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     

Relationships among map resolution, fish assemblages, and habitat variables in a coral reef ecosystem

Benthic maps are broad-scale characterizations that lack the detailed environmental attributes which have been the focus of most prior empirical studies linking reef fish and habitat. We used multivariate analyses to quantify correlations among fish assemblages, local habitat variables, and reef types depicted in benthic maps. Benthic maps of a study system in the U.S. Virgin Islands with high (100 m² minimum mapping unit) and low (4,048 m² minimum mapping unit) spatial resolution, respectively, were evaluated. Benthic maps depicted six reef types and two shelf positions (lagoon vs. shelf). Fish assemblages and local environmental variables were quantified at random sites throughout the landscape by diver surveys. Multivariate ordination based on either fish assemblages or environmental data did not result in well-separated groups of sites. Mapped reef types were not associated with distinct values of either local environmental variables or fish assemblages. Reef types exhibited substantial overlap in ordination plots based on benthic characteristics with groupings based on fish assemblages showing even greater overlap. Ordination patterns involving reef type were largely the same for both low- and high-resolution maps. In contrast, sites showed clear groups for lagoon and shelf in ordinations based on both environmental variables and fish assemblage composition, respectively. These results suggest that knowledge of the overall fish assemblage or fine-scale environmental characteristics could not be used to predict reef type depicted in benthic maps or vice versa. In contrast, reef zone could be used to predict fish assemblage or fine-scale environmental variables and vice versa.     

Does increased habitat complexity reduce predation and competition in coral reef fish assemblages?

Greater habitat complexity is often associated with a greater abundance and diversity of organisms. High complexity habitats may reduce predation and competition, thereby allowing more individuals to occupy a given area. Using 16 spatially isolated reefs in the Bahamas, I tested whether increased habitat complexity reduced the negative effects of resident predators and competitors on recruitment and survival of a common damselfish. Two levels of habitat complexity were cross-factored with the presence or absence of two guilds of resident fishes: predators (sea basses and moray eels) and interference competitors (large territorial damselfishes). I monitored subsequent recruitment and recruit mortality for 60 days. Residents had strong negative effects on recruitment regardless of habitat complexity. In the presence of residents, recruits suffered high mortality immediately after settlement that was similar on low and high complexity reefs, although high complexity reduced mortality of recruits that survived this early postsettlement period. Comparisons between shelter hole diameters and the sizes of residents suggest that territorial damselfishes and small resident predators could access most shelter holes, whereas large resident predators were excluded from many shelter holes. This study demonstrates that whether habitat complexity reduces predation and competition may depend on several key factors, such as the availability of appropriate shelter, behavioral attributes of interactors, and developmental stage of prey/inferior competitors.     

Interactions between coral restoration and fish assemblages: implications for reef management

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.     

Predator-induced demographic shifts in coral reef fish assemblages

In recent years, it has become apparent that human impacts have altered community structure in coastal and marine ecosystems worldwide. Of these, fishing is one of the most pervasive, and a growing body of work suggests that fishing can have strong effects on the ecology of target species, especially top predators. However, the effects of removing top predators on lower trophic groups of prey fishes are less clear, particularly in highly diverse and trophically complex coral reef ecosystems. We examined patterns of abundance, size structure, and age-based demography through surveys and collection-based studies of five fish species from a variety of trophic levels at Kiritimati and Palmyra, two nearby atolls in the Northern Line Islands. These islands have similar biogeography and oceanography, and yet Kiritimati has ～10,000 people with extensive local fishing while Palmyra is a US National Wildlife Refuge with no permanent human population, no fishing, and an intact predator fauna. Surveys indicated that top predators were relatively larger and more abundant at unfished Palmyra, while prey functional groups were relatively smaller but showed no clear trends in abundance as would be expected from classic trophic cascades. Through detailed analyses of focal species, we found that size and longevity of a top predator were lower at fished Kiritimati than at unfished Palmyra. Demographic patterns also shifted dramatically for 4 of 5 fish species in lower trophic groups, opposite in direction to the top predator, including decreases in average size and longevity at Palmyra relative to Kiritimati. Overall, these results suggest that fishing may alter community structure in complex and non-intuitive ways, and that indirect demographic effects should be considered more broadly in ecosystem-based management.     

Thermal stress induces persistently altered coral reef fish assemblages

Ecological communities are reorganizing in response to warming temperatures. For continuous ocean habitats this reorganization is characterized by large‐scale species redistribution, but for tropical discontinuous habitats such as coral reefs, spatial isolation coupled with strong habitat dependence of fish species imply that turnover and local extinctions are more significant mechanisms. In these systems, transient marine heatwaves are causing coral bleaching and profoundly altering habitat structure, yet despite severe bleaching events becoming more frequent and projections indicating annual severe bleaching by the 2050s at most reefs, long‐term effects on the diversity and structure of fish assemblages remain unclear. Using a 23‐year time series spanning a thermal stress event, we describe and model structural changes and recovery trajectories of fish communities after mass bleaching. Communities changed fundamentally, with the new emergent communities dominated by herbivores and persisting for >15 years, a period exceeding realized and projected intervals between thermal stress events on coral reefs. Reefs which shifted to macroalgal states had the lowest species richness and highest compositional dissimilarity, whereas reefs where live coral recovered exceeded prebleaching fish richness, but remained dissimilar to prebleaching compositions. Given realized and projected frequencies of bleaching events, our results show that fish communities historically associated with coral reefs will not re‐establish, requiring substantial adaptation by managers and resource users.     

Predators target rare prey in coral reef fish assemblages

Predation can result in differing patterns of local prey diversity depending on whether predators are selective and, if so, how they select prey. A recent study comparing the diversity of juvenile fish assemblages among coral reefs with and without predators concluded that decreased prey diversity in the presence of predators was most likely caused by predators actively selecting rare prey species. We used several related laboratory experiments to explore this hypothesis by testing: (1) whether predators prefer particular prey species, (2) whether individual predators consistently select the same prey species, (3) whether predators target rare prey, and (4) whether rare prey are more vulnerable to predation because they differ in appearance/colouration from common prey. Rare prey suffered greater predation than expected and were not more vulnerable to predators because their appearance/colouration differed from common prey. Individual predators did not consistently select the same prey species through time, suggesting that prey selection behaviour was flexible and context dependent rather than fixed. Thus, selection of rare prey was unlikely to be explained by simple preferences for particular prey species. We hypothesize that when faced with multiple prey species predators may initially focus on rare, conspicuous species to overcome the sensory confusion experienced when attacking aggregated prey, thereby minimizing the time required to capture prey. This hypothesis represents a community-level manifestation of two well-documented and related phenomena, the “confusion effect” and the “oddity effect”, and may be an important, and often overlooked, mechanism by which predators influence local species diversity.     

Wave energy and swimming performance shape coral reef fish assemblages

Physical factors often have an overriding influence on the distribution patterns of organisms, and can ultimately shape the long-term structure of communities. Although distribution patterns in sessile marine organisms have frequently been attributed to functional characteristics interacting with wave-induced water motion, similar evidence for mobile organisms is lacking. Links between fin morphology and swimming performance were examined in three diverse coral reef fish families from two major evolutionary lineages. Among-habitat variation in morphology and performance was directly compared with quantitative values of wave-induced water motion from seven coral reef habitats of different depth and wave exposure on the Great Barrier Reef. Fin morphology was strongly correlated with both field and experimental swimming speeds in all three families. The range of observed swimming speeds coincided closely with the magnitude of water velocities commonly found on coral reefs. Distribution patterns in all three families displayed highly congruent relationships between fin morphology and wave-induced water motion. Our findings indicate a general functional relationship between fin morphology and swimming performance in labriform-swimming fishes, and provide quantitative evidence that wave energy may directly influence the assemblage structure of coral reef fishes through interactions with morphology and swimming performance.     

Within and between day variability in coral reef fish assemblages: Implications for fish community surveys

Coral reef ecosystems are under increasing anthropogenic pressures making it ever more important to monitor changes in fish communities to implement appropriate management. In contrast to long-term spatial and temporal variation which has been extensively documented, little work has been carried out to identify variability in fish assemblages on short time scales, with few studies testing patterns of fish assemblages between and within days. Here we investigated the diurnal changes in species richness, relative abundance and assemblage composition in a shallow coral reef fish community in Egypt. To do so, a section of coral reef was filmed during the morning (0600 h), midday (1000 and 1400 h) and afternoon (1800 h) over eleven days. Dusk (0600 h) and dawn samples (1800 h) showed higher species richness compared to late morning (1000 h) and mid-day samples (1400 h) and borderline significantly higher numbers of total individuals, likely associated with feeding activity and predator avoidance. Assemblage composition varied across days and time-of-day, showing greater variability during dusk and dawn associated with a transition between day-time and night-time assemblages. Our results have implications for designing coral reef fish surveys, emphasising that short-term changes in fish communities should be considered when designing experiments to monitor fish assemblages over time. Where possible, we suggest increasing replication within sites and time scales or randomising data within a specific time window at all sites, looking to exclude dusk and dawn.     

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.     

The importance of structural complexity in coral reef ecosystems

The importance of structural complexity in coral reefs has come to the fore with the global degradation of reef condition; however, the limited scale and replication of many studies have restricted our understanding of the role of complexity in the ecosystem. We qualitatively and quantitatively (where sufficient standardised data were available) assess the literature regarding the role of structural complexity in coral reef ecosystems. A rapidly increasing number of publications have studied the role of complexity in reef ecosystems over the past four decades, with a concomitant increase in the diversity of methods used to quantify structure. Quantitative analyses of existing data indicate a strong negative relationship between structural complexity and algal cover, which may reflect the important role complexity plays in enhancing herbivory by reef fishes. The cover of total live coral and branching coral was positively correlated with structural complexity. These habitat attributes may be creating much of the structure, resulting in a collinear relationship; however, there is also evidence of enhanced coral recovery from disturbances where structural complexity is high. Urchin densities were negatively correlated with structural complexity; a relationship that may be driven by urchins eroding reef structure or by their gregarious behaviour when in open space. There was a strong positive relationship between structural complexity and fish density and biomass, likely mediated through density-dependent competition and refuge from predation. More variable responses were found when assessing individual fish families, with all families examined displaying a positive relationship to structural complexity, but only half of these relationships were significant. Although only corroborated with qualitative data, structural complexity also seems to have a positive effect on two ecosystem services: tourism and shoreline protection. Clearly, structural complexity is an integral component of coral reef ecosystems, and it should be incorporated into monitoring programs and management objectives.     

The influence of coral reef benthic condition on associated fish assemblages

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

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

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

Environmental factors affecting large-bodied coral reef fish assemblages in the Mariana Archipelago

Large-bodied reef fishes represent an economically and ecologically important segment of the coral reef fish assemblage. Many of these individuals supply the bulk of the reproductive output for their population and have a disproportionate effect on their environment (e.g. as apex predators or bioeroding herbivores). Large-bodied reef fishes also tend to be at greatest risk of overfishing, and their loss can result in a myriad of either cascading (direct) or indirect trophic and other effects. While many studies have investigated habitat characteristics affecting populations of small-bodied reef fishes, few have explored the relationship between large-bodied species and their environment. Here, we describe the distribution of the large-bodied reef fishes in the Mariana Archipelago with an emphasis on the environmental factors associated with their distribution. Of the factors considered in this study, a negative association with human population density showed the highest relative influence on the distribution of large-bodied reef fishes; however, depth, water temperature, and distance to deep water also were important. These findings provide new information on the ecology of large-bodied reef fishes can inform discussions concerning essential fish habitat and ecosystem-based management for these species and highlight important knowledge gaps worthy of additional research.     

Environmental variability, early life-history traits, and survival of new coral reef fish recruits

For benthic marine organisms with complex life cycles, conditionsexperienced by pelagic larvae can influence juvenile survival.Trait-specific selective mortality has been documented in thelaboratory and field, yet our knowledge of the factors contributingto the existence, strength, and consistency of natural selectivemortality is limited. We compiled previously published and unpublisheddata on the common Caribbean coral reef fish, Thalassoma bifasciatum,recruiting to Barbados, West Indies, and the upper Florida Keysto examine how environmental variability during pelagic larvallife influences the distribution of early life-history traitsexhibited by new recruits. We explored how the scope of variabilityin otolith-derived traits such as larval growth, pelagic larvalduration (PLD), size and condition at settlement, and earlyjuvenile growth influences the degree to which mortality ofjuveniles is selective. At both locations, contrasting oceanographicconditions (periodic passage of large low-salinity North BrazilCurrent [NBC] rings near Barbados and seasonal variation inwater temperature at Florida) led to significant differencesin the early life-history traits of recruits. Mortality wasmost frequently selective for the two most variable traits,condition at settlement and early juvenile growth. Environmentalvariability (including variation in predation pressure and stress-inducingconditions) also likely influences juvenile mortality and consequentlythe degree to which selective loss of particular traits occurs.As we begin to better understand the spatial, temporal, andspecies-specific circumstances in which events occurring duringlarval life influence juvenile performance, studies must alsobe extended to examine how these processes are translated intoadult fitness.     

Kenyan coral reef lagoon fish: effects of fishing,substrate complexity,and sea urchins

Population density, number of species, diversity, and species-area relationships of fish species in eight common coral reef-associated families were studied in three marine parks receiving total protection from fishing, four sites with unregulated fishing, and one reef which recently received protection from fishing (referred to as a transition reef). Data on coral cover, reef topographic complexity, and sea urchin abundance were collected and correlated with fish abundance and species richness. The most striking result of this survey is a consistent and large reduction in the population density and species richness of 5 families (surgeonfish, triggerfish, butterflyfish, angelfish, and parrotfish). Poor recovery of parrotfish in the transition reef, relative to other fish families, is interpreted as evidence for competitive exclusion of parrotfish by sea urchins. Reef substrate complexity is significantly associated with fish abundance and diversity, but data suggest different responses for protected versus fished reefs, protected reefs having higher species richness and numbers of individuals than unprotected reefs for the same reef complexity. Sea urchin abundance is negatively associated with numbers of fish and fish species but the interrelationship between sea urchins, substrate complexity, coral cover, and management make it difficult to attribute a set percent of variance to each factor-although fishing versus no fishing appears to be the strongest variable in predicting numbers of individuals and species of fish, and their community similarity. Localized species extirpation is evident for many species on fished reefs (for the sampled area of 1.0 ha). Fifty-two of 110 species found on protected reefs were not found on unprotected reefs.     

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.