Relationships between coral reef substrata and fish

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

Maintenance of fish diversity on disturbed coral reefs

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

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

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

Mass coral bleaching causes biotic homogenization of reef fish assemblages

Global climate change is altering community composition across many ecosystems due to nonrandom species turnover, typically characterized by the loss of specialist species and increasing similarity of biological communities across spatial scales. As anthropogenic disturbances continue to alter species composition globally, there is a growing need to identify how species responses influence the establishment of distinct assemblages, such that management actions may be appropriately assigned. Here, we use trait‐based analyses to compare temporal changes in five complementary indices of reef fish assemblage structure among six taxonomically distinct coral reef habitats exposed to a system‐wide thermal stress event. Our results revealed increased taxonomic and functional similarity of previously distinct reef fish assemblages following mass coral bleaching, with changes characterized by subtle, but significant, shifts toward predominance of small‐bodied, algal‐farming habitat generalists. Furthermore, while the taxonomic or functional richness of fish assemblages did not change across all habitats, an increase in functional originality indicated an overall loss of functional redundancy. We also found that prebleaching coral composition better predicted changes in fish assemblage structure than the magnitude of coral loss. These results emphasize how measures of alpha diversity can mask important changes in the structure and functioning of ecosystems as assemblages reorganize. Our findings also highlight the role of coral species composition in structuring communities and influencing the diversity of responses of reef fishes to disturbance. As new coral species configurations emerge, their desirability will hinge upon the composition of associated species and their capacity to maintain key ecological processes in spite of ongoing disturbances.     

Multispecies spawning sites for fishes on a low‐latitude coral reef: spatial and temporal patterns

Spawning sites used by one or more species were located by intensively searching nearshore coral reefs of Kimbe Bay (New Britain, Papua New Guinea). Once identified, the spawning sites were surveyed repeatedly within fixed 5 m radius circular areas, for > 2000 h of observations ranging from before dawn to after dusk spanning 190 days between July 2001 and May 2004. A total of 38 spawning sites were identified on the seven study reefs distributed at an average of one site every 60 m of reef edge. Pelagic spawning was observed in 41 fish species from six families. On three intensively studied reefs, all 17 spawning sites identified were used by at least three species, with a maximum of 30 different species observed spawning at a single site. Spawning was observed during every month of the study, on all days of the lunar month, at all states of the tide and at most hours of the day studied. Nevertheless, the majority of species were observed spawning on proportionately more days from December to April, on more days around the new moon and in association with higher tides. The strongest temporal association, however, was with species‐specific diel spawning times spanning < 3 h for most species. While dawn spawning, afternoon spawning and dusk spawning species were differentiated, the time of spawning for the striated surgeonfish Ctenochaetus striatus also differed significantly among sites. The large number of species spawning at the same restricted locations during predictable times suggests that these sites are extremely important on this low‐latitude coral reef.     

Evidence of artisanal fishing impacts and depth refuge in assemblages of Fijian reef fish

Protection from fishing generally results in an increase in the abundance and biomass of species targeted by fisheries within marine reserve boundaries. Natural refuges such as depth may also protect such species, yet few studies in the Indo Pacific have investigated the effects of depth concomitant with marine reserves. We studied the effects of artisanal fishing and depth on reef fish assemblages in the Kubulau District of Vanua Levu Island, Fiji, using baited remote underwater stereo-video systems. Video samples were collected from shallow (5–8 m) and deep (25–30 m) sites inside and outside of a large old marine reserve (60.6 km², 13 years old) and a small new marine reserve (4.25 km², 4 years old). Species richness tended to be greater in the shallow waters of the large old reserve when compared to fished areas. In the deeper waters, species richness appeared to be comparable. The difference in shallow waters was driven by species targeted by fisheries, indicative of a depth refuge effect. In contrast, differences in the abundance composition of the fish assemblage existed between protected and fished areas for deep sites, but not shallow. Fish species targeted by local fisheries were 89% more abundant inside the large old reserve than surrounding fished areas, while non-targeted species were comparable. We observed no difference in the species richness or abundance of species targeted by fisheries inside and outside of the small new reserve. This study suggests that artisanal fishing impacts on the abundance and species richness of coral reef fish assemblages and effects of protection are more apparent with large reserves that have been established for a long period of time. Observed effects of protection also vary with depth, highlighting the importance of explicitly incorporating multiple depth strata in studies of marine reserves.     

Species composition,habitat configuration and seasonal changes of coral reef fish assemblages in western Mexico

In spite of their ecological and economic importance, reef fishes from the coast of Oaxaca, Mexico are rarely studied, therefore precluding their management and conservation. In order to identify the set of habitat characteristics/environmental conditions that predict major shifts in fish assemblages in space and time, a stationary census (5′, φ = 5 m) was conducted on a semi‐monthly basis from 2006 to 2009 at patch reefs along the coast. Habitat configuration was gathered using 25 m long point‐intersect transects (data every 25 cm), recording all underlying coral species and substrate characteristics (rocks, sand, algal mats, rubble or dead corals). Recorded were 65 452 fishes grouped in 11 orders, 36 families, 65 genera and 89 species. Labridae (nine species), Pomacentridae (eight species) and Serranidae (seven species) were the most frequent families. Abundance is severely skewed among species; four species Thalassoma lucasanum, Chromis atrilobata, Apogon pacificus and Stegastes acapulcoensis comprise nearly 59% of the fish abundance, 11 species contribute 30%, whereas most of the species (75) can be considered as rare since they contribute <1% each to the total. Species richness and family‐level assemblage composition are similar to those recorded elsewhere in the eastern Pacific. Non‐parametric multivariate analysis of variance demonstrated that changes of diversity metrics might be associated with environmental differences on the scale of hundreds of meters to kilometers, as well as coupled with major changes on oceanographic variables throughout time, exerting meaningful changes on reef‐related fish assemblages.     

Spatial predictability of juvenile fish species richness and abundance in a coral reef environment

Juvenile reef fish communities represent an essential component of coral reef ecosystems in the current focus of fish population dynamics and coral reef resilience. Juvenile fish survival depends on habitat characteristics and is, following settlement, the first determinant of the number of individuals within adult populations. The goal of this study was to provide methods for mapping juvenile fish species richness and abundance into spatial domains suitable for micro and meso-scale analysis and management decisions. Generalized Linear Models predicting juvenile fish species richness and abundance were developed according to spatial and temporal environmental variables measured from 10 m up to 10 km in the southwest lagoon of New Caledonia. The statistical model was further spatially generalized using a 1.5-m resolution, independently created, remotely sensed, habitat map. This procedure revealed that : (1) spatial factors at 10 to 100-m scale explained up to 71% of variability in juvenile species richness, (2) a small improvement (75%) was gained when a combination of environmental variables at different spatial and temporal scales was used and (3) the coupling of remotely sensed data, geographical information system tools and point-based ecological data showed that the highest species richness and abundance were predicted along a narrow margin overlapping the coral reef flat and adjacent seagrass beds. Spatially explicit models of species distribution may be relevant for the management of reef communities when strong relationships exist between faunistic and environmental variables and when models are built at appropriate scales.     

Geography and island geomorphology shape fish assemblage structure on isolated coral reef systems

We quantify the relative importance of multi‐scale drivers of reef fish assemblage structure on isolated coral reefs at the intersection of the Indian and Indo‐Pacific biogeographical provinces. Large (>30 cm), functionally‐important and commonly targeted species of fish, were surveyed on the outer reef crest/front at 38 coral reef sites spread across three oceanic coral reef systems (i.e. Christmas Island, Cocos (Keeling) Islands and the Rowley Shoals), in the tropical Indian Ocean (c. 1.126 x 106 km²). The effects of coral cover, exposure, fishing pressure, lagoon size and geographical context, on observed patterns of fish assemblage structure were modelled using Multivariate Regression Trees. Reef fish assemblages were clearly separated in space with geographical location explaining ~53 % of the observed variation. Lagoon size, within each isolated reef system was an equally effective proxy for explaining fish assemblage structure. Among local‐scale variables, ‘distance from port’, a proxy for the influence of fishing, explained 5.2% of total variation and separated the four most isolated reefs from Cocos (Keeling) Island, from reefs with closer boating access. Other factors were not significant. Major divisions in assemblage structure were driven by sister taxa that displayed little geographical overlap between reef systems and low abundances of several species on Christmas Island corresponding to small lagoon habitats. Exclusion of geographical context from the analysis resulted in local processes explaining 47.3% of the variation, highlighting the importance of controlling for spatial correlation to understand the drivers of fish assemblage structure. Our results suggest reef fish assemblage structure on remote coral reef systems in the tropical eastern Indian Ocean reflects a biogeographical legacy of isolation between Indian and Pacific fish faunas and geomorphological variation within the region, more than local fishing pressure or reef condition. Our findings re‐emphasise the importance that historical processes play in structuring contemporary biotic communities.     

Island biogeography of Caribbean coral reef fish

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

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

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

Diurnal variability in a fish assemblage of a Bahamian coral reef

Synopsis Variation in the diurnal composition of a fish assemblage of a Bahamian coral reef was investigated by comparing visual counts of fishes taken along a 100 × 4 m wide fixed transect at four times: 0900, 1200, 1500 and 1800 hours during the summer of 1979. One sample per day was obtained at each time period over a span of 20 consecutive days. Forty-two species were recorded in these samples, with 25 occurring frequently enough to permit statistical analysis of diurnal variations in abundance. Of the 25 species compared, nearly one fourth (6 species) displayed significant variation in abundance patterns among the four time periods tested. It is suggested, because of the strong possibility of bias which might otherwise be introduced because of these variations, that repetitive quantitative visual censusing of coral reef fishes be undertaken at about the same time each day.     

Local ecological impacts of regional biodiversity on reef fish assemblages

Aim We examined comparative data for cryptobenthic reef fishes to determine how variation in regional species richness relates to local species richness, abundance, and taxonomic and trophic composition, and to test whether systems with higher species richness exhibit finer habitat partitioning. Locations Lizard Island, Great Barrier Reef (GBR), Australia; Bahía de Loreto, Gulf of California (GoC), Mexico. Methods Cryptobenthic reef fish assemblages from four habitats (coral heads, rubble, and horizontal and vertical surfaces of boulders) were collected using clove oil. Differences in density, species richness and biomass were examined between regions and among habitats. Habitat associations were identified for each habitat/location based on multivariate ordination, and the statistical significance of patterns was tested using analysis of similarity (ANOSIM). In addition, the trophic group composition of the assemblages for both regions was examined. Results A total of 91 species in 20 families were recorded (GBR, 66 species; GoC, 25 species). Total and habitat species richness were higher on the GBR, whereas biomass was higher in the GoC. No difference in fish density between regions was found. Habitat division among assemblages was greater in the depauperate GoC. Only coral head associations proved to be distinctive on the GBR, whereas three sample groups were found in the GoC (coral heads, horizontal boulders and vertical boulders/rubble). Trophic composition in the two regions was markedly different, with omnivores dominating the GBR fauna and planktivores the GoC. Main conclusions A positive regional–local relationship in fish diversity was found between regions, but fish abundance in both regions remained similar. Contrary to expectations, habitat partitioning, at a community level, was greater in the depauperate GoC. Differences in trophic composition and patterns of habitat use appear to reflect the disparate history of the regions, whereas patterns of abundance may reflect the influence of fundamental relationships between size and abundance in communities. This study highlights the potential of reef faunas to conform to universal numerical trends while maintaining an ability to respond ecologically to local/evolutionary influences. The GoC fauna appears to be exceptionally vulnerable to natural and anthropogenic disturbance owing to the high numerical dominance of habitat‐specific species and to the limited potential for functional redundancy within the system.     

Diurnal temporal patterns of the diversity and the abundance of reef fishes in a branching coral patch in New Caledonia

Small‐scale spatial and temporal variability in animal abundance is an intrinsic characteristic of marine ecosystems but remains largely unknown for most animals, including coral reef fishes. In this study, we used a remote autonomous unbaited video system and recorded reef fish assemblages during daylight hours, 10 times a day for 34 consecutive days in a branching coral patch of the lagoon of New Caledonia. In total, 50 031 fish observations belonging to 114 taxa, 66 genera and 31 families were recorded in 256 recorded videos. Carnivores and herbivore‐detritus feeders dominated the trophic structure. We found significant variations in the composition of fish assemblages between times of day. Taxa richness and fish abundance were greater in the early morning and in the late afternoon than during the day. Fourteen taxa displayed well‐defined temporal patterns in abundance with one taxon influenced by time of day, six influenced by tidal state and seven influenced by both time of day and tidal state. None of these 14 taxa were piscivores, 10 were herbivore‐detritus feeders, three were carnivores and one was plankton feeder. Our results suggest a diel migration from feeding grounds to shelter areas and highlight the importance of taking into account small‐scale temporal variability in animal diversity and abundance when studying connectivity between habitats and monitoring communities.     

Structure of Caribbean coral reef communities across a large gradient of fish biomass

The collapse of Caribbean coral reefs has been attributed in part to historic overfishing, but whether fish assemblages can recover and how such recovery might affect the benthic reef community has not been tested across appropriate scales. We surveyed the biomass of reef communities across a range in fish abundance from 14 to 593 g m⁻², a gradient exceeding that of any previously reported for coral reefs. Increased fish biomass was correlated with an increased proportion of apex predators, which were abundant only inside large marine reserves. Increased herbivorous fish biomass was correlated with a decrease in fleshy algal biomass but corals have not yet recovered.     

Disturbance and the structure of coral reef fish communities on the reef slope

Three levels of physical disturbance were applied to corals in permanent 10x10 m quadrats along a section of fringing reef at Lizard Island on the Great Barrier Reef to investigate the response of fish assemblages. Tabular and corymbose corals were overturned and left in situ, reducing total hard coral cover from ˜55% to ˜47%, ˜43%, and ˜34%. Despite pre-existing associations with benthic cover, all fish groups examined (pomacentrids, labrids, chaetodontids, and acanthurids) were resistent to benthic disturbances at the level and scale at which they were applied. Partial Mantel's tests, in combination with partial Canonical Correspondence Analysis enabled spatial and temporal variation to be factored out from experimental effects. Most of the variation in the fish community could be assigned to spatio-temporal variables, indicating that spatial structure over the reef landscape may moderate localised disturbance effects. This study indicates that coral reef fish assemblages may be more resistant to disturbance than many correlative studies would suggest, and highlights a need for further information on levels and scales of natural habitat disturbance in order to apply a structured approach to the experimental investigation of the importance of habitat in structuring coral-reef fish assemblages.     

Anthropogenic Stressors,Inter-Specific Competition and ENSO Effects on a Mauritian Coral Reef

Much of the western Indian Ocean suffered widespread loss of live coral in 1998 and interest is now focussed on the indirect effects of this coral loss on other components of the ecosystem, in particular fishes. However, it is just as important to identify changes in fish assemblages at locations that did not suffer coral mortality to understand local versus regional drivers. We surveyed benthic and fish communities on a reef flat in Mauritius five times between 1994 and 2005. The design allowed for comparison through time, along the coast and between inshore and offshore reef locations. The benthic community demonstrates a clear trend along the coast, likely in response to a dredged water ski lane, but little change through time. Branching Acropora colonies dominate much of the live coral and best explain patterns in the fish assemblage (P < 0.01). Few changes in overall fish species richness through time were identified, and observed changes were within fishery target families rather than species reliant on live coral. Departure from expected levels of taxonomic distinctness suggests degradation in the community associated with the dredged ski lane. Non-metric multi-dimensional scaling of the fish assemblage demonstrates a similar pattern to that seen in the benthos; greater differences along the coast (Global R = 0.34) than through time (Global R = 0.17) and no trend between reef positions. SIMPER analysis identified two species of Stegastes as the main drivers of trends in the MDS plot and the most dominant of these, S. lividus, appears to be reducing species richness of the remaining fish community. The study highlights Mauritius as a regional refugia of thermally-sensitive corals and specialised fish, suggesting a need for careful management.     

Meta-population structure in a coral reef fish demonstrated by genetic data on patterns of migration,extinction and re-colonisation

Background  

Management strategies for coral reefs are dependant on information about the spatial population structure and connectivity of reef organisms. Genetic tools can reveal important information about population structure, however, this information is lacking for many reef species. We used a mitochondrial molecular marker to examine the population genetic structure and the potential for meta-population dynamics in a direct developing coral reef fish using 283 individuals from 15 reefs on the Great Barrier Reef, Australia. We employed a hierarchical sampling design to test genetic models of population structure at multiple geographical scales including among regions, among shelf position and reefs within regions. Predictions from island, isolation-by-distance and meta-population models, including the potential for asymmetric migration, local extinction and patterns of re-colonisation were examined.     

Evidence of an original scale development during the settlement phase of a coral reef fish (Acanthurus triostegus)

As the majority of coral reef fishes, the Convict Surgeonfish Acanthurus triostegus (Acanthuridae) has a complex life cycle that involves an ontogenetic change in morphology, physiology and behaviour as its pelagic larval stage colonizes the benthic habitat. Few studies are devoted to the changes in skeleton during the settlement phase of coral reef fishes. In the present study, we highlighted an unexpected scales development in A. trisostegus just after the reef settlement. At settlement (t₀), A. triostegus showed calcified and very thin vertical plates, lying in the dermis on the whole body. During the first 9 days after settlement, thin vertical plates regressed and adult scales began to appear simultaneously. At 12 days post‐settlement, the whole body was covered with small scales. Overall, such a rapid skeletal transformation is an example of morphological changes dealing with ‘metamorphosis’ of coral reef fishes.     

Co-occurrence of herbivorous fish functional groups correlates with enhanced coral reef benthic state

Aim

Biodiversity loss is impacting essential ecosystem functions and services across the globe. Recently, our interest in the benefits of biodiversity for ecosystem function has shifted focus from measurements of species richness to functional diversity and composition. However, the additional importance of other community characteristics, such as species evenness and co-occurrence, for diversity-driven ecosystem function is less known. We used herbivorous coral reef fish as a model system to investigate how co-occurrence of different functional groups, rather than purely functional diversity, within an assemblage might affect the coral reef benthic state.

Location

Western Atlantic.

Time period

2007–2017.

Major taxa studied

Herbivorous reef fish.

Methods

We analysed benthic and fish assemblage data from 601 sites across 12 countries in the western Atlantic. Using diversity–interaction models, we investigated how the composition and relative abundances of reef fish functional groups were correlated with benthic cover and estimates of coral calcification rates. We used statistical interactions to explore the importance of co-occurrence of herbivorous fish functional groups for the coral reef benthic state.

Results

We found that co-occurrence of herbivorous fish functional groups, in addition to functional diversity, was correlated with reduced algal cover and increased coral accretion. Moreover, pairwise statistical interactions between functional groups were significantly correlated with an improvement in the coral reef benthic state.

Main conclusions

Our results support the idea that functional group co-occurrence, in addition to functional diversity, within herbivorous fish offers additional benefits to the coral reef benthic state. We identify farming damselfish and excavating parrotfish as potential key determinants of the coral reef benthic state and highlight that co-occurrence of cropping and scraping herbivores might promote coral accretion. Our findings support the argument that protecting herbivore abundance without regard to the species and functional groups present is not enough to preserve coral reef health and that fine-scale community composition must be considered.