Full-Sibs in Cohorts of Newly Settled Coral Reef Fishes

Reef fishes exhibit a bipartite life cycle where a benthic adult stage is preceded by a pelagic dispersal phase during which larvae are presumed to be mixed and transported by oceanic currents. Genetic analyses based on twelve microsatellite loci of 181 three-spot dascyllus (Dascyllus trimaculatus) that settled concurrently on a small reef in French Polynesia revealed 11 groups of siblings (1 full sibs and 10 half-sibs). This is the first evidence that fish siblings can journey together throughout their entire planktonic dispersal phase (nearly a month long for three-spot dascyllus). Our findings have critical implications for the dynamics and genetic structure of fish populations, as well as for the design of marine protected areas and management of fisheries.     

Evolution and Diversification of Antarctic Notothenioid Fishes

Antarctica supported fossil ichthyofaunas during the Devonian,Jurassic, Cretaceous and Eocene/Oligocene. These faunas arenot ancestral to each other, nor are they related to any componentof the modern fauna. About one hundred species of notothenioidsdominate a modern fauna of over 200 species of bottom fishes.This highly endemic perciform suborder is not representedinthe fossil record of Antarctica. Notothenioids may have evolvedin situ on the margins of the Antarctic continent while graduallyadapting to cooling conditions during the Tertiary. Cladisticstudies indicate that notothenioids are a monophyletic group,but a sister group has not been identified among perciform fishes.With relatively few non-notothenioid fishes in Antarctic waters,notothenioids fill ecological roles normally occupied by taxonomicallydiverse fishes in temperate waters. There are six notothenioidfamilies: Bovichtidae, Nototheniidae, Harpagiferidae, Artedidraconidae,Bathydraconidae and Channichthyidae. Aspects of theirbiologyare briefly considered with emphasis on the Nototheniidae, themost speciose family. Evolutionary diversification within thisfamily allows recognition of species which are pelagic, cryopelagic,benthopelagic and benthic.     

Counter-Gradient Variation in Respiratory Performance of Coral Reef Fishes at Elevated Temperatures

The response of species to global warming depends on how different populations are affected by increasing temperature throughout the species'' geographic range. Local adaptation to thermal gradients could cause populations in different parts of the range to respond differently. In aquatic systems, keeping pace with increased oxygen demand is the key parameter affecting species'' response to higher temperatures. Therefore, respiratory performance is expected to vary between populations at different latitudes because they experience different thermal environments. We tested for geographical variation in respiratory performance of tropical marine fishes by comparing thermal effects on resting and maximum rates of oxygen uptake for six species of coral reef fish at two locations on the Great Barrier Reef (GBR), Australia. The two locations, Heron Island and Lizard Island, are separated by approximately 1200 km along a latitudinal gradient. We found strong counter-gradient variation in aerobic scope between locations in four species from two families (Pomacentridae and Apogonidae). High-latitude populations (Heron Island, southern GBR) performed significantly better than low-latitude populations (Lizard Island, northern GBR) at temperatures up to 5°C above average summer surface-water temperature. The other two species showed no difference in aerobic scope between locations. Latitudinal variation in aerobic scope was primarily driven by up to 80% higher maximum rates of oxygen uptake in the higher latitude populations. Our findings suggest that compensatory mechanisms in high-latitude populations enhance their performance at extreme temperatures, and consequently, that high-latitude populations of reef fishes will be less impacted by ocean warming than will low-latitude populations.     

Monitoring Herbivorous Fishes as Indicators of Coral Reef Resilience in American Samoa

Resilience-based management aims to promote or protect processes and species that underpin an ecosystem''s capacity to withstand and recover from disturbance. The management of ecological processes is a developing field that requires reliable indicators that can be monitored over time. Herbivory is a key ecological process on coral reefs, and pooling herbivorous fishes into functional groups based on their feeding mode is increasingly used as it may quantify herbivory in ways that indicate resilience. Here we evaluate whether the biomass estimates of these herbivore functional groups are good predictors of reef benthic assemblages, using data from 240 sites from five island groups in American Samoa. Using an information theoretic approach, we assembled a candidate set of linear and nonlinear models to identify the relations between benthic cover and total herbivore and non-herbivore biomass and the biomass of the aforementioned functional groups. For each benthic substrate type considered (encrusting algae, fleshy macroalgae, hard coral and turf algae), the biomass of herbivorous fishes were important explanatory variables in predicting benthic cover, whereas biomass of all fishes combined generally was not. Also, in all four cases, variation in cover was best explained by the biomass of specific functional groups rather than by all herbivores combined. Specifically: 1) macroalgal and turf algal cover decreased with increasing biomass of ‘grazers/detritivores’; and 2) cover of encrusting algae increased with increasing biomass of ‘grazers/detritivores’ and browsers. Furthermore, hard coral cover increased with the biomass of large excavators/bio-eroders (made up of large-bodied parrotfishes). Collectively, these findings emphasize the link between herbivorous fishes and the benthic community and demonstrate support for the use of functional groups of herbivores as indicators for resilience-based monitoring.     

Predators Exacerbate Competitive Interactions and Dominance Hierarchies between Two Coral Reef Fishes

Predation and competition are critical processes influencing the ecology of organisms, and can play an integral role in shaping coral reef fish communities. This study compared the relative and interacting effects of competition and predation on two competing species of coral reef fish, Pomacentrus amboinensis and P. moluccensis (Pomacentridae), using a multifactorial experiment. Fish were subjected to the sight and smell of a known predator (Pseudochromis fuscus), the presence of the heterospecific competitor (i.e., P. amboinensis vs. P. moluccensis), or a combination of the two for a period of 19 days. The sub-lethal effects of predator/competitor treatments were compared with controls; a combination of otolith microstructure analysis and observations were used to determine otolith growth patterns and behaviour. We predicted that the stress of competition and/or predation would result in strong sub-lethal impacts, and act synergistically on growth and behavioural patterns. We found strong evidence to support this prediction, but only for P. amboinensis, which suffered reductions in growth in both predator and competitor treatments, with the largest reductions occurring when subjected to both predation and competition concurrently. There was strong evidence of asymmetrical competition between the two damselfish species, with P. moluccensis as the dominant competitor, displaying strong aggressive behaviour towards P. amboinensis. Growth reductions for P. amboinensis in predator/competitor treatments appeared to come about primarily due to increases in shelter seeking behaviour, which significantly reduced the foraging rates of individuals compared with controls. These data highlight the importance of predator/competitor synergisms in influencing key behaviours and demographic parameters for juvenile coral reef fishes.     

Determining Methods of Underwater Visual Census for Estimating the Abundance of Coral Reef Fishes

We aimed to determine optimal methods of underwater visual census (UVC) for estimating the abundance of coral reef fishes exploited by fisheries in the tropical Pacific. Two main methods were tested using SCUBA: strip transect and stationary point counts. We assessed their relative accuracy, precision, power and efficiency (cost), and compared different census area dimensions, observer swimming speeds, and number of replicates. Twenty-five comparisons of the two UVC methods were conducted for 73 species from five families of coral reef fishes, on reefs in Australia and Fiji. Species were grouped within families based on their mobility. Few significant differences were found, either among or between strip transects and stationary point counts. The data were characterised by high variability, low precision and low power. A trend for greater accuracy in density estimates of small sedentary species with smaller census area was apparent, which probably reflects searching efficiency. Only one species group, the sedentary Acanthuridae showed differences between transects and point counts. Higher, hence presumably more accurate, density estimates were obtained with 50m×5m transects. Notably, point counts could be deployed in 70% of the time of transects. A bootstrapping procedure demonstrated a consistent improvement in precision of density estimates with increasing number of replicates, but no appreciable change in precision was found beyond 10 to 15 replicates, in all species groups and for both transects and point counts. Consequently, and because of the high variability inherent in fish density estimates, we recommend that at least 10 replicates be used to quantify the species considered here. The power calculations showed that only large differences in density will be detected with the replication levels typical of UVC surveys. Power was greatest for the roving serranids, with a decrease in density of about 50% detectable for this species group. We discuss the importance of carefully defining the behavioural attributes of species prior to selecting a UVC method. A method of temporal stratification in a count is described for censusing a range of species of varying mobilities. We suggest that fish mobility and search efficiency are key factors in optimising UVC methods.     

Body Fineness Ratio as a Predictor of Maximum Prolonged-Swimming Speed in Coral Reef Fishes

The ability to sustain high swimming speeds is believed to be an important factor affecting resource acquisition in fishes. While we have gained insights into how fin morphology and motion influences swimming performance in coral reef fishes, the role of other traits, such as body shape, remains poorly understood. We explore the ability of two mechanistic models of the causal relationship between body fineness ratio and endurance swimming-performance to predict maximum prolonged-swimming speed (U_max) among 84 fish species from the Great Barrier Reef, Australia. A drag model, based on semi-empirical data on the drag of rigid, submerged bodies of revolution, was applied to species that employ pectoral-fin propulsion with a rigid body at U_max. An alternative model, based on the results of computer simulations of optimal shape in self-propelled undulating bodies, was applied to the species that swim by body-caudal-fin propulsion at U_max. For pectoral-fin swimmers, U_max increased with fineness, and the rate of increase decreased with fineness, as predicted by the drag model. While the mechanistic and statistical models of the relationship between fineness and U_max were very similar, the mechanistic (and statistical) model explained only a small fraction of the variance in U_max. For body-caudal-fin swimmers, we found a non-linear relationship between fineness and U_max, which was largely negative over most of the range of fineness. This pattern fails to support either predictions from the computational models or standard functional interpretations of body shape variation in fishes. Our results suggest that the widespread hypothesis that a more optimal fineness increases endurance-swimming performance via reduced drag should be limited to fishes that swim with rigid bodies.     

Reef Fishes in Biodiversity Hotspots Are at Greatest Risk from Loss of Coral Species

Coral reef ecosystems are under a variety of threats from global change and anthropogenic disturbances that are reducing the number and type of coral species on reefs. Coral reefs support upwards of one third of all marine species of fish, so the loss of coral habitat may have substantial consequences to local fish diversity. We posit that the effects of habitat degradation will be most severe in coral regions with highest biodiversity of fishes due to greater specialization by fishes for particular coral habitats. Our novel approach to this important but untested hypothesis was to conduct the same field experiment at three geographic locations across the Indo-Pacific biodiversity gradient (Papua New Guinea; Great Barrier Reef, Australia; French Polynesia). Specifically, we experimentally explored whether the response of local fish communities to identical changes in diversity of habitat-providing corals was independent of the size of the regional species pool of fishes. We found that the proportional reduction (sensitivity) in fish biodiversity to loss of coral diversity was greater for regions with larger background species pools, reflecting variation in the degree of habitat specialization of fishes across the Indo-Pacific diversity gradient. This result implies that habitat-associated fish in diversity hotspots are at greater risk of local extinction to a given loss of habitat diversity compared to regions with lower species richness. This mechanism, related to the positive relationship between habitat specialization and regional biodiversity, and the elevated extinction risk this poses for biodiversity hotspots, may apply to species in other types of ecosystems.     

Patterns of Embryo Mortality in a Demersally Spawning Coral Reef Fish and the Role of Predatory Fishes

The biological significance of embryo mortality in demersally spawning coral reef fishes is poorly understood. Here we describe patterns of variation in embryo mortality in Pomacentrus amboinensis (Pomacentridae) at Lizard Island (Great Barrier Reef). The aim was to determine whether numbers of embryos hatched substantially differed from egg production, and if so, identify whether predatory fishes were a source of embryo mortality. Spawning success (number of eggs laid), embryo mortality (proportion of embryos that died prior to hatching) and number of embryos hatching were estimated from daily maps of clutches laid on artificial surfaces (PVC tiles) defended by nesting males. Patterns of variation in eggs laid, embryo mortality and numbers of embryos hatched were examined at three spatial scales: (1) among widely-spaced locations around the island; (2) between adjacent reef slope and patch reef habitats occupied by P. amboinensis at a single location; and (3) among different males within these two habitats. The embryo mortality was extremely high, with a mean of 25.9pm ± 6.2% (S.E.) for 4 locations examined in 1994 and a mean of 69.2pm ± 2.9% for two habitats surveyed in 1995. There were no significant differences in embryo mortality among locations or habitats in either year. This meant that spatial patterns in the number of embryos hatching reflected differences in the number of eggs laid on tiles. Embryo mortality was extremely variable on the scale of individual territories, with embryo mortality commonly ranging from 0% to 100%. Much of the mortality could be attributed to diurnal predatory fishes, especially the wrasse Thalassoma lunare. However, variation in predator densities did not explain spatial patterns in embryo mortality rates. Both solitary and group predatory behaviour was observed, with groups often causing 100% embryo mortality. The level of embryo mortality observed suggests that predation prior to hatching may have a substantial effect on the reproductive output of populations of this demersal-nesting fish.     

Comparative Phylogeography in Fijian Coral Reef Fishes: A Multi-Taxa Approach towards Marine Reserve Design

Delineating barriers to connectivity is important in marine reserve design as they describe the strength and number of connections among a reserve''s constituent parts, and ultimately help characterize the resilience of the system to perturbations at each node. Here we demonstrate the utility of multi-taxa phylogeography in the design of a system of marine protected areas within Fiji. Gathering mtDNA control region data from five species of coral reef fish in five genera and two families, we find a range of population structure patterns, from those experiencing little (Chrysiptera talboti, Halichoeres hortulanus, and Pomacentrus maafu), to moderate (Amphiprion barberi, Φ_st = 0.14 and Amblyglyphidodon orbicularis Φ_st = 0.05) barriers to dispersal. Furthermore estimates of gene flow over ecological time scales suggest species-specific, asymmetric migration among the regions within Fiji. The diversity among species-specific results underscores the limitations of generalizing from single-taxon studies, including the inability to differentiate between a species-specific result and a replication of concordant phylogeographic patterns, and suggests that greater taxonomic coverage results in greater resolution of community dynamics within Fiji. Our results indicate that the Fijian reefs should not be managed as a single unit, and that closely related species can express dramatically different levels of population connectivity.     

Population Dynamics and Spatial Distribution of Coral Reef Fishes: Comparison Between Continuous and Isolated Habitats

Recent studies have shown that there are high degrees of spatial and temporal stability in coral reef fish assemblage structures in a continuous habitat, in contrast to results of observations in isolated habitats. In order to determine the reason for the difference in temporal stability of fish assemblage structures in a continuous habitat site and an isolated habitat site, population dynamics and spatial distributions of coral reef fishes (six species of pomacentrids and two species of apogonids) in the two habitat site were investigated over a 2-year period in an Okinawan coral reef. The population densities of pomacentrid and apogonid species increased in juvenile settlement periods at both sites, but the magnitude of seasonal fluctuation in population density was significantly greater at the isolated habitat site, indicating that the rate of juvenile settlement and mortality rate in the isolated habitat were greater than those in the continuous habitat. The magnitude of aggregation of fishes, which affects density-dependent biological interactions that modify population density such as competition and predation, was also significantly greater at the isolated habitat site, especially in the juvenile settlement season. Most of the fishes at the isolated habitat site exhibited more generalized patterns of microhabitat selection because of less coral coverage and diversity. The seasonal stability in the species composition of fishes was greater at the continuous habitat site than that at the isolated habitat. Our findings suggest that the relative importance of various ecological factors responsible for regulation of the population density of coral reef fishes (e.g., competition, predation, microhabitat selection and post-settlement movement) in a continuous habitat site and the isolated habitat site are different.     

Morphology of the eye and visual acuities in the settlement-intervals of some Coral Reef Fishes (Labridae, Scaridae)

The morphology of the retina and photoreceptors of settlement-interval larvae and early juveniles and some adults of 12 species of Caribbean labrids and scarids were examined using histological techniques. The retinal structure is described in these species and life intervals. Larvae have a pure cone retina and unorganized mosaic and organization into a square mosaic pattern occurs during metamorphosis. Early post-settlement juveniles have an organized mosaic with structures that may enable them to detect polarized and UV light. Visual acuities were calculated for all species and life intervals and acuities ranged between 86.6–29.4min of arc in the settlement-intervals and 16–1.8min of arc in the adults. The visual abilities of the settlement-interval fishes and the possibility of the use of vision during settlement are discussed.     

Sporadic Disturbances in Fluctuating Coral Reef Environments: El Nino and Coral Reef Development in the Eastern Pacific

The disastrous effects of the intense 1982–83 El Niño-SouthernOscillation (ENSO) bring new insight into the long-term developmentof eastern Pacific coral reefs. The 1988–83 ENSO sea surfacewarming event caused extensive reef coral bleaching (loss ofsymbiotic zooxanthellae), resulting in up to 70–95% coralmortality on reefs in Costa Rica, Panama, Colombia and Ecuador.In the Galapagos Islands (Ecuador), most coral reefs experienced>95% coral mortality. Also, several coral species experiencedextreme reductions in population size, and local and regionalextinctions. The El Niño event spawned secondary disturbances,such as increased predation and bioerosion, that continue toimpact reef-building corals. The death of Pocillopora colonieswith their crustacean guards eliminated coral barriers now allowingthe corallivore Acanthaster planci access to formerly protectedcoral prey. Sea urchins and other organisms eroded disturbedcorals at rates that exceed carbonate production, potentiallyresulting in the elimination of existing reef buildups. In otherreefbuilding regions following extensive, catastrophic coralmortality, rapid recovery often occurs through the growth ofsurviving corals, recruitment of new corals from nearby sourcepopulations, and survival of consolidated reef surfaces. Inthe eastern Pacific, however, the return of upwelling conditionsand the survival of coral predators and bioeroders hamper coralreef recovery by reducing recruitment success and eroding coralreef substrates. Thus, coral reef growth that occurs betweendisturbance events is not conserved. Repeated El Niñodisturbances, which have occurred throughout the recent geologichistory of the eastern Pacific, prevent coral communities fromincreasing in diversity and limit the development and persistenceof significant reef features. The poor development of easternPacific coral reefs throughout Holocene and perhaps much ofPleistocene time may result from recurrent thermal disturbancesof the intensity of the 1982–83 El Niño event.     

Ecological Physiology of a Coral Pathogen and the Coral Reef Environment

Laboratory studies on the ecological physiology of a coral pathogen were carried out to investigate growth potential in terms of environmental factors that may control coral diseases on reefs. The disease chosen for this study, white plague type II, is considered to be one of the major diseases of Caribbean scleractinian corals, affecting a wide range of coral hosts and causing rapid and widespread tissue loss. It is caused by a single pathogen, the bacterium Aurantimonas coralicida. A series of laboratory experiments using a pure culture of the pathogen was carried out to examine the roles of temperature, pH, and O₂ concentration on growth rate. Results revealed optimal growth between 30 and 35°C, and between pH values of 6 and 8. There was a distinctive synergistic relationship between pH and temperature. Increasing temperature from 25 to 35°C expanded the range of pH tolerance from a minimum of 6.0 down to 5.0. O₂ concentration directly affected growth rate, which increased with increasing O₂. The combined effects of increasing O₂ and increasing temperature resulted in a synergistic effect of more rapid growth. These laboratory results are discussed in terms of the coral host and the range of the environmental factors that occur on coral reefs. We conclude that changing environmental conditions in the reef environment, in particular observed increases in water temperature, may be promoting coral diseases by allowing coral pathogens to expand their ecological niches. In the case of the white plague type II pathogen, elevated temperature would allow A. coralicida to colonize the low pH environment of the coral surface mucopolysaccharide layer as an initial stage of infection. The synergistic effect between temperature and oxygen concentration appeared to be less environmentally relevant for this coral pathogen.     

Coral Reef Communities as Prime Resources for Analysis of Evolution and Physiology of Behavior

The abundance and great diversity of life on coral reef ecosystemsprovides many good opportunities for studying the evolutionand specializations of neurophysiological systems and behavior.Crucial stages in the evolution of nervous systems appear tohave occurred in the Precambrian, as revealed in Ediacaran fossilsand their closest living relatives. By the Ordovician, whenChazy "reefs’ exemplify some of the earliest complex animalcommunities fixed in one place, more elaborate neurologicalmechanisms for orientation, predation, and escape reactionsare indicated. With the evolution offish, the behavioral richnessof reef communities became further enhanced. Elaborate specializationsof feeding, defensive, aggressive, signaling, schooling, andreproductive behaviors are common in fish. Several examplesof behavioral studies on reef organisms are used to illustrateresearch methodologies and the types of conclusions which maybe drawn. These examples include: (1) analysis of symbioticbehavior of an invertebrate and a vertebrate—sea anemoneand clownfish; (2) signaling behavior of a fish—the sailfishblenny; and (3) a combined electrophysiological and behavioralanalysis of orientation and feeding/attack behavior—sharks.An almost endless number of possibilities for similar analysismakes the organisms of coral reefs especially useful, and challenging,for teaching purposes as well as further research.     

A Morphospace for Reef Fishes: Elongation Is the Dominant Axis of Body Shape Evolution

Tropical reef fishes are widely regarded as being perhaps the most morphologically diverse vertebrate assemblage on earth, yet much remains to be discovered about the scope and patterns of this diversity. We created a morphospace of 2,939 species spanning 56 families of tropical Indo-Pacific reef fishes and established the primary axes of body shape variation, the phylogenetic consistency of these patterns, and whether dominant patterns of shape change can be accomplished by diverse underlying changes. Principal component analysis showed a major axis of shape variation that contrasts deep-bodied species with slender, elongate forms. Furthermore, using custom methods to compare the elongation vector (axis that maximizes elongation deformation) and the main vector of shape variation (first principal component) for each family in the morphospace, we showed that two thirds of the families diversify along an axis of body elongation. Finally, a comparative analysis using a principal coordinate analysis based on the angles among first principal component vectors of each family shape showed that families accomplish changes in elongation with a wide range of underlying modifications. Some groups such as Pomacentridae and Lethrinidae undergo decreases in body depth with proportional increases in all body regions, while other families show disproportionate changes in the length of the head (e.g., Labridae), the trunk or caudal region in all combinations (e.g., Pempheridae and Pinguipedidae). In conclusion, we found that evolutionary changes in body shape along an axis of elongation dominates diversification in reef fishes. Changes in shape on this axis are thought to have immediate implications for swimming performance, defense from gape limited predators, suction feeding performance and access to some highly specialized habitats. The morphological modifications that underlie changes in elongation are highly diverse, suggesting a role for a range of developmental processes and functional consequences.     

Sex, Symbiosis and Coral Reef Communities

SYNOPSIS. Questions about how today's corals and coral reefswill fare in a future that holds not only increasing directanthropogenic impacts, but also global change, cannot be satisfactorilyanswered if we do not understand the relations of corals andreef systems to today's environmental conditions. This paperdiscusses four aspects of modern reef biology: coral reproduction,coral population biology, the coral-zooxanthella symbiosis,and reef community ecology. Conclusions of this survey of currentknowledge are that complexities of cnidarian reproductive biology,and our rudimentary knowledge of reproductive patterns in reefcnidarians, make forecasting based on current knowledge uncertainat best; new discoveries about the coral algal symbiotic systemsuggest a possible mode of adjustment to environmental changethat warrants a strong research effort; coral communities ofthe future may well be unlike what we are familiar with today;and these new assemblages will be shaped by the interactionof novel environmental conditions and the characteristics ofindividual reef species.