Relationships between structural complexity,coral traits,and reef fish assemblages

With the ongoing loss of coral cover and the associated flattening of reef architecture, understanding the links between coral habitat and reef fishes is of critical importance. Here, we investigate whether considering coral traits and functional diversity provides new insights into the relationship between structural complexity and reef fish communities, and whether coral traits and community composition can predict structural complexity. Across 157 sites in Seychelles, Maldives, the Chagos Archipelago, and Australia’s Great Barrier Reef, we find that structural complexity and reef zone are the strongest and most consistent predictors of reef fish abundance, biomass, species richness, and trophic structure. However, coral traits, diversity, and life histories provided additional predictive power for models of reef fish assemblages, and were key drivers of structural complexity. Our findings highlight that reef complexity relies on living corals—with different traits and life histories—continuing to build carbonate skeletons, and that these nuanced relationships between coral assemblages and habitat complexity can affect the structure of reef fish assemblages. Seascape-level estimates of structural complexity are rapid and cost effective with important implications for the structure and function of fish assemblages, and should be incorporated into monitoring programs.     

Relative Importance of Coral Cover,Habitat Complexity and Diversity in Determining the Structure of Reef Fish Communities

The structure of coral reef habitat has a pronounced influence on the diversity, composition and abundance of reef-associated fishes. However, the particular features of the habitat that are most critical are not always known. Coral habitats can vary in many characteristics, notably live coral cover, topographic complexity and coral diversity, but the relative effects of these habitat characteristics are often not distinguished. Here, we investigate the strength of the relationships between these habitat features and local fish diversity, abundance and community structure in the lagoon of Lizard Island, Great Barrier Reef. In a spatial comparison using sixty-six 2m² quadrats, fish species richness, total abundance and community structure were examined in relation to a wide range of habitat variables, including topographic complexity, habitat diversity, coral diversity, coral species richness, hard coral cover, branching coral cover and the cover of corymbose corals. Fish species richness and total abundance were strongly associated with coral species richness and cover, but only weakly associated with topographic complexity. Regression tree analysis showed that coral species richness accounted for most of the variation in fish species richness (63.6%), while hard coral cover explained more variation in total fish abundance (17.4%), than any other variable. In contrast, topographic complexity accounted for little spatial variation in reef fish assemblages. In degrading coral reef environments, the potential effects of loss of coral cover and topographic complexity are often emphasized, but these findings suggest that reduced coral biodiversity may ultimately have an equal, or greater, impact on reef-associated fish communities.     

Response of fish-based metrics to anthropogenic pressures in temperate rocky reefs

The increasing degradation of marine ecosystems as a result of increasing impact caused by anthropogenic pressures, urges for well-founded knowledge to develop efficient tools to appraise the quality status of fish assemblages, as required by the “Marine Strategy Framework Directive”. This study analyzed the structural and functional response of rocky fish assemblages to several pressures on the Portuguese coast, i.e. fishing, sewage discharges, port activities and thermal effluent, by selecting fish-based metrics that best distinguished disturbed from control areas. One of the novel aspects of this research is the integrated assessment made through the analysis of several metrics representing numerous attributes of fish assemblages (namely diversity, abundance, trophic structure, mobility, resilience, habitat association, nursery function), which contrasts with the most commonly used approaches that in general focus on fish species/families. PERMANOVA results showed significant differences on metrics composition for all pressures with the exception of the thermal effluent. Moreover, two major patterns of stress were identified: (1) selective pressure, which affects differentially the fish assemblages (fishing); (2) broad-range pressure, which affects the entire fish assemblage with metrics of several attributes (e.g. structure, resilience, trophic guilds, nursery function) responding to its presence (sewage discharges, port activities). Taking into account the sensitivity results (discriminant analysis and Mann–Whitney test), biological meaning and redundancy with other metrics (Spearman correlations), the following metrics were selected as the most suitable to detect changes on temperate reef fish assemblages: “density of generalist individuals”, “density of territorial individuals”, “density of large individuals with medium to high commercial value (>20 cm)”, “density of juveniles” and metrics relative to trophic guild (except zooplanktivores). Since metrics grouped species that have some degree of functional overlap, the present approach was useful to understand human-induced changes at the assemblage level, contributing for the future use of marine fishes as biological indicators.     

Temperate macroalgae impacts tropical fish recruitment at forefronts of range expansion

Warming waters and changing ocean currents are increasing the supply of tropical fish larvae to temperature regions where they are exposed to novel habitats, namely temperate macroalgae and barren reefs. Here, we use underwater surveys on the temperate reefs of south-eastern (SE) Australia and western Japan (~33.5°N and S, respectively) to investigate how temperate macroalgal and non-macroalgal habitats influence recruitment success of a range of tropical fishes. We show that temperate macroalgae strongly affected recruitment of many tropical fish species in both regions and across three recruitment seasons in SE Australia. Densities and richness of recruiting tropical fishes, primarily planktivores and herbivores, were over seven times greater in non-macroalgal than macroalgal reef habitat. Species and trophic diversity (K-dominance) were also greater in non-macroalgal habitat. Temperate macroalgal cover was a stronger predictor of tropical fish assemblages than temperate fish assemblages, reef rugosities or wave exposure. Tropical fish richness, diversity and density were greater on barren reef than on reef dominated by turfing algae. One common species, the neon damselfish (Pomacentrus coelestis), chose non-macroalgal habitat over temperate macroalgae for settlement in an aquarium experiment. This study highlights that temperate macroalgae may partly account for spatial variation in recruitment success of many tropical fishes into higher latitudes. Hence, habitat composition of temperate reefs may need to be considered to accurately predict the geographic responses of many tropical fishes to climate change.

     

Cross-scale Habitat Structure Drives Fish Body Size Distributions on Coral Reefs

Despite a large number of studies focusing on the complexity of coral reef habitats and the characteristics of associated fish assemblages, the relationship between reef structure and fish assemblages remains unclear. The textural discontinuity hypothesis, which proposes that multi-modal body size distributions of organisms are driven by discontinuous habitat structure, provides a theoretical basis that may explain the influence of habitat availability on associated organisms. In this study we use fractal techniques to characterize patterns of cross-scale habitat complexity, and examine how this relates to body-depth abundance distributions of associated fish assemblages over corresponding spatial scales. Our study demonstrates that: (1) Reefs formed from different underlying substrata exhibit distinct patterns of cross-scale habitat complexity; (2) The availability of potential refuges at different scales correlates with patterns in fish body depth distributions, but habitat structure is more strongly related to the relative abundance of fish in the body depth modes, rather than to the number of modes; (3) As reefs change from coral- to algal-dominated states, the complexity of the underlying reef substratum may change, presenting a more homogenous environment to associated assemblages; (4) Individual fish body depth distributions may be multi-modal, however, these distributions are not static characteristics of the fish assemblage and may change to uni-modal forms in response to changing habitat condition. In light of predicted anthropogenic changes, there is a clear need to improve our understanding of the scale of ecological relationships to anticipate future changes and vulnerabilities.     

Community Structure of Fishes and Habitat Complexity on a Tropical Rocky Shore

The relationship between the variables of reef fish community structure (fish richness, fish diversity and total number of fishes) and those of habitat complexity (total surface area, substratum diversity, topographic complexity, number of holes, percent cover of turf algae, zoanthids, millepores, massive corals, bare rock, encrusting calcareous algae, urchins, other sessile organisms and Sargassum) were examined on three different rocky shores (FA, FB and FT) at Arraial do Cabo, a tropical region located on Brazilian southeastern coast (23° S, 42° W). Fish abundance and vertical distribution were assessed by a visual census technique using strip transects. Percentage cover of benthic organisms and other substratum types were calculated by replicated transects using a chain laid down on the substratum. Topographic complexity was determined by the chain link method and number of holes were estimated by direct counts on replicated transects. More than 91 fishes belonging to 37 families were seen in all study sites during one year of visual census effort. FA and FB sites had similar fish community structure and habitat complexity characteristics, while FT showed different habitat characteristics and higher fish diversity and richness, plus total number of fishes. Vertical distribution of fishes along the rocky shores studied seemed to be predictable and was determined by factors such as feeding habits and behavior, availability of refuges and social interactions. The habitat variables that best explained the higher diversity and number of fishes observed in FT site were total surface area of rocky shores and the abundance of benthic sessile invertebrates; these conditions were typical of rocky shores more exposed to wave surge.     

Interactions between coral restoration and fish assemblages: implications for reef management

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.     

Relationships between the spread of Caulerpa filiformis and fish communities on temperate rocky reefs

The previously sub‐dominant native marine macrophyte Caulerpa filiformis is now dominant on many sub‐tidal rocky reefs in New South Wales (NSW), Australia and is expanding its distribution. As C. filiformis is highly chemically defended and structurally different to co‐occurring habitat‐forming macrophytes, two key attributes that govern fish assemblages, we hypothesized that fish assemblages, particularly herbivorous fishes, would be different at sites where C. filiformis occurred from where it was previously absent and within sites, fish community structure would be correlated to the cover of C. filiformis. We investigated these hypotheses by determining reef‐associated fish assemblage attributes (assemblage structure, species richness, total abundance, Shannon‐Weiner diversity, abundance of herbivorous species) along transects within sites where C. filiformis was present and absent. Surprisingly, despite large patches and very high densities of C. filiformis on the reefs we sampled, at larger spatial scales (i.e., among sites) no fish assemblage metrics differed between sites with large stands of C. filiformis and sites without the alga. Moreover the abundance of one dominant herbivore, the rock cale Aplodactylus lophodon, was greater at sites within large beds of C. filiformis. At smaller spatial scales, however, i.e. within sites where C. filiformis was present, fish assemblages did vary as a function of C. filiformis cover along transects, although this was not consistent across sampling times. Overall, our results suggest that the potential effects of the spread of this alga on faunal communities warrants further investigation.     

Drivers of reef fish assemblages in an upwelling region from the Eastern Tropical Pacific Ocean

Reef fish assemblages are exposed to a wide range of anthropogenic threats as well as chronic natural disturbances. In upwelling regions, for example, there is a seasonal influx of cool nutrient-rich waters that may shape the structure and composition of reef fish assemblages. Given that climate change may disrupt the natural oceanographic processes by altering the frequency and strength of natural disturbances, understanding how fish assemblages respond to upwelling events is essential to effectively manage reef ecosystems under changing ocean conditions. This study used the baited remote underwater video stations (BRUVS) and the traditional underwater visual census (UVC) to investigate the spatiotemporal patterns of reef fish assemblages in an upwelling region in the North Pacific of Costa Rica. A total of 183 reef fish species from 60 families were recorded, of which 166 species were detected using BRUVS and 122 using UVC. Only 66% of all species were detected using both methods. This study showed that the upwelling had an important role in shaping reef fish assemblages in this region, but there was also a significant interaction between upwelling and location. In addition, other drivers such as habitat complexity and habitat composition had an effect on reef fish abundances and species. To authors’ knowledge, this is the first study in the Eastern Tropical Pacific that combines BRUVS and UVC to monitor reef fish assemblages in an upwelling region, which provides more detailed information to assess the state of reef ecosystems in response to multiple threats and changing ocean conditions.     

Reef fish and habitat relationships in a Caribbean seascape: the importance of reef context

Marine protected area (MPA) effectiveness is contingent on understanding key ecological patterns and processes at appropriate spatial scales and may depend upon maintaining critical linkages among essential habitat patches to conserve reef-fish communities. Hypotheses were tested to investigate the importance of habitat linkages in the US Virgin Islands. As expected, reef context (the spatial pattern of surrounding habitat patches) was a strong predictor of reef fish assemblage structure. Specific relationships were functionally consistent with the ecology of the fishes of interest. For example, reefs with large amounts of seagrass nearby harbored the greatest numerical abundance of fishes, particularly mobile invertebrate feeders and the exploited fish families of Haemulidae (grunts) and Lutjanidae (snappers). Species richness for the entire fish community and within these fish groups was also strongly associated with reef context. Furthermore, reef fish mobility influenced how fishes related to reef context. Fish-habitat relationships were detected as far as 1 km from study reefs, suggesting that fish movements result in habitat encounter rates that may influence their patterns of distribution. Consequently, functional habitat connectivity of habitat patches appears important in structuring reef-fish assemblages, and suggests that landscape-scale metrics may provide insights useful to managers in the design of MPAs.     

Effects of coral restoration on fish communities: snapshots of long‐term,multiregional responses and implications for practice

Coral restoration is widely used around the world to address dramatic declines in coral cover; however, very few studies have looked specifically at the temporal response of fish assemblages (i.e. abundance and diversity) to coral restoration. Several critical reef functions and processes are driven by fishes, thereby making their recovery and responses around restoration structures key indicators of success. This study evaluates fish abundance and community composition on restoration plots following 8–12 years of restoration activity, in four locations (two Caribbean and two Indo‐Pacific). Responses were very complex with region‐, site‐, and body size‐specific patterns. Overall, fish abundance only increased in Indo‐Pacific sites where damselfish responded positively to increased coral cover and topographic complexity. Restoration effects on other fish families and particularly on larger bodied reef fish were negative or neutral at all locations. If restoration initiatives are going to substantively improve the condition and recovery of degraded reef fish communities, restoration efforts need to be planned, designed, and monitored based on fish‐specific habitat requirements and locally specific community dynamics.     

LIDAR optical rugosity of coral reefs in Biscayne National Park,Florida

The NASA Experimental Advanced Airborne Research Lidar (EAARL), a temporal waveform-resolving, airborne, green wavelength LIDAR (light detection and ranging), is designed to measure the submeter-scale topography of shallow reef substrates. Topographic variability is a prime component of habitat complexity, an ecological factor that both expresses and controls the abundance and distribution of many reef organisms. Following the acquisition of EAARL coverage over both mid-platform patch reefs and shelf-margin bank reefs within Biscayne National Park in August 2002, EAARL-based optical indices of topographic variability were evaluated at 15 patch reef and bank reef sites. Several sites were selected to match reefs previously evaluated in situ along underwater video and belt transects. The analysis used large populations of submarine topographic transects derived from the examination of closely spaced laser spot reflections along LIDAR raster scans. At all 15 sites, each LIDAR transect was evaluated separately to determine optical rugosity (Ro_tran), and the average elevation difference between adjacent points (Av(E_ap)). Further, the whole-site mean and maximum values of Ro_tran and Av(E_ap) for the entire population of transects at each analysis site, along with their standard deviations, were calculated. This study revealed that the greater habitat complexity of inshore patch reefs versus outer bank reefs results in relative differences in topographic complexity that can be discerned in the laser returns. Accordingly, LIDAR sensing of optical rugosity is proposed as a complementary new technique for the rapid assessment of shallow coral reefs.     

Spatial variation and effects of habitat on temperate reef fish assemblages in northeastern New Zealand

Reef-associated fishes can respond to changes in habitat structure and the nature of their response can change with different spatial scales of observation. A structured hierarchical mensurative sampling design was used to sample temperate reef fish assemblages in northeastern New Zealand at several spatial scales over 2 years. The three spatial scales examined were tens of meters (transects), hundreds to thousands of meters (sites) and hundreds of kilometers (locations). We tested the hypothesis that fish assemblages differed between kelp forest habitat (relatively dense stands of the kelp, Ecklonia radiata (C. Agardh) J. Agardh, median depth=13.5 m) and barrens habitat (rocky reef dominated by turfing and encrusting red algae and the grazing urchin, Evechinus chloroticus (Valenciennes), median depth=6.7 m). Recently developed multivariate techniques were used to test for and quantify multivariate variation at different spatial scales. There were significant effects of habitat on the spatial distribution of fish assemblages, characterised by greater abundances or frequencies of Parika scaber, Chromis dispilus, Trachurus novaezelandiae, Nemadactylus douglasii, Bodianus unimaculatus, Odax pullus and Pseudolabrus miles in kelp forest habitat, and greater abundances or frequencies of Notolabrus celidotus, Notolabrus fucicola, Girella tricuspidata, Coris sandageri, Chironemus marmoratus, Parma alboscapularis, Scorpis violaceus and Kyphosus sydneyanus in barrens habitat. Some of the more common species, including Upeneichthys lineatus, Scorpis lineolatus and Cheilodactylus spectabilis showed no strong consistent effects of these two differing habitats on their distributions. There was, however, a significant Habitat×Locations interaction: effects of habitat did not occur at all locations. Variability was highest at the scale of individual transects and variability from site to site and from location to location was comparable. Spatial variation was large compared to inter-annual variation, which was minimal, and spatial patterns were consistent in the 2 years examined. Further experiments, including manipulations, are required to understand what mechanisms and processes might be driving these patterns. This study, coupled with results from previous studies, suggests that there may be a dynamic inter-play between effects of habitat on fish and effects of fish on biogenic habitat, such as kelp forests.     

Habitat type and complexity drive fish assemblages in a tropical seascape

Inshore marine seascapes support a diversity of interconnected habitats and are an important focus for biodiversity conservation. This study examines the importance of habitat attributes to fish assemblages across a mosaic of inshore habitats: coral reefs, rocky reefs, macroalgae beds and sand/rubble beds. Fishes and benthic habitats were surveyed at 34 sites around continental islands of the central Great Barrier Reef using baited remote underwater video stations (BRUVS). Species richness was influenced foremost by habitat type and also by structural complexity within habitat types. The most speciose assemblages occurred in coral and rocky reef habitats with high structural complexity, provided by the presence of coral bommies/overhangs, boulders and rock crevices. Nonetheless, macroalgae and sand/rubble beds also supported unique species, and therefore contributed to the overall richness of fish assemblages in the seascape. Most trophic groups had positive associations with complexity, which was the most important predictor for abundance of piscivorous fishes and mobile planktivores. There was significant differentiation of fish assemblages among habitats, with the notable exception of coral and rocky reefs. Species assemblages overlapped substantially between coral and rocky reefs, which had 60% common species, despite coral cover being lower on rocky reefs. This suggests that, for many species, rocky and coral substrates can provide equivalent habitat structure, emphasizing the importance of complexity in providing habitat refuges, and highlighting the contribution of rocky reefs to habitat provision within tropical seascapes. The results of this study support an emerging recognition of the collective value of habitat mosaics in inshore marine ecosystems.     

Temporal trends in algae, benthic invertebrate, and fish assemblages in streams and rivers draining basins of varying land use in the south-central United States, 1993–2007

Site-specific temporal trends in algae, benthic invertebrate, and fish assemblages were investigated in 15 streams and rivers draining basins of varying land use in the south-central United States from 1993–2007. A multivariate approach was used to identify sites with statistically significant trends in aquatic assemblages which were then tested for correlations with assemblage metrics and abiotic environmental variables (climate, water quality, streamflow, and physical habitat). Significant temporal trends in one or more of the aquatic assemblages were identified at more than half (eight of 15) of the streams in the study. Assemblage metrics and abiotic environmental variables found to be significantly correlated with aquatic assemblages differed between land use categories. For example, algal assemblages at undeveloped sites were associated with physical habitat, while algal assemblages at more anthropogenically altered sites (agricultural and urban) were associated with nutrient and streamflow metrics. In urban stream sites results indicate that streamflow metrics may act as important controls on water quality conditions, as represented by aquatic assemblage metrics. The site-specific identification of biotic trends and abiotic–biotic relations presented here will provide valuable information that can inform interpretation of continued monitoring data and the design of future studies. In addition, the subsets of abiotic variables identified as potentially important drivers of change in aquatic assemblages provide policy makers and resource managers with information that will assist in the design and implementation of monitoring programs aimed at the protection of aquatic resources.     

Habitat specialization in tropical continental shelf demersal fish assemblages

The implications of shallow water impacts such as fishing and climate change on fish assemblages are generally considered in isolation from the distribution and abundance of these fish assemblages in adjacent deeper waters. We investigate the abundance and length of demersal fish assemblages across a section of tropical continental shelf at Ningaloo Reef, Western Australia, to identify fish and fish habitat relationships across steep gradients in depth and in different benthic habitat types. The assemblage composition of demersal fish were assessed from baited remote underwater stereo-video samples (n = 304) collected from 16 depth and habitat combinations. Samples were collected across a depth range poorly represented in the literature from the fringing reef lagoon (1-10 m depth), down the fore reef slope to the reef base (10-30 m depth) then across the adjacent continental shelf (30-110 m depth). Multivariate analyses showed that there were distinctive fish assemblages and different sized fish were associated with each habitat/depth category. Species richness, MaxN and diversity declined with depth, while average length and trophic level increased. The assemblage structure, diversity, size and trophic structure of demersal fishes changes from shallow inshore habitats to deeper water habitats. More habitat specialists (unique species per habitat/depth category) were associated with the reef slope and reef base than other habitats, but offshore sponge-dominated habitats and inshore coral-dominated reef also supported unique species. This suggests that marine protected areas in shallow coral-dominated reef habitats may not adequately protect those species whose depth distribution extends beyond shallow habitats, or other significant elements of demersal fish biodiversity. The ontogenetic habitat partitioning which is characteristic of many species, suggests that to maintain entire species life histories it is necessary to protect corridors of connected habitats through which fish can migrate.     

How will coral reef fish communities respond to climate-driven disturbances? Insight from landscape-scale perturbations

Global climate change is rapidly altering disturbance regimes in many ecosystems including coral reefs, yet the long-term impacts of these changes on ecosystem structure and function are difficult to predict. A major ecosystem service provided by coral reefs is the provisioning of physical habitat for other organisms, and consequently, many of the effects of climate change on coral reefs will be mediated by their impacts on habitat structure. Therefore, there is an urgent need to understand the independent and combined effects of coral mortality and loss of physical habitat on reef-associated biota. Here, we use a unique series of events affecting the coral reefs around the Pacific island of Moorea, French Polynesia to differentiate between the impacts of coral mortality and the degradation of physical habitat on the structure of reef fish communities. We found that, by removing large amounts of physical habitat, a tropical cyclone had larger impacts on reef fish communities than an outbreak of coral-eating sea stars that caused widespread coral mortality but left the physical structure intact. In addition, the impacts of declining structural complexity on reef fish assemblages accelerated as structure became increasingly rare. Structure provided by dead coral colonies can take up to decades to erode following coral mortality, and, consequently, our results suggest that predictions based on short-term studies are likely to grossly underestimate the long-term impacts of coral decline on reef fish communities.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.