Habitat associations of juvenile fish at Ningaloo Reef, Western Australia: the importance of coral and algae

Habitat specificity plays a pivotal role in forming community patterns in coral reef fishes, yet considerable uncertainty remains as to the extent of this selectivity, particularly among newly settled recruits. Here we quantified habitat specificity of juvenile coral reef fish at three ecological levels; algal meadows vs. coral reefs, live vs. dead coral and among different coral morphologies. In total, 6979 individuals from 11 families and 56 species were censused along Ningaloo Reef, Western Australia. Juvenile fishes exhibited divergence in habitat use and specialization among species and at all study scales. Despite the close proximity of coral reef and algal meadows (10's of metres) 25 species were unique to coral reef habitats, and seven to algal meadows. Of the seven unique to algal meadows, several species are known to occupy coral reef habitat as adults, suggesting possible ontogenetic shifts in habitat use. Selectivity between live and dead coral was found to be species-specific. In particular, juvenile scarids were found predominantly on the skeletons of dead coral whereas many damsel and butterfly fishes were closely associated with live coral habitat. Among the coral dependent species, coral morphology played a key role in juvenile distribution. Corymbose corals supported a disproportionate number of coral species and individuals relative to their availability, whereas less complex shapes (i.e. massive & encrusting) were rarely used by juvenile fish. Habitat specialisation by juvenile species of ecological and fisheries importance, for a variety of habitat types, argues strongly for the careful conservation and management of multiple habitat types within marine parks, and indicates that the current emphasis on planning conservation using representative habitat areas is warranted. Furthermore, the close association of many juvenile fish with corals susceptible to climate change related disturbances suggests that identifying and protecting reefs resilient to this should be a conservation priority.     

Interactions Between Habitat Use and Patterns of Abundance in Coral-dwelling Fishes of the Genus <Emphasis Type="Italic">Gobiodon</Emphasis>

Coral-dwelling fishes from the genus Gobiodon are some of the most habitat specialised fishes on coral reefs. Consequently, we might expect that their population dynamics will be closely associated with the abundance of host corals. I used a combination of log-linear modelling and resource selection ratios to examine patterns of habitat use among eight species of Gobiodon in Kimbe Bay, Papua New Guinea. I then used multiple regression analysis to investigate relationships between the abundance of each species of Gobiodon and the abundance of the corals they inhabited. Each species of Gobiodon used one or more species of coral more frequently than expected by chance. The pattern of habitat use exhibited by each species of Gobiodon did not vary among reef zones or among reefs with different exposures to prevailing winds, despite changes in the relative abundances of corals among reef zones. This consistency in habitat use might be expected if the coral species inhabited confer considerable fitness advantages and, therefore, are strongly preferred. For most species of Gobiodon, abundances among reef zones and exposure regimes were correlated with the abundance of the coral species usually inhabited. Therefore, it appears that habitat availability helps determine abundances of most species of Gobiodon in Kimbe Bay. In addition to correlations with habitat availability, the abundances of G. histrio, G. quinquestrigatus, G. rivulatus (dark form) and the group others were also associated with particular reef zones and exposure regimes. Therefore, in these species, reef type appears to influence patterns of abundance independently of coral availability. In contrast to other species of Gobiodon, the abundance of the most specialised species, Gobiodon sp.A, was not closely associated with the abundance of the only coral species it inhabited. This study demonstrates that even for habitat specialised species, the relationship between habitat availability and abundance varies widely and is multiscale.     

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.     

Fish-habitat associations across multiple spatial scales

Important to the study of reef fish ecology is understanding the degree to which fish community structure varies across space, what factors can account for such variation, and whether these factors are scale dependent. This study examined the structure of reef fish communities across four spatial scales (1, 10 100, and 200 m²) visually censused from seven sites within Tague Bay, St. Croix, U.S. Virgin Islands. Relative differences in the number of individuals and species among sites indicated a pattern that was consistent across spatial scales. Spearmans rank correlation revealed significant positive correlation in site rankings, in terms of species richness, between 1 and 10 m², and 100 and 200 m²; and for the abundance of individuals between 100 and 200 m². In order to understand the degree to which quantified habitat variables account for patterns in the abundance of individuals and species, and whether these fish-habitat relationships were consistent regardless of spatial scale, separate canonical correlation analyses were conducted at each scale. Independent of scale, the total number of individuals and species were correlated with specific habitat variables, either negatively (with areas of pavement, sand, no algae, and low structural complexity) or positively (with areas of Amphiroa rigida, Halimeda incrassata, high structural complexity, and diverse algae/seagrass communities). These habitat variables explained 31–81% (at scales of 1–200 m²) of the variation in the number of individuals and species. Similar analyses were also performed on the abundances of the nine most common species, and whether their specific habitat associations were independent of scale. Results indicated that habitat variables explained 19–73% (at scales of 1–200 m²) of the variation in abundances of each species. Unique fish-habitat relationships were observed for each species, and most such relationships were consistent across spatial scales. The structure of reef fish communities of Tague Bay was explained in large part by the composition of coral and algae communities present. Both the spatial variation in community structure and the fish-habitat relationships, at the community and population level, appeared to be largely independent of the spatial scale examined. This suggests that generalizations across Tague Bay are possible. Similar habitat associations reported in the literature are discussed with regard to the possibility for generalizations across regions.     

Spatial and temporal patterns of abundance of coral reef gobies (Teleostei: Gobiidae)

This study investigated the spatial and temporal patterns of abundance of four substratum-associated species of Gobiidae on a heterogeneous reef flat comprised of four distinct habitat zones, and examined microhabitat use within each zone. Asterropteryx semipunctatus had the widest distribution and was the most abundant species in each habitat zone, followed by Amblygobius bynoensis , Valenciennea muralis and Amblygobius phalaena . Significant temporal and spatial differences in mean density were evident. The highest density of A. semipunctatus (312 individuals 10 m^–2) was recorded in a habitat zone dominated by algal-covered rubble, whereas A. bynoensis and V. muralis were most abundant (mean summer density 5·5–5·8 individuals 10 m^–2) in habitats containing both sand and hard substrata. In contrast, A. phalaena was uncommon (mean density ≤ 0·4 individuals 10 m^–2) in all four habitat zones. Significant seasonal differences in abundance were due to the large influx of recruits in summer. Ontogenetic shifts in habitat use were not evident at either the macrohabitat ( i.e. among habitat zones) or microhabitat scale ( i.e. substratum use within zones). At the microhabitat scale, V. muralis consistently exhibited a strong positive association with sand and was rarely associated with hard substrata. In contrast, the two Amblygobius species were commonly associated with both sand and hard substrata, but patterns of microhabitat use differed among habitat zones. Substratum composition at the microhabitat scale may influence spatial patterns of abundance at larger spatial scales by providing essential resources and, or influencing carrying capacity and predation risk.     

Habitat selection and aggression as determinants of spatial segregation among damselfish on a coral reef

Adults of many closely related coral reef fish species are segregated along gradients of depth or habitat structure. Both habitat selection by new settlers and subsequent competitive interactions can potentially produce such patterns, but their relative importance is unclear. This study examines the potential roles of habitat selection and aggression in determining the spatial distribution of adults and juveniles of four highly aggressive damselfishes at Lizard Island, northern Great Barrier Reef. Dischistodus perspicillatus, D. prosopotaenia, D. melanotus, and D. pseudochrysopoecilus maintain almost non-overlapping distributions across reef zones, with adults of one species dominating each reef zone. Juveniles exhibit slightly broader distributional patterns suggesting that subsequent interactions reduce overlap among species. Although habitat choice experiments in aquaria suggest that associations between juveniles and substrata types in the field are partly due to habitat selection, large overlaps in the use of substrata by the different species were also found, suggesting that substratum selection alone is insufficient in explaining the discrete spatial distributions of adults. The strength of aggressive interactions among all four species was tested by a "bottle" experiment, in which an adult or juvenile of each species was placed in the territories of adult fish on the reef. The greatest levels of interspecific aggression were directed against adults and juveniles of neighbouring species. The highest levels of aggression were associated with species exhibiting the greatest levels of overlap in resource use. Evidently both habitat selection and interspecific aggression combine to determine the adult distributions of these species.     

Long-lived groupers require structurally stable reefs in the face of repeated climate change disturbances

Benthic recovery from climate-related disturbances does not always warrant a commensurate functional recovery for reef-associated fish communities. Here, we examine the distribution of benthic groupers (family Serranidae) in coral reef communities from the Lakshadweep archipelago (Arabian Sea) in response to structural complexity and long-term habitat stability. These coral reefs that have been subject to two major El Niño Southern Oscillation-related coral bleaching events in the last decades (1998 and 2010). First, we employ a long-term (12-yr) benthic-monitoring dataset to track habitat structural stability at twelve reef sites in the archipelago. Structural stability of reefs was strongly driven by exposure to monsoon storms and depth, which made deeper and more sheltered reefs on the eastern aspect more stable than the more exposed (western) and shallower reefs. We surveyed groupers (species richness, abundance, biomass) in 60 sites across the entire archipelago, representing both exposures and depths. Sites were selected along a gradient of structural complexity from very low to high. Grouper biomass appeared to vary with habitat stability with significant differences between depth and exposure; sheltered deep reefs had a higher grouper biomass than either sheltered shallow or exposed (deep and shallow) reefs. Species richness and abundance showed similar (though not significant) trends. More interestingly, average grouper biomass increased exponentially with structural complexity, but only at the sheltered deep (high stability) sites, despite the availability of recovered structure at exposed deep and shallow sites (lower-stability sites). This trend was especially pronounced for long-lived groupers (life span >10 yrs). These results suggest that long-lived groupers may prefer temporally stable reefs, independent of the local availability of habitat structure. In reefs subject to repeated disturbances, the presence of structurally stable reefs may be critical as refuges for functionally important, long-lived species like groupers.     

Transformation of algal turf by echinoids and scarid fishes on French Polynesian coral reefs

The respective roles of regular echinoids and scarid fishes in the transformation of turf algae, the main food resource for reef herbivores, were investigated on French Polynesian coral reefs. The role of one species of parrotfish (Scarus sordidus) was compared with that of four species of echinoids. The degree and ways of degradation of the algal matter were determined by the organic matter percentage, the composition of the sugar fraction, and the concentration and composition of chlorophylltype pigments as assayed by HPLC analysis. Chemical analyses were performed on anterior and posterior intestines for scarids, intestinal contents and faeces for echinoids, and on fresh algal turf as a control of initial food quality. A decrease in mean percentage of organic matter in gut content was observed from intestine (9.7%) to faeces (7%) in sea urchins, but not in parrotfishes. The total sugar fraction decreased from fresh algal turf (32% of total organic matter) to echinoid (28%) to scarid (18%) gut contents. The ratio of insoluble to soluble sugars (I/S ratios) was higher in echinoids (2.6) than in scarid gut contents (1.0). A decrease in the total pigment concentration was measured from fresh algal turf to echinoid and it was found to be even lower in scarid gut contents. Chromatograms showed that the composition of chlorophyll-type pigments in scarid intestines was very similar to fresh algal turf, with a dominance of native forms, mainly chlorophyll a and b. On the contrary, degraded pigment forms dominated in echinoids. The main degraded products were pheophorbides in sea urchins, and chlorophyllides in parrotfishes. These results provided evidence for differentiation in digestive processes occurring in the two types of grazers. Echinoids released higher degraded algal material than did scarids. Thus, these two types of grazers play different roles in the recycling of organic matter on coral reefs.     

The Structures and Dynamics of Fish Communities in an Okinawan Coral Reef: Effects of Coral-based Habitat Structures at Sites with Rocky and Sandy Sea Bottoms

Although there have been many studies on ecological factors responsible for the organization of reef fish communities, most of the studies have focused on isolated habitats. However, findings from isolated habitats cannot necessarily be applied to fish communities in other habitats (e.g., a continuous habitat). In this study, therefore, we examined the structures of fish communities (abundance, species richness and species composition), and the dynamics of fish communities (seasonal changes in abundance, species richness and species composition) over a 2-year period in two different habitats (continuous habitat and isolated habitat) in an Okinawan coral reef. We established eight permanent quadrats (8m × 8m) on a rocky reef flat (continuous habitat) and rock reef patches surrounded by a sandy sea bottom (isolated habitat). The abundance and species richness of fishes such as pomacentrids, labrids, chaetodontids and acanthurids were greater in quadrats located in the continuous habitat, whereas those of blenniids, gobiids and mullids were greater in quadrats located in the isolated habitat. This caused marked differences between the fish community structure at the two sites. Seasonal and annual changes in fish community structure were relatively small at the continuous habitat site (>0.7 similarity based on C index) but were large at the isolated habitat site (C < 0.4), indicating that the fish community structure was relatively stable at the continuous habitat site but unstable at the isolated habitat site throughout the study period. Spatial differences between the fish community structures in the four quadrats at the continuous habitat site were small (C > 0.65 in most sites), but these differences were large at the isolated habitat site (C < 0.4). Our findings suggest that habitat structure (spatial arrangement of habitats) affects both spatial and seasonal differences in the reef fish community structure. The results also suggest that the main mechanisms underlying organization of reef fish communities in continuous and isolated habitats are different.     

Empirical relationships among resilience indicators on Micronesian reefs

A process-orientated understanding of ecosystems usually starts with an exploratory analysis of empirical relationships among potential drivers and state variables. While relationships among herbivory, algal cover, and coral recruitment, have been explored in the Caribbean, the nature of such relationships in the Pacific appears to be variable or unclear. Here, we examine potential drivers structuring the benthos and herbivorous fish assemblages of outer-shelf reefs in Micronesia (Palau, Guam and Pohnpei). Surveys were stratified by wave exposure and protection from fishing. High biomass of most herbivores was favoured by high wave exposure. High abundance of large-bodied scarids was associated with low turf abundance, high coral cover, and marine reserves. The remaining herbivores were more abundant in reefs with low coral cover, possibly because space and hence food limitation occur in high-coral-cover reefs. Rugosity had no detectable effect on herbivorous fish abundance once differences in exposure and coral cover were accounted for. At identical depths, high wave exposure was associated with greater volumes (cover × canopy height) of macroalgae and algal turfs, which most likely resulted from high primary productivity driven by flow. In exposed areas, macroalgal cover declined as the acanthurid biomass increased. The volume of algal turfs was negatively associated with coral cover and herbivore biomass. In turn, high coral cover and herbivore biomass are likely to intensify grazing. The density of juvenile corals was variable where macroalgal cover was low but was confined to lower densities where macroalgal cover was high. High coral cover and density of juvenile corals were favoured in sheltered habitats. While a weak positive relationship was found between scarid biomass and juvenile coral density, we hypothesise that high scarid densities may hinder juvenile density through increased corallivory. New hypotheses emerged that will help clarify the role of acanthurids, wave exposure, and corallivory in driving the recovery of Pacific coral communities.     

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.     

Distribution and habitat associations of the California moray (<Emphasis Type="Italic">Gymnothorax mordax</Emphasis>) within Two Harbors,Santa Catalina Island,California

While kelp forests are some of the best-surveyed ecosystems in California, information on cryptic inhabitants and their role within the community are lacking. Kelp itself provides overall structure to the habitat; however the rocky reef to which the kelp attaches is known to provide additional structure for cryptic species. Gymnothorax mordax, the California moray, is an elusive predatory species that is considered abundant in the waters around Catalina Island. However, no life history data exists for this species. We examined habitat composition, relative abundance, size pattern distributions, and biomass of G. mordax within Two Harbors, Catalina Island. Habitats were sampled using a combination of baited trap collection and transect surveys using SCUBA. A total of 462 G. mordax were captured, primarily in shallow (< 10 m) waters. Individuals of G. mordax were associated with mostly boulder and cobble substrates. Measurements of relative abundance and density indicate that G. mordax is more prevalent than reported in previous studies. We also discovered that the 6 trapping sites from which all morays were collected, differed in size structuring and density while the relatively high biomass did not change across sites. In general, southern facing sites exhibited higher densities of morays, while northern facing sites showed more size structuring. We show how the structural complexity of the rocky reef habitat in an already diverse kelp forest ecosystem, can support a high biomass of a cryptic elongate predatory fish.     

Herbivore abundance and grazing intensity on a Caribbean coral reef

Herbivory is a primary factor in determining the structure of coral reef communities. Spatial variation among reef habitats in the intensity of herbivory has been documented, but underlying variation in species composition and abundance within the herbivore guild has received little attention. The distribution and relative abundances of herbivorous fishes and sea urchins across several habitats were studied on the Belizean barrier reef off the Caribbean coast of Central America. Marked variation in total herbivore density as well as major changes in the composition of the herbivore guild were found across reef habitats. Acanthurids (surgeonfishes) predominated in shallow areas (< 5 m) while scarids (parrotfishes) were dominant in deeper habitats. Significant differences among habitats in an experimental assay of grazing intensity were strongly correlated with herbivore abundance. The spatial distribution of herbivorous fishes across reef habitats does not appear to be simply explained by differences in reef topography, but may depend on complex interactions among proximity to nearby shelter, predator abundance, density of territorial competitors, and local availability of food resources.     

Shelters and their use by fishes on fringing coral reefs

Coral reef fish density and species richness are often higher at sites with more structural complexity. This association may be due to greater availability of shelters, but surprisingly little is known about the size and density of shelters and their use by coral reef fishes. We quantified shelter availability and use by fishes for the first time on a Caribbean coral reef by counting all holes and overhangs with a minimum entrance diameter ≥3 cm in 30 quadrats (25 m(2)) on two fringing reefs in Barbados. Shelter size was highly variable, ranging from 42 cm(3) to over 4,000,000 cm(3), with many more small than large shelters. On average, there were 3.8 shelters m(-2), with a median volume of 1,200 cm(3) and a total volume of 52,000 cm(3) m(-2). The number of fish per occupied shelter ranged from 1 to 35 individual fishes belonging to 66 species, with a median of 1. The proportion of shelters occupied and the number of occupants increased strongly with shelter size. Shelter density and total volume increased with substrate complexity, and this relationship varied among reef zones. The density of shelter-using fish was much more strongly predicted by shelter density and median size than by substrate complexity and increased linearly with shelter density, indicating that shelter availability is a limiting resource for some coral reef fishes. The results demonstrate the importance of large shelters for fish density and support the hypothesis that structural complexity is associated with fish abundance, at least in part, due to its association with shelter availability. This information can help identify critical habitat for coral reef fishes, predict the effects of reductions in structural complexity of natural reefs and improve the design of artificial reefs.     

Additive partitioning of reef fish diversity variation: a promising marine biodiversity management tool

Additive partitioning was applied to variation in reef fish spatial diversity at Isla Isabel National Park, Nayarit state, Mexico, and to identify the environmental and spatial variables that best explains it. Analyses included expected and observed species curves, rare species analysis, additive partitioning of alpha- and beta-diversity, and canonical redundancy analysis. A total of 10,517 individuals were recorded from 75 species and 33 reef fish families, representing 85% of expected richness. Species richness beta-diversity was dependent on the site scale, while the alpha-diversity of the Shannon diversity was most significant at the transect scale. Canonical partitioning showed species richness and Shannon diversity was explained by spatially-structured environmental components. Variation in species composition and abundance was explained by a purely environmental component. Therefore, elements of habitat structure (especially corals), topographic complexity, and refuge availability determine fish species diversity. Our results suggest that greater emphasis is required to conserve sites that promote β-diversity, increasing fish spatial diversity. In Isla Isabel, these sites would be mostly those located at eastern and southern of protected sides, where coral reef patches are well represented. The results of this multi-scale analysis are valuable and useful as an addition and complement to the holistic management strategies implemented at Isla Isabel.     

Community structure and habitat associations of parrotfishes on Oahu,Hawaii

Despite their ecological importance as bioeroders and their economic importance in commercial, artisanal, and recreational fisheries, there have been relatively few studies on parrotfish (Scaridae) ecology in Hawaii. Belt transects were conducted around the island of Oahu to survey current parrotfish distributions, size structure, species composition and associated habitats. Scarid communities in this heavily fished region are dominated by smaller species and smaller individuals within all species. Specific habitat characteristics such as rugosity, substrate diversity, and percent live coral cover were positively correlated with scarid numerical abundance. Scarids, however, were patchily distributed and were often absent from preferable habitats, suggesting that intense fishing pressure may be an important factor preventing these fish from fully exploiting available habitats. This study is the first thorough, broad-scale investigation of scarid community structure in Hawaii, and provides important information that has management and conservation implications for parrotfish in Hawaii and throughout tropical coral reef ecosystems.     

Spatial patterns of epilithic algal and detrital resources on a windward coral reef

Algae and detritus co-occur on reef substrata, but their relative importance to consumers, in terms of biomass and nutrient content, is poorly understood. Epilithic algae, sediment and detritus were sampled among four windward zones at Lizard Island, Australia, using a vacuum sampler. Total N, P and biomass (total organic C) were quantified for algae and detritus. Algal and detrital biomass varied among zones and both were positively correlated with sediment load. Algal biomass was approximately six times higher than detritus on the reef crest. In other zones, the two components were more similar in abundance. Particulates on the reef crest had the highest organic:inorganic ratios, a feature likely to be favoured by herbivorous and detritivorous fishes. C:N ratios of algae and detritus, while locally variable, were broadly comparable among zones. C:P ratios of both components decreased in leeward zones. Although locally variable, detritus had a higher content of both N and P. Overall, detritus is relatively abundant on windward reef substrata and of equal or greater nutritional quality than algae. It represents a potentially important resource for both detritivorous and herbivorous fishes.     

At Palmyra Atoll,the fish-community environmental DNA signal changes across habitats but not with tides

At Palmyra Atoll, the environmental DNA (eDNA) signal on tidal sand flats was associated with fish biomass density and captured 98%–100% of the expected species diversity there. Although eDNA spilled over across habitats, species associated with reef habitat contributed more eDNA to reef sites than to sand-flat sites, and species associated with sand-flat habitat contributed more eDNA to sand-flat sites than to reef sites. Tides did not disrupt the sand-flat habitat signal. At least 25 samples give a coverage >97.5% at this diverse, tropical, marine system.