Predation on juvenile coral reef fishes: an exclusion experiment

The densities of recruits on caged and uncaged areas were compared in an experiment done to show the extent of predation on recently metamorphosed coral reef fishes. The design was unlike typical caging experiments, however, in that areas were caged only for short periods of 20–30 days and several independent trials, testing the same null hypothesis, were run. This was done to avoid confounding the effects of excluding herbivorous fishes with the effects of excluding piscivorous fishes. A third treatment, partially-meshed cages, revealed that the experiment was complicated by several other factors. Some prey species were attracted to the high relief offered by the experimental structures. Others responded to the differences in shelter from predators by redispersing themselves among the treatments shortly after settlement. There was also at least one significant edge effect caused by fishes preferring to settle near the boundaries of all treatments. In spite of these difficulties, observations on known individuals revealed that rates of mortality were age-dependent and decreased rapidly after metamorphosis. More than 25% of such fishes disappeared during their first five days in the benthic habitat compared with >10% of fishes aged 6–10 days and no losses of fishes aged 11–15 days. These early losses are the greatest instantaneous rates of mortality yet documented for recruited reef fishes. The experiment also suggested different rates of early mortality for various groupings of species: individuals of solitary, sedentary species disappeared approximately half as fast as individuals of the more mobile, and the more gregarious, species. This is probably a true reflection of the different vulnerability of these groups to predation and it may be caused by the different ways in which these fishes use the coral substratum. Our experience suggests that caging artifacts can have major impacts on the results obtained from this type of experiment and they must be controlled for adequately. We conclude that studies of predation on reef fishes may be done more easily using other methods.     

Direct evaluation of macroalgal removal by herbivorous coral reef fishes

Few studies have examined the relative functional impacts of individual herbivorous fish species on coral reef ecosystem processes in the Indo-Pacific. This study assessed the potential grazing impact of individual species within an inshore herbivorous reef fish assemblage on the central Great Barrier Reef (GBR), by determining which fish species were able to remove particular macroalgal species. Transplanted multiple-choice algal assays and remote stationary underwater digital video cameras were used to quantify the impact of local herbivorous reef fish species on 12 species of macroalgae. Macroalgal removal by the fishes was rapid. Within 3 h of exposure to herbivorous reef fishes there was significant evidence of intense grazing. After 12 h of exposure, 10 of the 12 macroalgal species had decreased to less than 15% of their original mass. Chlorodesmis fastigiata (Chlorophyta) and Galaxaura sp. (Rhodophyta) showed significantly less susceptibility to herbivorous reef fish grazing than all other macroalgae, even after 24 h exposure. Six herbivorous and/or nominally herbivorous reef fish species were identified as the dominant grazers of macroalgae: Siganus doliatus, Siganus canaliculatus, Chlorurus microrhinos, Hipposcarus longiceps, Scarus rivulatus and Pomacanthus sexstriatus. The siganid S. doliatus fed heavily on Hypnea sp., while S. canaliculatus fed intensively on Sargassum sp. Variation in macroalgal susceptibility was not clearly correlated with morphological and/or chemical defenses that have been previously suggested as deterrents against herbivory. Nevertheless, the results stress the potential importance of individual herbivorous reef fish species in removing macroalgae from coral reefs.     

The ontogeny of home ranges: evidence from coral reef fishes

The concept of home ranges is fundamental to ecology. Numerous studies have quantified how home ranges scale with body size across taxa. However, these relationships are not always applicable intraspecifically. Here, we describe how the home range of an important group of reef fish, the parrotfishes, scales with body mass. With masses spanning five orders of magnitude, from the early postsettlement stage through to adulthood, we find no evidence of a response to predation risk, dietary shifts or sex change on home range expansion rates. Instead, we document a distinct ontogenetic shift in home range expansion with sexual maturity. Juvenile parrotfishes displayed rapid home range growth until reaching approximately 100–150 mm length. Thereafter, the relationship between home range and mass broke down. This shift reflected changes in colour patterns, social status and reproductive behaviour associated with the transition to adult stages. While there is a clear relationship between body mass and home ranges among adult individuals of different species, it does not appear to be applicable to size changes within species. Ontogenetic changes in parrotfishes do not follow expected mass–area scaling relationships.     

New aspects of sex change among reef fishes: recent studies in Japan

New aspects of sex change in reef fishes are reviewed with special emphasis on recent studies in Japan. For protogyny, studies on both monandric and diandric species have been conducted, but the distinction of primary males from prematurational secondary males seems to need further examination. For protandry, detailed field studies on anemonefishes have revealed alternative life-history styles associated with movements between hosts before or after maturation. The most interesting new aspect has been the discovery of 2-way sex change within a species. Conditions for evolution of 2-way sex change are examined in relation to the size-advantage model and social control mechanisms. A fish may change sex when it becomes dominant in a mating group, but a dominant fish may also change sex in the reverse direction when its social status changes to subordinate through inter-group movement. Two-way sex change has hitherto been reported only from basically protogynous fishes (e.g., Gobiidae, Pomacanthidae, Cirrhitidae, Epinephelinae). Possibilities of the reverse sex change in the protandrous anemonefishes are discussed with data from some unpublished studies.     

Phylogeography and the conservation of coral reef fishes

Here we present a review of how the study of the geographic distribution of genetic lineages (phylogeography) has helped identify management units, evolutionary significant units, cryptic species, and areas of endemism, and how this information can help efforts to achieve effective conservation of coral reefs. These studies have confirmed the major biogeographic barriers that were originally identified by tropical species distributions. Ancient separations, identified primarily with mtDNA sequence comparisons, became apparent between populations on each side of the barriers. The general lack of correlation between pelagic larval duration and genetic connectivity across barriers indicates that life history and ecology can be as influential as oceanography and geography in shaping evolutionary partitions within ocean basins. Hence, conservation strategies require a recognition of ecological hotspots, those areas where habitat heterogeneity promotes speciation, in addition to more traditional approaches based on biogeography. Finally, the emerging field of genomics will add a new dimension to phylogeography, allowing the study of genes that are pertinent to recent and ongoing differentiation, and ultimately providing higher resolution to detect evolutionary significant units that have diverged in an ecological time scale.     

Growth impacts in a changing ocean: insights from two coral reef fishes in an extreme environment

Coral Reefs - Determining the life-history consequences for fishes living in extreme and variable environments will be vital in predicting the likely impacts of ongoing climate change on reef fish...     

The stability of P in coral reef fishes

The constancy of phenotypic variation and covariation is an assumption that underlies most recent investigations of past selective regimes and attempts to predict future responses to selection. Few studies have tested this assumption of constancy despite good reasons to expect that the pattern of phenotypic variation and covariation may vary in space and time. We compared phenotypic variance-covariance matrices (P) estimated for populations of six species of distantly related coral reef fishes sampled at two locations on Australia's Great Barrier Reef separated by more than 1000 km. The intraspecific similarity between these matrices was estimated using two methods: matrix correlation and common principal component analysis. Although there was no evidence of equality between pairs of P, both statistical approaches indicated a high degree of similarity in morphology between the two populations for each species. In general, the hierarchical decomposition of the variance-covariance structure of these populations indicated that all principal components of phenotypic variance-covariance were shared but that they differed in the degree of variation associated with each of these components. The consistency of this pattern is remarkable given the diversity of morphologies and life histories encompassed by these species. Although some phenotypic instability was indicated, these results were consistent with a generally conserved pattern of multivariate selection between populations.     

Hybridization in coral reef fishes: introgression and bi-directional gene exchange in Thalassoma (family Labridae)

Hybrids in coral reef fishes have traditionally been described based on external features using meristic characters and colouration to identify putative parental contributors. This study utilised molecular genetic techniques to verify hybrid status and identify putative parental species for five hybrid specimens (Labridae: Thalassoma) collected from Holmes Reef in the Coral Sea. Phylogenetic analyses support hybrid origins of the specimens. Mitochondrial COI gene, nuclear S7 (intron 1) and nuclear copy of mitochondrial (NUMT) D-loop region corroborate the identity of T. quinquevittatum as the maternal and T. jansenii as the paternal contributor. Backcrossing to parental species by hybrids and bi-directional gene exchange between the Holmes Reef populations of T. jansenii and T. quinquevittatum was detected, suggesting that hybrids are fertile and able to reproduce successfully. F(1) hybrids display a mixture of the colouration attributes of the two parental species, but subsequent backcrossed individuals were unrecognisable as hybrids and displayed colouration of either parental species. A large numerical imbalance exists between the putative parental species at Holmes Reef, with T. quinquevittatum outnumbering T. jansenii by approximately 25:1. In this case study, hybridization appears to be driven by ecological rather than evolutionary factors.     

Social determination of sex in reef fishes

Many fishes living in reef environments display remarkable flexibility in sexuality with social interactions determining their sex either during juvenile development or in adulthood. The evolutionary advantages of such flexibility are relatively well established. By contrast, the mechanisms by which social cues guide development of the sexual phenotype are less well understood. This paper reviews our understanding of these processes for some well-studied reef fish groups at the gonadal and neuroendocrine levels as well as proposing promising directions for future study.     

Hypoxia in paradise: widespread hypoxia tolerance in coral reef fishes

Using respirometry, we examined the hypoxia tolerance of 31 teleost fish species (seven families) inhabiting coral reefs at a 2-5 m depth in the lagoon at Lizard Island (Great Barrier Reef, Australia). All fishes studied maintained their rate of oxygen consumption down to relatively severe hypoxia (20-30% air saturation). Indeed, most fishes appeared unaffected by hypoxia until the oxygen level fell below 10% of air saturation. This, hitherto unrecognized, hypoxia tolerance among coral reef fishes could reflect adaptations to nocturnal hypoxia in tide pools. It may also be needed to enable fishes to reside deep within branching coral at night to avoid predation. Widespread hypoxia tolerance in a habitat with such an extreme biodiversity as coral reefs indicate that there is a wealth of hypoxia related adaptations to be discovered in reef fishes.     

Otolith microstructure and the demography of coral reef fishes

The ecology of planktonic larvae - particularly the extent to which larvae are transported between habitat patches - is a poorly known aspect of the life history of tropical reef-associated fishes. The logistic difficulties of studying larvae in the field and of relating their behavior in captivity to field conditions have greatly impeded the progress of research. However, a recently developed technique, based on analysis of the microstructural growth elements in otoliths ('ear stones'), may prove a powerful means of assessing the early life history of reef fishes and relating it to population and community dynamics.     

On the spatial scale of dispersal in coral reef fishes

Marine biologists have gone through a paradigm shift, from the assumption that marine populations are largely ‘open’ owing to extensive larval dispersal to the realization that marine dispersal is ‘more restricted than previously thought’. Yet, population genetic studies often reveal low levels of genetic structure across large geographic areas. On the other side, more direct approaches such as mark‐recapture provide evidence of localized dispersal. To what extent can direct and indirect studies of marine dispersal be reconciled? One approach consists in applying genetic methods that have been validated with direct estimates of dispersal. Here, we use such an approach—genetic isolation by distance between individuals in continuous populations—to estimate the spatial scale of dispersal in five species of coral reef fish presenting low levels of genetic structure across the Caribbean. Individuals were sampled continuously along a 220‐km transect following the Mesoamerican Barrier Reef, population densities were estimated from surveys covering 17 200 m² of reef, and samples were genotyped at a total of 58 microsatellite loci. A small but positive isolation‐by‐distance slope was observed in the five species, providing mean parent‐offspring dispersal estimates ranging between 7 and 42 km (CI 1–113 km) and suggesting that there might be a correlation between minimum/maximum pelagic larval duration and dispersal in coral reef fishes. Coalescent‐based simulations indicate that these results are robust to a variety of dispersal distributions and sampling designs. We conclude that low levels of genetic structure across large geographic areas are not necessarily indicative of extensive dispersal at ecological timescales.     

Community assembly and diversification in Indo-Pacific coral reef fishes

Theories of species coexistence have played a central role in ecology and evolutionary studies of the origin and maintenance of biodiversity in highly diverse communities. The concept of niche and associated theories predict that competition for available ecological space leads to a ceiling in species richness that influences further diversification patterns. By contrast, the neutral theory supports that speciation is stochastic and diversity independent. We examined the phylogenetic community structure and diversification rates in three families and 14 sites within coral reef fish communities from the Indian and Pacific oceans. Using the phylogenetic relationships among 157 species estimated with 2300 bp of mitochondrial DNA, we tested predictions in terms of species coexistence from the neutral and niche theories. At the regional scale, our findings suggest that phylogenetic community structure shifts during community assembly to a pattern of dispersion as a consequence of allopatric speciation in recent times but overall, variations in diversification rates did not relate with sea level changes. At the local scale, the phylogenetic community structure is consistent with a neutral model of community assembly since no departure from a random sorting of species was observed. The present results support a neutral model of community assembly as a consequence of the stochastic and unpredictable nature of coral reefs favoring generalist and sedentary species competing for living space rather than trophic resources. As a consequence, the observed decrease in diversification rates may be seen as the result of a limited supply of living space as expected in a finite island model.     

Auditory sensitivity in settlement-stage larvae of coral reef fishes

The larval phase of most species of coral reef fishes is spent away from the reef in the pelagic environment. At the time of settlement, these larvae need to locate a reef, and recent research indicates that sound emanating from reefs may act as a cue to guide them. Here, the auditory abilities of settlement-stage larvae of four species of coral reef fishes (families Pomacentridae, Lutjanidae and Serranidae) and similar-sized individuals of two pelagic species (Carangidae) were tested using an electrophysiological technique, auditory brainstem response (ABR). Five of the six species heard frequencies in the 100–2,000 Hz range, whilst one carangid species did not detect frequencies higher than 800 Hz. The audiograms of the six species were of similar shape, with best hearing at lower frequencies between 100 and 300 Hz. Strong within-species differences were found in hearing sensitivity both among the coral reef species and among the pelagic species. Larvae of the coral reef species had significantly more sensitive hearing than the larvae of the pelagic species. The results suggest that settlement-stage larval reef fishes may be able to detect reef sounds at distances of a few 100 m. If true hearing thresholds are lower than ABR estimates, as indicated in some comparisons of ABR and behavioural methods, the detection distances would be much larger.     

Zoogeography of the coral reef fishes of the Socotra Archipelago

Fish communities and habitats were studied at the Socotra archipelago (Gulf of Aden, ≈12°N 54°E). Extensive and unexpected hermatypic coral communities were recorded, at the centre of a 2200 km gap in knowledge of species and habitat distributions which coincides with a change from a western Indian Ocean coral reef fauna to an Arabian one. The fish assemblage associated with the Socotra archipelago corals is predominantly south Arabian. An east African influence, minimal on the mainland coasts of Arabia, is more evident here, and results in previously unrecorded sympatry between Arabian endemic species and their Indian Ocean sister taxa. A study of distributions of Chaetodontidae (butterflyfishes) in the north-western Indian Ocean reveals a number of distinct patterns, with a trend for species replacement along a track from the northern Red Sea to the Indian Ocean. A major feature of the reef fish zoogeography of the region is found to be a distinct south Arabian area, characterized by a 'pseudo-high latitude effect' which results from seasonal cold water upwelling along the Arabian sea coasts of Yemen and Oman and the Indian Ocean coast of Somalia. This south Arabian feature is consistent across a wide range of fish families. It is most pronounced in Oman and Yemen, and although it is the dominant influence at Socotra it is slightly 'diluted' here by the east African influence. The south Arabian area wholly or partly accounts for most of the major marine zoogeographic features around Arabia, and is the principal feature fragmenting Arabian coastal fish assemblages, and separating them from those of the wider Indo-west Pacific.