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.

C and N stable isotopes enlighten the trophic behaviour of the dugong (Dugong dugon).

The dugong (Dugong dugon), a large marine mammal herbivore of the Indo-Pacific, is vulnerable to extinction at a global scale due to a combination of human-related threats including habitat degradation. The species forages on seagrass habitats (marine phanerogams) and plays a key role in the functioning and sensitivity of these declining coastal ecosystems. The trophic behaviour and plasticity of dugong populations in response to extrinsic and intrinsic factors are therefore crucial features to both dugong and seagrass conservation. Yet, this knowledge remains limited to few visual observations and analyses of mouth, stomach or faecal contents of stranded individuals. We take advantage of a long-term monitoring of stranded individuals from the endangered New Caledonian population to depict features of dugongs' trophic ecology from Carbon and Nitrogen stable isotopes. A total of 59 dugong skin samples were used to portrait the stable isotope niche of dugongs according to their sex and maturity. In light of previous work conducted in New Caledonia, a subset of these samples was used to model the trophic mix of dugong males and females. Our stable isotope mixing models used C and N isotope values of 10 taxa bbelonging to five divisions of metazoans, plants, and chromists. Our results represent the first estimate of the species dietary niche in the isotopic space. They suggest that the diet of dugong calves overlaps more with that of adult females (δ13C: - 6.38 ± 1.13 ‰; δ15N: 2.49 ± 1.10 ‰) than males (δ13C: - 5.92 ± 1.10 ‰; δ15N: 3.69 ± 1.28 ‰). Further, we highlight differences in the expected trophic mix of dugong adult males and females. From these, we formulate a sex-specific foraging behaviour hypothesis in dugongs, whereby lactating females could forage over smaller spatial ranges but more diverse food sources thanmales. The study emphasizes the importance of long-term stranding monitoring programs to study the ecology of marine mammals.. Finally, it depicts an ecological feature that may contribute to the sensitivity of vulnerable dugongs to ongoing changes on tropical coastal ecosystems.

Metallic trace elements in marine sponges living in a semi-enclosed tropical lagoon.

The ability of marine filter feeders to accumulate metals could help monitor the health of the marine environment. This study examined the concentration of metallic trace elements (MTE) in two marine sponges, Rhabdastrella globostellata and Hyrtios erectus, from three sampling zones of the semi-enclosed Bouraké Lagoon (New Caledonia, South West Pacific). MTE in sponge tissues, seawater, and surrounding sediments was measured using inductively coupled plasma with optical emission spectroscopy. The variability in sponge MTE concentrations between species and sampling zones was visually discriminated using a principal component analysis (PCA). Sponges showed Fe, Mn, Cr, Ni, and Zn concentrations 2 to 10 times higher than in the surrounding sediments and seawater. Hyrtios erectus accumulated 3 to 20 times more MTE than R. globostellata, except for Zn. Average bioconcentration factors in sponge tissues were (in decreasing order) Zn > Ni > Mn > Fe > Cr relate to sediments and Fe > Ni > Mn > Cr > Zn relate to seawater. The PCA confirmed higher MTE concentrations in H. erectus compared to R. globostellata. Our results confirm that marine sponges can accumulate MTE to some extent and could be used as a tool for assessing metals contamination in lagoon ecosystems, particularly in New Caledonia, where 40% of the lagoon is classified as a UNESCO World Heritage Site.

Isotopic and biochemical trophic markers reveal the complexity of interactions at the base of pelagic food webs (Mediterranean sea).

To gain insight into the impact of bottom-up changes in the plankton community on planktivorous fish in the context of the decline of small pelagic fisheries in the Northwestern Mediterranean Sea, we have conducted an extensive year-long study. The investigation combined biochemical analyses (proteins, carbohydrates, and lipids) with C and N stable isotope analyses (SIA) to simultaneously study phytoplankton, zooplankton, and eight planktivorous fish species (Engraulis encrasicolus, Sardina pilchardus, Sardinella aurita, Sprattus sprattus, Cepola macrophthalma, Chromis chromis, Boops boops, and Spicara maena). This study is the first to analyze both stable isotope and biochemical compositions in coastal particulate organic matter (POM) size classes (0.7-2.7 µm, 2.7-20 µm, and 20-200 µm), zooplankton size classes (200-300 µm, 300-500 µm, 500-1000 µm, 1000-2000 µm, and >2000 µm), and taxonomic groups. We demonstrated that: (1) POM stable isotope compositions varied based on its spatial origin, the taxonomic composition of its biota, and its biochemical content; (2) δ15N values increased with zooplankton size classes and groups, indicating different trophic levels; (3) Phytoplankton exhibited a lipid-rich composition (∼55%), while zooplankton and fish muscles were protein-rich (∼61% and ∼66%, respectively). Bayesian stable isotope mixing models revealed that, on average: (1) POM from oceanic waters contributed the most to the POM in the bay (>51%), with a dominance of pico-POM (∼43%); (2) The 200-1000 µm zooplankton primarily consumed nano-POM, the 1000-2000 µm zooplankton mostly consumed micro-POM (∼64%), and the >2000 µm zooplankton also mostly consumed micro-POM; (3) Mesozooplankton (200-2000 µm) constituted the main portion (∼42%) of the diet for planktivorous fish species, while macrozooplankton organisms (>2000 µm) were the primary food resource (∼43%) for both B. boops and S. sprattus. Our study underscores the complexity of the pelagic food web and highlights the bottom-up transfer of organic matter from the smallest phytoplankton size fractions to planktivorous fish.

Sponge organic matter recycling: Reduced detritus production under extreme environmental conditions.

Sponges are a key component of coral reef ecosystems and play an important role in carbon and nutrient cycles. Many sponges are known to consume dissolved organic carbon and transform this into detritus, which moves through detrital food chains and eventually to higher trophic levels via what is known as the sponge loop. Despite the importance of this loop, little is known about how these cycles will be impacted by future environmental conditions. During two years (2018 and 2020), we measured the organic carbon, nutrient recycling, and photosynthetic activity of the massive HMA, photosymbiotic sponge Rhabdastrella globostellata at the natural laboratory of Bouraké in New Caledonia, where the physical and chemical composition of seawater regularly change according to the tide. We found that while sponges experienced acidification and low dissolved oxygen at low tide in both sampling years, a change in organic carbon recycling whereby sponges stopped producing detritus (i.e., the sponge loop) was only found when sponges also experienced higher temperature in 2020. Our findings provide new insights into how important trophic pathways may be affected by changing ocean conditions.

Biological trade-offs underpin coral reef ecosystem functioning.

Human impact increasingly alters global ecosystems, often reducing biodiversity and disrupting the provision of essential ecosystem services to humanity. Therefore, preserving ecosystem functioning is a critical challenge of the twenty-first century. Coral reefs are declining worldwide due to the pervasive effects of climate change and intensive fishing, and although research on coral reef ecosystem functioning has gained momentum, most studies rely on simplified proxies, such as fish biomass. This lack of quantitative assessments of multiple process-based ecosystem functions hinders local and regional conservation efforts. Here we combine global coral reef fish community surveys and bioenergetic models to quantify five key ecosystem functions mediated by coral reef fishes. We show that functions exhibit critical trade-offs driven by varying community structures, such that no community can maximize all functions. Furthermore, functions are locally dominated by few species, but the identity of dominant species substantially varies at the global scale. In fact, half of the 1,110 species in our dataset are functionally dominant in at least one location. Our results reinforce the need for a nuanced, locally tailored approach to coral reef conservation that considers multiple ecological functions beyond the effect of standing stock biomass.

Multi-trophic markers illuminate the understanding of the functioning of a remote, low coral cover Marquesan coral reef food web.

We studied the food web structure and functioning of a coral reef ecosystem in the Marquesas Islands, French Polynesia, characterized by low coral cover, high sea surface temperature and meso- to eutrophic waters. The Marquesas constitute a relevant ecosystem to understand the functioning of low diversity reefs that are also subject to global change. A multi-tracer assessment of organic matter pathways was run to delineate ecosystem functioning, using analysis of fatty acids, bulk and compound specific stable isotope analysis and stable isotopes mixing models. Macroalgae and phytoplankton were the two major food sources fueling this food web with, however, some marked seasonal variations. Specifically, zooplankton relied on phytoplankton-derived organic matter and herbivorous fishes on macroalgae-derived organic matter to a much higher extent in summer than in winter (~ 75% vs. ~ 15%, and ~ 70 to 75% vs. ~ 5 to 15%, respectively) . Despite remarkably high δ15N values for all trophic compartments, likely due to local dynamics in the nitrogen stock, trophic levels of consumers were similar to those of other coral reef ecosystems. These findings shed light on the functioning of low coral cover systems, which are expected to expand worldwide under global change.

Phylogeny, body morphology, and trophic level shape intestinal traits in coral reef fishes.

Trait-based approaches are increasingly used to study species assemblages and understand ecosystem functioning. The strength of these approaches lies in the appropriate choice of functional traits that relate to the functions of interest. However, trait-function relationships are often supported by weak empirical evidence.Processes related to digestion and nutrient assimilation are particularly challenging to integrate into trait-based approaches. In fishes, intestinal length is commonly used to describe these functions. Although there is broad consensus concerning the relationship between fish intestinal length and diet, evolutionary and environmental forces have shaped a diversity of intestinal morphologies that is not captured by length alone.Focusing on coral reef fishes, we investigate how evolutionary history and ecology shape intestinal morphology. Using a large dataset encompassing 142 species across 31 families collected in French Polynesia, we test how phylogeny, body morphology, and diet relate to three intestinal morphological traits: intestinal length, diameter, and surface area.We demonstrate that phylogeny, body morphology, and trophic level explain most of the interspecific variability in fish intestinal morphology. Despite the high degree of phylogenetic conservatism, taxonomically unrelated herbivorous fishes exhibit similar intestinal morphology due to adaptive convergent evolution. Furthermore, we show that stomachless, durophagous species have the widest intestines to compensate for the lack of a stomach and allow passage of relatively large undigested food particles.Rather than traditionally applied metrics of intestinal length, intestinal surface area may be the most appropriate trait to characterize intestinal morphology in functional studies.

Isotopic discrimination in helminths infecting coral reef fishes depends on parasite group, habitat within host, and host stable isotope value.

Stable isotopes of carbon and nitrogen characterize trophic relationships in predator-prey relationships, with clear differences between consumer and diet (discrimination factor Δ13C and Δ15N). However, parasite-host isotopic relationships remain unclear, with Δ13C and Δ15N remaining incompletely characterized, especially for helminths. In this study, we used stable isotopes to determine discrimination factors for 13 parasite-host pairings of helminths in coral reef fish. Differences in Δ15N values grouped according to parasite groups and habitat within the host with positive Δ15N values observed for trematodes and nematodes from the digestive tract and variable Δ15N values observed for cestodes and nematodes from the general cavity. Furthermore, Δ13C values showed more complex patterns with no effect of parasite group or habitat within host. A negative relationship was observed between Δ15N and host δ15N values among different host-parasite pairings as well as within 7 out of the 13 pairings, indicating that host metabolic processing affects host-parasite discrimination values. In contrast, no relationships were observed for Δ13C values. Our results indicate that parasite group, habitat within host, and host stable isotope value drive Δ15N of helminths in coral reef fish while their effect on Δ13C is more idiosyncratic. These results call for use of taxon- or species-specific and scaled framework for bulk stable isotopes in the trophic ecology of parasites.

Individual back-calculated size-at-age based on otoliths from Pacific coral reef fish species.

Somatic growth is a critical biological trait for organismal, population, and ecosystem-level processes. Due to its direct link with energetic demands, growth also represents an important parameter to estimate energy and nutrient fluxes. For marine fishes, growth rate information is most frequently derived from sagittal otoliths, and most of the available data stems from studies on temperate species that are targeted by commercial fisheries. Although the analysis of otoliths is a powerful tool to estimate individual growth, the time-consuming nature of otolith processing is one barrier for collection of comprehensive datasets across multiple species. This is especially true for coral reef fishes, which are extremely diverse. Here, we provide back-calculated size-at-age estimates (including measures of uncertainty) based on sagittal otoliths from 710 individuals belonging to 45 coral reef fish species from French Polynesia. In addition, we provide Von Bertalanffy growth parameters which are useful to predict community level biomass production.

Behavioral and trophic segregations help the Tahiti petrel to cope with the abundance of wedge-tailed shearwater when foraging in oligotrophic tropical waters.

Two species breeding in sympatry are more likely to coexist if their ecological niches are segregated either in time, space or in trophic habits. Here, we combined GPS-tracking, stable isotope analysis and DNA metabarcoding analysis to understand how the rare Tahiti petrel Pseudobulweria rostrata (TP) copes with the very abundant (i.e. 500,000 breeding pairs) wedge-tailed shearwater Ardenna pacifica (WTS) when breeding in sympatry in a tropical area. WTS foraged in restricted areas along their path, while TP predominantly foraged using extensive search behavior, suggesting a more opportunistic foraging strategy. Interspecific overlap of foraging areas was higher than intraspecific overlap. Breeding seasons largely overlap between species during the study, but TP seems to be asynchronous breeders. TP fed upon prey with higher δ15N values than WTS, and their diet was mainly composed of deep-sea organisms. TP could feed upon dead prey floating at the surface while WTS preyed mainly upon fish species that generally move in schools. Our study highlights several segregating mechanisms (temporal, behavioral and trophic) that could facilitate the coexistence of the two species despite the predominant number of WTS, and provides the very first information on the foraging and trophic ecology of the poorly-known TP.

A Model of Mercury Distribution in Tuna from the Western and Central Pacific Ocean: Influence of Physiology, Ecology and Environmental Factors.

Information on ocean scale drivers of methylmercury levels and variability in tuna is scarce, yet crucial in the context of anthropogenic mercury (Hg) inputs and potential threats to human health. Here we assess Hg concentrations in three commercial tuna species (bigeye, yellowfin, and albacore, n = 1000) from the Western and Central Pacific Ocean (WCPO). Models were developed to map regional Hg variance and understand the main drivers. Mercury concentrations are enriched in southern latitudes (10°S-20°S) relative to the equator (0°-10°S) for each species, with bigeye exhibiting the strongest spatial gradients. Fish size is the primary factor explaining Hg variance but physical oceanography also contributes, with higher Hg concentrations in regions exhibiting deeper thermoclines. Tuna trophic position and oceanic primary productivity were of weaker importance. Predictive models perform well in the Central Equatorial Pacific and Hawaii, but underestimate Hg concentrations in the Eastern Pacific. A literature review from the global ocean indicates that size tends to govern tuna Hg concentrations, however regional information on vertical habitats, methylmercury production, and/or Hg inputs are needed to understand Hg distribution at a broader scale. Finally, this study establishes a geographical context of Hg levels to weigh the risks and benefits of tuna consumption in the WCPO.

Differential uses of coral reef habitats by a poorly-known cryptic fish predator.

This study used otolith microchemistry to evaluate whether the moray eel Gymnothorax chilospilus uses different habitats throughout its life (mainly juvenile and adult phases). Of the most informative trace elements within otoliths (the twelve isotopes 23 Na, 25 Mg, 43 Ca, 55 Mn, 59 Co, 60 Ni, 63 Cu, 66 Zn, 86 Sr, 111 Cd, 138 Ba and 208 Pb) only three ratios of Ca (Na:Ca, Sr:Ca and Ba:Ca) were informative and therefore used in a multivariate regression-tree analysis. Using a multivariate partitioning, three main phases were described from profiles, including the larval life phase (leptocephali), the intermediate phase (longest section between the larval life phase and the terminal phase) and the terminal phase (final section i.e., the most recent months preceding the death of fish). According to concentrations of the three ratios to Ca, G. chilospilus can be separated into three groups during their larval life stage (very different in Sr and Na), four groups during the intermediate phase (few differences in Sr and Na) and three groups during the terminal phase (differences in Sr), illustrating that G. chilospilus inhabit different habitats during these three phases. Our results showed that the leptocephali encountered different oceanic water masses with fluctuating Sr:Ca ratios during the early larval phase. During the intermediate phase (main part of their life-span), they lived in lagoonal waters such as fringing reefs or reef flats of lagoonal islets, characterized by a lower Sr:Ca ratio. During the latter part of their life, approximately one third of G. chilospilus encountered more oceanic waters close to or at barrier reefs, suggesting possible movements of these fish along a coast-to-ocean gradient.

Tracking trace elements into complex coral reef trophic networks.

The integration, accumulation and transfer of trace elements across the main tropic levels of many food webs are poorly documented. This is notably the case for the complex trophic webs of coral reef ecosystems. Our results show that in the south-west lagoon of New Caledonia both abiotic (i.e. sediments) and biotic (i.e. primary producers, consumers and predators) compartments are contaminated by trace elements. However, our analyses revealed different contamination patterns from the sources of organic matter to the predators. The trophic levels involved in the sedimentary benthic food web (S-BFW, based on the sedimentary organic matter) and to a lesser extent in the reef benthic food web (R-BFW, based on algal turf) were mainly contaminated by trace elements that originate from mining activities like Ni and associated trace elements (Co, Cr, Fe, and Mn). Trace elements linked to agro-industrial (As, Hg, and Zn) and urban (Ag, Cd, Cu, Pb, Se, and V) activities were also integrated into the S-BFW, but preferentially into the R-BFW, and to a lesser extent into the detrital benthic food web (D-BFW, supplied by sea-grass plants). Most of the trace elements were biodiminished with increasing trophic levels along food webs. However, a marked biomagnification was observed for Hg, and suspected for Se and Zn. These results provide important baseline information to better interpret trace element contamination in the different organisms and trophic levels in a highly diversified coral reef lagoon.

Major Sources of Organic Matter in a Complex Coral Reef Lagoon: Identification from Isotopic Signatures (δ13C and δ15N).

A wide investigation was conducted into the main organic matter (OM) sources supporting coral reef trophic networks in the lagoon of New Caledonia. Sampling included different reef locations (fringing, intermediate and barrier reef), different associated ecosystems (mangroves and seagrass beds) and rivers. In total, 30 taxa of macrophytes, plus pools of particulate and sedimentary OM (POM and SOM) were sampled. Isotopic signatures (C and N) of each OM sources was characterized and the composition of OM pools assessed. In addition, spatial and seasonal variations of reef OM sources were examined. Mangroves isotopic signatures were the most C-depleted (-30.17 ± 0.41 ‰) and seagrass signatures were the most C-enriched (-4.36 ± 0.72 ‰). Trichodesmium spp. had the most N-depleted signatures (-0.14 ± 0.03 ‰) whereas mangroves had the most N-enriched signatures (6.47 ± 0.41 ‰). The composition of POM and SOM varied along a coast-to-barrier reef gradient. River POM and marine POM contributed equally to coastal POM, whereas marine POM represented 90% of the POM on barrier reefs, compared to 10% river POM. The relative importance of river POM, marine POM and mangroves to the SOM pool decreased from fringing to barrier reefs. Conversely, the relative importance of seagrass, Trichodesmium spp. and macroalgae increased along this gradient. Overall, spatial fluctuations in POM and SOM were much greater than in primary producers. Seasonal fluctuations were low for all OM sources. Our results demonstrated that a large variety of OM sources sustain coral reefs, varying in their origin, composition and role and suggest that δ13C was a more useful fingerprint than δ15N in this endeavour. This study also suggested substantial OM exchanges and trophic connections between coral reefs and surrounding ecosystems. Finally, the importance of accounting for environmental characteristics at small temporal and spatial scales before drawing general patterns is highlighted.

The great melting pot. Common sole population connectivity assessed by otolith and water fingerprints.

Quantifying the scale and importance of individual dispersion between populations and life stages is a key challenge in marine ecology. The common sole (Solea solea), an important commercial flatfish in the North Sea, Atlantic Ocean and the Mediterranean Sea, has a marine pelagic larval stage, a benthic juvenile stage in coastal nurseries (lagoons, estuaries or shallow marine areas) and a benthic adult stage in deeper marine waters on the continental shelf. To date, the ecological connectivity among these life stages has been little assessed in the Mediterranean. Here, such an assessment is provided for the first time for the Gulf of Lions, NW Mediterranean, based on a dataset on otolith microchemistry and stable isotopic composition as indicators of the water masses inhabited by individual fish. Specifically, otolith Ba/Ca and Sr/Ca profiles, and δ(13)C and δ(18)O values of adults collected in four areas of the Gulf of Lions were compared with those of young-of-the-year collected in different coastal nurseries. Results showed that a high proportion of adults (>46%) were influenced by river inputs during their larval stage. Furthermore Sr/Ca ratios and the otolith length at one year of age revealed that most adults (∼70%) spent their juvenile stage in nurseries with high salinity, whereas the remainder used brackish environments. In total, data were consistent with the use of six nursery types, three with high salinity (marine areas and two types of highly saline lagoons) and three brackish (coastal areas near river mouths, and two types of brackish environments), all of which contributed to the replenishment of adult populations. These finding implicated panmixia in sole population in the Gulf of Lions and claimed for a habitat integrated management of fisheries.

Anguilliform fish reveal large scale contamination by mine trace elements in the coral reefs of New Caledonia.

Due to intensive mining activity, increasing urbanization and industrialization, vast amounts of contaminants are discharged into the lagoon of New Caledonia, one of the largest continuous coral reef systems and a major biodiversity hotspot. The levels of 11 trace element concentrations were examined in the muscles of predator fish in the south-western lagoon (moray eels and congers). These species are sedentary, widespread, abundant, and they are easily collected using a sea snake sampling technique. We found the highest mean and maximal concentrations of different trace elements ever found in coral fish, notably regarding trace elements typical from mining activity (e.g., mean values for Cr and Ni, respectively: 5.53 ± 6.99 µg g(-1) [max, 35.7 µg g(-1)] and 2.84 ± 3.38 µg g(-1) [max, 18.0 µg g(-1)]). Results show that important trace element contamination extends throughout the lagoon to the barrier reef, following a concentration gradient from the oldest nickel factory (Nouméa).

The coral sea: physical environment, ecosystem status and biodiversity assets.

The Coral Sea, located at the southwestern rim of the Pacific Ocean, is the only tropical marginal sea where human impacts remain relatively minor. Patterns and processes identified within the region have global relevance as a baseline for understanding impacts in more disturbed tropical locations. Despite 70 years of documented research, the Coral Sea has been relatively neglected, with a slower rate of increase in publications over the past 20 years than total marine research globally. We review current knowledge of the Coral Sea to provide an overview of regional geology, oceanography, ecology and fisheries. Interactions between physical features and biological assemblages influence ecological processes and the direction and strength of connectivity among Coral Sea ecosystems. To inform management effectively, we will need to fill some major knowledge gaps, including geographic gaps in sampling and a lack of integration of research themes, which hinder the understanding of most ecosystem processes.

Global human footprint on the linkage between biodiversity and ecosystem functioning in reef fishes.

Difficulties in scaling up theoretical and experimental results have raised controversy over the consequences of biodiversity loss for the functioning of natural ecosystems. Using a global survey of reef fish assemblages, we show that in contrast to previous theoretical and experimental studies, ecosystem functioning (as measured by standing biomass) scales in a non-saturating manner with biodiversity (as measured by species and functional richness) in this ecosystem. Our field study also shows a significant and negative interaction between human population density and biodiversity on ecosystem functioning (i.e., for the same human density there were larger reductions in standing biomass at more diverse reefs). Human effects were found to be related to fishing, coastal development, and land use stressors, and currently affect over 75% of the world's coral reefs. Our results indicate that the consequences of biodiversity loss in coral reefs have been considerably underestimated based on existing knowledge and that reef fish assemblages, particularly the most diverse, are greatly vulnerable to the expansion and intensity of anthropogenic stressors in coastal areas.