The Eocene fishes of Monte Bolca: the earliest coral reef fish assemblage

The fish assemblage from the Eocene deposits of Monte Bolca, Northern Italy, are compared with those of Recent coral reefs. A family-level taxonomic definition of a Recent coral reef fish assemblage is formulated to permit direct comparisons. On this basis, the Monte Bolca fishes represent the earliest clearly defined coral reef fish assemblage. Quantitative analyses of the relative abundance of fish families revealed significant differences between the two assemblages. The Bolca assemblage has Mesozoic links (Pycnodontiformes) and non-perciform taxa are relatively abundant, particularly the Beryciformes (Holocentridae). However, Bolca represents the earliest record of a perciform-dominated benthic fish assemblage (68.4% of all non-clupeid taxa). Within the Perciformes, the abundance of the major reef fish lineages (higher squamipinnes and Labroidei) differs markedly between the two assemblages. The numerical dominance of labroid fishes on coral reefs appears to have been a relatively recent occurrence.     

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     

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.     

Herbivorous fishes,ecosystem function and mobile links on coral reefs

Understanding large-scale movement of ecologically important taxa is key to both species and ecosystem management. Those species responsible for maintaining functional connectivity between habitats are often called mobile links and are regarded as essential elements of resilience. By providing connectivity, they support resilience across spatial scales. Most marine organisms, including fishes, have long-term, biogeographic-scale connectivity through larval movement. Although most reef species are highly site attached after larval settlement, some taxa may also be able to provide rapid, reef-scale connectivity as adults. On coral reefs, the identity of such taxa and the extent of their mobility are not yet known. We use acoustic telemetry to monitor the movements of Kyphosus vaigiensis, one of the few reef fishes that feeds on adult brown macroalgae. Unlike other benthic herbivorous fish species, it also exhibits large-scale (>2 km) movements. Individual K. vaigiensis cover, on average, a 2.5 km length of reef (11 km maximum) each day. These large-scale movements suggest that this species may act as a mobile link, providing functional connectivity, should the need arise, and helping to support functional processes across habitats and spatial scales. An analysis of published studies of home ranges in reef fishes found a consistent relationship between home range size and body length. K. vaigiensis is the sole herbivore to depart significantly from the expected home range–body size relationship, with home range sizes more comparable to exceptionally mobile large pelagic predators rather than other reef herbivores. While the large-scale movements of K. vaigiensis reveal its potential capacity to enhance resilience over large areas, it also emphasizes the potential limitations of small marine reserves to protect some herbivore populations.     

The influence of coral reef benthic condition on associated fish assemblages

Accumulative disturbances can erode a coral reef's resilience, often leading to replacement of scleractinian corals by macroalgae or other non-coral organisms. These degraded reef systems have been mostly described based on changes in the composition of the reef benthos, and there is little understanding of how such changes are influenced by, and in turn influence, other components of the reef ecosystem. This study investigated the spatial variation in benthic communities on fringing reefs around the inner Seychelles islands. Specifically, relationships between benthic composition and the underlying substrata, as well as the associated fish assemblages were assessed. High variability in benthic composition was found among reefs, with a gradient from high coral cover (up to 58%) and high structural complexity to high macroalgae cover (up to 95%) and low structural complexity at the extremes. This gradient was associated with declining species richness of fishes, reduced diversity of fish functional groups, and lower abundance of corallivorous fishes. There were no reciprocal increases in herbivorous fish abundances, and relationships with other fish functional groups and total fish abundance were weak. Reefs grouping at the extremes of complex coral habitats or low-complexity macroalgal habitats displayed markedly different fish communities, with only two species of benthic invertebrate feeding fishes in greater abundance in the macroalgal habitat. These results have negative implications for the continuation of many coral reef ecosystem processes and services if more reefs shift to extreme degraded conditions dominated by macroalgae.     

The relationship between habitat structure and fish faunas on New Zealand reefs

Reef fish abundances were sampled at 11 shallow reef localities extending over 1000 km of coastline in northern New Zealand. Sampling was restricted to the 4–10-m depth stratum and included six coastal and five island localities. These were either coralline reef flats dominated by echinoids, or algal reefs with high densities of laminarian and fucoid algae. Reefs dominated by macroscopic algae supported large numbers of small fishes, mainly labrids, and few large benthic-feeding fishes. Echinoid-dominated reefs supported a different fish fauna with more large benthic-feeding species. Additional sampling of echinoiddominated reefs and algal stands in deeper water provided confirmation of these findings. A second sampling programme was carried out at a series of eight sites within a single locality covering 5 km of coastline. These spanned a moderate exposure gradient and ranged from algal dominated reefs to typical coralline reef flats with high densities of grazing invertebrates. The relationship between habitat structure and reef fish species composition and size frequency was similar to that of the large-scale sampling programme. Thirdly, observations on reef fish foraging and feeding patterns within a single reef site suggested that larger benthic-feeding reef fishes were less likely to feed within macroscopic algal stands. Experimental reductions of grazing invertebrates designed to produce brown algal stands on echinoid-dominated reef flats supported these observations. Larger individuals capable of removing echinoids and grazing gastropods did not frequent or feed in laminarian and fucoid algal stands. This pattern is discernible at several spatial scales. Our conclusion is that the type of shallow reef habitat, echinoid- as opposed to algal-dominated, will have an important rôle in determining the associated reef fish fauna.     

A preliminary distributional study of fish larvae near a ribbon coral reef in the Great Barrier Reef

Fish larvae from horizontal plankton tows along a single transect near outer ribbon reefs of the Great Barrier Reef in spring 1979 and summer 1980 had persistent distributional patterns. Larvae were identified to family and divided into young (preflexion) and old (postflexion) larvae, thus giving 28 taxa abundant enough for analysis. Non-uniform larval distributions were found for 81% of the 16 reef fish taxa with non-pelagic eggs, but for only 17% of the six reef fish taxa with pelagic eggs. Most differences in larval concentration were between the lagoonal and seaward sides of the reef. Only tripterygiid larvae had highest concentration just seaward of the reef, while larvae of 12 reef and three oceanic fish taxa occurred in highest concentrations on the lagoonal side of the reef. In five taxa of reef fishes, higher larval concentrations were found in the lagoonal backreef compared with the mid-lagoon habitat; but the reverse was not found in any taxon. Eleven taxa had indeterminate distributions, (i.e. no difference in concentration between stations). Mechanisms responsible for the distribution remain unknown, but we suggest that the view which considers fish larvae to be passively-drifting particles is unjustified without more information on larval behaviour.     

The relative influence of local to regional drivers of variation in reef fishes

In this study, fishes and habitat attributes were quantified, four times over 1 year, on three reefs within four regions encompassing a c. 6° latitudinal gradient across south-western Australia. The variability observed was partitioned at these spatio-temporal scales in relation to reef fish variables and the influence of environmental drivers quantified at local scales, i.e. at the scale of reefs (the number of small and large topographic elements, the cover of kelp, fucalean and red algae, depth and wave exposure) and at the scale of regions (mean and maximum nutrient concentrations and mean seawater temperature) with regard to the total abundance, species density, species diversity and the multivariate structure of reef fishes. Variation in reef fish species density and diversity was significant at the regional scale, whereas variation in the total abundance and assemblage structure of fishes was also significant at local scales. Spatial variation was greater than temporal variation in all cases. A systematic and gradual species turnover in assemblage structure was observed between adjacent regions across the latitudinal gradient. The cover of red algae within larger patches of brown macroalgae (a biological attribute of the reef) and the number of large topographic elements (a structural attribute of the reef) were correlated with variation observed at local scales, while seawater temperature correlated with variation at the scale of regions. In conclusion, conservation efforts on reef fishes need to incorporate processes operating at regional scales with processes that shape local reef fish communities at local scales.     

Non-Random Variability in Functional Composition of Coral Reef Fish Communities along an Environmental Gradient

Changes in the coral reef complex can affect predator-prey relationships, resource availability and niche utilisation in the associated fish community, which may be reflected in decreased stability of the functional traits present in a community. This is because particular traits may be favoured by a changing environment, or by habitat degradation. Furthermore, other traits can be selected against because degradation can relax the association between fishes and benthic habitat. We characterised six important ecological traits for fish species occurring at seven sites across a disturbed coral reef archipelago in Indonesia, where reefs have been exposed to eutrophication and destructive fishing practices for decades. Functional diversity was assessed using two complementary indices (FRic and RaoQ) and correlated to important environmental factors (live coral cover and rugosity, representing local reef health, and distance from shore, representing a cross-shelf environmental gradient). Indices were examined for both a change in their mean, as well as temporal (short-term; hours) and spatial (cross-shelf) variability, to assess whether fish-habitat association became relaxed along with habitat degradation. Furthermore, variability in individual traits was examined to identify the traits that are most affected by habitat change. Increases in the general reef health indicators, live coral cover and rugosity (correlated with distance from the mainland), were associated with decreases in the variability of functional diversity and with community-level changes in the abundance of several traits (notably home range size, maximum length, microalgae, detritus and small invertebrate feeding and reproductive turnover). A decrease in coral cover increased variability of RaoQ while rugosity and distance both inversely affected variability of FRic; however, averages for these indices did not reveal patterns associated with the environment. These results suggest that increased degradation of coral reefs is associated with increased variability in fish community functional composition resulting from selective impacts on specific traits, thereby affecting the functional response of these communities to increasing perturbations.     

Influence de la distance à l'océan sur les peuplements ichtyologiques des récifs frangeants de Nouvelle-Calédonie

Coral reef fish are bound to the pelagic environment throughout their larval stage, although they spend most of their adult life on reefs. Thus, the morphology of a lagoon and the location of a reef within the lagoon may influence the nature of its fish assemblage. Because of its asymetrical shape, the SW lagoon of New Caledonia allows us to test such a hypothesis. Several fringing reefs located 4–34 km from the open ocean were compared with one another. No influence of the reef-ocean distance could be found on the global values of diversity, density or biomass. However, presence-absence data of the 343 taxa encountered are related to this distance. Similarly, ecological structures based on diet, size and mobility of fish present strong correlations with reef—ocean distance. Thus, the major effects of an increase in reef—ocean distance concern both taxinomical and ecological structures. Suggested explicative hypotheses are linked to the fish feeding habits closely related to the environment but also to their utilisation of space. Territorial behaviour seems indeed to be important in structuring reef fish assemblage.     

Marine reserves indirectly affect fine‐scale habitat associations,but not overall densities,of small benthic fishes

Many large, fishery‐targeted predatory species have attained very high relative densities as a direct result of protection by no‐take marine reserves. Indirect effects, via interactions with targeted species, may also occur for species that are not themselves targeted by fishing. In some temperate rocky reef ecosystems, indirect effects have caused profound changes in community structure, notably the restoration of predator–urchin–macroalgae trophic cascades. Yet, indirect effects on small benthic reef fishes remain poorly understood, perhaps because of behavioral associations with complex, refuge‐providing habitats. Few, if any, studies have evaluated any potential effects of marine reserves on habitat associations in small benthic fishes. We surveyed densities of small benthic fishes, including some endemic species of triplefin (Tripterygiidae), along with fine‐scale habitat features in kelp forests on rocky reefs in and around multiple marine reserves in northern New Zealand over 3 years. Bayesian generalized linear mixed models were used to evaluate evidence for (1) main effects of marine reserve protection, (2) associations with habitat gradients, including complexity, and (3) differences in habitat associations inside versus outside reserves. No evidence of overall main effects of marine reserves on species richness or densities of fishes was found. Both richness and densities showed strong associations with gradients in habitat features, particularly habitat complexity. In addition, some species exhibited reserve‐by‐habitat interactions, having different associations with habitat gradients inside versus outside marine reserves. Two species (Ruanoho whero and Forsterygion flavonigrum) showed stronger positive associations with habitat complexity inside reserves. These results are consistent with the presence of a behavioral risk effect, whereby prey fishes are more strongly attracted to habitats that provide refuge from predation in areas where predators are more abundant. This work highlights the importance of habitat structure and the potential for fishing to affect behavioral interactions and the interspecific dynamic attributes of community structure beyond simple predator–prey consumption and archetypal trophic cascades.     

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.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Fish assemblages on sunken vessels and natural reefs in southeast Florida,USA

Derelict ships are commonly deployed as artificial reefs in the United States, mainly for recreational fishers and divers. Despite their popularity, few studies have rigorously examined fish assemblages on these structures and compared them to natural reefs. Six vessel-reefs off the coast of southeast Florida were censused quarterly (two ships per month) to characterize their associated fish assemblages. SCUBA divers used a non-destructive point-count method to visually assess the fish assemblages over 13- and 12-month intervals (March 2000 to March 2001 and March 2002 to February 2003). During the same intervals, fish assemblages at neighboring natural reefs were also censused. A total of 114,252 fishes of 177 species was counted on natural and vessel-reefs combined. Mean fish abundance and biomass were significantly greater on vessel-reefs in comparison to surrounding natural reef areas. Haemulidae was the most abundant family on vessel-reefs, where it represented 46% of total fish abundance. The most abundant family on natural reefs was Labridae, where it accounted for 24% of total fish abundance. Mean species richness was significantly greater on vessel-reefs than neighboring natural reefs and also differed among vessel-reefs. Both mean fish abundance and mean species richness were not significantly different between natural reefs neighboring vessel-reefs and natural reefs with no artificial structures nearby. This suggests the vessel-reefs are not, in general, attracting fish away from neighboring natural reefs in our area. Additionally, economically important fish species seem to prefer vessel-reefs, as there was a greater abundance of these species on vessel-reefs than surrounding natural reef areas. Fish assemblage structure on natural versus artificial reefs exhibited a low similarity (25.8%). Although no one species was responsible for more than 6% of the total dissimilarity, fish assemblage trophic structure differed strikingly between the two reef types. Planktivores dominated on vessel-reefs, accounting for 54% of the total abundance. Conversely, planktivores only made up 27% of total abundance on natural reefs. The results of this study indicate vessel-reef fish assemblages are unique and that these fishes may be utilizing food resources and habitat characteristics not accessible from or found at natural reefs in our area. Production may also be occurring at vessel-reefs as the attraction of fish species from nearby natural reefs seems to be minimal. Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users     

Herbivorous fish assemblages and herbivory pressure on Laamu Atoll, Republic of Maldives

While herbivory is recognized as a fundamental process structuring coral reef communities, herbivore assemblages and processes are poorly described for reefs in the Indian Ocean region. We quantified herbivorous fish assemblage structure (abundance and diversity) in Laamu Atoll, Republic of Maldives, in four reef habitat types: faro reef flats, faro reef slopes, inner and outer atoll reef slopes (20 sites in total). Herbivorous fish assemblages, representing a total of 30 species, grouped strongly by habitat type, with the highest absolute abundance observed on faro reef flats and lowest abundance on inside atoll rim reef slopes. Removal of Thalassia seagrass blades by ambient herbivore assemblages was used in a bioassay to assess relative herbivory pressure among four habitat types (eight sites). Also, at one site a choice herbivory assay was performed to assess herbivore preference among four benthic plants across three depth zones. Relative herbivory, as indicated by Thalassia assays, was highest on inside atoll rim reef slopes and lowest on outside atoll rim reef slopes. Thalassia consumption did not correspond to overall herbivorous fish abundance, but corresponded more closely with parrotfish abundance. In the choice assays, herbivores showed strong preferences among plant types and consumption of most plant types was higher at mid-depth than in the shallow reef flat or deep reef knoll zones.     

Using GIS mapping of the extent of nearshore rocky reefs to estimate the abundance and reproductive output of important fishery species

Kelp Bass (Paralabrax clathratus) and California Sheephead (Semicossyphus pulcher) are economically and ecologically valuable rocky reef fishes in southern California, making them likely indicator species for evaluating resource management actions. Multiple spatial datasets, aerial and satellite photography, underwater observations and expert judgment were used to produce a comprehensive map of nearshore natural rocky reef habitat for the Santa Monica Bay region (California, USA). It was then used to examine the relative contribution of individual reefs to a regional estimate of abundance and reproductive potential of the focal species. For the reefs surveyed for fishes (i.e. 18 out of the 22 in the region, comprising 82% the natural rocky reef habitat <30 m depth, with a total area of 1850 ha), total abundance and annual egg production of California Sheephead were 451 thousand fish (95% CI: 369 to 533 thousand) and 203 billion eggs (95% CI: 135 to 272 billion). For Kelp Bass, estimates were 805 thousand fish (95% CI: 669 to 941 thousand) and 512 billion eggs (95% CI: 414 to 610 billion). Size structure and reef area were key factors in reef-specific contributions to the regional egg production. The size structures of both species illustrated impacts from fishing, and results demonstrate the potential that relatively small increases in the proportion of large females on larger reefs could have on regional egg production. For California Sheephead, a substantial proportion of the regional egg production estimate (>30%) was produced from a relatively small proportion of the regional reef area (c. 10%). Natural nearshore rocky reefs make up only 11% of the area in the newly designated MPAs in this region, but results provide some optimism that regional fisheries could benefit through an increase in overall reproductive output, if adequate increases in size structure of targeted species are realized.     

Coexistence of Low Coral Cover and High Fish Biomass at Farquhar Atoll,Seychelles

We report a reef ecosystem where corals may have lost their role as major reef engineering species but fish biomass and assemblage structure is comparable to unfished reefs elsewhere around the world. This scenario is based on an extensive assessment of the coral reefs of Farquhar Atoll, the most southern of the Seychelles Islands. Coral cover and overall benthic community condition at Farquhar was poor, likely due to a combination of limited habitat, localized upwelling, past coral bleaching, and cyclones. Farquhar Atoll harbors a relatively intact reef fish assemblage with very large biomass (3.2 t ha⁻¹) reflecting natural ecological processes that are not influenced by fishing or other local anthropogenic factors. The most striking feature of the reef fish assemblage is the dominance by large groupers, snappers, and jacks with large (>1 m) potato cod (Epinephelus tukula) and marbled grouper (E. polyphekadion), commonly observed at many locations. Napoleon wrasse (Cheilinus undulatus) and bumphead parrotfish (Bolbometopon muricatum) are listed as endangered and vulnerable, respectively, but were frequently encountered at Farquhar. The high abundance and large sizes of parrotfishes at Farquhar also appears to regulate macroalgal abundance and enhance the dominance of crustose corallines, which are a necessary condition for maintenance of healthy reef communities. Overall fish biomass and biomass of large predators at Farquhar are substantially higher than other areas within the Seychelles, and are some of the highest recorded in the Indian Ocean. Remote islands like Farquhar Atoll with low human populations and limited fishing pressure offer ideal opportunities for understanding whether reefs can be resilient from global threats if local threats are minimized.     

Coral and fish distribution patterns in Mafia Island Marine Park, Tanzania: fish–habitat interactions

We examined coral reef communities at 11 sites within Mafia Island Marine Park using a point count method for substrate and visually censused belt transects for fish populations. Multivariate ordinations showed that the benthic habitat differed among reefs. The patterns were mainly attributed to variations in depth, hydrodynamics and benthic composition. In total, the substratum was dominated by dead coral (49%) and algae (25%), with a live coral cover of only 14%. Three hundred and ninety-four fish species belonging to 56 families were recorded. According to MDS-ordinations and RELATE procedures, fish assemblage composition varied among sites in concordance with the habitats provided. Sites with highest proportion of dead coral exhibited highest degree of dispersion in the multivariate ordinations of fish assemblages. Stepwise multiple regression was used to determine the proportion of variance among sites which could be explained by depth, exposure, rugosity, substrate diversity, branching substrate, live coral cover, dead coral cover and different types of algae. The results showed that habitat variables explained up to 92% of the variation in species numbers and in total, and taxon-specific, abundance. Live coral cover was the foremost predictor of both numerical and species abundance, as well as of corallivores, invertivores, planktivores and of the families Pomacentridae, Chaetodontidae and Pomacanthidae. Our results suggest that habitat characteristics play a dominant role in determining fish assemblage composition on coral reefs.     

Multiscale habitat associations of detrivorous blennies (Blenniidae: Salariini)

The distribution and habitat associations of detrivorous blennies on a tropical coral reef were investigated at several spatial scales and compared with other fish that feed on the epilithic algal matrix to assess density and biomass contributions of small detrivorous fishes to these assemblages. At broad spatial scales total blenny abundance and biomass were highest on the tops of reefs exposed to prevailing winds. On the finer scale of microhabitat use, all species showed a preference for non-living corals, although the type of coral utilised differed between species. The high abundance of blennies on reef tops and non-living corals may be partially related to the quality and availability of detritus in these habitats. Comparisons of total blenny abundance and biomass with other territorial detrivores found that blennies accounted for approximately 60% of this functional group's density and 21% of their biomass on exposed reef tops. Overall, territorial detrivores were found to constitute approximately 37% of the density and 26% of the biomass of the detrivorous/herbivorous fish assemblage on exposed reef tops. Small detrivorous fish therefore represent a substantial proportion of fish assemblages that feed on epilithic algae and associated detritus on coral reefs.