Reef Invertebrates Research Articles

Occurrence of plastic additives in coral-reef invertebrates on natural and plastic substrates

Numerous studies have investigated the occurrence of plastic additives in marine biota. Yet, their main vector of transfer into organisms tissues remains unknown. We explored seven common additives in benthic coral reef invertebrates residing on natural/plastic substrates in a protected marine reserve versus an unprotected reef to ascertain whether additives transfer by substrate leaching. Samples of three coral-reef species were extracted and analyzed by GCMS and HPLC. Of the seven chemical additives investigated, dibenzylamine and bis(2-ethylhexyl) phthalate were detected. No significant association was found between additives and substrate type, possibly because these plastics have been submerged for years, and the majority of additives within them have leached. The marine reserve had fewer samples with additives, highlighting the importance of active management. Understanding the transfer vectors of plastic additives into biota is essential for assessing the risk they pose and devising effective management tools for protecting coral reefs.

Challenging trophic position assessments in complex ecosystems: Calculation method, choice of baseline, trophic enrichment factors, season and feeding guild do matter: A case study from Marquesas Islands coral reefs.

Assessments of ecosystem functioning are a fundamental ecological challenge and an essential foundation for ecosystem-based management. Species trophic position (TP) is essential to characterize food web architecture. However, despite the intuitive nature of the concept, empirically estimating TP is a challenging task due to the complexity of trophic interaction networks. Various methods are proposed to assess TPs, including using different sources of organic matter at the base of the food web (the 'baseline'). However, it is often not clear which methodological approach and which baseline choices are the most reliable. Using an ecosystem-wide assessment of a tropical reef (Marquesas Islands, with available data for 70 coral reef invertebrate and fish species), we tested whether different commonly used TP estimation methods yield similar results and, if not, whether it is possible to identify the most reliable method. We found significant differences in TP estimates of up to 1.7 TPs for the same species, depending on the method and the baseline used. When using bulk stable isotope data, the choice of the baseline significantly impacted TP values. Indeed, while nitrogen stable isotope (δ15N) values of macroalgae led to consistent TP estimates, those using phytoplankton generated unrealistically low TP estimates. The use of a conventional enrichment factor (i.e. 3.4‰) or a 'variable' enrichment factor (i.e. according to feeding guilds) also produced clear discrepancies between TP estimates. TPs obtained with δ15N values of source amino acids (compound-specific isotope analysis) were close to those assessed with macroalgae. An opposite seasonal pattern was found, with significantly lower TPs in winter than in summer for most species, with particularly pronounced differences for lower TP species. We use the observed differences to discuss possible drivers of the diverging TP estimates and the potential ecological implications.

Opportunistic consumption of coral spawn by the ruby brittle star (Ophioderma rubicundum).

Many reef invertebrates reproduce through simultaneous broadcast spawning, with an apparent advantage of overwhelming potential predators and maximizing propagule survival. Although reef fish have been observed to consume coral gamete bundles during spawning events, there are few records of such predation by benthic invertebrates. Here, we document several instances of the ruby brittle star, Ophioderma rubicundum, capturing and consuming egg-sperm bundles of the mountainous star coral, Orbicella faveolata, and the symmetrical brain coral, Pseudodiploria strigosa, during spawning events in the Cayman Islands in 2012 and the Florida Keys in 2022. These observations are widely separated in space and time (>600 km, 10 years), suggesting that this behavior may be prevalent on western Atlantic reefs. Since O. rubicundum spawns on the same or subsequent nights as these coral species, we hypothesize that this opportunistic feeding behavior takes advantage of lipid-rich coral gamete bundles to recover energy reserves expended by the brittle star during gametogenesis. The consumption of coral gametes by adult brittle stars suggests an underexplored trophic link between reef invertebrates and also provides evidence that ophiuroid-coral symbioses may oscillate between commensalism and parasitism depending on the ontogeny and reproductive status of both animals. Our observations provide insights into the nuanced, dynamic associations between coral reef invertebrates and may have implications for coral reproductive success and resilience.

Effects of sponge-to-sponge contact on the microbiomes of three spatially competing Caribbean coral reef species.

Sponges perform important ecosystem functions, host diverse microbial symbiont communities (microbiomes), and have been increasing in density on Caribbean coral reefs over the last decade. Sponges compete for space in coral reef communities through both morphological and allelopathic strategies, but no studies of microbiome impacts during these interactions have been conducted. Microbiome alterations mediate spatial competition in other coral reef invertebrates and may similarly impact competitive outcomes for sponges. In this study, we characterized the microbiomes of three common Caribbean sponges (Agelas tubulata, Iotrochota birotulata, and Xestospongia muta) observed to naturally interact spatially in Key Largo, Florida (USA). For each species, replicate samples were collected from sponges in contact with neighbors at the site of contact (contact) and distant from the site of contact (no contact), and from sponges spatially isolated from neighbors (control). Next-generation amplicon sequencing (V4 region of 16S rRNA) revealed significant differences in microbial community structure and diversity among sponge species, but no significant effects were observed within sponge species across all contact states and competitor pairings, indicating no large community shifts in response to direct contact. At a finer scale, particular symbiont taxa (operational taxonomic units at 97% sequence identity, OTUs) were shown to decrease significantly in some interaction pairings, suggesting localized effects for specific sponge competitors. Overall, these results revealed that direct contact during spatial competition does not significantly alter microbial community composition or structure of interacting sponges, suggesting that allelopathic interactions and competitive outcomes are not mediated by microbiome damage or destabilization.

Observations of coral and cryptobenthic sponge fluorescence and recruitment on autonomous reef monitoring structures (ARMS)

Fluorescence imaging of benthic communities is a widely used tool for determining the rate of hard coral recruitment in tropical reefs. Whilst fluorescent proteins are well-studied in scleractinian corals, less is understood about their distribution and function in other sessile reef invertebrates. This short study examines fluorescence images of benthic communities on Autonomous Reef Monitoring Structures (ARMS) from a remote and protected Indian Ocean reef system. We compare the abundance of adult and juvenile hard corals across three sites and between the topside and underside of ARMS recruitment plates. We also discuss observations of skeletal fluorescence in sponges, as well as uneven green fluorescent protein (GFP) concentrations across adult coral colonies. Our findings provide an insight into the recovery of shallow reefs previously hit by severe bleaching events and highlight the potential of ARMS fluorescence imaging for the analysis of cryptobenthic communities.

Effects of low pH on the coral reef cryptic invertebrate communities near CO2 vents in Papua New Guinea.

Small cryptic invertebrates (the cryptofauna) are extremely abundant, ecologically important, and species rich on coral reefs. Ongoing ocean acidification is likely to have both direct effects on the biology of these organisms, as well as indirect effects through cascading impacts on their habitats and trophic relationships. Naturally acidified habitats have been important model systems for studying these complex interactions because entire communities that are adapted to these environmental conditions can be analyzed. However, few studies have examined the cryptofauna because they are difficult to census quantitatively in topographically complex habitats and are challenging to identify. We addressed these challenges by using Autonomous Reef Monitoring Structures (ARMS) for sampling reef-dwelling invertebrates >2 mm in size and by using DNA barcoding for taxonomic identifications. The study took place in Papua New Guinea at two reef localities, each with three sites at varying distances from carbon dioxide seeps, thereby sampling across a natural gradient in acidification. We observed sharp overall declines in both the abundance (34–56%) and diversity (42–45%) of organisms in ARMS under the lowest pH conditions sampled (7.64–7.75). However, the overall abundance of gastropods increased slightly in lower pH conditions, and crustacean and gastropod families exhibited varying patterns. There was also variability in response between the two localities, despite their close proximity, as one control pH site displayed unusually low diversity and abundances for all invertebrate groups. The data illustrate the complexity of responses of the reef fauna to pH conditions, and the role of additional factors that influence the diversity and abundance of cryptic reef invertebrates.

Accumulation of commonly used agricultural herbicides in coral reef organisms from iSimangaliso Wetland Park, South Africa

Coral reefs are amongst the most biodiverse ecosystems on earth, but are significantly impacted by agricultural runoff. Despite herbicides being commonly detected in coastal waters, the possibility of herbicide accumulation in coral reef species has largely been overlooked. We investigate the accumulation of several herbicides in five species of coral reef invertebrates collected from ten sites along the Maputaland coast, South Africa. Multiple herbicide residues were detected in 95% of the samples, with total average concentrations across sites ranging between 25.2 ng g−1 to 51.3 ng g−1 dw. Acetochlor, alachlor and hexazinone were the predominant herbicides detected at all sites, with atrazine and simazine detected less frequently. Significant interactive effects were detected between sites nested in reef complex crossed with species, based on multiple and total herbicide concentrations. In general, multivariate herbicide concentrations varied significantly between species within and across most sites. Contrastingly, the concentrations of the different herbicides and that of total herbicide did not differ between conspecifics at most sites nested in their respective reef complexes. On average, highest total herbicide concentrations were measured in soft coral (Sarcophyton glaucum; 90.4 ± 60 ng g−1 and Sinularia gravis; 42.7 ± 25 ng g−1) and sponge (Theonela swinhoei; 39.0 ± 40 ng g−1) species, while significantly lower concentrations were detected in hard corals (Echinopora hirsutissima; 10.5 ± 5.9 ng g−1 and Acropora austera; 5.20 ± 4.5 ng g−1) at most sites. Agricultural runoff entering the ocean via the uMfolozi-St Lucia Estuary and Maputo Bay are likely sources of herbicide contamination to coral reefs in the region. There is an urgent need to assess the long-term effects of herbicide exposure on coral reef communities.

Testing cophylogeny between coral reef invertebrates and their bacterial and archaeal symbionts.

Marine invertebrates harbour a complex suite of bacterial and archaeal symbionts, a subset of which are probably linked to host health and homeostasis. Within a complex microbiome it can be difficult to tease apart beneficial or parasitic symbionts from nonessential commensal or transient microorganisms; however, one approach is to detect strong cophylogenetic patterns between microbial lineages and their respective hosts. We employed the Procrustean approach to cophylogeny (PACo) on 16S rRNA gene derived microbial community profiles paired with COI, 18S rRNA and ITS1host phylogenies. Second, we undertook a network analysis to identify groups of microbes that were co-occurring within our host species. Across 12 coral, 10 octocoral and five sponge species, each host group and their core microbiota (50% prevalence within host species replicates) had a significant fit to the cophylogenetic model. Independent assessment of each microbial genus and family found that bacteria and archaea affiliated to Endozoicomonadaceae, Spirochaetaceae and Nitrosopumilaceae have the strongest cophylogenetic signals. Further, local Moran's I measure of spatial autocorrelation identified 14 ASVs, including Endozoicomonadaceae and Spirochaetaceae, whose distributions were significantly clustered by host phylogeny. Four co-occurring subnetworks were identified, each of which was dominant in a different host group. Endozoicomonadaceae and Spirochaetaceae ASVs were abundant among the subnetworks, particularly one subnetwork that was exclusively comprised of these two bacterial families and dominated the octocoral microbiota. Our results disentangle key microbial interactions that occur within complex microbiomes and reveal long-standing, essential microbial symbioses in coral reef invertebrates.

Microbiome-mediated mechanisms contributing to the environmental tolerance of reef invertebrate species

Coral reefs globally are increasingly impacted by climate change. High temperature and pCO2 levels disrupt multiple physiological and biochemical pathways in marine organisms, often leading to disease and mortality in sensitive reef species. Host-associated microbes contribute critical functions that underpin host health, and environmentally induced changes in microbial communities represent a potential source for new metabolic features within holobiont systems. However, whether the acquisition of new beneficial microbial functions contributes to environmental acclimatisation of reef species is currently unknown. Using extensively studied model systems, we identify potential direct and indirect microbiome-mediated mechanisms that may contribute to environmental acclimatisation in reef invertebrate species. These mechanisms include increasing energy metabolism in the host, reduction of oxidative stress, regulation of nutrients in host cells, and increased pathogen resistance. We also propose a robust experimental strategy to test how microbial metabolic pathways may facilitate environmental acclimatisation of reef taxa. Understanding the mechanisms of microbiome-mediated acclimatisation and the timescales over which it can occur will be critical for predicting reef ecosystem dynamics under future climate scenarios and applying effective reef conservation strategies.

Correction to: Isolation and genotyping of novel T4 cyanophages associated with diverse coral reef invertebrates

This erratum is published due to formatting discrepancies noticed with figures and tables overlooked by vendor during proofing.

Tracing trophic pathways through the marine ecosystem of Rapa Nui (Easter Island)

Abstract The structure of food webs provides important insight into biodiversity, organic matter (OM) pathways, and ecosystem functioning. Stable isotope analysis (δ13C and δ15N) was used to characterize the trophic structure and the main OM pathways supporting food webs in the Rapa Nui coastal marine ecosystem. The trophic position of consumers and isotopic niche metrics were estimated for different assemblages (i.e. mesozooplankton, emergent zooplankton, reef invertebrates, reef fishes, pelagic fishes, and seabirds). Furthermore, the relative importance of different OM sources (i.e. macroalgae, zooxanthellate corals, and particulate OM [POM]) was assessed for heterotrophic consumers using Bayesian mixing model (MixSIAR). Results show a clear pattern of 13C and 15N enrichment from small‐sized pelagic and benthic invertebrates, to reef and pelagic fishes, and seabirds. Most invertebrates were classified as primary consumers, reef fishes as secondary consumers and pelagic predators and seabirds as tertiary and quaternary consumers. Isotopic niche metrics indicate a low trophic diversity for pelagic assemblages (mesozooplankton and pelagic fishes), in contrast to reef fauna (invertebrates and fishes), whose higher trophic diversity suggest the exploitation of a wider range of trophic resources. Overlapping of standard ellipses areas between reef invertebrates and reef fishes indicates that both assemblages could be sharing trophic resources. Mixing models results indicate that POM is the main trophic pathway for mesozooplankton, macroalgae (Rhodophyta) for emergent zooplankton, and a mix of coral‐derived OM and Rhodophyta for coral reef assemblages such as macrobenthos and reef invertebrates. In contrast, POM contribution was notably more important for some pelagic fishes and seabirds from upper trophic levels. This study provides key elements for conservation efforts on coral reefs, management planning and full‐implementation of the recently created Rapa Nui Multiple Use Marine Protected Area.

Isolation and genotyping of novel T4 cyanophages associated with diverse coral reef invertebrates

In an effort to facilitate virus isolation-based studies on coral reefs, we describe here a simple holobiont virus extraction protocol that is effective at separating and concentrating virus particles from coral reef invertebrates. We demonstrate the application of this protocol by isolating and barcoding cyanophages from invertebrate holobionts associated with coral reefs in southwest Puerto Rico. Cyanophages were also isolated and barcoded from adjacent coral reef seawater. Barcoding of cyanophage isolates was carried out with the cyanomyovirus DNA polymerase (g43) or major capsid protein (g23) marker genes. We then utilized cyanophage sequences from Puerto Rico, along with those published previously associated with Rhode Island and Bermuda seawater, to assess the presence of cyanophage-like sequences in reef invertebrate virome, metagenome, and transcriptome sequencing libraries. The detailed holobiont virus extraction protocol successfully separated and concentrated virus particles from the tissue of 20 different species of coral reef invertebrate. Cyanophages were isolated and barcoded from 15 of these species: three scleractinian corals, a gorgonian, a corallimorpharian, a zoantharian, two hydrozoans, four species of sponges, two tunicates, and a nudibranch. In total, there were 146 cyanophages isolated and barcoded from seawater (n = 46) or invertebrate tissue (n = 100). The majority (69%) of cyanomyovirus sequences reported in this study were novel, sharing rather low similarity (< 98% nucleotide similarity) with publicly available sequences in the NCBI nucleotide database. Sequence library mining efforts revealed evidence of cyanophage-like sequences in 23 next-generation sequencing datasets, representing 17 species of coral reef invertebrates which included seven species of stony corals, one scyphozoan, seven species of sponges, and two species of copepods. This is the first investigation into cyanophage diversity on Puerto Rico reefs and is a relevant step in coral reef virology, ideally stimulating holobiont-associated isolation efforts to further explore virus genetic and functional diversity within invertebrate tissue on the reef.

Dimethylsulfoniopropionate concentration in coral reef invertebrates varies according to species assemblages

Dimethylsulfoniopropionate (DMSP) is a key compound in the marine sulfur cycle, and is produced in large quantities in coral reefs. In addition to Symbiodiniaceae, corals and associated bacteria have recently been shown to play a role in DMSP metabolism. Numerous ecological studies have focused on DMSP concentrations in corals, which led to the hypothesis that increases in DMSP levels might be a general response to stress. Here we used multiple species assemblages of three common Indo-Pacific holobionts, the scleractinian corals Pocillopora damicornis and Acropora cytherea, and the giant clam Tridacna maxima and examined the DMSP concentrations associated with each species within different assemblages and thermal conditions. Results showed that the concentration of DMSP in A. cytherea and T. maxima is modulated according to the complexity of species assemblages. To determine the potential importance of symbiotic dinoflagellates in DMSP production, we then explored the relative abundance of Symbiodiniaceae clades in relation to DMSP levels using metabarcoding, and found no significant correlation between these factors. Finally, this study also revealed the existence of homologs involved in DMSP production in giant clams, suggesting for the first time that, like corals, they may also contribute to DMSP production. Taken together, our results demonstrated that corals and giant clams play important roles in the sulfur cycle. Because DMSP production varies in response to specific species-environment interactions, this study offers new perspectives for future global sulfur cycling research.

Diverse coral reef invertebrates exhibit patterns of phylosymbiosis.

Microbiome assemblages of plants and animals often show a degree of correlation with host phylogeny; an eco-evolutionary pattern known as phylosymbiosis. Using 16S rRNA gene sequencing to profile the microbiome, paired with COI, 18S rRNA and ITS1 host phylogenies, phylosymbiosis was investigated in four groups of coral reef invertebrates (scleractinian corals, octocorals, sponges and ascidians). We tested three commonly used metrics to evaluate the extent of phylosymbiosis: (a) intraspecific versus interspecific microbiome variation, (b) topological comparisons between host phylogeny and hierarchical clustering (dendrogram) of host-associated microbial communities, and (c) correlation of host phylogenetic distance with microbial community dissimilarity. In all instances, intraspecific variation in microbiome composition was significantly lower than interspecific variation. Similarly, topological congruency between host phylogeny and the associated microbial dendrogram was more significant than would be expected by chance across all groups, except when using unweighted UniFrac distance (compared with weighted UniFrac and Bray–Curtis dissimilarity). Interestingly, all but the ascidians showed a significant positive correlation between host phylogenetic distance and associated microbial dissimilarity. Our findings provide new perspectives on the diverse nature of marine phylosymbioses and the complex roles of the microbiome in the evolution of marine invertebrates.

Comatulids (Crinoidea, Comatulida) chemically defend against coral fish by themselves, without assistance from their symbionts

Symbiotic associations between small animals and relatively large sessile invertebrates that use taste deterrents for protection are widespread in the marine environment. To determine whether the symbionts are involved in the chemical protection of their hosts, the palatability of ten species of comatulids and six species of their symbionts was evaluated. Taste attractiveness was determined by offering agar pellets flavoured with extracts of comatulids and their symbionts for four coral reef fish species. Five species of symbiont were highly palatable, and one was indifferent to the taste. Almost all comatulids were distasteful, while their aversiveness was different for different fish. These findings indicate that comatulids chemically defend themselves without assistance from symbionts, and the taste deterrents are not universal and can only be effective against particular predators. The presence of tasteful symbionts reduces the security of their hosts by provoking attacks of predators and may impact on the individual and population fitness of comatulids. However, the chemical protection of comatulids is useful for symbionts and undoubtedly increases their survival. Obtained results allows the relationship between comatulids and their symbionts considered commensalism. Most likely, similar relationships can be established in many other associations, where symbionts inhabit chemically defended coral reef invertebrates.

A mesophotic hotel: the octocoral Bebryce cf. grandicalyx as a host.

Mesophotic coral ecosystems are widespread and harbour rich coral reef communities below 30 m and down to 150 m depth (Puglise et al. 2009). Although in recent years the knowledge of these charismatic ecosystems has increased (Khang et al. 2014), many aspects are still to be explored, including for instance symbiotic associations. Sessile reef invertebrates are known to host a multitude of organisms in shallow waters (e.g. Hoeksema et al. 2012), and this may be likewise extended to mesophotic organisms. For instance, both shallow and deep-water octocorals are known to form a large variety of associations with microorganisms, invertebrates, and vertebrates (Goh et al. 1999, Buhl-Mortensen and Mortensen, 2004), including host-specific relationships (e.g. Montano et al. 2017a). In a few cases, these associations include commensalistic, mutualistic, and parasitic interactions (Watling et al. 2011).

A non-lethal SPME-LC/MS method for the analysis of plastic-associated contaminants in coral reef invertebrates

A non-lethal SPME-LC/MS procedure to detect plastic associated contaminants in coral reef invertebrates.

Coral endosymbionts (Symbiodiniaceae) emit species-specific volatilomes that shift when exposed to thermal stress

Biogenic volatile organic compounds (BVOCs) influence organism fitness by promoting stress resistance and regulating trophic interactions. Studies examining BVOC emissions have predominantly focussed on terrestrial ecosystems and atmospheric chemistry – surprisingly, highly productive marine ecosystems remain largely overlooked. Here we examined the volatilome (total BVOCs) of the microalgal endosymbionts of reef invertebrates, Symbiodiniaceae. We used GC-MS to characterise five species (Symbiodinium linucheae, Breviolum psygmophilum, Durusdinium trenchii, Effrenium voratum, Fugacium kawagutii) under steady-state growth. A diverse range of 32 BVOCs were detected (from 12 in D. trenchii to 27 in S. linucheae) with halogenated hydrocarbons, alkanes and esters the most common chemical functional groups. A thermal stress experiment on thermally-sensitive Cladocopium goreaui and thermally-tolerant D. trenchii significantly affected the volatilomes of both species. More BVOCs were detected in D. trenchii following thermal stress (32 °C), while fewer BVOCs were recorded in stressed C. goreaui. The onset of stress caused dramatic increases of dimethyl-disulfide (98.52%) in C. goreaui and nonanoic acid (99.85%) in D. trenchii. This first volatilome analysis of Symbiodiniaceae reveals that both species-specificity and environmental factors govern the composition of BVOC emissions among the Symbiodiniaceae, which potentially have, as yet unexplored, physiological and ecological importance in shaping coral reef community functioning.

Climate and fishing drive regime shifts in consumer-mediated nutrient cycling in kelp forests.

Globally, anthropogenic pressures are reducing the abundances of marine species and altering ecosystems through modification of trophic interactions. Yet, consumer declines also disrupt important bottom-up processes, like nutrient recycling, which are critical for ecosystem functioning. Consumer-mediated nutrient dynamics (CND) is now considered a major biogeochemical component of most ecosystems, but lacking long-term studies, it is difficult to predict how CND will respond to accelerating disturbances in the wake of global change. To aid such predictions, we coupled empirical ammonium excretion rates with an 18-year time series of the standing biomass of common benthic macroinvertebrates in southern California kelp forests. This time series of excretion rates encompassed an extended period of extreme ocean warming, disease outbreaks, and the abolishment of fishing at two of our study sites, allowing us to assess kelp forest CND across a wide range of environmental conditions. At their peak, reef invertebrates supplied an average of 18.3±3.0µmolNH4 + m-2 hr-1 to kelp forests when sea stars were regionally abundant, but dropped to 3.5±1.0µmolNH4 + m-2 hr-1 following their mass mortality due to disease during a prolonged period of extreme warming. However, a coincident increase in the abundance of the California spiny lobster, Palinurus interupptus (Randall, 1840), likely in response to both reduced fishing and a warmer ocean, compensated for much of the recycled ammonium lost to sea star mortality. Both lobsters and sea stars are widely recognized as key predators that can profoundly influence community structure in benthic marine systems. Our study is the first to demonstrate their importance in nutrient cycling, thus expanding their roles in the ecosystem. Climate change is increasing the frequency and severity of warming events, and rising human populations are intensifying fishing pressure in coastal ecosystems worldwide. Our study documents how these projected global changes can drive regime shifts in CND and fundamentally alter a critical ecosystem function.

Marine reserves drive both taxonomic and functional change in coral reef invertebrate communities

A leading argument for no-take marine protected area (marine reserve) establishment is their contribution to the conservation of biodiversity, but the impacts of reserves on ecosystem functioning have seldom been quantified. This is unusual given the value of services provided by ocean ecosystems to human well-being. While no single index can describe ecosystem function, a set of life-history attributes possessed by taxa can be used to infer differences in ecosystem function across space and time. In this study, we use biological trait analysis to determine whether the attributes of invertebrate taxa differ between inside of six no-take marine reserves and outside, in fished areas in the Central Philippines. Using permutational multivariate analyses, we found that the composition of traits and taxa were significantly different between reserve and non-reserve areas. Habitat use, morphology and mobility traits were the biggest contributors to dissimilarity, indicating that reserves can have community-wide effects that change the functional composition of invertebrate assemblages. Notably, traits associated with coral habitat use, bearing a shell, lacking mobility and filter feeding are the most important traits associated with differences in community structure between reserve and non-reserve areas. At the taxa composition level, small shrimps, three families of bivalve, two families of burrowing snails and brittle stars are the most important contributors to differences in taxonomic community composition. The addition of organismal attributes to traditional taxa composition approaches provides richer insight into how ecosystems respond to protection and has the potential to inform practitioners on conserving for ecosystem traits.