawah Island in the United Arab Emirates in addition to the shallow waters between Saudi Arabia, Qatar, and United Arab Emirates. [ABSTRACT FROM AUTHOR]

Plant–herbivore interactions are often regulated by a series of direct and indirect buffer mechanisms (compensatory or defensive plant responses, population control, alternative resources) that can determine the relative stability of the system. In plant‐dominated marine environments, these mechanisms are particularly important given how vulnerable these systems are to strong consumer pressure. Here, we evaluate the presence and, where possible, the strength, of five mechanisms that get activated under high herbivory pressure and are capable of dampening their effects in a seagrass meadow ecosystem: 1) seagrass compensatory growth, 2) increase in plant resistance, 3) availability of alternative resources, 4) predatory control and 5) density‐dependent control. We assessed these mechanisms for the interaction between the Mediterranean seagrass Posidonia oceanica and the sea urchin Paracentrotus lividus through a short‐term series of controlled field and laboratory experiments after simulated or natural herbivory events. Of the five mechanisms tested, we found that three mechanisms – availability of alternative resources, increased predation rate and reduction in sea urchin numbers – effectively dampened herbivory and worked as potential buffering mechanisms. In contrast, plant compensatory growth and resistance did not show clear responses. While compensatory growth and plant resistance are direct plant mechanism to tolerate the effects of herbivory, the rest are indirect mechanisms that begin with a modification of a plant trait (i.e. canopy height) that influences other species preference and/or behaviour, which in turn influences plant consumption. These adaptive behaviours may be a crucial and often overlooked factor in the remarkable resilience that Mediterranean seagrass ecosystems show to herbivory. [ABSTRACT FROM AUTHOR]

Predicted sea-level rise and increased storm frequency caused by climate change drastically threaten low-lying inhabited coral atolls. Coral reef frameworks are the atolls' primary defence from these changes. The growth and integrity of these frameworks is reflected in their carbonate budgets, a dynamic balance between biogenic accretion through coral growth and erosional forces, both of which are affected by factors acting at different spatial scales. We explored how carbonate budgets, estimated using the ReefBudget methodology, vary between three inhabited atolls that face differing anthropogenic stressors in the Lakshadweep Archipelago in the Northern Indian Ocean. We surveyed ten reefs, at two depths each, across the three islands. Overall, net carbonate budgets of reefs across all atolls were below optimal production rates needed to continue protecting shorelines (5 G measured on healthy reefs). This was a result of repeated mass bleaching events as well as local impacts. Carbonate production was influenced by a recent mass bleaching event in 2016 and a cyclone in 2018, and varied between depths and exposures, potentially due to differential recovery and mortality dynamics. Erosional processes were locally mediated with both urchin and parrotfish density showing large differences between islands, possibly linked to nutrient outflow and fishing intensity. We also find that by the year 2100, a large proportion of shallow sites will experience an increase in water depth above half a metre under moderate and high emissions scenarios, but none will breach this threshold under a low emissions scenario. Our results show that patterns of carbonate production were largely mediated by the history of global/regional disturbances, while erosional rates were much more dependent on local factors. [ABSTRACT FROM AUTHOR]

A colonial ascidian of the genus Distaplia caused a mass mortality of the pen shell Atrina maura (Sowerby, 1835) during June 2016 in the southwest of the Gulf of California (Mexico), with a significant socioeconomic cost. Tentatively identified in previous works as Distaplia cf. stylifera, a precise taxonomic determination was still lacking. In the present work, based on a detailed morphological study, it is confirmed that this aggressive species is Distaplia stylifera (Kowalevsky, 1874). Originally described from the Red Sea, the species currently has a wide circumtropical distribution (with the exception of the Eastern Pacific to date) and is reported as introduced in parts of its range. The present account thus represents an important range extension of this species. However, when revising the original description and later observations, the reported variability of several characters makes it likely that the binomen is in fact a complex of species, as is common in other ascidians with wide distributions. A complete morphological and genetic study including populations from the entire range of distribution would be necessary to settle the status of D. stylifera. Taxonomic uncertainties hinder a correct interpretation of biogeographical patterns and inference on the origin of the studied population. Nevertheless, the known introduction potential of the species, coupled with an explosive growth in an anthropized environment, and the lack of any previous reports in the Eastern Pacific, strongly suggest that the investigated population represents yet another instance of ascidian introduction. From the point of view of management, its invasive behavior is cause for great concern and warrants mitigation measures. [ABSTRACT FROM AUTHOR]

In the last three decades, quantitative approaches that rely on organism traits instead of taxonomy have advanced different fields of ecological research through establishing the mechanistic links between environmental drivers, functional traits, and ecosystem functions. A research subfield where trait-based approaches have been frequently used but poorly synthesized is the ecology of seagrasses; marine angiosperms that colonized the ocean 100M YA and today make up productive yet threatened coastal ecosystems globally. Here, we compiled a comprehensive trait-based response-effect framework (TBF) which builds on previous concepts and ideas, including the use of traits for the study of community assembly processes, from dispersal and response to abiotic and biotic factors, to ecosystem function and service provision. We then apply this framework to the global seagrass literature, using a systematic review to identify the strengths, gaps, and opportunities of the field. Seagrass trait research has mostly focused on the effect of environmental drivers on traits, i.e., “environmental filtering” (72%), whereas links between traits and functions are less common (26.9%). Despite the richness of trait-based data available, concepts related to TBFs are rare in the seagrass literature (15% of studies), including the relative importance of neutral and niche assembly processes, or the influence of trait dominance or complementarity in ecosystem function provision. These knowledge gaps indicate ample potential for further research, highlighting the need to understand the links between the unique traits of seagrasses and the ecosystem services they provide. [ABSTRACT FROM AUTHOR]

Abiotic environmental conditions can significantly influence the way species interact. In particular, plant–herbivore interactions can be substantially dependent on temperature and nutrients. The overall product of these relationships is critical for the fate and stability of vegetated ecosystems like marine forests. The last few decades have seen a rapid spread of barrens on temperate rocky reefs mainly as a result of overgrazing. The ecological feedbacks that characterize the barren state involve a different set of interactions than those occurring in vegetated habitats. Reversing these trends requires a proper understanding of the novel feedbacks and the conditions under which they operate. Here, we explored the role of a secondary herbivore in reinforcing the stability of barrens formed by sea urchin overgrazing under different nutrient conditions. Combining comparative and experimental studies in two Mediterranean regions characterized by contrasting nutrient conditions, we assessed: (i) if the creation of barren areas enhances limpet abundance, (ii) the size‐specific grazing impact by limpets, and (iii) the ability of limpets alone to maintain barrens. Our results show that urchin overgrazing enhanced limpet abundance. The effects of limpet grazing varied with nutrient conditions, being up to five times more intense under oligotrophic conditions. Limpets were able to maintain barrens in the absence of sea urchins only under low‐nutrient conditions, enhancing the stability of the depauperate state. Overall, our study suggests a greater vulnerability of subtidal forests in oligotrophic regions of the Mediterranean and demonstrates the importance of environment conditions in regulating feedbacks mediated by plant–herbivore interactions. [ABSTRACT FROM AUTHOR]

Climate-driven species redistributions are reshuffling the composition of marine ecosystems. How these changes alter ecosystem functions, however, remains poorly understood. Here we examine how impacts of herbivory change across a gradient of tropicalization in the Mediterranean Sea, which includes a steep climatic gradient and marked changes in plant nutritional quality and fish herbivore composition. We quantified individual feeding rates and behaviour of 755 fishes of the native Sarpa salpa, and non-native Siganus rivulatus and Siganus luridus. We measured herbivore and benthic assemblage composition across 20 sites along the gradient, spanning 30° of longitude and 8° of latitude. We coupled patterns in behaviour and composition with temperature measurements and nutrient concentrations to assess changes in herbivory under tropicalization. We found a transition in ecological impacts by fish herbivory across the Mediterranean from a predominance of seagrass herbivory in the west to a dominance of macroalgal herbivory in the east. Underlying this shift were changes in both individual feeding behaviour (i.e. food choice) and fish assemblage composition. The shift in feeding selectivity was consistent among temperate and warm-affiliated herbivores. Our findings suggest herbivory can contribute to the increased vulnerability of seaweed communities and reduced vulnerability of seagrass meadows in tropicalized ecosystems. [ABSTRACT FROM AUTHOR]

As invasive species spread, the ability of local communities to resist invasion depends on the strength of biotic interactions. Evolutionarily unused to the invader, native predators or herbivores may be initially wary of consuming newcomers, allowing them to proliferate. However, these relationships may be highly dynamic, and novel consumer–resource interactions could form as familiarity grows. Here, we explore the development of effective biotic resistance towards a highly invasive alga using multiple space‐for‐time approaches. We show that the principal native Mediterranean herbivore learns to consume the invader within less than a decade. At recently invaded sites, the herbivore actively avoided the alga, shifting to distinct preference and high consumptions at older sites. This rapid strengthening of the interaction contributed to the eventual collapse of the alga after an initial dominance. Therefore, our results stress the importance of conserving key native populations to allow communities to develop effective resistance mechanisms against invaders. [ABSTRACT FROM AUTHOR]

Summary: The prevalence of local adaptation and phenotypic plasticity among populations is critical to accurately predicting when and where climate change impacts will occur. Currently, comparisons of thermal performance between populations are untested for most marine species or overlooked by models predicting the thermal sensitivity of species to extirpation.Here we compared the ecological response and recovery of seagrass populations (Posidonia oceanica) to thermal stress throughout a year‐long translocation experiment across a 2800‐km gradient in ocean climate. Transplants in central and warm‐edge locations experienced temperatures > 29°C, representing thermal anomalies > 5°C above long‐term maxima for cool‐edge populations, 1.5°C for central and < 1°C for warm‐edge populations.Cool‐edge, central and warm‐edge populations differed in thermal performance when grown under common conditions, but patterns contrasted with expectations based on thermal geography. Cool‐edge populations did not differ from warm‐edge populations under common conditions and performed significantly better than central populations in growth and survival.Our findings reveal that thermal performance does not necessarily reflect the thermal geography of a species. We demonstrate that warm‐edge populations can be less sensitive to thermal stress than cooler, central populations suggesting that Mediterranean seagrasses have greater resilience to warming than current paradigms suggest. [ABSTRACT FROM AUTHOR]

Humans are rapidly transforming the structural configuration of the planet's ecosystems, but these changes and their ecological consequences remain poorly quantified in underwater habitats. Here, we show that the loss of forest‐forming seaweeds and the rise of ground‐covering 'turfs' across four continents consistently resulted in the miniaturization of underwater habitat structure, with seascapes converging towards flattened habitats with smaller habitable spaces. Globally, turf seascapes occupied a smaller architectural trait space and were structurally more similar across regions than marine forests, evidencing habitat homogenization. Surprisingly, such habitat convergence occurred despite turf seascapes consisting of vastly different species richness and with different taxa providing habitat architecture, as well as across disparate drivers of marine forest decline. Turf seascapes contained high sediment loads, with the miniaturization of habitat across 100s of km in mid‐Western Australia resulting in reefs retaining an additional ~242 million tons of sediment (four orders of magnitude more than the sediments delivered fluvially annually). Together, this work demonstrates that the replacement of marine forests by turfs is a generalizable phenomenon that has profound consequences for the ecology of temperate reefs. [ABSTRACT FROM AUTHOR]

Background: Classic ecological formulations of predator–prey interactions often assume that predators and prey interact randomly in an information-limited environment. In the field, however, most prey can accurately assess predation risk by sensing predator chemical cues, which typically trigger some form of escape response to reduce the probability of capture. Here, we explore under laboratory-controlled conditions the long-term (minutes to hours) escaping response of the sea urchin Paracentrotus lividus, a key species in Mediterranean subtidal macrophyte communities. Methods: Behavioural experiments involved exposing a random sample of P. lividus to either one of two treatments: (i) control water (filtered seawater) or (ii) predator-conditioned water (with cues from the main P. lividus benthic predator—the gastropod Hexaplex trunculus). We analysed individual sea urchin trajectories, computed their heading angles, speed, path straightness, diffusive properties, and directional entropy (as a measure of path unpredictability). To account for the full picture of escaping strategies, we followed not only the first instants post-predator exposure, but also the entire escape trajectory. We then used linear models to compare the observed results from control and predators treatments. Results: The trajectories from sea urchins subjected to predator cues were, on average, straighter and faster than those coming from controls, which translated into differences in the diffusive properties and unpredictability of their movement patterns. Sea urchins in control trials showed complex diffusive properties in an information-limited environment, with highly variable trajectories, ranging from Brownian motion to superdiffusion, and even marginal ballistic motion. In predator cue treatments, variability reduced, and trajectories became more homogeneous and predictable at the edge of ballistic motion. Conclusions: Despite their old evolutionary origin, lack of cephalization, and homogenous external appearance, the trajectories that sea urchins displayed in information-limited environments were complex and ranged widely between individuals. Such variable behavioural repertoire appeared to be intrinsic to the species and emerged when the animals were left unconstrained. Our results highlight that fear from predators can be an important driver of sea urchin movement patterns. All in all, the observation of anomalous diffusion, highly variable trajectories and the behavioural shift induced by predator cues, further highlight that the functional forms currently used in classical predator–prey models are far from realistic. [ABSTRACT FROM AUTHOR]

As species struggle to cope with rising ocean temperatures, temperate marine assemblages are facing major reorganization. Many benthic species have a brief but critical period dispersing through the plankton, when they are particularly susceptible to variations in temperature. Impacts of rising temperatures can thus ripple through the population with community‐wide consequences. However, responses are highly species‐specific, making it difficult to discern assemblage‐wide patterns in the life histories of different fish species.Here, we evaluate the responses to temperature in the early life histories of several fish species using otolith reconstructive techniques. We also assess the consequences of future warming scenarios to this assemblage.We sampled recent settlers of nine common species across a temperature gradient in the Mediterranean Sea and obtained environmental data for each individual. Using otolith microstructure, we measured early life traits including pelagic larval duration (PLD), growth rate, settlement size, hatching and settlement dates. We used a GLM framework to examine how environmental variables influenced early life‐history parameters.We show that increasing temperature results in considerable reduction in the dispersal potential of temperate fish. We find a nearly universal, assemblage‐wide decline in pelagic larval duration (PLD) of between 10% and 25%. This was because, with increasing temperature, larvae grew quicker to their settlement size. Settlement size itself was less affected by temperature and appears to be an ontogenetically fixed process.Given current estimates of ocean warming, there could be an assemblage‐wide reduction in larval dispersal of up to 50 km across the Mediterranean, reducing connectivity and potentially isolating populations as waters warm. [ABSTRACT FROM AUTHOR]

The global distribution of primary production and consumption by humans (fisheries) is well-documented, but we have no map linking the central ecological process of consumption within food webs to temperature and other ecological drivers. Using standardized assays that span 105° of latitude on four continents, we show that rates of bait consumption by generalist predators in shallow marine ecosystems are tightly linked to both temperature and the composition of consumer assemblages. Unexpectedly, rates of consumption peaked at midlatitudes (25 to 35°) in both Northern and Southern Hemispheres across both seagrass and unvegetated sediment habitats. This pattern contrasts with terrestrial systems, where biotic interactions reportedly weaken away from the equator, but it parallels an emerging pattern of a subtropical peak in marine biodiversity. The higher consumption at midlatitudes was closely related to the type of consumers present, which explained rates of consumption better than consumer density, biomass, species diversity, or habitat. Indeed, the apparent effect of temperature on consumption was mostly driven by temperature-associated turnover in consumer community composition. Our findings reinforce the key influence of climate warming on altered species composition and highlight its implications for the functioning of Earth's ecosystems. [ABSTRACT FROM AUTHOR]

The relative benefits of group foraging change as animals grow. Metabolic requirements, competitive abilities and predation risk are often allometric and influenced by group size. How individuals optimise costs and benefits as they grow can strongly influence consumption patterns. The shoaling fish Sarpa salpa is the principal herbivore of temperate Posidonia oceanica seagrass meadows. We used in-situ observations to describe how ontogeny influenced S. salpa individual feeding behaviour, shoaling behaviour and group foraging strategies, and its potential consequences to seagrass meadows. Shoaling was strongly influenced by body length: shoals were highly length-assorted and there was a clear positive relationship between body length and shoal size. Foraging strategies changed dramatically with shoal size. Small shoals foraged simultaneously and scattered over large areas. In contrast, larger shoals (made of larger individuals) employed a potentially cooperative strategy where individuals fed rotationally and focused in smaller areas for longer times (spot feeding). Thus, as individuals grew, they increased their potential impact as well, not merely because they consumed more, but because they formed larger shoals capable of considerably concentrating their grazing within the landscape. Our results indicate that ontogenetic shifts in group foraging strategies can have large ecosystem-wide consequences when the species is an important ecosystem modifier. [ABSTRACT FROM AUTHOR]

Harsh environmental conditions limit how species use the landscape, strongly influencing the way assemblages are distributed. In the wake of repeated coral bleaching mortalities in Lakshadweep, we examined how wave exposure influences herbivory in exposed and sheltered reefs. We used a combination of i. field observations of fish herbivore composition, abundance and activity across 6 exposed and 6 sheltered reefs; ii. experimental manipulations in a subset of these reefs (herbivore exclosures); and iii. opportunistic observations of fish recruitment, to determine how exposure influences herbivore biomass and herbivory. Species richness, biomass, abundance, total bite rates and species-specific per capita bite rates were lower in exposed compared to sheltered reefs, linked to strong environmental filtering of species composition, abundance and behaviour. For some critical species, this environmental filtering begins with differential recruitment and post-recruitment processes between exposures. Bite rates at sheltered sites were dominated by just a few species, most being laterally compressed surgeonfish that may find it difficult accessing or surviving in wave-battered shallow reefs. Exclosure experiments confirmed that exposed reefs were less controlled by herbivores than sheltered reefs. In post-disturbed reefs like Lakshadweep, environmental gradients appear to be key mediators of critical functions like herbivory by determining species composition, abundance and behaviour. [ABSTRACT FROM AUTHOR]

Herbivore outbreaks often trigger catastrophic overgrazing events in marine macrophyte ecosystems. The sea urchin Paracentrotus lividus, the dominant herbivore of shallow Mediterranean seascapes, is capable of precipitating shifts to barrens when its populations explode. Paracentrotus lividus is found ubiquitously in rocky macroalgal communities and in sandy seagrass meadows of Posidonia oceanica, two of the most important subtidal habitats in the Mediterranean. We explored if habitat‐specific regulation across the principal stages of the urchin life cycle could help explain the persistence of these populations in connected mosaics. We measured each of three relevant ecological processes (i.e. settlement, post‐settlement survival and predation) across a wide stretch of the Mediterranean coast (ca 600 km). Our results show that habitat‐specific regulation is critical in determining urchin populations: each habitat limited urchin sub‐populations at different life stages. Settlement was never limiting; urchins settled at similar rates in both habitats across the coast. Post‐settlement survival was a clear bottleneck, particularly in seagrass meadows where no juvenile urchins were recorded. Despite this bottleneck in seagrasses, adult urchin populations were very similar in both seagrass and macroalgal habitats indicating that other processes (potentially migration) could be key in determining adult distributions across the mosaic. The fact that population regulation is clearly habitat‐specific suggests that sea urchin populations may be significantly buffered from bottlenecks in mixed seascapes where both habitats co‐occur. Sea urchin populations can therefore persist across the seascape despite strong habitat‐specific regulation either by maintaining reproductive output in one habitat or by migrating between them. By affording these regulatory escapes to habitat‐modifying species, patchy mosaics may be much more prone to herbivore outbreaks and a host of cascading effects that come in their wake. [ABSTRACT FROM AUTHOR]

Abstract: By modifying how critical ecosystem functions are distributed across the landscape, the spatial configuration and characteristics of patches can play a strong role in structuring communities. In strongly predator‐controlled ecosystems, this patchy distribution of function can have complex downstream consequences, subjecting some areas to disproportionately high rates of predation, leaving other areas susceptible to herbivore outbreaks. In this study, we assess how spatial attributes at patch and landscape scales potentially influence the spatial and temporal distribution of predation on a functionally important herbivore in a patchy Mediterranean marine macrophyte community characterized by strong top‐down control. We experimentally tracked how predation risk of tethered sea urchins varied across space over a 10‐day period in a patchy seagrass meadow. We related these patterns with patch and landscape‐level attributes across the habitat mosaic. At the level of the patch, predation risk was the highest in seagrass patches with low canopies, without access to sheltering rocks. Scaling up to the landscape mosaic however, predation risk increased in dense aggregations of patches with high perimeter‐to‐area ratios close to rocky habitats. Predation aggregated in spatially explicit hotspots and coldspots that were maintained through time. Interestingly, this pattern of predation risk correlated well with the natural abundance of sea urchins. Our results show that spatial patch configuration can be a strong mediator of top trophic functions in marine ecosystems, causing significant clumping in the way predation—and therefore herbivory—are distributed across space. Given the importance of top‐down control for these shallow marine ecosystems, it is crucial to incorporate landscape attributes in understanding the impact of functionally important herbivores on highly fragmented habitats. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Predicting where state-changing thresholds lie can be inherently complex in ecosystems characterized by nonlinear dynamics. Unpacking the mechanisms underlying these transitions can help considerably reduce this unpredictability. We used empirical observations, field and laboratory experiments, and mathematical models to examine how differences in nutrient regimes mediate the capacity of macrophyte communities to sustain sea urchin grazing. In relatively nutrient-rich conditions, macrophyte systems were more resilient to grazing, shifting to barrens beyond 1 800 g m-2 (urchin biomass), more than twice the threshold of nutrient-poor conditions. The mechanisms driving these differences are linked to how nutrients mediate urchin foraging and algal growth: controlled experiments showed that low-nutrient regimes trigger compensatory feeding and reduce plant growth, mechanisms supported by our consumer=resource model. These mechanisms act together to halve macrophyte community resilience. Our study demonstrates that by mediating the underlying drivers, inherent conditions can strongly influence the buffer capacity of nonlinear systems. [ABSTRACT FROM AUTHOR]

Increasingly frequent and intense ocean warming events seriously test the buffer and recovery capacities of tropical coral reefs. Post-disturbance, available settlement structures on a reef (often dead coral skeletons) vary considerably in their mechanical stability and substrate composition, critically influencing coral recruit settlement choice and fate. In the wake of a coral mass mortality in the Lakshadweep archipelago, we examine (1) the relative availability of recruit settlement structures (from stable to unstable: reef platform, dead massive coral, consolidated rubble, dead corymbose coral, dead tabular coral, and unconsolidated rubble) in 12 recovering reefs across three atolls in the archipelago, (2) the substrate composition [crustose coralline algae (CCA), mixed turf, macroalgae] of these structural forms, and (3) whether the choice and fate of young coral are mediated by the substrate and stability of different structural forms. For this, we measured the abundance and distribution of recruit (<1 cm), juvenile (1-5 cm), and young adult (5-10) corals of 24 common coral genera. Four years after the mass mortality, reefs differed considerably in composition of settlement structures. The structures themselves varied significantly in substrate cover with dead tables largely covered in CCA [60 ± 6.05 % (SE)] and dead corymbose coral dominated by mixed turf (61.83 ± 3.8 %). The youngest visible recruits (<1 cm) clearly preferred CCA-rich structures such as dead massives and tables. However, older size classes were rarely found on unstable structures (strongly 'avoiding' tables, Ivlev's electivity index, E = −0.5). Our results indicate that while substrate cover might mediate coral choice, the mechanical stability of settlement structures is critical in determining post-settlement coral survival. The composition and availability of settlement structures on a reef may serve as a characteristic signature of its recovery potential, aiding in assessments of reef resilience. [ABSTRACT FROM AUTHOR]

While large herbivores can have strong impacts on terrestrial ecosystems, much less is known of their role in aquatic systems. We reviewed the literature to determine: 1) which large herbivores (> 10 kg) have a (semi-)aquatic lifestyle and are important consumers of submerged vascular plants, 2) their impact on submerged plant abundance and species composition, and 3) their ecosystem functions. We grouped herbivores according to diet, habitat selection and movement ecology: 1) Fully aquatic species, either resident or migratory (manatees, dugongs, turtles), 2) Semi-aquatic species that live both in water and on land, either resident or migratory (swans), 3) Resident semi-aquatic species that live in water and forage mainly on land (hippopotamuses, beavers, capybara), 4) Resident terrestrial species with relatively large home ranges that frequent aquatic habitats (cervids, water buffalo, lowland tapir). Fully aquatic species and swans have the strongest impact on submerged plant abundance and species composition. They may maintain grazing lawns. Because they sometimes target belowground parts, their activity can result in local collapse of plant beds. Semi-aquatic species and turtles serve as important aquatic-terrestrial linkages, by transporting nutrients across ecosystem boundaries. Hippopotamuses and beavers are important geomorphological engineers, capable of altering the land and hydrology at landscape scales. Migratory species and terrestrial species with large home ranges are potentially important dispersal vectors of plant propagules and nutrients. Clearly, large aquatic herbivores have strong impacts on associated species and can be critical ecosystem engineers of aquatic systems, with the ability to modify direct and indirect functional pathways in ecosystems. While global populations of large aquatic herbivores are declining, some show remarkable local recoveries with dramatic consequences for the systems they inhabit. A better understanding of these functional roles will help set priorities for the effective management of large aquatic herbivores along with the plant habitats they rely on. [ABSTRACT FROM AUTHOR]

Spatial and temporal heterogeneity in flowering occur in many plant species with abiotic pollination and may confer fitness advantages through mechanisms such as predator satiation or pollination efficiency. Environmental factors such as light quality or quantity and temperature play an important role in inducing synchronization on wide geographic scales. On a smaller geographic scale, external factors such as resource availability and herbivory are theorized to trigger flowering, while genetic factors may also play an important role., In this study, we assessed the importance of ecological and genetic factors in shaping seascape-level spatial heterogeneity in flowering of the seagrass Posidonia oceanica. By investigating spatially close sites (<20 km) with similar seascape configurations and depth, we assume that major environmental drivers (temperature and light) were equivalent., We assessed four ecological factors (productivity, leaf nitrogen and carbon content and herbivory) and three genetic factors (heterozygosity, relatedness and clonality) to assess three hypotheses for synchronized flowering in P. oceanica: (i) clone synchronization (internal clock hypothesis), (ii) variation in nutrient availability, potentially caused by spatial heterogeneity in herbivory rates or nutrient translocation via clonal integration (resource budget hypothesis) or (iii) kin selection and sibling synchronization., Internal relatedness and heterozygosity had a significant positive effect on the abundance of flowers. Moreover, productivity and genotypic richness (clonality) were negatively associated with flower density, although at a lower level of significance. In addition, we found that clones were almost exclusively shared among mass-flowering patches and patches without mass-flowering, respectively., Synthesis. The results shed new light on seagrass flowering patterns and on the mechanisms of flower synchronization at the patch level within a wider spatial scale. We found support for the kin selection hypothesis and indirect evidence for the resource budget hypothesis. Thus, a combination of mainly genetic but also ecological factors causes the observed heterogeneous flowering patterns in Posidonia oceanica seascapes. In addition, we found a strong positive relationship between the number of flowers and heterozygosity, adding evidence to the controversial association between heterozygosity and fitness when a limited number of loci are used. To our knowledge, this study is the first to link both ecological and genetic factors with flower abundance in a species with a presumed masting strategy. [ABSTRACT FROM AUTHOR]

Herds of dugong, a largely tropical marine megaherbivore, are known to undertake long-distance movements, sequentially overgrazing seagrass meadows in their path. Given their drastic declines in many regions, it is unclear whether at lower densities, their grazing is less intense, reducing their need to travel between meadows. We studied the effect of the feeding behaviour of a small dugong population in the Andaman and Nicobar archipelago, India to understand how small isolated populations graze seagrasses. In the seven years of our observation, all recorded dugongs travelled either solitarily or in pairs, and their use of seagrasses was limited to 8 meadows, some of which were persistently grazed. These meadows were relatively large, contiguous and dominated by short-lived seagrasses species. Dugongs consumed approximately 15% of meadow primary production, but there was a large variation (3–40% of total meadow production) in consumption patterns between meadows. The impact of herbivory was relatively high, with shoot densities c. 50% higher inside herbivore exclosures than in areas exposed to repeated grazing. Our results indicate that dugongs in the study area repeatedly graze the same meadows probably because the proportion of primary production consumed reduces shoot density to levels that are still above values that can trigger meadow abandonment. This ability of seagrasses to cope perhaps explains the long-term site fidelity shown by individual dugongs in these meadows. The fact that seagrass meadows in the archipelago are able to support dugong foraging requirements allows us to clearly identify locations where this remnant population persists, and where urgent management efforts can be directed. [ABSTRACT FROM AUTHOR]

To sustainably manage naturally scarce resources, island communities often evolve complex mechanisms including customary laws, belief systems, and reciprocity arrangements among others, to prevent overharvest. Their effectiveness depends largely on the extent to which resource users comply with the rules. We examined patterns of compliance with traditional marine management in the Nicobar Archipelago, India, before, and six years after, the 2004 tsunami. We used interview-based surveys to document marine harvest regulations, and changes in compliance patterns. Our results indicate that pre-tsunami, complex harvest rules existed, including spatio-temporal closures, gear restrictions and species bans; many reefs were subject to multiple, overlapping restrictions. Post-tsunami, compliance weakened considerably; younger individuals (19-35 years) and individuals receiving tsunami aid (boats, gear, etc.) were the most likely non-compliers. Around 84 % of interviewees attributed declining compliance directly to post-tsunami changes in resource availability and a perceived decline in traditional authority. Changes in resource availability can interact strongly with institutional decline, eroding the resilience of traditional management. [ABSTRACT FROM AUTHOR]

Since Gleason and Clements, our understanding of community dynamics has been influenced by theories emphasising either dispersal or niche assembly as central to community structuring. Determining the relative importance of these processes in structuring real-world communities remains a challenge. We tracked reef fish community reassembly after a catastrophic coral mortality in a relatively unfished archipelago. We revisited the stochastic model underlying MacArthur and Wilson's Island Biogeography Theory, with a simple extension to account for trophic identity. Colonisation and extinction rates calculated from decadal presence-absence data based on (1) species neutrality, (2) trophic identity and (3) site-specificity were used to model post-disturbance reassembly, and compared with empirical observations. Results indicate that species neutrality holds within trophic guilds, and trophic identity significantly increases overall model performance. Strikingly, extinction rates increased clearly with trophic position, indicating that fish communities may be inherently susceptible to trophic downgrading even without targeted fishing of top predators. [ABSTRACT FROM AUTHOR]

The functioning of ecosystems can be strongly driven by landscape attributes. Despite its importance, however, our understanding of how landscape influences ecosystem function derives mostly from species richness and abundance patterns, with few studies assessing how these relate to actual functional rates., We examined the influence of landscape attributes on the rates of herbivory in seagrass meadows, where herbivory has been identified as a key process structuring these relatively simple systems. The study was conducted in three representative Posidonia oceanica meadows. The principal herbivores in these meadows are the fish Sarpa salpa and the sea urchin Paracentrotus lividus, and we hypothesized that differences in their interaction with landscape attributes would significantly influence herbivory rates., We measured herbivore abundance, herbivory rates, primary production and plant quality (C:N) in seagrass patches embedded either in rock or in sand (matrix attribute), in patches either near or far from a rocky reef (distance attribute) and at the edges and interior of meadows., Our results show that matrix and meadow edges significantly affected the actual levels of herbivory. Herbivory rates were higher in seagrass patches embedded in a rocky matrix compared to those on sand, and herbivory at the centre of seagrass meadows was higher than at the edges. In contrast, patch distance to rocky reefs did not affect herbivory. Neither herbivore abundance nor food quality explained the patterns across different landscape attributes. This suggests that variation in herbivory across the landscape may be related much more to behavioural differences between species in their evaluation of risk, movement and food preference in relation to the landscape structure., Our results indicate that richness and abundance patterns may mask critical interactions between landscape attributes and species responses, which result in considerable heterogeneity in the way key functional processes like herbivory are distributed across the ecosystem mosaic. [ABSTRACT FROM AUTHOR]

Structural complexity strongly influences the outcome of predator-prey interactions in benthic marine communities affecting both prey concealment and predator hunting efficacy. How habitat structure interacts with species-specific differences in predatory style and antipredatory strategies may therefore be critical in determining higher trophic functions. We examined the role of structural complexity in mediating predator-prey interactions across several macrophyte habitats along a gradient of structural complexity in three different bioregions: western Mediterranean Sea (WMS), eastern Indian Ocean (EIO) and northern Gulf of Mexico (NGM). Using sea urchins as model prey, we measured survival rates of small (juveniles) and medium (young adults) size classes in different habitat zones: within the macrophyte habitat, along the edge and in bare sandy spaces. At each site we also measured structural variables and predator abundance. Generalised linear models identified biomass and predatory fish abundance as the main determinants of predation intensity but the efficiency of predation was also influenced by urchin size class. Interestingly though, the direction of structure-mediated effects on predation risk was markedly different between habitats and bioregions. In WMS and NGM, where predation by roving fish was relatively high, structure served as a critical prey refuge, particularly for juvenile urchins. In contrast, in EIO, where roving fish predation was low, predation was generally higher inside structurally complex environments where sea stars were responsible for much of the predation. Larger prey were generally less affected by predation in all habitats, probably due to the absence of large predators. Overall, our results indicate that, while the structural complexity of habitats is critical in mediating predator-prey interactions, the direction of this mediation is strongly influenced by differences in predator composition. Whether the regional pool of predators is dominated by visual roving species or chemotactic benthic predators may determine if structure dampens or enhances the influence of top-down control in marine macrophyte communities. [ABSTRACT FROM AUTHOR]

Prioritizing efforts for conserving rare and threatened species with limited past data and lacking population estimates is predicated on robust assessments of their occupancy rates. This is particularly challenging for elusive, long-lived and wide-ranging marine mammals. In this paper we estimate trends in long-term (over 50years) occupancy, persistence and extinction of a vulnerable and data-poor dugong (Dugong dugon) population across multiple seagrass meadows in the Andaman and Nicobar archipelago (India). For this we use hierarchical Bayesian dynamic occupancy models accounting for false negatives (detection probability<1), persistence and extinction, to two datasets: a) fragmentary long-term occurrence records from multiple sources (1959–2004, n = 40 locations), and b) systematic detection/non-detection data from current surveys (2010–2012, n = 57). Dugong occupancy across the archipelago declined by 60% (from 0.45 to 0.18) over the last 20 years and present distribution was largely restricted to sheltered bays and channels with seagrass meadows dominated by Halophila and Halodule sp. Dugongs were not found in patchy meadows with low seagrass cover. In general, seagrass habitat availability was not limiting for dugong occupancy, suggesting that anthropogenic factors such as entanglement in gillnets and direct hunting may have led to local extinction of dugongs from locations where extensive seagrass meadows still thrive. Effective management of these remnant dugong populations will require a multi-pronged approach, involving 1) protection of areas where dugongs still persist, 2) monitoring of seagrass habitats that dugongs could recolonize, 3) reducing gillnet use in areas used by dugongs, and 4) engaging with indigenous/settler communities to reduce impacts of hunting. [ABSTRACT FROM AUTHOR]

Historical declines of marine megaherbivores have led to a view of seagrass communities structured largely by abiotic disturbance and plant competition. There is, however, growing recognition of the significance of top-down control through herbivory, on seagrass ecosystem processes, raising the question of how meadows functioned under historically high populations of megaherbivores., We assess the impacts of such intense herbivory on seagrass meadow composition in the Lakshadweep islands (India), where high-density feeding aggregations of green turtles have persisted for over a decade. We use a series of complementary approaches: (i) natural herbivory exclosures (ii) published data on seagrass composition before and after turtles established (at one atoll: Agatti) and (iii) present species composition along a turtle herbivory gradient over multiple atolls., Long-term natural exclosures in Agatti indicated that sustained turtle grazing caused clear shifts in species dominance from the long-lived, higher-successional Thalassia hemprichii to the relatively short-lived, pioneering species Cymodocea rotundata (dominant in grazed areas). T. hemprichii was the dominant species c. 20 years ago but is now restricted to areas within exclosures in Agatti, and to the least grazed meadows (<5%) in other atolls., We conducted field experiments to identify possible mechanisms by which herbivory mediated direct or apparent competitive interactions between seagrass species. To verify if grazing reduced growth rates of T. hemprichii in comparison with C. rotundata, we conducted clipping experiments in 1m2 plots, simulating turtle herbivory on equal shoot proportions of both species. After 4 months, T. hemprichii shoot density showed major declines in clipped vs. control plots, but C. rotundata shoot density remained relatively unaffected., To test whether selective grazing on T. hemprichii facilitated C. rotundata, we established paired seagrass preference experiments. Turtles had clear preferences for T. hemprichii (64%), but also grazed on C. rotundata., Synthesis. Taken together, our results show that high-impact turtle herbivory changes seagrass composition, precipitating dominance shifts in grazed meadows by mediating direct and apparent competition. Given the crucial role of megaherbivores in seagrass meadow functioning, our results suggest that past meadows may have had natural functional limits to megaherbivore densities that they could sustainably support. [ABSTRACT FROM AUTHOR]

Catastrophic storms have been observed to be one of the major elements in shaping the standing structure of marine benthic ecosystems. Yet, little is known about the effect of catastrophic storms on ecosystem processes. Specifically, herbivory is the main control mechanism of macrophyte communities in the Mediterranean, with two main key herbivores: the sea urchin Paracentrotus lividus and the fish Sarpa salpa. Consequently, the effects of extreme storm events on these two herbivores (at the population level and on their behaviour) may be critical for the functioning of the ecosystem. With the aim of filling this gap, we took advantage of two parallel studies that were conducted before, during and after an unexpected catastrophic storm event. Specifically, fish and sea urchin abundance were assessed before and after the storm in monitored fixed areas (one site for sea urchin assessment and 3 sites for fish visual transects). Additionally, we investigated the behavioural response to the disturbance of S. salpa fishes that had been tagged with acoustic transmitters. Given their low mobility, sea urchins were severely affected by the storm (ca. 50% losses) with higher losses in those patches with a higher density of sea urchins. This may be due to a limited availability of refuges within each patch. In contrast, fish abundance was not affected, as fish were able to move to protected areas (i.e. deeper) as a result of the high mobility of this species. Our results highlight that catastrophic storms differentially affect the two dominant macroherbivores of rocky macroalgal and seagrass systems due to differences in mobility and escaping strategies. This study emphasises that under catastrophic disturbances, the presence of different responses among the key herbivores of the system may be critical for the maintenance of the herbivory function. [ABSTRACT FROM AUTHOR]

Identifying and quantifying the factors that contribute to the potential misclassification of the ecological status of water bodies is a major challenge of the Water Framework Directive (WFD). The present study compiles extensive biomonitoring data from a range of macrophyte-based classification methods developed by several European countries. The data reflect spatial and temporal variation as well as inter-observer variation. Uncertainty analysis identified that factors related to the spatial scale of sampling generally contributed most to the uncertainty in classifying water bodies to their ecological status, reflecting the high horizontal and depth-related heterogeneity displayed by macrophyte communities. In contrast, the uncertainty associated with temporal variation was low. In addition, inter-observer variation, where assessed, did not contribute much to overall uncertainty, indicating that these methods are easily transferable and insensitive to observer error. The study, therefore, suggests that macrophyte-based sampling schemes should prioritize large spatial replication over temporal replication to maximize the effectiveness and reliability of water body classification within the WFD. We encourage conducting similar uncertainty analyses for new/additional ecological indicators to optimize sampling schemes and improve the reliability of classification of ecological status. [ABSTRACT FROM AUTHOR]

Seagrasses are key components of coastal marine ecosystems and many monitoring programmes worldwide assess seagrass health and apply seagrasses as indicators of environmental status. This study aims at identifying the diversity and characteristics of seagrass indicators in use within and across European ecoregions in order to provide an overview of seagrass monitoring effort in Europe. We identified 49 seagrass indicators used in 42 monitoring programmes and including a total of 51 metrics. The seagrass metrics represented 6 broad categories covering different seagrass organizational levels and spatial scales. The large diversity is particularly striking considering that the pan-European Water Framework Directive sets common demands for the presence and abundance of seagrasses and related disturbance-sensitive species. The diversity of indicators reduces the possibility to provide pan-European overviews of the status of seagrass ecosystems. The diversity can be partially justified by differences in species, differences in habitat conditions and associated communities but also seems to be determined by tradition. Within each European region, we strongly encourage the evaluation of seagrass indicator-pressure responses and quantification of the uncertainty of classification associated to the indicator in order to identify the most effective seagrass indicators for assessing ecological quality of coastal and transitional water bodies. [ABSTRACT FROM AUTHOR]

The dominant paradigm of top-down control of ecological communities through direct consumption pathways is giving way to a more nuanced understanding of trophic interactions with the recognition that subtler indirect effects can often play an important role in structuring communities. Direct and indirect trophic and non-trophic processes could well be acting simultaneously within the same food web, and their overall effects may even interact with each other., We studied indirect interactions in a relatively simple trophic system comprising a single producer, two herbivores and a guild of predators. In particular, we assessed whether (i) the principal herbivore fish, Sarpa salpa, is capable of modifying a seagrass trait, canopy height, by grazing and (ii) whether grazing-induced habitat alteration can trigger an environment-mediated modification of the interaction between herbivorous sea urchins Paracentrotus lividus and their predators., We tested these hypotheses with field experiments including fish herbivore-exclusion experiments (to examine the ability of S. salpa to modify seagrass habitats) and predation experiments using tethered sea urchins in a meadow with varying canopy heights (to test whether habitat modification can mediate urchin predation risk)., The effect of S. salpa herbivory was highly significant. Canopy height in herbivore-excluded plots was more than 3·5 times higher than in uncaged control plots. In addition, adult sea urchin predation risk in the most highly grazed plots was 4-5 times higher than plots with higher canopy heights. In contrast, predation risk on juvenile urchins was not influenced by canopy height., Our results show that predation pressure on a key herbivore can be modified both by the environmental context within which it finds itself and by the actions of another herbivore that modifies the plant traits that create this environmental context. These act as two discrete pathways that interact in potentially nonlinear ways, mediating top-down control in these ecosystems. Herbivores, particularly when acting as ecosystem engineers, may have the potential to mediate and increase predation risk, as they substantially modify habitat structure, with consequences for refuge availability, among others. Assessing these indirect interactions is not just important to understand the functioning of the system itself but may have important management and conservation consequences. [ABSTRACT FROM AUTHOR]

The European Water Framework Directive commits partner countries to evolve uniform protocols for monitoring the environmental condition of natural water bodies, crucially integrating biological and ecological criteria from the associated ecosystems. This has encouraged considerable research on the development of bioindicator-based systems of water quality monitoring. A critical step towards this end is providing evidence that the proposed bioindicator system adequately reflects the human pressures to which a specific water body is submitted. Here we investigate the utility of pulse-amplitude-modulated (PAM) fluorometry, a fast, non-destructive and increasingly popular bioindicator-based method, in assessing water quality based on the widespread Mediterranean seagrass Posidonia oceanica, an important constituent of submersed benthic vegetation. Specifically, we evaluated the ability of PAM to discriminate between sites along a pre-established gradient of anthropogenic pressures and the consistency and reliability of PAM parameters across spatial scales. Our results show that the maximum quantum yield (Fv/Fm), representing the structural photosynthetic efficiency of the plant, responds significantly to the degree of site-level anthropogenic pressure. However, Fv/Fm values in our study increased with increasing pressure, in striking contrast with other studies that report declines in Fv/Fm values with increasing stress. A potential explanation for this discrepancy is that our study sites were influenced by multiple diffuse stressors (characteristic of most coastal waters) that could potentially interact with each other to influence Fv/Fm values in often unpredictable ways. The photosynthetic variables calculated from rapid light curves (ETR, maximum electron transport rate; α, initial slope of the curve; I, saturating irradiance), which represent an instant picture of the photosynthetic activity of the plant, were unable to clearly discriminate between sites subject to different anthropogenic pressures due to considerable small-scale variability. Taken together, these results suggest that even though PAM fluorometry may be a good candidate tool for monitoring water bodies in terms of costs and applicability, considerably more needs to be understood about how its parameters respond to real-world stressors, particularly when they act in concert with each other. With our present understanding of seagrass photosynthetic responses to anthropogenic stress, it would be ill advised to employ PAM as anything but a complementary tool to validate environmental stress derived with other, more robust methodologies. [ABSTRACT FROM AUTHOR]

Demographic processes exert different degrees of control as individuals grow, and in species that span several habitats and spatial scales, this can influence our ability to predict their population at a particular life-history stage given the previous life stage. In particular, when keystone species are involved, this relative coupling between demographic stages can have significant implications for the functioning of ecosystems. We examined benthic and pelagic abundances of the sea urchin Paracentrotus lividus in order to: 1) understand the main life-history bottlenecks by observing the degree of coupling between demographic stages; and 2) explore the processes driving these linkages. P. lividus is the dominant invertebrate herbivore in the Mediterranean Sea, and has been repeatedly observed to overgraze shallow beds of the seagrass Posidonia oceanica and rocky macroalgal communities. We used a hierarchical sampling design at different spatial scales (100 s, 10 s and <1 km) and habitats (seagrass and rocky macroalgae) to describe the spatial patterns in the abundance of different demographic stages (larvae, settlers, recruits and adults). Our results indicate that large-scale factors (potentially currents, nutrients, temperature, etc.) determine larval availability and settlement in the pelagic stages of urchin life history. In rocky macroalgal habitats, benthic processes (like predation) acting at large or medium scales drive adult abundances. In contrast, adult numbers in seagrass meadows are most likely influenced by factors like local migration (from adjoining rocky habitats) functioning at much smaller scales. The complexity of spatial and habitat-dependent processes shaping urchin populations demands a multiplicity of approaches when addressing habitat conservation actions, yet such actions are currently mostly aimed at managing predation processes and fish numbers. We argue that a more holistic ecosystem management also needs to incorporate the landscape and habitat-quality level processes (eutrophication, fragmentation, etc.) that together regulate the populations of this keystone herbivore. [ABSTRACT FROM AUTHOR]

Major coastal storms, associated with strong winds, high waves and intensified currents, and occasionally with heavy rains and flash floods, are mostly known because of the serious damage they can cause along the shoreline and the threats they pose to navigation. However, there is a profound lack of knowledge on the deep-sea impacts of severe coastal storms. Concurrent measurements of key parameters along the coast and in the deep-sea are extremely rare. Here we present a unique data set showing how one of the most extreme coastal storms of the last decades lashing the Western Mediterranean Sea rapidly impacted the deep-sea ecosystem. The storm peaked the 26th of December 2008 leading to the remobilization of a shallow-water reservoir of marine organic carbon associated with fine particles and resulting in its redistribution across the deep basin. The storm also initiated the movement of large amounts of coarse shelf sediment, which abraded and buried benthic communities. Our findings demonstrate, first, that severe coastal storms are highly efficient in transporting organic carbon from shallow water to deep water, thus contributing to its sequestration and, second, that natural, intermittent atmospheric drivers sensitive to global climate change have the potential to tremendously impact the largest and least known ecosystem on Earth, the deep-sea ecosystem. [ABSTRACT FROM AUTHOR]

Herbivory is now recognized as an important structuring agent in seagrass meadows but the attack pattern and tissue damage of consumers are highly variable. Nutritional preferences of herbivores and/or easy access to resources may cause differences in biomass loss among tissues that damage the plant in functionally distinctive ways. The two main Mediterranean herbivores, the fish Sarpa salpa (L.) and the sea urchin Paracentrotus lividus (Lmk.), remove higher amounts of intermediate and external shoot leaves, respectively. To test whether this selective feeding can have different consequences on the allocation patterns of nutrient within plants, we simulated the effect of both herbivores by clipping external and intermediate leaves (plus unclipped controls) of Posidonia oceanica (L.) and we measured plant tolerance in terms of shoot growth and leaf nutrient supply to new tissue using isotopic markers. As expected, control treatments displayed high carbon and nutrient supply from external leaves (83% of the total N and 84% of the total C incorporated by the shoot). When subjected to clipping, the remaining leaves enhanced carbon and nitrogen supply compared with the control by 16% of N and 36% of C-in the intermediate clipping-and by over 100% of N and 200% of C-in the external clipping-to compensate for the nutrient lost. However, only in the case of fish herbivory (intermediate clipping), enhanced supply alone was able to fully compensate for the nutrient losses. In contrast, this mechanism is not completely effective when external leaves are clipped (urchin herbivory). Yet, the consequences of this nutrient loss under sea urchin herbivory are not apparent from the nutrient content of the new tissue, suggesting that there are other sources of nitrogen income (uptake or reallocation from rhizomes). Our study does not only confirm the tolerance of P. oceanica to herbivory, but also constitutes the first evidence of leaf-specific, compensatory nutrient supply in seagrasses. [ABSTRACT FROM AUTHOR]

Within-plant variation in susceptibility to herbivory can significantly influence the ecological and evolutionary consequences of plant-herbivore interactions. Seagrasses are marine angiosperms characterised by substantial intra-individual differences in multiple traits, such as nutrients, chemical and structural defences and epibiotic load, all of which can strongly influence herbivore preferences. We quantified the within-plant feeding choices of the two main consumers of the temperate seagrass Posidonia oceanica--the fish Sarpa salpa and the sea urchin Paracentrotus lividus--and determined the plant traits that explained their foraging strategies. We found strong within-plant heterogeneity in both seagrass susceptibility to herbivory and chemical composition, but different consumers exhibited contrasting feeding choices. S. salpa preferred the most nutritious and chemically defended younger leaves, suggesting a full adaptation to consuming this macrophyte and a greater impact of this herbivore on the plant. In contrast, P. lividus consistently preferred the older leaves covered by epibionts, probably attenuating the relative impact of this consumer and helping to explain the weak effects usually recorded for this echinoid in undisturbed meadows. Artificial diet experiments showed that morphology and fine-scale structural defences were the primary determinant of urchin feeding choices, with nutrient content and chemical defences being of secondary importance. Epibiosis did not strongly influence fish feeding, but it did have a strong 'shared-doom' effect on urchin consumption. This effect was driven by a distinct preference towards a mixed diet that included both host tissues and their epibiotic community. [ABSTRACT FROM AUTHOR]

Biotic indices have become key assessment tools in most recent national and trans-national policies aimed at improving the quality of coastal waters and the integrity of their associated ecosystems. In this study we analyzed 90 published biotic indices, classified them into four types, and analyzed the strengths and weaknesses of each type in relation to the requirements of these policies. We identified three main type-specific weaknesses. First, the problems of applicability, due to practical and conceptual difficulties, which affect most indices related to ecosystem function. Second, the failure of many indices based on structural attributes of the community (e.g.taxonomic composition) to link deterioration with causative stressors, or to provide an early-detection capacity. Third, the poor relevance to the ecological integrity of indices based on attributes at the sub-individual level (e.g.multi-biomarkers). Additionally, most indices still fail on two further aspects: the broad-scale applicability and the definition of reference conditions. Nowadays, the most promising approach seems to be the aggregation of indices with complementary strengths, and obtained from different biological communities. [ABSTRACT FROM AUTHOR]

Nitrogen inputs to coastal environments can considerably alter the abundance of primary producers. However, how herbivores modify their trophic signatures and adjust to changes in food resource conditions remains controversial. Here, we assess the effect of nutrient availability on the diet shifts of the two main Mediterranean herbivores, the Sparid fish Sarpa salpa L. and the sea urchin Paracentrotus lividus (Lmk.) that feed mostly on the seagrass Posidonia oceanica L. (Delile), epiphytes and benthic macroalgae. To do this, we (1) investigate the patterns of isotopic composition (δ13C and δ15N signatures) of the two herbivores and their potential food sources in three areas of contrasting nutrient conditions and, (2) we assess the diet shift along this nutrient gradient by estimating the isotopic nutrient enrichment (i.e., the contribution of δ13C and δ15N signatures in consumers’ tissues relative to potential food sources). Food web signatures of δ13C were similar among the three study sites, and no patterns of δ13C shift were observed in their diets. In contrast, there was a consistent increase in N contents and δ15N along the nutrient gradient for all primary producers and their consumers. The rate of δ15N enrichment was also clearly distinctive between the two herbivores: in P. lividus it increased by 61% along the nutrient gradient, while in S. salpa it remained constant. Our results suggest that sea urchins behave as facultative omnivores and feed on vegetable or mixed diets depending on the trophic status of the system. It is unclear, however, if this modification is behavioral or the consequence of mere changes in the availability of food items, as animal epiphytes (e.g., hydrozoans, bryozoans and ascidians) can also became more abundant on seagrass leaves under increased nutrient conditions. In contrast, adult fish appear to feed on vegetal material independent of nutrient availability in the ecosystem. [ABSTRACT FROM AUTHOR]

Herbivory can induce changes in plant traits that may involve both tolerance mechanisms that compensate for biomass loss and resistance traits that reduce herbivore preference. Seagrasses are marine vascular plants that possess many attributes that may favour tolerance and compensatory growth, and they are also defended with mechanisms of resistance such as toughness and secondary metabolites. We quantified phenotypic changes induced by herbivore damage on the temperate seagrass Posidonia oceanica in order to identify specific compensatory and resistance mechanisms in this plant, and to assess any potential trade-offs between these two strategies of defence. We simulated three natural levels of fish herbivory by repeatedly clipping seagrass leaves during the summer period of maximum herbivory. Compensatory responses were determined by measuring shoot-specific growth, photosynthetic rate, and the concentration of nitrogen and carbon resources in leaves and rhizomes. Induced resistance was determined by measuring the concentration of phenolic secondary metabolites and by assessing the long-term effects of continued clipping on herbivore feeding preferences using bioassays. Plants showed a significant ability to compensate for low and moderate losses of leaf biomass by increasing aboveground growth of damaged shoots, but this was not supported by an increase in photosynthetic capacity. Low levels of herbivory induced compensatory growth without any measurable effects on stored resources. In contrast, nitrogen reserves in the rhizomes played a crucial role in the plant’s ability to compensate and survive herbivore damage under moderate and high levels of herbivory, respectively. We found no evidence of inducibility of long-term resistance traits in response to herbivory. The concentration of phenolics decreased with increasing compensatory growth despite all treatments having similar carbon leaf content, suggesting reallocation of these compounds towards primary functions such as cell-wall construction. [ABSTRACT FROM AUTHOR]

Plant–herbivore interactions have strong ecological and evolutionary consequences, but have been traditionally overlooked in marine higher plants. Despite recent advances in seagrass ecology that highlight the importance of herbivory, the mechanisms that regulate the feeding behaviour of seagrass consumers remain largely unknown. Herbivores have been shown to reduce the sexual reproductive success of seagrasses through direct consumption of inflorescences and seeds, but we know little about intraspecific variation in susceptibility to grazing of different seagrass tissues. We contrasted the relative palatability of reproductive and vegetative tissues of the temperate seagrass Posidonia oceanica in the field, and we assessed the feeding preferences among these tissues of the main consumers of the plant, the fish Sarpa salpa and the urchin Paracentrotus lividus. Moreover, we identified the plant traits that explained the observed feeding behaviour. We provide strong evidence for herbivore selectivity among seagrass tissues. In the field, 70–90% of inflorescences were damaged by herbivores compared to 3–60% of leaves of similar age. In feeding assays, the urchin P. lividus showed over a twofold preference for reproductive tissue at various stages of development. By contrast, we detected no feeding activity on either leaves or inflorescences from the fish S. salpa, which is known to migrate to deeper waters soon after flowering starts and during the period of fruit maturation. Despite being the preferred food of urchins, inflorescences were chemically defended, had higher levels of phenolics and lower nutrient and calorific content than leaves. We experimentally demonstrated that leaf structural defences are the primary factor in determining urchin feeding preferences. Removal of plant structure results in a drastic shift in urchin selectivity towards the most nutritious and less chemically defended leaf tissue, indicating that multiple mechanisms of defence to herbivory may coexist in seagrasses. [ABSTRACT FROM AUTHOR]

Lophocladia lallemandii, an Indo-Pacific alga that is currently invading several Mediterranean areas, is especially aggressive when colonizing meadows of Posidonia oceanica in Formentera (Balearic Islands). L. lallemandii settles on rhizomes and old leaves situated mainly at meadow edges and in low density Posidonia patches, growing rapidly, producing disc-like holdfasts along the thalli that allow formation of a mat of the red algal filaments intermingled with P. oceanica leaves. This mat can get so thick and dense that new leaves become confined inside the web of red algal filaments. The seagrass leaves display chlorosis and sometimes die. Invasion by L. lallemandii significantly reduces seagrass shoot size, leaf biomass, and percentage of living shoots. A lower content of sucrose in shoots from invaded plots points to a reduction of photosynthetic activity. However, other causes such as enhanced sediment trapping, organic and nutrient enrichment inside the algal carpet, and increased oxygen consumption may jointly or individually contribute to observed seagrass shoot demise. [ABSTRACT FROM AUTHOR]

Eutrophication is one of the most relevant man-induced changes occurring in coastal waters. The identification and assessment of specific responses to eutrophication in seagrasses can provide a useful tool for the detection of changes in the water quality in coastal zones, given the wide range of distribution of these organisms. In this study, we combine a correlational (across-sites comparison) and a manipulative (fertilization experiment) approach to evaluate the usefulness and potential of alkaline phosphatase activity (APA) in the endemic Mediterranean seagrass Posidonia oceanica as an eutrophication biomarker. Our results showed that APA decreases promptly following nutrient additions, the response being maintained except during the winter period. APA also varies across natural meadows under different levels of nutrient discharges at scales relevant for monitoring purposes. AP activity seems to be an optimal ‘physiological biomarker’ that responds promptly and reliably to a pulse of eutrophication exposure. However, other considerations, such as the seasonality (the response disappears in winter), suggest its use with some caution and, as far as possible, as a complement of other bio-indicators. [ABSTRACT FROM AUTHOR]

While known to be uniformly non-feeding, short-lived, and potentially short dispersing, sponge larvae display different behaviours (swimming ability and taxis). Our aim was to show whether sponge larvae with different behaviours exhibit different dispersal strategies under variable intensity of water movements. We first assessed the distribution of larvae of six taxa: Dictyoceratida spp., Dysidea avara, Crambe crambe, Phorbas tenacior, Scopalina lophyropoda, and Cliona viridis, collected through plankton sampling, and the abundance of the corresponding adult sponges across three hard bottom communities and a sandy bottom from a north-west Mediterranean rocky shore. We then tested adult–larvae couplings (abundance of larvae vs abundance of adults) under increasing levels of water movements (surge) to assess the importance of this environmental factor in driving differences in dispersal strategies. Adults of Dictyoceratida spp., D. avara, and P. tenacior were most abundant in semi-dark caves (SDC), C. crambe and C. viridis in communities of sciaphilic algae (SA), whereas the distribution of S. lophyropoda was extremely patchy, being present almost only in the SA community of one of the five stations studied. Larvae of Dictyoceratida spp. and P. tenacior were more abundant in the SDC, whereas D. avara and C. crambe were homogeneously distributed across the communities. The larvae of C. viridis were more abundant in the SA communities and the S. lophyropoda larvae were mostly present in one station and one community (SA). Increased water movement did not modify the adult–larvae coupling for Dictyoceratida spp., D. avara, and C. crambe, whereas it broke up the positive association for P. tenacior and to some extent S. lophyropoda. For C. viridis, possible variability in adult–larvae coupling was not tested because the larvae were collected on only one day under calm sea conditions. We confirm that efficient-swimming larvae with some cue response can actively counteract hydrodynamic forces and highlight the importance of both larval behaviour and environmental conditions in determining small-scale patterns of dispersal. [ABSTRACT FROM AUTHOR]

Mediterranean populations of Schizobrachiella sanguinea seem to span a wide range of contrasting benthic habitats. We have studied adult, larval, and recruit populations of S. sanguinea from sublittoral (approximately 10 m depth) communities at Blanes (Spain, NW Mediterranean). At the study site, the species colonises semi-obscure caves and, to a lesser extent, communities of sciaphilic algae. Our first goal was to show the periods of brooding, larval release, and recruitment. Plankton was collected about twice a week between July 1998 and June 2000 over an artificial reef. A total of 303 larvae were collected in 23 of the 102 days of sampling. At the study site the species shows a single, annual larval release period, which occurs from late March to June–July every year. Recruits were found from late May to late August 1999 and in late May 2000. A recruitment peak was observed in July 1999 and again in late May 2000. The presence of brooding adults corresponded to the periods of larval supply and recruitment. Our second goal was to investigate the causes of the variable distribution of the species along several communities in the north-west Mediterranean Sea. We describe adult distribution and the dynamics of larval supply, recruitment of early (≤15 days from settlement) and late (≤4 months from settlement) recruits. Patterns of larval presence and early recruit distribution along the communities on the reef did not reflect the adult distributions, whereas this was closely matched by the distribution of late recruits. Post-recruitment mortality strongly affected recruits of the PA (photophilic algae) and SA (sciaphilic algae) communities, whereas it was very low in the communities of SOC (semi-obscure caves). Mortality of recruits was density independent and colony survivorship in the SA communities, where recruitment was the highest, appeared strongly affected by competition with brown algae. Like the majority of cheilostomate bryozoans, S. sanguinea releases a coronate larva capable of active swimming and positive reaction to light. Such swimming abilities may allow larvae to actively select the substrate on which to settle and to avoid unsuitable substrates. It seems that larval behaviour does not explain the preference of S. sanguinea for sciaphilic habitats. High post-recruitment mortality due to various factors, especially competition with fast-growing algae, seems particularly relevant in determining the adult distributions observed. [ABSTRACT FROM AUTHOR]