Quantifying the role of photoacclimation and self-facilitation for seagrass resilience to light deprivation.

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 different 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.

Tolerance to aerial exposure influences distributional patterns in multi-species intertidal seagrass meadows.

Multi-specific seagrass meadow assemblages dominate most tropical intertidal regions but the relative role of environmental stress in determining distribution patterns is still uncertain. Here we combine observational and experimental approaches to examine aerial exposure as a factor driving species occurrence patterns in intertidal meadows of the Andaman archipelago, where up to 6 seagrass species co-occur. In the studied meadow, patterns of exposure did not map onto distance from the coast, instead creating a patchy matrix of exposure, based on fine-scale bathymetric differences. Distributional surveys showed that seagrass species were similarly patchy, often tracking the degree of aerial exposure during low tide. While some species (Halophila ovalis, Halophila minor, and Thalassia hemprichii) frequently occurred in submerged or subtidal areas and were rarely found in completely exposed areas, other species (Cymodocea rotundata, Halophila beccarii, and Halodule uninervis) also occupied areas that were subject to partial or complete aerial exposure during low tide. To confirm this pattern, we used field-based transplant experiments, employing a natural gradient of tidal exposure to subject six seagrass species to different desiccation exposure times. After a month, H. beccarii and H. uninervis transplants survived in areas that sustained more than 3 h of aerial tidal exposure without significant mortality, compared with other species (H. ovalis, H. minor, T. hemprichii, C. rotundata) that showed dramatic shoot mortality at the same exposure regimes. For all species, 4 h represented the upper limit of exposure, in both experimental and distributional studies. However, despite their wider tolerance of exposure to air, H. beccarii and H. uninervis did not dominate the entire meadow. This could be a result either of their poor tolerance to other environmental factors or their lower competitive abilities among other mechanisms. This suggests that in tropical multi-specific meadows, strong environmental filters could override clear intertidal zonation to create patchy matrices based on species tolerances.

Taxonomic identity of Distapliastylifera (Tunicata, Ascidiacea), a new arrival to the eastern Pacific displaying invasive behavior in the Gulf of California, Mexico.

A colonial ascidian of the genus Distaplia caused a mass mortality of the pen shell Atrinamaura (Sowerby, 1835) during June 2016 in the southwest of the Gulf of California (Mexico), with a significant socio-economic cost. Tentatively identified in previous works as Distapliacf.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 Distapliastylifera (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.

A trait-based framework for seagrass ecology: Trends and prospects.

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.

Nutrient conditions determine the strength of herbivore-mediated stabilizing feedbacks in barrens.

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.

Tropicalization shifts herbivore pressure from seagrass to rocky reef communities.

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.

Learning takes time: Biotic resistance by native herbivores increases through the invasion process.

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.

Identifying conservation priorities for a widespread dugong population in the Red Sea: Megaherbivore grazing patterns inform management planning.

Extensive home ranges of marine megafauna present a challenge for systematic conservation planning because they exceed spatial scales of conventional management. For elusive species like dugongs, their management is additionally hampered by a paucity of basic distributional information across much of their range. The Red Sea is home to a wide-spread, globally important but data-poor population of dugongs. We surveyed the north-eastern Red Sea in the waters of NEOM, Kingdom of Saudi Arabia, to locate feeding sites and determine priority areas for dugong conservation. We conducted large-scale in-water surveys of dugong feeding trails across 27 seagrass meadows that span 0.7 degree of latitude and recorded nine seagrass species and 13 dugong feeding sites. Spread over ∼4'061 km2 of nearshore and offshore waters, many of these sites clustered around five main core feeding areas. Dugong feeding trails were mostly recorded at sites dominated by the fast-growing pioneer seagrasses Halodule uninervis, Halophila ovalis and/or H. stipulacea. Multispecific meadows with pioneer seagrasses tended to be sheltered and shallow, reflecting a similar spatial pattern to the identified dugong feeding sites. Often close to hotels and fishing harbours, these high-use dugong areas are subject to high boat traffic, fishing, and coastal development which places considerable pressures on this vulnerable mammal and its seagrass habitat. The rapidly accelerating coastal development in the northern Red Sea directly threatens the future of its dugong population. Although our sampling focuses on feeding signs in early successional seagrasses, the results are valuable to spatial conservation planning as they will trigger overdue conservation interventions for a globally threatened species in a data-poor area. Urgent dugong conservation management actions in the northern Red Sea should focus on shallow waters sheltered by coastal lagoons, bays and the lee of large islands.

Understanding the depth limit of the seagrass Cymodocea nodosa as a critical transition: Field and modeling evidence.

Changes in light and sediment conditions can sometimes trigger abrupt regime shifts in seagrass meadows resulting in dramatic and unexpected die-offs of seagrass. Light attenuates rapidly with depth, and in seagrass systems with non-linear behaviours, can serve as a sharp boundary beyond which the meadow transitions to bare sand. Determining system behaviour is therefore essential to ensuring resilience is maintained and to prevent stubborn critical ecosystem transitions caused by declines in water quality. Here we combined field and modelling studies to explore the transition from meadow to bare sand in the seagrass Cymodocea nodosa at the limit of its depth distribution in a shallow, light-limited bay. We first describe the relationship between light availability and seagrass density along a depth gradient in an extensive unfragmented meadow (Alfacs bay, NE Spain). We then develop a simple mechanistic model to characterise system behaviour. In the field, we identified sharp decline in shoot density beyond a threshold of ∼1.9 m depth, shifting from a vegetated state to bare sand. The dynamic population model we developed assumes light-dependent growth and an inverse density-dependent mortality due to facilitation between shoots (mortality rate decreases as shoot density increases). The model closely tracked our empirical observations, and both the model and the field data showed signs of bistability. This strongly suggests that the depth limit of C. nodosa is a critical transition driven by photosynthetic light requirements. While the mechanisms still need to be confirmed with experimental evidence, recognizing the non-linear behaviour of C. nodosa meadows is vital not only in improving our understanding of light effects on seagrass dynamics, but also in managing shallow-water meadows. Given the shallow threshold (<2m), light-limited systems may experience significant and recalcitrant meadow retractions with even small changes in sediment and light conditions. Understanding the processes underlying meadow resilience can inform the maintenance and restoration of meadows worldwide.

Exploring coexistence mechanisms in a three-species assemblage.

Interactions among species are essential in shaping ecological communities, although it is not always clear under what conditions they can persist when the number of species involved is higher than two. Here we describe a three-species assemblage involving the seagrass Cymodocea nodosa, the pen shell Pinna nobilis and the herbivore sea urchin Paracentrotus lividus, and we explore the mechanisms allowing its persistence through field observations and manipulative experiments. The abundance of pen shells was higher in seagrass beds than in bare sand, suggesting a recruitment facilitation. The presence of sea urchins, almost exclusively attached or around pen shells, indicated habitat facilitation for sea urchins, which overgrazed the meadow around the pen shells forming seagrass-free halos. Our results suggest that this system persists thanks to: (i) the behavioral reluctance of sea urchins to move far from pen shells, making their impact on seagrass strictly local, (ii) the sparse distribution of pen shells and (iii) the plant's resistance mechanisms to herbivory. Unpacking these mechanisms allows a better understanding of how ecological communities are assembled.

Resilience of seagrass populations to thermal stress does not reflect regional differences in ocean climate.

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.

The scent of fear makes sea urchins go ballistic.

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.

Selection of parameters for seagrass management: Towards the development of integrated indicators for French Antilles.

Seagrass beds are increasingly impacted by human activities in coastal areas, particularly in tropical regions. The objective of this research program was to study seagrass beds characteristics under various environmental conditions in the French Antilles (FA, Caribbean Sea). A total of 61 parameters, from plant physiology to seagrass ecosystem, were tested along a gradient of anthropogenic conditions, distributed across 11 sites and 3 islands of the FA. A selection of 7 parameters was identified as relevant for the monitoring of seagrass meadows in the framework of public policies. They combined "early warning indicators" (e.g. nutrients and some trace metals) and long-term responding parameters (e.g. shoot density) adapted to management time scales. The ecological status of seagrass meadows was evaluated using a PCA. This work is a first step towards monitoring and management of seagrass meadows in the FA.

Homogenization and miniaturization of habitat structure in temperate marine forests.

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.

Temperature reduces fish dispersal as larvae grow faster to their settlement size.

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.

Warming intensifies the interaction between the temperate seagrass Posidonia oceanica and its dominant fish herbivore Sarpa salpa.

Apart from directly influencing individual life histories of species, climate change is altering key biotic interactions as well, causing community processes to unravel. With rising temperatures, disruptions to producer-consumer relationships can have major knock-on effects, particularly when the producer is a habitat-forming species. We studied how sea surface temperature (SST) modifies multiple pathways influencing the interaction between the foundational seagrass species, Posidonia oceanica, and its main consumer, the fish Sarpa salpa in the Mediterranean Sea. We used a combination of a field-based temperature gradient approaches and experimental manipulations to assess the effect of temperature on seagrass performance (growth) and fish early life history (larval development) as well as on the interaction itself (seagrass palatability and fish foraging activity). Within the range of temperatures assessed, S. salpa larvae grew slightly faster at warmer conditions but maintained their settlement size, resulting in a relatively small reduction in pelagic larval duration (PLD) and potentially reducing dispersion. Under warmer conditions (>24 °C), P. oceanica reduced its growth rate considerably and seemed to display fewer deterring mechanisms as indicated by a disproportionate consumption in choice experiments. However, our field-based observations along the temperature gradient showed no change in fish foraging time, or in other aspects of feeding behaviour. As oceans warm, our results indicate that, while S. salpa may show little change in early life history, its preference towards P. oceanica might increase, which, together with reduced seagrass growth, could considerably intensify the strength of herbivory. It is unclear if P. oceanica meadows can sustain such an intensification, but it will clearly add to the raft of pressures this threatened ecosystem already faces from global and local environmental change.

Climate drives the geography of marine consumption by changing predator communities.

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.

The dominant seagrass herbivore Sarpa salpa shifts its shoaling and feeding strategies as they grow.

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.

Wave exposure reduces herbivory in post-disturbed reefs by filtering species composition, abundance and behaviour of key fish herbivores.

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.