Geographical gradients of marine herbivorous fishes: patterns and processes

We present new data and the first rigorous analysis of latitudinal and thermal gradients of diversity, density and biomass of marine herbivorous fishes and review proposed explanatory mechanisms. Consistently, negative relationships between latitude, and positive relationships between sea surface temperature (SST), and relative richness and relative abundance of herbivorous fishes were found worldwide. Significant differences in the strength of gradients of richness and abundance with latitude and SST between tropical and extratropical zones were found consistently across ocean basins. Standardized sampling along the western Atlantic also showed negative relationships between latitude and total density and biomass. The trends, however, are driven by different components of the fish assemblages (i.e., scarids in the Caribbean and acanthurids in Brazil). Patterns of abundance along thermal gradients, generally associated with extensive latitudinal gradients, also were found at the local scale. Feeding rate of the ocean surgeonfish Acanthurus bahianus decreases with temperature more rapidly than the mean metabolic rate of teleost fishes. This relationship suggests a temperature-related physiological constraint. From the new standardized and comparative data presented and the review of the explanatory hypotheses, we conclude that temperature-related feeding and digestive processes are most likely involved in the distribution patterns of herbivorous fishes. Electronic Supplementary Material Supplementary material is available for this article at     

Latitudinal variation in abundance of herbivorous fishes: a comparison of temperate and tropical reefs

The aim of the study was to provide comparable estimates of abundance of herbivorous reef fishes at temperate and tropical localities using a standardized methodology. Faunas of herbivorous fish were sampled on the rocky reefs of temperate northern New Zealand and on the coral reefs of the northern Great Barrier Reef (GBR), Australia, and the San Blas Archipelago in the Caribbean. A pilot study established the most appropriate habitat setting and the scale and magnitude of replication for the sampling program in temperate waters. Herbivorous fishes, including members of families endemic to the southern hemisphere (Odacidae and Aplodactylidae), were most abundant in turbulent, shallow water (0 to 6 m) and had patchy distributions within this habitat. A hierarchical sampling program using 10-min transect counts within the 0 to 6 m depth stratum examined abundance patterns at a range of spatial scales including mainland and island coasts, localities separated by up to 100 km and sites separated by up to 10 km. This program identified a characteristic fauna of seven species of herbivorous fishes with mean total abundances ranging from 23 to 30 individuals per 10-min transect. Species composition of the fauna varied between islands and coasts. A similar methodology was used to sample the major families of herbivorous fish in a number of sites in each of the tropical regions. These sampling programs revealed a fauna dominated by acanthurids and scarids in both the GBR and Caribbean localities. Estimates of abundance from these regions were similar, with a mean of 108 individuals recorded on the GBR and 129 per 10-min transect in the Caribbean. Species richness varied between each region, with 44 taxa recorded from the GBR and 11 from the Caribbean. Abundances of temperate water herbivores in New Zealand were found to be 75 to 80% lower than those recorded from shallow water habitats sampled on coral reefs. This was not related to species richness, since both New Zealand and the Caribbean locality had patterns of low richness. We suggest that the differences in abundance found by our study between temperate and tropical regions are not restricted to herbivorous fishes, but are representative of general latitudinal trends in reef fish faunas. Received: 4 November 1996 / Accepted: 15 December 1996     

Zur Morphologie,Histochemie, Tagesperiodik und Biologie der Operculardrüse bei Labriden und Scariden (Pisces)

A mucous gland is described, which is responsible for the production of the slime envelope in scarids and labrids. For this gland the name ‘opercular gland’ is suggested. The gland is developed, to a greater or lesser extent, in most of the genera examined. Morphological and histochemical studies indicate that the form of the gland is characteristic of the genus. Labrids and scarids can be divided, histochemically, into 3 groups: (1) opercular gland secretes only neutral MPS; (2) opercular gland secretes both neutral and weakly acidic MPS; (3) opercular gland secretes neutral MPS, weakly acidic and strongly acidic MPS (the strongly acidic MPS are probably sulphated). Species are found in all 3 groups with granular cells which secrete a tyrosine containing glycoprotein. The fish can be made to produce the mucous envelope by stimulation of the parasympathetic nervous system using pilocarpine. It is, therefore, assumed that the gland is controlled by the nervous system. Morphological studies of the gland show that the mucous is normally synthesised only in the evening, i.e. when it is needed; no storage takes place. Continuous light experiments have shown the gland to possess an endogenous rhythm of the same frequency as the fish's activity rhythm. Even those labrids which sleep under sand produce mucous. The average area of the longitudinal section of an opercular gland cell is more than 500 µ² in those species which produce a complete envelope, and usually somewhat less in the other species examined. The possible importance of the opercular gland in taxonomical and phylogenetical studies is discussed.     

Stream ecosystem response to limestone treatment in acid impacted watersheds of the Allegheny Plateau.

Restoration programs are expanding worldwide, but assessments of restoration effectiveness are rare. The objectives of our study were to assess current acid-precipitation remediation programs in streams of the Allegheny Plateau ecoregion of West Virginia (USA), identify specific attributes that could and could not be fully restored, and quantify temporal trends in ecosystem recovery. We sampled water chemistry, physical habitat, periphyton biomass, and benthic macroinvertebrate and fish community structure in three stream types: acidic (four streams), naturally circumneutral (eight streams), and acidic streams treated with limestone sand (eight streams). We observed no temporal trends in ecosystem recovery in treated streams despite sampling streams that ranged from 2 to 20 years since initial treatment. Our results indicated that the application of limestone sand to acidic streams was effective in fully recovering some characteristics, such as pH, alkalinity, Ca2+, Ca:H ratios, trout biomass and density, and trout reproductive success. However, recovery of many other characteristics was strongly dependent upon spatial proximity to treatment, and still others were never fully recovered. For example, limestone treatment did not restore dissolved aluminum concentrations, macroinvertebrate taxon richness, and total fish biomass to circumneutral reference conditions. Full recovery may not be occurring because treated streams continue to drain acidic watersheds and remain isolated in a network of acidic streams. We propose a revised stream restoration plan for the Allegheny Plateau that includes restoring stream ecosystems as connected networks rather than isolated reaches and recognizes that full recovery of acidified watersheds may not be possible.     

Wave exposure,swimming performance,and the structure of tropical and temperate reef fish assemblages

We examined the relationship between swimming performance, wave exposure, and the distribution patterns of labrids on temperate rocky reefs, in comparison with previous functional analyses of a tropical assemblage. Visual censuses of the distribution and abundance of labrids across two major gradients of wave exposure (depth and aspect to prevailing winds) were made at two offshore islands near Port Stephens, New South Wales, Australia. Distinct shifts in species composition and abundance were evident between high and low wave exposure habitats on temperate rocky reefs, particularly between deep and shallow habitats on exposed reef fronts. The swimming performances of temperate labrids were assessed through examination of pectoral fin shape (aspect ratio) and in situ swimming speeds. A diversity of pectoral fin morphologies was exhibited within this temperate assemblage, ranging from rounded to tapered fins (aspect ratios of 0.52 and 1.43, respectively). Fin shape was strongly correlated (Pearsons correlation 0.884, P<0.001) with swimming speed (ranging from 1.05 and 3.06 body lengths s^–1), in a relationship comparable to that observed in tropical labrids. Inter-specific differences in swimming ability provided some explanation for differences in the distribution and abundance of temperate labrids in relation to wave exposure. However, our findings suggest that although coral reef labrids appear to predominantly use high aspect-ratio fins to successfully occupy wave-exposed habitats, temperate labrids appear to be using an enhanced swimming ability through increased body size.Communicated by G.F. Humphrey, Sydney     

Long-term variation of organochlorine residues and assemblages of epibenthic organisms in a shallow North Florida (USA) estuary

A 4-year study (1972–1976) determined long-term trends of organochlorine residues (DDT, DDE, DDD, PCB's, mirex) and trawl-susceptible organisms in a shallow, river-dominated estuary in North Florida (Apalachicola Bay, USA). Moderate levels of such compounds were found in various species prior to the restricted use of DDT in 1972. A subsequent precipitous decline in organochlorine besidues was attributed to decreased upland usage, major flushing of the river basin in early 1973, and various factors associated with estuarine function. No mirex was found in sediments or aquatic organisms. Apparently, the half-life of organochlorines is relatively short in this bay system. Various statistical methods were used to test the relationships of different physico-chemical and biological parameters. During the 4-year study period, seasonal river flow fluctuations dominated water color, turbidity, salinity, nutrients (NO₃), chlorophyll a, and the temporal succession of fishes in the bay. Certain long-term trends of fish associations were noted; relative dominance of key fish species declined and stabilized while bay-wide species richness and diversity increased with time. Qualitative changes in species representation determined the long-term pattern of community variability. This was consonant with a distinctive fish fauna during the first year of sampling. The bay anchovy Anchoa mitchilli was dominant during 9 of the first 12 monts of the project; this influenced the time-related changes in community indices. Temporally clustered fish associations reflected the importance of river flow in the estuarine environment. Direct correlation of fish distribution with the rapid disappearance of organochlorine compounds was complicated by aperiodic natural phenomena such as storms and river fluctuations. Population and community trends appeared consistent with other studies showing similar patterns of dominance of stress-resistant fish populations and related changes in community parameters. In this case, the relatively predictable annual succession of fish associations allowed an appraisal of key forcing functions. Due to the high level of seasonal and annual biological variability in this estuary, there were some problems in the application of linear statistical models to the data base. Although the long-term trend of relative species representation is useful as an index of stress, new techniques are needed to analyze extensive field data so that functions such as trophic interactions are included in the estimation of causal relationships. There are indications that such effects could be related to the impact of organochlorine compounds on estuarine systems.     

Recovery of Faunal Communities During Tropical Forest Regeneration

Abstract:  As mature tropical forests are cleared, secondary forests may play an important role in the conservation of animal species, depending on how fast animal communities recover during forest regeneration. I reviewed published studies on the recovery of animal species richness and composition during tropical forest regeneration. In 38 of the 39 data sets I examined, conversion of forest to agriculture or pasture substantially reduced species richness. Given suitable conditions for forest recovery, the species richness of the animal taxa considered can be predicted to resemble that of mature forests roughly 20–40 years after land abandonment. At least for ants and birds, however, recovery of species composition appears to take substantially longer than recovery of species richness. Because species richness for many taxa appears to recover relatively rapidly in secondary forests, conservation of secondary forests may be an effective investment in future diversity. The slower recovery of species composition indicates, however, that some species will require stands of mature forest to persist.     

Marine reserves,fisheries ban,and 20 years of positive change in a coral reef ecosystem

By 2004, Belize was exhibiting classic fishing down of the food web. Groupers (Serranidae) and snappers (Lutjanidae) were scarce and fisheries turned to parrotfishes (Scarinae), leading to a 41% decline in their biomass. Several policies were enacted in 2009–2010, including a moratorium on fishing parrotfish and a new marine park with no-take areas. Using a 20-year time series on reef fish and benthos, we evaluated the impact of these policies approximately 10 years after their implementation. Establishment of the Southwater Caye Marine Reserve led to a recovery of snapper at 2 out of 3 sites, but there was no evidence of recovery outside the reserve. Snapper populations in an older reserve continued to increase, implying that at least 9 years is required for their recovery. Despite concerns over the feasibility of banning parrotfish harvest once it has become a dominant fin fishery, parrotfishes returned and exceeded biomass levels prior to the fishery. The majority of these changes involved an increase in parrotfish density; species composition and adult body size generally exhibited little change. Recovery occurred equally well in reserves and areas open to other forms of fishing, implying strong compliance. Temporal trends in parrotfish grazing intensity were strongly negatively associated with the cover of macroalgae, which by 2018 had fallen to the lowest levels observed since measurements began in 1998. Coral populations remained resilient and continued to exhibit periods of net recovery after disturbance. We found that a moratorium on parrotfish harvesting is feasible and appears to help constrain macroalgae, which can otherwise impede coral resilience.     

Occurrence and characteristics of unusual protistan symbionts from surgeonfishes (Acanthuridae) of the Great Barrier Reef,Australia

The occurrence of unusual symbiotic microorganisms was examined in the intestines of a range of fish from the Great Barrier Reef, Australia. The fish taxa examined included 26 species of the family Acanthuridae, as well as representatives of phylogenetically related and herbivorous taxa. The microorganisms, referred to as protists, were only found in herbivorous and detritivorous members of the Acanthuridae. Protists were not found in planktivorous acanthurids, nor in any members of the families Kyphosidae, Pomacentridae, Scaridae, Zanclidae, Siganidae and Bleniidae we examined. In addition, protists were absent from the herbivorous acanthurids A. xanthopterus and A. nigricans. A range of protist forms, characterized by differences in size (8 to 417 m), shape and mode of cell division (daughter-cell production and binary fission), was observed. The occurrence of these forms appeared to be correlated with host feedingecology. Large forms (>100 m) of the protists were only found in acanthurids which fed over hard-reef substrata. Smaller forms were found in sand-grazing and detritivorous species. One of the protist forms appears identical to protists previously reported from Red Sea acanthurids.     

Analysis of the influence of substrate variables on coral reef fish communities

A simple field technique to obtain a gross estimate of the surface area of a quadrat on a coral reef is described. This measure, termed the substrate rugosity index, was determined, in conjunction with two other substrate variables (vertical relief and coral species richness), in a series of 4 quadrats (10 to 40 m depth) along 4 transects. The mean substrate rugosity and vertical relief of a quadrat were highly correlated. A correlation analysis was made of the substrate variables and several reef fish community parameters (species richness, number of fishes and diversity). Species richness was highly correlated with substrate rugosity. This relationship was tested in two experimental quadrats and the results were generally in accord with those predicted. Stratification of the fish communities by body size revealed that the correlation with substrate rugosity was scale-dependent. The fish community parameters were poorly correlated with percentage substrate cover by corals (ramose and glomerate) and by sand. A significant area effect was determined for two species of sand-dwelling goby.     

Response to organic enrichment of infaunal macrobenthic communities under salmonid seacages

The response of infaunal macrobenthic communities beneath salmonid seacages to solid organic wastes (food pellets and fish excreta) was tested by means of the ABC-method (abundance, biomass comparison method: Warwick 1986) on samples collected from the SAFCOL seafarm at Badger Cove, southeast Tasmania, from February to July 1988. The intermittent nature of the addition of organic waste allowed us to monitor decline and recovery of the macrofauna, i.e., harvesting of a cage permitted a period of recovery whilst restocking precipitated a decline. The ABC-method proved to be a sensitive indicator of community health. Under a normal feeding regime the macrofaunal community structure indicated a moderately disturbed condition. Only 7 wk after the cage was harvested, species richness had increased markedly and the community adopted an undisturbed condition. Further improvement was apparent 14 wk post-harvest. Similarly, a decline to a moderately disturbed condition was apparent 7 wk after restocking and species richness had declined. No such changes occurred under a cage which contained fish continuously over the same period.     

Effects of Desertification Caused by Lithophaga lithophaga (Mollusca) Fishery on Littoral Fish Assemblages along Rocky Coasts of Southeastern Italy

Abstract:  We surveyed shallow, rocky reefs in southwestern Apulia (Mediterranean Sea) to assess the effects on coastal fish assemblages of the date mussel (  Lithophaga lithophaga ) fishery, an illegal practice that strips the rocky reef bare. We visually sampled fish four times over 15 months at three locations, one affected by date-mussel fishery and two controls. The fish assemblage at the affected location differed significantly from those at the control locations over all sampling times. Herbivorous fishes, sparids, and labrids (genus Symphodus ) contributed most to the differences between the affected location and controls. Lower densities of Symphodus spp. were observed at the affected location, whereas detritivorous fishes were recorded exclusively at control sites. Small serranids and sparids showed temporal trends that differed between the affected location and the control locations. Our results suggest that the date-mussel fishery affects fish assemblages chiefly through reduction of arborescent macroalgae (contributing to habitat complexity and primary production) and emphasize the need for more effective policing against this destructive practice.     

The role of sponge competition on coral reef alternative steady states

Sponges constitute an abundant and functionally important component of coral reef systems. Given their demonstrated resistance to environmental stress, it might be expected that the role of sponges in reef systems under modern regimes of frequent and severe disturbance may become even more substantial. Disturbances have recently reshaped the community structure of many Caribbean coral reefs shifting them towards a state of persistent low coral cover and often a dominance of macroalgae. Using competition and growth rates recorded from Glover's Atoll in Belize, we parameterise a mathematical model used to simulate the three-way competition between sponges, macroalgae and coral. We use the model to determine the range of parameters in which each of the three species might be expected to dominate. Emergent properties arise from our simple model of this complex system, and these include a special case in which heightened competitive ability of macroalgae versus coral may counter-intuitively prove to be advantageous to the persistence of corals. Importantly, we show that even under scenarios whereby sponges fail to invade the system, inclusion of this third antagonist can qualitatively affect the likelihood of alternative stable states - generally in favour of macroalgal dominance. The interplay between multi-species competition and predation is complex, but our efforts highlight a key process that has, until now, remained unexplored: the extent to which sponges dissipate algal grazing pressure by providing generalist fish with an alternative food source. We highlight the necessity of identifying the extent by which this process takes place in tropical systems in order to improve projections of alternative stable states for Caribbean coral reefs.     

Relative influence of environmental factors and fishing on coral reef fish assemblages

Understanding whether assemblages of species respond more strongly to bottom-up (availability of trophic resources or habitats) or top-down (predation pressure) processes is important for effective management of resources and ecosystems. We determined the relative influence of environmental factors and predation by humans in shaping the density, biomass, and species richness of 4 medium-bodied (10–40 cm total length [TL]) coral reef fish groups targeted by fishers (mesopredators, planktivores, grazer and detritivores, and scrapers) and the density of 2 groups not targeted by fishers (invertivores, small fish ≤10 cm TL) in the central Philippines. Boosted regression trees were used to model the response of each fish group to 21 predictor variables: 13 habitat variables, 5 island variables, and 3 fishing variables (no-take marine reserve [NTMR] presence or absence, NTMR size, and NTMR age). Targeted and nontargeted fish groups responded most strongly to habitat variables, then island variables. Fishing (NTMR) variables generally had less influence on fish groups. Of the habitat variables, live hard coral cover, structural complexity or habitat complexity index, and depth had the greatest effects on density, biomass, and species richness of targeted fish groups and on the density of nontargeted fishes. Of the island variables, proximity to the nearest river and island elevation had the most influence on fish groups. The NTMRs affected only fishes targeted by fishers; NTMR size positively correlated with density, biomass, and species richness of targeted fishes, particularly mesopredatory, and grazing and detritivorous fishes. Importantly, NTMRs as small as 15 ha positively affected medium-bodied fishes. This finding provides reassurance for regions that have invested in small-scale community-managed NTMRs. However, management strategies that integrate sound coastal land-use practices to conserve adjacent reef fish habitat, strategic NTMR placement, and establishment of larger NTMRs will be crucial for maintaining biodiversity and fisheries.     

A comparison of temperate reef fish assemblages recorded by three underwater stereo-video techniques

Three underwater stereo-video techniques were used to sample the relative densities and species richness of temperate reef fish assemblages at three reef locations and two habitats (high- and low-relief reef) within Hamelin Bay, south-western Australia. The three techniques compared were diver-operated stereo-video strip transects, baited remote stereo-video and unbaited remote stereo-video. While unbaited remote stereo-video and diver-operated stereo-video transects recorded greater species richness at high compared to low-relief reefs, baited remote stereo-video recorded similar species richness at the two habitat types. The diver-operated stereo-video system was manoeuvred through caves and under overhangs recording small, cryptic, cave-dwelling species that were not recorded by either remote video techniques (Trachinops noarlungae, Trachinops brauni, Chromis klunzingeri, Trachichthys australis). Both remote video techniques recorded greater species richness and relative density of the most common species of Labridae, Ophthalmolepsis lineolatus. Baited remote video recorded the rarer, large predatory fish species (e.g. Seriola hippos, Glaucosoma hebraicum, Heterodontus portusjacksoni). None of the techniques sampled small cryptic fish families such as Gobiidae or Blenniidae. A combination of survey techniques is recommended for comprehensive fishery-independent studies that aim to sample broad components of fish assemblages.An erratum to this article can be found at     

Changing Species Richness and Composition in Canadian National Parks

Abstract: Canada's national parks and their surrounding areas differ markedly in size, climate, vegetation, and extent of human development. We tested the extent to which total species richness, native species richness, and the number of extirpations and introductions of terrestrial vertebrates were correlated with each of these factors. To do this, we used surveys of park fauna from the present and from the time of park establishment. Richness, extirpations, and introductions were all strongly related to climate. After we controlled for climate, smaller parks had higher rates of species loss than larger parks. Land-use patterns (forest cover and fragmentation, roads, etc.) within parks were strongly correlated with land use in the regions surrounding the parks, showing that parks have not been isolated from regional development. Richness and extirpations within parks were generally more strongly related to regional characteristics than to the characteristics of the parks themselves. Species richness and numbers of introduced species were higher in parks found in landscapes with greater fragmentation. Frequencies of extirpations were less clearly related to human-influenced habitat characteristics. Introductions and extinctions most often involved game species or species directly associated with human activities. There is little evidence of subtle ecological effects being responsible for species loss. Our results suggest that management should focus on direct human interventions, such as hunting, introduction of game species, and habitat fragmentation, in parks and surrounding areas.     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

Trait groups as management entities in a complex,multispecies reef fishery

Localized stressors compound the ongoing climate-driven decline of coral reefs, requiring natural resource managers to work with rapidly shifting paradigms. Trait-based adaptive management (TBAM) is a new framework to help address changing conditions by choosing and implementing management actions specific to species groups that share key traits, vulnerabilities, and management responses. In TBAM maintenance of functioning ecosystems is balanced with provisioning for human subsistence and livelihoods. We first identified trait-based groups of food fish in a Pacific coral reef with hierarchical clustering. Positing that trait-based groups performing comparable functions respond similarly to both stressors and management actions, we ascertained biophysical and socioeconomic drivers of trait-group biomass and evaluated their vulnerabilities with generalized additive models. Clustering identified 7 trait groups from 131 species. Groups responded to different drivers and displayed divergent vulnerabilities; human activities emerged as important predictors of community structuring. Biomass of small, solitary reef-associated species increased with distance from key fishing ports, and large, solitary piscivores exhibited a decline in biomass with distance from a port. Group biomass also varied in response to different habitat types, the presence or absence of reported dynamite fishing activity, and exposure to wave energy. The differential vulnerabilities of trait groups revealed how the community structure of food fishes is driven by different aspects of resource use and habitat. This inherent variability in the responses of trait-based groups presents opportunities to apply selective TBAM strategies for complex, multispecies fisheries. This approach can be widely adjusted to suit local contexts and priorities.     

Natural and anthropogenic influences on the diversity structure of reef fish communities in the Tuamotu Archipelago (French Polynesia)

Despite the rapid rate of human-induced species losses, the relative influence of natural and anthropogenic factors on the functional diversity of species assemblages remains unknown for most ecosystems. A model was previously developed to predict the diversity structure of coral reef fish assemblages in 10 atolls of low human pressure and contrasting morphology of the Tuamotu Archipelago (French Polynesia). This existing model predicted smoothed histograms (spectra) of species richness according to size classes, diet classes and life-history classes of fish assemblages using a combination of environmental characteristics at different spatial scales. The present study applied the model to Tikehau, another atoll of the same archipelago where commercial fishing is practiced and where the same sampling strategy was reproduced. Significant differences appeared between predicted and observed species richness in several size, diet and life-history classes of fish assemblages in Tikehau. Two parameters which were not accounted for in the initial model, i.e. fishing pressure and atoll position within the archipelago, explained together 63% of variance in model residuals, >60% being explained by fishing pressure only. The respective effects of fishing and atoll position on the diversity of coral reef fish assemblages are discussed, with the potential of such modelling approach to assess the relative importance of factors affecting functional diversity within communities.