The determinants of species richness of a relatively young coral-reef ichthyofauna

Aim Unique topographic features left the Red Sea and its north‐eastern extension into the Gulf of Aqaba practically devoid of coral‐reef‐based organisms during the last glacial maximum. The current ichthyofauna in these two ‘regions’ thus represents the product of relatively recent colonization by species found in the Arabian Sea, which adjoins the Red Sea at its southern tip. We used this system to test why some marine species seemingly fail to extend their geographic range, thereby generating spatial heterogeneity in biodiversity. Location The Arabian Sea, Red Sea, and the Gulf of Aqaba. Methods A list of coral‐reef‐associated fish species, belonging to the 10 most speciose families, was compiled for each region using published sources. The data were analysed (major axis regression, randomization tests) for taxonomic and body‐size‐dependent biases in colonization probabilities. A simple probabilistic model was used to examine the potential contribution of local (within‐region) extinctions to determining species composition in the Red Sea. Results Of the 462 reef‐associated species that inhabit the Arabian Sea, 69% have crossed successfully into the Red Sea; of these, 55% have crossed into the Gulf of Aqaba. A species’ probability of being found in either ‘target’ was independent of presumed innate differences, i.e. ecological correlates of taxonomic affiliation and body size. Similarly, local extinctions were found unlikely to have been of consequence over the past several thousand years. Main conclusions Present‐day differences in the species richness of reef‐associated fish species among the Arabian Sea, Red Sea and Gulf of Aqaba appear to be the product of external, non‐selective constraints on colonization. The random nature of the colonization process is suggestive of ecological redundancy among coral‐reef fish species. Importantly, the study places a time frame on the processes that determine spatial patterns of biodiversity in reef fish.     

Zoogeography of the coral reef fishes of the Socotra Archipelago

Fish communities and habitats were studied at the Socotra archipelago (Gulf of Aden, ≈12°N 54°E). Extensive and unexpected hermatypic coral communities were recorded, at the centre of a 2200 km gap in knowledge of species and habitat distributions which coincides with a change from a western Indian Ocean coral reef fauna to an Arabian one. The fish assemblage associated with the Socotra archipelago corals is predominantly south Arabian. An east African influence, minimal on the mainland coasts of Arabia, is more evident here, and results in previously unrecorded sympatry between Arabian endemic species and their Indian Ocean sister taxa. A study of distributions of Chaetodontidae (butterflyfishes) in the north-western Indian Ocean reveals a number of distinct patterns, with a trend for species replacement along a track from the northern Red Sea to the Indian Ocean. A major feature of the reef fish zoogeography of the region is found to be a distinct south Arabian area, characterized by a 'pseudo-high latitude effect' which results from seasonal cold water upwelling along the Arabian sea coasts of Yemen and Oman and the Indian Ocean coast of Somalia. This south Arabian feature is consistent across a wide range of fish families. It is most pronounced in Oman and Yemen, and although it is the dominant influence at Socotra it is slightly 'diluted' here by the east African influence. The south Arabian area wholly or partly accounts for most of the major marine zoogeographic features around Arabia, and is the principal feature fragmenting Arabian coastal fish assemblages, and separating them from those of the wider Indo-west Pacific.     

Fish assemblages on artificial and natural reefs in the Flower Garden Banks National Marine Sanctuary, USA

 Visual censusing was used to characterize fish assemblages on artificial and natural reefs located within the boundaries of the Flower Garden Banks National Marine Sanctuary (FGBNMS) in the northwestern Gulf of Mexico. Emphasis was placed on determining spatial and temporal patterns in habitat utilization by fishes on an offshore artificial reef (Mobil Platform HI-A389A). Overall, 43 species were observed during diurnal surveys in the upper 24 m of the artificial reef. Midwater pelagic fishes (i.e., carangids and scombrids) accounted for over 50% of all taxa enumerated on the artificial reef; however, these taxa were transient members of the assemblage and were observed infrequently. Labrids, pomacentrids, and serranids were the dominant reef-dependent taxa. Distinct trends in vertical, diel, and seasonal abundances were observed for juvenile and adult fishes. Of the three designated depth zones (upper 1.5–9.0, middle 9.0–16.5; lower 16.5–24.0 m), abundance and species diversity were lowest in the upper zone. Nocturnal counts were characterized by a marked reduction or complete absence of most species, due in part to twilight cover-seeking and movement activities. Seasonal variation in community composition and species abundance (May versus September) was primarily due to recruitment of juveniles (0-age fishes) to the artificial reef in late summer. Increases in total fish abundance (all taxa combined) coincided with both increasing habitat rugosity and degree of fouling. Species richness on natural coral reefs in the FGBNMS was higher than on the artificial reef. Unlike the artificial reef, fish assemblages on the natural reefs were dominated by a single family (Pomacentridae) which accounted for over 50% of all individuals observed. Accepted: 1 August 1996     

Characterization of the mesophotic reef fish community in south Florida,USA

The southeastern coast of Florida, USA supports a substantial recreational fishery, yet little is known of the coral reef ecosystem or fisheries resources past 50 m depth. Fish assemblages associated with low‐relief substrate and three vessel reefs between 50 and 120 m depth off southeast Florida were surveyed by remotely operated vehicles providing the first characterization of the mesophotic fish assemblages in the region. Two distinct assemblages were observed on low‐relief substrate and high‐relief vessel reefs. A total of 560 fishes of 42 species was recorded on 27 dives on low‐relief substrate, and 50 152 fishes of 65 species were recorded on 24 dives on three vessel reefs. Small planktivorous Anthiinae fishes and several economically valuable species were common on vessel reefs but rare on low‐relief substrate. Fish assemblages on vessel reefs more closely resembled those found at similar depths in high‐relief natural areas off east‐central Florida and the Gulf of Mexico than those associated with adjacent low‐relief habitat or nearby coral reef tracts. From a fisheries perspective, these results provide limited support to the hypothesis that in deep‐water regions with limited relief, vessel reefs may provide an opportunity to increase fish diversity and abundance by creating high‐relief habitat without compromising adjacent fish assemblages.     

Maintenance of fish diversity on disturbed coral reefs

Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.     

Vertical zonation of coral reef fishes in the Sudanese Red Sea

The vertical zonation of 38 species of fish inhabiting coral reefs in the Sudanese Red Sea is described. A marked degree of vertical zonation of the fishes is shown to exist. The usefulness of a depth oriented survey method for studying reef fish is discussed in relation to these results.General relationships between vertical zonation and species diversity are noted.     

Relationships between coral reef substrata and fish

 The objective of this work is to identify which substrata characteristics (such as coral morphology, coral diversity, coral species richness, percentage coverage by live coral or by algae) influence the structure and abundance of fish communities. The study was carried out at Reunion Island, Indian Ocean, where six sites were sampled in two zones (reef flat and outer reef slope). Quantitative data were collected by visual census techniques, along a linear transect of 50 m for the substratum, and a belt of 50×2 m for the fish communities. Correspondence analysis (CA) and an optimising cluster analysis, called dynamic clustering method (DCM) were used to describe and compare fish assemblages with the benthic composition. The relationships between benthic and fish communities were examined using the classes revealed by the partitioning of the substratum with DCM. This partitioning allowed us to derive four classes of substratum: the non-disturbed reef flat, the non-disturbed outer reef slope, the perturbed reef habitat and the reef pass. The analysis of the partitioning based on the coral variables suggests that there are significant relationships between benthic and fish assemblages. Accepted: 26 July 1996     

Modelling sponge species diversity using a morphological predictor: a tropical test of a temperate model

The contribution of sponges to marine surveys is often underestimated due to problems of identification, synonymous species and limited numbers of specialists in the field. Bell & Barnes (2001) illustrated how sponge morphological diversity (diversity of body forms) might be used as a predictor of sponge species diversity and richness. This study investigated these relationships at six tropical West Indian Ocean localities in a number of habitat types. These habitats included tropical coral reefs, soft substratum (seagrass, mangrove and sand), caves and boulders. Sampling was undertaken at three depth zones in coral reef habitats only (intertidal, 10–15 m and 20–25 m), with the other habitats sampled in less than 10m of water. Species diversity and richness were significantly correlated (P < 0.05) with morphological diversity at all localities and depths in coral reef and soft substratum habitats. However, no significant correlation was found between these variables in cave or boulder habitats. The slope of the linear regression found between morphological diversity and species diversity did not significantly differ between coral reef, soft substratum and temperate reef (data taken from Bell & Barnes 2001) habitats. Similarly coral reefs showed the same relationship between morphological diversity and species richness as temperate reefs, however the relationship between morphological diversity and species richness was significantly different at both habitats compared with soft substratum environments. Sponge morphological diversity therefore may be more useful as a predictor of sponge species diversity, rather than species richness, as the former relationship is common between more habitats than the latter.     

Different Surrounding Landscapes may Result in Different Fish Assemblages in East African Seagrass Beds

Few studies have considered how seagrass fish assemblages are influenced by surrounding habitats. This information is needed for a better understanding of the connectivity between tropical coastal ecosystems. To study the effects of surrounding habitats on the composition, diversity and densities of coral reef fish species on seagrass beds, underwater visual census surveys were carried out in two seagrass habitat types at various locations along the coast of Zanzibar (Tanzania) in the western Indian Ocean. Fish assemblages of seagrass beds in a marine embayment with large areas of mangroves (bay seagrasses) situated 9 km away from coral reefs were compared with those of seagrass beds situated on the continental shelf adjacent to coral reefs (reef seagrasses). No differences in total fish density, total species richness or total juvenile fish density and species richness were observed between the two seagrass habitat types. However, at species level, nine species showed significantly higher densities in bay seagrasses, while eight other species showed significantly higher densities in reef seagrasses. Another four species were exclusively observed in bay seagrasses. Since seagrass complexity could not be related to these differences, it is suggested that the arrangement of seagrass beds in the surrounding landscape (i.e. the arrangement on the continental shelf adjacent to the coral reef, or the arrangement in an embayment with mangroves situated away from reefs) has a possible effect on the occurrence of various reef-associated fish species on seagrass beds. Fish migration from or to the seagrass beds and recruitment and settlement patterns of larvae possibly explain these observations. Juvenile fish densities were similar in the two types of seagrass habitats indicating that seagrass beds adjacent to coral reefs also function as important juvenile habitats, even though they may be subject to higher levels of predation. On the contrary, the density and species richness of adult fish was significantly higher on reef seagrasses than on bay seagrasses, indicating that proximity to the coral reef increases density of adult fish on reef seagrasses, and/or that ontogenetic shifts to the reef may reduce adult density on bay seagrasses.     

Length–weight relationships of cryptic reef fishes from the southwestern Gulf of California, Mèxico

Length–weight relationships of 33 cryptic reef fishes from the southwestern Gulf of California were estimated. The most common families were Gobiidae (seven species), Labrisomidae (four), Pomacentridae (three), and Tripterygiidae (three). These are the first length–weight parameters reported for 32 of these species. Specimens were sampled during a comparative ecological study of community structure of fish assemblages associated with coral heads, rocky walls, and artificial reefs in Bahía de La Paz, Baja California Sur. The b values varied between 2.63 and 3.61. New maximum length records for five of the species were established in the study.     

Effect of Phase Shift from Corals to Zoantharia on Reef Fish Assemblages

Consequences of reef phase shifts on fish communities remain poorly understood. Studies on the causes, effects and consequences of phase shifts on reef fish communities have only been considered for coral-to-macroalgae shifts. Therefore, there is a large information gap regarding the consequences of novel phase shifts and how these kinds of phase shifts impact on fish assemblages. This study aimed to compare the fish assemblages on reefs under normal conditions (relatively high cover of corals) to those which have shifted to a dominance of the zoantharian Palythoa cf. variabilis on coral reefs in Todos os Santos Bay (TSB), Brazilian eastern coast. We examined eight reefs, where we estimated cover of corals and P. cf. variabilis and coral reef fish richness, abundance and body size. Fish richness differed significantly between normal reefs (48 species) and phase-shift reefs (38 species), a 20% reduction in species. However there was no difference in fish abundance between normal and phase shift reefs. One fish species, Chaetodon striatus, was significantly less abundant on normal reefs. The differences in fish assemblages between different reef phases was due to differences in trophic groups of fish; on normal reefs carnivorous fishes were more abundant, while on phase shift reefs mobile invertivores dominated.     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Recruitment of coral reef fishes along a cross-shelf gradient in the Red Sea peaks outside the hottest season

Knowledge on the early life history, ecology, and biology of marine species is crucial for future projections of the resilience of coral reef ecosystems and for adequate management strategies. A fundamental component of population dynamics is the recruitment of new individuals, and in some marine populations, this may be a limiting factor. Recruitment peaks of coral reef fishes commonly occur during the warmer months of the year in many subtropical and temperate locations worldwide. In the Red Sea, very little is known about the influence of temperature on reproductive patterns of coral reef fishes and studies on recruitment are missing. The Red Sea is one of the hottest and most isolated tropical seas in the world. We hypothesized that sea surface temperatures (SSTs) during the Red Sea’s hottest season may exceed the optimum for successful recruitment of some coral reef fishes, which therefore has to occur during other, cooler seasons, unlike recruitment among coral reef ecosystems around the world. We identified taxa among fish recruits by matching mitochondrial DNA sequences (using COI, commonly known as “barcoding”) and assessed potential biological and environmental drivers of recruitment. We studied three reefs located along a cross-shelf gradient for 12 consecutive months in the central Red Sea to capture seasonal changes in biotic and abiotic parameters along this gradient. Our results indicated that recruitment peaks did not occur during the hottest SSTs for most taxa, especially at the hottest inshore and mid-shelf reefs, and identified fish recruitment to be mainly and strongly correlated with the biomass of planktonic invertebrates. Moreover, temporal patterns of fish recruitment differed within and among taxonomic families among the reefs.

     

Consequences of marine barriers for genetic diversity of the coral‐specialist yellowbar angelfish from the Northwestern Indian Ocean

Ocean circulation, geological history, geographic distance, and seascape heterogeneity play an important role in phylogeography of coral‐dependent fishes. Here, we investigate potential genetic population structure within the yellowbar angelfish (Pomacanthus maculosus) across the Northwestern Indian Ocean (NIO). We then discuss our results with respect to the above abiotic features in order to understand the contemporary distribution of genetic diversity of the species. To do so, restriction site‐associated DNA sequencing (RAD‐seq) was utilized to carry out population genetic analyses on P. maculosus sampled throughout the species’ distributional range. First, genetic data were correlated to geographic and environmental distances, and tested for isolation‐by‐distance and isolation‐by‐environment, respectively, by applying the Mantel test. Secondly, we used distance‐based and model‐based methods for clustering genetic data. Our results suggest the presence of two putative barriers to dispersal; one off the southern coast of the Arabian Peninsula and the other off northern Somalia, which together create three genetic subdivisions of P. maculosus within the NIO. Around the Arabian Peninsula, one genetic cluster was associated with the Red Sea and the adjacent Gulf of Aden in the west, and another cluster was associated with the Arabian Gulf and the Sea of Oman in the east. Individuals sampled in Kenya represented a third genetic cluster. The geographic locations of genetic discontinuities observed between genetic subdivisions coincide with the presence of substantial upwelling systems, as well as habitat discontinuity. Our findings shed light on the origin and maintenance of genetic patterns in a common coral reef fish inhabiting the NIO, and reinforce the hypothesis that the evolution of marine fish species in this region has likely been shaped by multiple vicariance events.     

Do coral-reef fish faunas have a distinctive taxonomic structure?

Do the highly diverse fish faunas that associate with coral reefs have distinguishing taxonomic and ecological characteristics, as proposed by Choat and Bellwood (1991) and Bellwood (1996)? Does a 50?my old (Eocene) fossil fish fauna from Italy represent a coral-reef fish assemblage that provides unique information about the evolution of such assemblages, as claimed by Bellwood (1996)? I compared the structure of the reef fish faunas of adjacent tropical regions rich and poor in coral reefs, in both America and Polynesia, and found that they exhibit no substantive differences in relative species richness among families of typical “coral-reef” fishes. While coral-rich regions have larger reef fish faunas, a variety of factors probably contribute to such differences. Thus coral-reef fish faunas may lack a distinctive taxonomic structure. A similar comparative approach would be useful for assessing whether assemblages of fishes on coral reefs have distinctive ecological characteristics. Based on patterns of habitat use by modern tropical shorefishes, the Italian Eocene fauna includes few definite reef fishes, and may well consist primarily of non-reef fishes preserved in a non-reef habitat. Until we know more about the environment in which those fossils were preserved, that fauna can contribute little to understanding how coral reef fish assemblages have evolved.     

An Analysis of Artificial Reef Fish Community Structure along the Northwestern Gulf of Mexico Shelf: Potential Impacts of “Rigs-to-Reefs” Programs

Artificial structures are the dominant complex marine habitat type along the northwestern Gulf of Mexico (GOM) shelf. These habitats can consist of a variety of materials, but in this region are primarily comprised of active and reefed oil and gas platforms. Despite being established for several decades, the fish communities inhabiting these structures remain poorly investigated. Between 2012 and 2013 we assessed fish communities at 15 sites using remotely operated vehicles (ROVs). Fish assemblages were quantified from standing platforms and an array of artificial reef types (Liberty Ships and partially removed or toppled platforms) distributed over the Texas continental shelf. The depth gradient covered by the surveys (30–84 m) and variability in structure density and relief also permitted analyses of the effects of these characteristics on fish richness, diversity, and assemblage composition. ROVs captured a variety of species inhabiting these reefs from large transient piscivores to small herbivorous reef fishes. While structure type and relief were shown to influence species richness and community structure, major trends in species composition were largely explained by the bottom depth where these structures occurred. We observed a shift in fish communities and relatively high diversity at approximately 60 m bottom depth, confirming trends observed in previous studies of standing platforms. This depth was also correlated with some of the largest Red Snapper captured on supplementary vertical longline surveys. Our work indicates that managers of artificial reefing programs (e.g., Rigs-to-Reefs) in the GOM should carefully consider the ambient environmental conditions when designing reef sites. For the Texas continental shelf, reefing materials at a 50–60 m bottom depth can serve a dual purpose of enhancing diving experiences and providing the best potential habitat for relatively large Red Snapper.     

Island biogeography of Caribbean coral reef fish

Aim The goal of our study was to test fundamental predictions of biogeographical theories in tropical reef fish assemblages, in particular relationships between fish species richness and island area, isolation and oceanographic variables (temperature and productivity) in the insular Caribbean. These analyses complement an analogous and more voluminous body of work from the tropical Indo‐Pacific. The Caribbean is more limited in area with smaller inter‐island distances than the Indo‐Pacific, providing a unique context to consider fundamental processes likely to affect richness patterns of reef fish. Location Caribbean Sea. Methods We compiled a set of data describing reef‐associated fish assemblages from 24 island nations across the Caribbean Sea, representing a wide range of isolation and varying in land area from 53 to 110,860 km². Regression‐based analyses compared the univariate and combined effects of island‐specific physical predictors on fish species richness. Results We found that diversity of reef‐associated fishes increases strongly with increasing island area and with decreasing isolation. Richness also increases with increasing nearshore productivity. Analyses of various subsets of the entire data set reveal the robustness of the richness data and biogeographical patterns. Main conclusions Within the relatively small and densely packed Caribbean basin, fish species richness fits the classical species–area relationship. Richness also was related negatively to isolation, suggesting direct effects of dispersal limitation in community assembly. Because oceanic productivity was correlated with isolation, however, the related effects of system‐wide productivity on richness cannot be disentangled. These results highlight fundamental mechanisms that underlie spatial patterns of biodiversity among Caribbean coral reefs, and which are probably also are functioning in the more widespread and heterogeneous reefs of the Indo‐Pacific.     

The abundance of herbivorous fish on an inshore Red Sea reef following a mass coral bleaching event

A healthy herbivore community is critical for the ability of a reef to resist and recover from severe disturbances and to regain lost coral cover (i.e., resilience). The densities of the two major herbivorous fish groups (the family Acanthuridae and scarine labrids) were comparatively studied for an inshore reef that was severely impacted by a mass coral bleaching event in 2010 and an unaffected reef within the same region. Densities were found to be significantly higher on the affected reef, most likely due to the high algal densities on that reef. However, densities of herbivores on both reefs were found to be on average about 1–2 orders of magnitude lower than previously published reports from some Pacific reefs and from Red Sea reefs in the Gulf of Aqaba and only slightly higher than Caribbean reefs. Thus, it is predicted that recovery for this reef and similarly affected reefs may be very slow. The protection of herbivores from overfishing and the introduction of other management strategies that maximize reef resilience in Saudi Arabian waters are highly recommended.     

Biogeography and the structure of coral reef fish communities on isolated islands

Aim To determine the applicability of biogeographical and ecological theory to marine species at two remote island locations. This study examines how biogeography, isolation and species geographic range size influence patterns of species richness, endemism, species composition and the abundance of coral reef fishes. Location Christmas Island and the Cocos (Keeling) Islands in the tropical eastern Indian Ocean. Methods Published species lists and underwater visual surveys were used to determine species richness, endemism, species composition and abundance of reef fishes at the islands. These data were statistically compared with patterns of species composition and abundance from the neighbouring ‘mainland’ Indonesian region. Results The two isolated reef fish communities were species‐poor and contained a distinct taxonomic composition with an overrepresentation of species with high dispersal potential. Despite low species richness, we found no evidence of density compensation, with population densities on the islands similar to those of species‐rich mainland assemblages. The mix of Indian and Pacific Ocean species and the proportional representations of the various regional faunas in the assemblages were not influenced by the relative proximity of the islands to different biogeographical provinces. Moreover, species at the edge of their range did not have a lower abundance than species at the centre of their range, and endemic species had substantially higher abundances than widespread species. At both locations, endemism was low (less than 1.2% of the community); this may be because the locations are not sufficiently isolated or old enough to promote the evolution of endemic species. Main conclusions The patterns observed generally conform to terrestrial biogeographical theory, suggesting that similar processes may be influencing species richness and community composition in reef fish communities at these remote islands. However, species abundances differed from typical terrestrial patterns, and this may be because of the life history of reef fishes and the processes maintaining isolated populations.     

Species-specific habitat distribution of coral reef fish assemblages in relation to habitat characteristics in an Okinawan coral reef

We determined the species-specific habitat associations of coral reef fishes and environmental characteristics in an Okinawan coral reef in Japan. We focused on three families (Pomacentridae, Gobiidae and Labridae) and attempted to determine differences in habitat utilization. We selected six sites along the coast of Amitori Bay, from the entrance to the innermost part, in order to cover a wide range of habitat characteristics (exposed habitat, semi-exposed habitat and sheltered habitat). The species diversity of coral assemblages was greater at the exposed and semi-exposed habitats, whereas branching coral mostly covered the sheltered habitat. The environmental factors that determine the species-specific spatial association in fishes differed among families. Both biological characteristics (coral morphology and coral species diversity) and physical characteristics (water depth and wave exposure) affected the spatial association of pomacentrids and gobiids. In contrast, physical characteristics such as substrate complexity and water depth affected the species-specific spatial association of labrid species. Further study is needed to determine the ecological factors that regulate the species-specific habitat preference in Okinawan coral reefs.