Phase-shift in coral reef communities in the Florida Keys National Marine Sanctuary (FKNMS), USA

Characterizing the Florida Keys National Marine Sanctuary (FKNMS), USA, has gained much attention over the past several decades because of apparent changes in the benthic community structure over space and time representative of patterns occurring in the Caribbean region. We used a 5-year dataset (1996–2000) of macroalgal and sponge cover and water quality measurements as predictor variables of hard coral community structure in the FKNMS. The 16 water quality variables were summarized into 4 groups by principal component analysis (PCA). Hierarchical agglomerative cluster analysis of the mean and standard deviation (SD) of the principal component scores of water quality variables separated the reef sites into two main groups (and five sub-groups), referred to as reefs of similar influence (RSI). The main groups corresponded with their geographical locations within the Florida Keys: the reefs in the Upper and Middle Keys being homogeneous and collectively, having lower water quality scores relative to reefs in the Lower Keys. Canonical correspondence analysis (CCA) between hard coral cover and key predictor variables (i.e., water quality, macroalgal cover and sponge cover) also separated the reefs in the Lower Keys from reefs in the Upper–Middle Keys, consistent with results of the cluster analysis, which categorized reefs based on RSI. These results suggest that the prevailing gradient of predictor variables may have influenced the structuring of coral reef communities at a spatial scale larger than the individual reef. Furthermore, it is conceivable that these predictor variables exerted influence for a long time rather than being a recent event. Results also revealed a pattern showing reduction in hard coral cover and species richness, and subsequent proliferation of macroalgae and sponges during the study period. Our analyses of the Florida Keys present a pattern that is consistent with the characteristics of a reef that has undergone a “phase-shift,” a phenomenon that is widely reported in the Caribbean region.     

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

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

Environmental influences on larval duration,growth and magnitude of settlement of a coral reef fish

The influence of environmental variables on the planktonic growth, pelagic larval duration and settlement magnitude was examined for the coral reef surgeonfish Acanthurus chirurgus. Newly settled fish were collected daily from patch reefs in the San Blas Archipelago, Caribbean Panama for 3.5 years. Environmental influences on growth were examined at three different life history stages: from 0 to 6 days, 7 to 25 days and from 26 to 50 days after hatching. Larval growth was correlated, using multiple regression techniques, with a combination of factors including solar radiation, rainfall, and along-shore winds. Depending on the life history stage, these accounted for 13–38% of the variation in growth rates when all the months were included in the analyses. Correlations between environmental variables and growth also varied among seasons and were stronger in the dry than in the wet season. During the dry season solar radiation, rainfall and along-shore winds described 57%, 86% and 74% of the variability in growth between 0 and 6 days, 7 and 25 days and 26 and 50 days, respectively. During the wet season rainfall, along-shore winds and temperature only described 38% of the variability in early growth and 27% of growth just before settlement. No significant model was found to describe growth 7–25 days after hatching during the wet season. Rainfall, solar radiation and along-shore winds were negatively correlated with growth up to 25 days after hatching but positively correlated as larvae approached settlement at a mean age of 52 days. Over 65% of the variability in pelagic larval duration was accounted for by a regression model that included solar radiation and along-shore winds. When data sets from wet and dry seasons were analysed separately, along-shore winds accounted for 67% of the change in larval duration in the dry season, and solar radiation accounted for 23% of the variation in larval duration in the wet season. Only 22% of the variability in settlement intensity could be described by solar radiation and temperature, when all months of the year were included in the analysis. Solar radiation and rainfall were included in a regression model that accounted for 40% of the variation in numbers of fish settling during the dry season. This study suggests that the levels of solar radiation, along-shore winds and rainfall during the early larval life can have important effects on the growth, larval duration and consequently, the settlement magnitude of marine fishes. Results also highlight the need to account for seasonality and ontogeny in studies of environmental influences.Communicated by G.F. Humphrey, Sydney     

Evaluating day–night changes in shallow Mediterranean rocky reef fish assemblages by visual census

Ecological information on coastal fish distribution patterns and habitat use can be greatly improved by nocturnal samplings and observations. To this purpose, the structure of a Mediterranean fish assemblage inhabiting the shallow rocky littoral of Linosa Island (Sicily Strait, Italy) was examined by using visual census to detect possible diel variations in species composition and abundance. Day–night fish distribution patterns were investigated by multivariate and univariate analyses. Overall, 42 fish taxa belonging to 19 families were recorded: 35 during the day and 24 during the night. Seventeen species were common to both diurnal and nocturnal assemblages. Within the diurnal assemblage, Chromis chromis was the most represented species (37.2%), followed by Thalassoma pavo (23.2%) and Sparisoma cretense (10.8%). Within the nocturnal assemblage, the most abundant taxon was Atherina spp. (33.9%), followed by Apogon imberbis (26.4%) and Boops boops (11.5%). Our results indicated wide variation in the abundance and species composition during the day and during the night. Multi Dimensional Scaling plot showed a clear-cut separation between the two assemblages and analysis of similarities found significant differences as well. SIMPER analysis revealed that ten species individually contributed by more than 2.5% to the dissimilarity between diurnal and nocturnal assemblages, T. pavo, C. chromis and S. cretense being the first three species in order of decreasing percentage. ANOVA performed on species richness and fish abundance detected significant differences between diurnal and nocturnal assemblages, the latter showing far lower average values for both variables.     

Larval supply and juvenile recruitment of coral reef fishes to marine reserves and non-reserves of the upper Florida Keys,USA

For marine organisms, decoupling between the planktonic larval stage and the benthic-associated juvenile stage can lead to variable patterns of population replenishment, which have the potential to influence the effectiveness of marine reserves. We measured spatial and temporal variability in larval supply and recruitment of fishes to coral reefs of different protection levels and tested whether protection level influenced the relationship between supply and recruitment. We sampled pre-settlement larvae and newly settled recruits from four reefs (two reserves and two non-reserves) in the Florida Keys National Marine Sanctuary, USA. Replicate point measures of larval supply over 14 months and 17 monthly measurements of recruitment varied significantly among months and sites. Sites with the same protection level had significantly different patterns of larval supply as well as larval and recruit diversity, but recruitment magnitude differed only by protection level, where densities were greater at reserves. Differences in larval supply among sites included two particularly large peaks in larval abundance at one site, possibly associated with the observed passage of small-scale oceanographic features. To examine whether relationships between larval supply and recruitment varied by protection level, we selected one species that was present in both the light trap samples and the monthly recruitment surveys. Recruitment of the bicolor damselfish Stegastes partitus was significantly and positively related to larval supply at three of the four sites thus, protection level did not influence this linkage. Since local variability among sites can lead to spatial differences in population replenishment, characterization of larval supply and recruitment to potential marine reserve sites may help to identify optimal locations in a region and contribute to more effective reserve design.     

Diurnal space utilization in coral reef fish communities

The diurnal use of space by 25 resident species of coral reef fishes was investigated along three depth (10 to 40 m) transects over an 18 month period. Emphasis was placed on the small, cryptic members of the communities. Three aspects of the use of space — temporal utilization, occupancy and time span — are defined and quantified for each of the species. In addition, an estimate of the diurnal home range volume of each species is provided. The territorial pomacentrids showed the highest level of temporal utilization, holding the same territories throughout the study. The consistent presence of the holocentrid species within the reef infrastructure had the potential to influence the space available to other species. Some species residing in tube-worm holes and sponges also exhibited constant use of space.     

Increased behavioural lateralization in parasitized coral reef fish

Preferential use of one side of the body for cognitive or behavioural tasks (lateralization) is common in many animals, including humans. However, few studies have demonstrated whether lateralization is phenotypically plastic, and varies depending on the ecological context. We studied lateralization (measured as a turning preference) in the bridled monocle bream (Scolopsis bilineatus). This coral reef fish is commonly infected by a large, ectoparasitic isopod (Anilocra nemipteri) that attaches to the left or right side of its host’s head. Fish that were parasitized showed no turning bias with respect to the side on which the parasite had attached. On average, however, parasitised fish were significantly more lateralized (i.e. had a strong side bias) than unparasitized fish. The extent of lateralization declined significantly when we experimentally removed the parasite. Our results indicate that lateralization can vary with the ecological context. One possible explanation is that lateralization shortens the response time until fish flee after encountering a predator. A stronger side bias might be advantageous for parasitized individuals to overcome their recently documented lower maximum swimming speed.     

Local and regional genetic connectivity in a Caribbean coral reef fish

Coupled bio-physical models of larval dispersal predict that the Costa Rica–Panama (CR–PAN) reefs should constitute a demographically isolated region in the western Caribbean. We tested the hypothesis that CR–PAN coral reef fish populations would be isolated from Mesoamerican Barrier Reef System (MBRS) populations. To test that, we assessed population genetic structure in bicolor damselfish (Stegastes partitus) from both regions. Adult fish were genotyped from five reefs in CR–PAN and from four reefs along the MBRS at 12 microsatellite loci. Between-region F _ST (F _ST = 0.0030, P < 0.005) and exact test (x ² = 74.34, df = 18, P < 0.0001) results indicated that there is weak but significant genetic differentiation between regions, suggesting some restriction in connectivity along the Central American coastline, as predicted by bio-oceanographic models. Additionally, there is among-site genetic structure in the CR–PAN region, relative to the MBRS and between regions, suggesting higher self-recruitment within CR–PAN. This finding may be explained by differences in habitat characteristics.     

Microhabitat utilisation patterns in cryptobenthic coral reef fish communities

The cryptobenthic reef fish communities from four microhabitats at Orpheus Island, central Great Barrier Reef are described. Eighty-four 0.4m² samples yielded a total of 368 individuals from 42 species in eight families, with a mean density of 11 individuals m^–2 (±1.7SE) and 2.9 species 0.4 m^–2 (±0.2SE). Caves contained the highest number of both individuals (120) and species (26), followed by sand/rubble, soft coral, and open reefs. Microhabitat associations included cave and soft coral specialists. Site fidelity in 71 tagged individuals of 4 species was high, with a mean recapture rate of 53% (±8.4SE) remaining within the ~0.4 m² sampling area after a 48-h period. Behavioural observations also reflect this limited movement, with the dominant mode of behaviour in 7 species being a motionless state (67.5% ±11.6SE), followed by feeding (21.8% ±8.7SE), hiding (6.3% ±1.6SE), and swimming (4.4% ±1.5SE). Two distinct behavioural groups are identified: (1) sedentary forms, characterised by long periods of immobility (5 species); and (2) winnowers, characterised by long feeding bouts (2 species). The fine-scale partitioning of microhabitats, restricted home ranges, and sedentary behaviour of many cryptobenthic reef fish species suggest that this reef fish community exhibits similar patterns of habitat utilisation to their larger reef-fish counterparts, but at a much finer scale.     

Acute effects of removing large fish from a near-pristine coral reef

Large animals are severely depleted in many ecosystems, yet we are only beginning to understand the ecological implications of their loss. To empirically measure the short-term effects of removing large animals from an ocean ecosystem, we used exclosures to remove large fish from a near-pristine coral reef at Palmyra Atoll, Central Pacific Ocean. We identified a range of effects that followed from the removal of these large fish. These effects were revealed within weeks of their removal. Removing large fish (1) altered the behavior of prey fish; (2) reduced rates of herbivory on certain species of reef algae; (3) had both direct positive (reduced mortality of coral recruits) and indirect negative (through reduced grazing pressure on competitive algae) impacts on recruiting corals; and (4) tended to decrease abundances of small mobile benthic invertebrates. Results of this kind help advance our understanding of the ecological importance of large animals in ecosystems.     

Ultraviolet photosensitivity and feeding in larval and juvenile coral reef fishes

The ability of young coral reef fishes to feed using solely ultraviolet-A (UV-A) radiation during ontogeny was examined using natural prey in experimental tanks. Larvae and juveniles of three coral reef fish species (Pomacentrus amboinensis, Premnas biaculeatus and Apogon compressus) are able to feed successfully using UV-A radiation alone during the later half of the pelagic larval phase. The minimum UV radiation intensities required for larval feeding occur in the field down to depths of 90–130 m in oceanic waters and 15–20 m in turbid inshore waters. There was no abrupt change in UV sensitivity after settlement, indicating that UV photosensitivity may continue to play a significant role in benthic juveniles on coral reefs. Tests of UV sensitivity in the field using light traps indicate that larval and juvenile stages of 16 coral reef fish families are able to detect and respond photopositively to UV wavelengths. These include representatives from families that are unlikely to possess UV sensitivity as adults due to the UV transmission characteristics of the ocular media. Functional UV sensitivity may be more widespread in young coral reef fishes than in the adults, and may play a significant role in detecting zooplanktonic prey.     

Stability and structure of a fish community on a coral patch reef in Hawaii

Two complete collections of the fishes residing on an isolated coral patch reef ( 1500 m²) at Oahu, Hawaii, were made 11 years apart. Of the 112 species of fishes in both collections combined, only 40% were in common, but these made up more than 85% of the wet biomass in each collection. The two assemblages of fishes were similar in trophic structure and standing crop. Many coral reef fish communities are dominated by carnivorous forms. In the present study, planktivorous fishes were the most important trophic group in the community; this was related to abundant zooplankton resources. Following the second collection in 1977, recolonization by fishes was followed for 1 year. Recolonization proceeded rapidly and was primarily by juvenile fishes well beyond larval metamorphosis. Within 6 months of the second collection, the trophic structure had been re-established. The MacArthur-Wilson model of insular colonization described the recolonization process and predicted an equilibrium situation in less than 2 years. The recolonization data suggested that chance factors may explain the colonization process on a small scale, but a relatively deterministic pattern emerged when considering the entire reef. Thus, at the community level the fishes are a persistent and predictable entity.     

Safety in numbers and the spatial scaling of density-dependent mortality in a coral reef fish

In coral reef fishes, density-dependent population regulation is commonly mediated via predation on juveniles that have recently settled from the plankton. All else being equal, strong density-dependent mortality should select against the formation of high-density aggregations, yet the juveniles of many reef fishes aggregate. In light of this apparent contradiction, we hypothesized that the form and intensity of density dependence vary with the spatial scale of measurement. Individual groups might enjoy safety in numbers, but predators could still produce density-dependent mortality at larger spatial scales. We investigated this possibility using recently settled juvenile bluehead wrasse, Thalassoma bifasciatum, a small, aggregating reef fish. An initial caging experiment demonstrated that juvenile bluehead wrasse settlers suffer high predation, and spatial settlement patterns indicated that bluehead wrasse juveniles preferentially settle in groups, although they are also found singly. We then monitored the mortality of recently settled juveniles at two spatial scales: microsites, occupied by individual fish or groups of fish and separated by centimeters, and sites, consisting of approximately 2400-m2 areas of reef and separated by kilometers. At the microsite scale, we measured group size and effective population density independently and found that per capita mortality decreased with group size but was not related to density. At the larger spatial scale, however, per capita mortality increased with settler density. This shift in the form of density dependence with spatial scale could reconcile the existence of small-scale aggregative behavior typical of many reef fishes with the population-scale density dependence that is essential to population stability and persistence.     

Spatial synchrony in coral reef fish populations and the influence of climate

We investigated spatial patterns of synchrony among coral reef fish populations and environmental variables over an eight-year period on the Great Barrier Reef, Australia. Our aims were to determine the spatial scale of intra- and interspecific synchrony of fluctuations in abundance of nine damselfish species (genus Pomacentrus) and assess whether environmental factors could have influenced population synchrony. All species showed intraspecific synchrony among populations on reefs separated by < or =100 km, and interspecific synchrony was also common at this scale. At greater spatial scales, only four species showed intraspecific synchrony, over distances ranging from 100-300 km to 500-800 km, and no cases of interspecific synchrony were recorded. The two mechanisms most likely to cause population synchrony are dispersal and environmental forcing through regionally correlated climate (the Moran effect). Dispersal may have influenced population synchrony over distances up to 100 km as this is the expected spatial range for ecologically significant reef fish dispersal. Environmental factors are also likely to have synchronized population fluctuations via the Moran effect for three reasons: (1) dispersal could not have caused interspecific synchrony that was common over distances < or =100 km because dispersal cannot link populations of different species, (2) variations in both sea surface temperature and wind speed were synchronized over greater spatial scales (>800 km) than fluctuations in damselfish abundance (< or =800 km) and were correlated with an index of global climate variability, the El Ni?o-Southern Oscillation (ENSO), and (3) synchronous population fluctuations of most damselfish species were correlated with ENSO; large population increases often followed ENSO events. We recorded regional variations in the strength of population synchrony that we suspect are due to spatial differences in geophysical, oceanographic, and population characteristics, which act to dilute or enhance the effects of synchronizing mechanisms. We conclude that synchrony is common among Pomacentrus populations separated by tens of kilometers but less prevalent at greater spatial scales, and that environmental variation linked to global climate is likely to be a driving force behind damselfish population synchrony at all spatial scales on the Great Barrier Reef.     

Vertical structure of very nearshore larval fish assemblages in a temperate rocky coast

Small-scale vertical patterns of larval distribution were studied at a very nearshore larval fish assemblage, during the spring–summer period of several years, at two depth strata (surface and bottom) using sub-surface and bottom trawls. A total of 4,589 larvae (2,016 from surface samples and 2,573 from bottom samples) belonging to 62 taxa included in 22 families were collected. Most larvae belonged to coastal species. Although inter-annual variations in larval density and diversity could be found, total larval abundance was always higher near the bottom whereas diversity was higher at the surface. A marked distinction between the structure of surface and bottom assemblages was found. Sixteen taxa explained 95% of the similarity among surface samples. Larvae which contributed most to this similarity included species like clupeiformes, sparids and serranids, and also blenniids, tripterygiids and some labrids. In the bottom samples, fewer species were present, with only six taxa, almost exclusively from species which lay demersal eggs, contributing to 95% of the similarity between samples. Larvae present at the surface were significantly smaller than at the bottom. For some of the most abundant species found at the bottom, only small larvae occurred at the surface while the whole range of sizes was present at the bottom, indicating that larvae may be completing the entire pelagic phase near the adults’ habitat. These results indicate that larval retention near the reefs probably occurs for these species, although for others dispersal seems to be the prevailing mechanism.     

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

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

The effects of mating system on male mate choice in a coral reef fish

Summary We show how mate limitation appears to be critical in determining whether or not males exercise mate choice among available females. Thalassoma bifasciatum is a Caribbean reef fish with two distinct mating patterns: group-spawning and pair-spawning. In both mating systems, female fecundity is variable and size dependent, and female availability is high. However, sperm competition among group-spawning males apparently limits the number of effective matings in which a male may engage, whereas territorial pair-spawning males have little or no such limitation. Group-spawning males should be discriminating in their choice of mates and our data confirm this: there is strong evidence for assortative mating in group-spawns, with more large males joining in mating groups around large females. In contrast, pair-spawning males show no indication of mate preferences, and spawn with all females who arrive at their territories.