Grazing pressure of herbivorous coral reef fishes on low coral-cover reefs

The impact of grazing by herbivorous fishes (Acanthuridae, Scaridae, and Pomacentridae) on low coral-cover reefs was assessed by measuring rates of benthic algal production and consumption on inshore and offshore reefs in the upper Florida Keys. Algal production rates, determined in situ with caged and uncaged experimental plates, were low (mean 1.05 g C m⁻² day⁻¹) and similar among reef types. Algal consumption rates were estimated using two different models, a detailed model incorporating fish bite rates and algal yield-per-bite for one species extrapolated to a guild-wide value, and a general regression relating fish biomass to algal consumption. Algal consumption differed among reef types: a majority of algal production was consumed on offshore reefs (55–100%), whereas consumption on inshore patch reefs was 31–51%. Spatial variation in algal consumption was driven by differences in herbivorous fish species composition, density, and size-structure among reef types. Algal consumption rates also varied temporally due to seasonal declines in bite rates and intermittent presence of large-bodied, vagile, schooling species. Spatial coherence of benthic community structure and temporal stability of algal turf over 3 years suggests that grazing intensity is currently sufficient to limit further spread of macroalgal cover on these low coral-cover reefs, but not to exclude it from the system.     

Varying responses of herbivorous and invertebrate-feeding fishes to macroalgal reduction on a coral reef

 The consequences of macroalgal overgrowth on reef fishes and means to reverse this condition have been little explored. An experimental reduction of macroalgae was conducted at a site in the Watamu Marine National Park in Kenya, where a documented increase in macroalgal cover has occurred over the last nine years. In four experimental 10 m by 10 m plots, macroalgae were greatly reduced (fleshy algal cover reduced by 84%) by scrubbing and shearing, while four similar plots acted as controls. The numerical abundance in all fish groups except wrasses and macroalgal-feeding parrotfishes (species in the genera Calotomus and Leptoscarus) increased in experimental algal reduction plots. Algal (Sargassum) and seagrass (Thalassia) assays, susceptible to scraping and excavating parrotfishes, were bitten more frequently in the algal reduction plots one month after the manipulation. Further, surgeonfish (Acanthurus leucosternon and A. nigrofuscus) foraging intensity increased in these algal reduction plots. The abundance of triggerfishes increased significantly in experimental plots relative to control plots, but densities remained low, and an index of sea urchin predation using tethered juvenile and adult Echinometra mathaei showed no differences between treatments following macroalgal reduction. Dominance of reefs by macrofleshy algae appears to reduce the abundance of fishes, mostly herbivores and their rates of herbivory, but also other groups such as predators of invertebrates (triggerfishes, butterflyfishes and angelfishes). Accepted: 2 February 1999     

Interactions between herbivorous fish guilds and their influence on algal succession on a coastal coral reef

Herbivory is an important mechanism affecting algal succession, particularly on coral reefs where the relationship between algae and corals is largely controlled by herbivores. However, different functional groups of herbivores may have contrasting effects on succession, which may explain different trajectories of coral reef recovery after disturbance. Here, the effects of different herbivore groups (roving herbivores = foragers and territorial damselfish = farmers) were isolated by a multi-factorial experiment carried out on a coastal coral reef with high macroalgal cover, high farmer densities and relatively low forager abundance. The effects of foragers and farmers were distinguished by monitoring algal succession on settlement tiles placed inside and outside exclusion cages, with orthogonal treatments established inside and outside damselfish territories (with appropriate cage controls). Within 12 months, algal assemblages on ungrazed tiles inside exclusion cages proceeded rapidly from fine filamentous turfs, to corticated algae, to tough erect (e.g. Amphiroa spp.) and foliose (e.g. Peyssonnellidae) calcified algae. Farmers had a dramatic impact on succession, essentially arresting the development of the algal community at a point where it was dominated by palatable filamentous algae of the genus Polysiphonia. Fleshy macroalgae such as Sargassum spp. were excluded from farmer territories. In contrast, foragers did not suppress fleshy macroalgae, but rather, appeared to decelerate succession and promote a relatively diverse assemblage. In contrast to forager-dominated reefs, farmer territories did not appear to function solely as forager exclusion areas or promote algal diversity as a result of intermediate grazing pressure. The relatively strong effects of farmers observed here may represent a future scenario for coral reefs that are increasingly subject to overfishing of large grazing fishes.     

The role of herbivorous fishes and urchins in coral reef communities

Synopsis Herbivorous fishes and invertebrates are conspicious elements of coral reef communities where they predominate both in numbers and biomass. Herbivores and the coral reef algae on which they feed represent a co-evolved system of defense and counter-defense. Algal species have developed toxic, structural, spatial and temporal defense or escape mechanisms, while the herbivores employ strategies that involve anatomical, physiological and behavioral adaptations. Current research demonstrates that many reef fishes are highly selective in the algae they consume. Food selection in these fishes may be correlated with their morphological and digestive capabilities to rupture algal cell walls. Sea urchins select more in accordance with relative abundance, although certain algal species are clearly avoided.The determinants of community structure on coral reefs have yet to be established but evidence indicates a strong influence by herbivores. Reef herbivores may reduce the abundance of certain competitively superior algae, thus allowing corals and cementing coralline algae to survive. We discuss how the foraging activities of tropical marine herbivores affect the distribution and abundance of algae and how these activities contribute to the development of coral reef structure and the fish assemblages which are intimately associated with reef structure.This paper forms a part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

Algae sediment dynamics are mediated by herbivorous fishes on a nearshore coral reef

Coral Reefs - Epilithic algae are a ubiquitous component of coral reefs. Components of the epilithic algal matrix (EAM) can have a significant influence on coral settlement and benthic feeding by...     

Global assessment of the status of coral reef herbivorous fishes: evidence for fishing effects

On coral reefs, herbivorous fishes consume benthic primary producers and regulate competition between fleshy algae and reef-building corals. Many of these species are also important fishery targets, yet little is known about their global status. Using a large-scale synthesis of peer-reviewed and unpublished data, we examine variability in abundance and biomass of herbivorous reef fishes and explore evidence for fishing impacts globally and within regions. We show that biomass is more than twice as high in locations not accessible to fisheries relative to fisheries-accessible locations. Although there are large biogeographic differences in total biomass, the effects of fishing are consistent in nearly all regions. We also show that exposure to fishing alters the structure of the herbivore community by disproportionately reducing biomass of large-bodied functional groups (scraper/excavators, browsers, grazer/detritivores), while increasing biomass and abundance of territorial algal-farming damselfishes (Pomacentridae). The browser functional group that consumes macroalgae and can help to prevent coral–macroalgal phase shifts appears to be most susceptible to fishing. This fishing down the herbivore guild probably alters the effectiveness of these fishes in regulating algal abundance on reefs. Finally, data from remote and unfished locations provide important baselines for setting management and conservation targets for this important group of fishes.     

Extinction vulnerability of coral reef fishes

With rapidly increasing rates of contemporary extinction, predicting extinction vulnerability and identifying how multiple stressors drive non-random species loss have become key challenges in ecology. These assessments are crucial for avoiding the loss of key functional groups that sustain ecosystem processes and services. We developed a novel predictive framework of species extinction vulnerability and applied it to coral reef fishes. Although relatively few coral reef fishes are at risk of global extinction from climate disturbances, a negative convex relationship between fish species locally vulnerable to climate change vs. fisheries exploitation indicates that the entire community is vulnerable on the many reefs where both stressors co-occur. Fishes involved in maintaining key ecosystem functions are more at risk from fishing than climate disturbances. This finding is encouraging as local and regional commitment to fisheries management action can maintain reef ecosystem functions pending progress towards the more complex global problem of stabilizing the climate.     

Choice of microhabitats by coral reef fishes at settlement

A set of small lagoonal patch reefs was searched every 1 to 3 days during the peak recruitment seasons of three summers and newly settled juvenile fishes were located. The majority of species remain rather sedentary during the first few days in the demersal environment, and we assumed that the site occupied was the site chosen at settlement. A series of characteristics of the occupied site were recorded, including percent cover of different types of substratum, and attributes related to the site's position on the patch reef. A set of null sites was randomly located on the same reefs for comparison with those selected by the fish. Sites chosen by individuals of eight common species were compared with these null sites, and sites chosen by fourteen species (including the eight) were compared with each other. Multiple discriminant analysis was used to assess the degree to which each species selected a unique type of site, and, for the eight species, the degree to which sites chosen by fish could be discriminated from randomly selected sites on the same patch reefs. Chosen sites were readily discriminated from null sites in seven of eight species, however the procedure was poor at discriminating among sites chosen by different species, and 8 pairs of species among the 14 chose sites which on average did not differ in the attributes measured. Attributes most important in discriminating sites chosen by each species are considered. Overall, the results indicate that while juvenile fish do not settle indiscriminantly onto lagoonal patch reefs, sites chosen by different species are often not very different from one another.     

The effect of coral morphology on shelter selection by coral reef fishes

While the loss of structural complexity causes declines in coral reef fish diversity, the processes leading to this decline are largely unexplained. To explore the role of coral morphology in providing shelter for fishes, tabular, branching and massive corals were filmed with video cameras and their usage by large reef fishes compared. Tabular corals were utilised more than the other two morphologies, with at least triple the abundance, biomass and residence times of large fishes. The preference of coral reef fishes for specific structural traits of tabular corals was also examined using artificial structural units. This experimental component showed that large reef fishes preferred opaque rather than translucent canopies. It appears that large fishes cue to tabular corals because of the concealment and/or shade provided. It is suggested that a loss of tabular corals as a result of climate change would have significant ecological impacts for the coral reef fishes that use these structures for shelter.     

Evaluation of algal regulation by herbivorous fishes on Caribbean coral reefs

The role of herbivorous fishes in maintaining low macroalgal cover was evaluated on coral reefs on several reef sites from Guadeloupe, either protected or not. Grazing by herbivorous fishes was assessed on different algal facies using fish-bite counts. Algal consumption by fish was estimated as well as algal production. Bite counts revealed that herbivorous fishes feed preferentially on algal turf and avoid brown macroalgae. The algal consumption varied between 0.4 and 2.8 g m⁻² days⁻¹ and was higher inside marine protected areas than outside. Comparison with algal production revealed that herbivorous fishes did not succeed in regulating algal growth. The insufficient number of grazers may lead to the dominance of stable assemblages of macroalgae on coral reefs, preventing the recovery of reef into previous coral-dominated ecosystems.     

Patterns and trends in coral reef macroalgae browsing: a review of browsing herbivorous fishes of the Indo-Pacific

Browsing fishes have been identified as an important component of coral reef resilience, because in contrast to other herbivorous fishes they are able to feed on established macroalgae. Climate change and local anthropogenic impacts have contributed to phase shifts in many coral reefs from coral to macroalgae dominance, and recent research suggests the potential ability of browsers to reverse such phase shifts. However, there is high variation among studies and some contradicting findings exist. Here, we review the relevant literature to assemble a list of species currently known to contribute to browsing in the Indo-Pacific. Furthermore, we identify spatial and temporal patterns, outline factors influencing browsing, and discuss the probability of phase shift reversal. We formulate research recommendations addressing the identified gaps in knowledge about the interactions of browsing fishes and their environment. To date, 37 species of fishes have been observed consuming macroalgae in the Indo-Pacific. The most important groups are the family Siganidae, the subfamily Scarinae (Labridae), and the subfamily Nasinae (Acanthuridae). Browsing species vary between studies depending on location, season and macroalgae species examined. Several influencing factors, such as structural complexity, palatability of macroalgae and ecosystem connectivity have been suggested to cause these discrepancies. The most promising avenues for future research are the effect of structural complexity, the importance of mobile link species and influences of food availability on the selectivity of browsing species. Increasing our knowledge in these fields will provide a better basis for successful management strategies directed at increasing the resilience of coral reefs.     

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.     

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.     

Effects of territorial damselfish on an algal-dominated coastal coral reef

Territorial damselfish are important herbivores on coral reefs because they can occupy a large proportion of the substratum and modify the benthic community to promote the cover of food algae. However, on coastal coral reefs damselfish occupy habitats that are often dominated by unpalatable macroalgae. The aim of this study was to examine whether damselfish can maintain distinctive algal assemblages on a coastal reef that is seasonally dominated by Sargassum (Magnetic Island, Great Barrier Reef). Here, three abundant species (Pomacentrus tripunctatus, P. wardi and Stegastes apicalis) occupied up to 60% of the reef substrata. All three species promoted the abundance of food algae in their territories. The magnitudes of the effects varied among reef zones, but patterns were relatively stable over time. Damselfish appear to readily co-exist with large unpalatable macroalgae as they can use it as a substratum for promoting the growth of palatable epiphytes. Damselfish territories represent patches of increased epiphyte load on macroalgae, decreased sediment cover, and enhanced cover of palatable algal turf.     

Nocturnal orientation to reefs by late pelagic stage coral reef fishes

 The nocturnal orientation behaviour of the late pelagic stages of two reef fish families (Apogonidae and Pomacentridae) was examined using behavioural cages deployed in the field. The behavioural cages enabled the fish to choose between swimming towards or away from the reef in response to natural cues. Overall, 55% of fish displayed a choice in the experiments, however, the proportion varied between the two families, with 67% of pomacentrids and 27% of apogonids displaying a choice. In both families, of the fish which displayed a choice, the proportion of fish swimming towards the reef was significantly greater than 50%, as random movement would predict (64% of pomacentrids and 67% of apogonids swam towards the reef). This proportion did not vary significantly among four field sites with different current regimes and geographic locations. The results suggest that the late pelagic stages of reef fish display nocturnal orientation behaviour, possibly in response to sound, which may aid in their settlement on reefs. Accepted: 18 August 1997     

Modelling larval dispersal and behaviour of coral reef fishes

Coral reef fish spend their first few weeks developing in the open ocean, where eggs and larvae appear merciless to tides and currents, before attempting to leave the pelagic zone and settle on a suitable reef. This pelagic dispersal phase is the process that determines population connectivity and allows replenishment of harvested populations across multiple coral reef habitats. Until recently this pelagic larval dispersal phase has been poorly understood and has often been referred to as the ‘black-box’ in the life-history of coral reef fishes. In this perspective article we highlight three areas where mathematical and computational approaches have been used to aid our understanding of this important ecological process. We discuss models that provide insights into the evolution of the pelagic larval phase in coral reef fish, an unresolved question which lends itself well to a modelling approach due to the difficulty in obtaining empirical data on this life history strategy. We describe how studies of fish hearing and physical sound propagation models can be used to predict the detection distance of reefs for settling larval fish, and the potential impact of anthropogenic noise. We explain how random walk models can be used to explore individual- and group-level behaviour in larval fish during the dispersal and settlement stage of their life-history. Finally, we discuss the mutual benefits that mathematical and computational approaches have brought to and gained from the field of larval behaviour and dispersal of reef fishes.     

Patch dynamics of coral reef macroalgae under chronic and acute disturbance

The patch dynamics (colonisation rate, growth rate, and extinction rate) are quantified for two dominant species of macroalgae on a Caribbean forereef in Belize: Lobophora variegata (Lamouroux) and Dictyota pulchella (Hörnig and Schnetter). Measurements were taken on time scales of days, weeks, months, and years during which three hurricanes occurred. All patches were followed on naturally occurring ramets of dead Montastraea annularis. The first hurricane (Mitch) caused massive coral mortality and liberated space for algal colonisation. The cover of Lobophora increased throughout the study and herbivores did not appear to limit its cover within a 4 year time frame. In contrast, the cover of D. pulchella fluctuated greatly and showed no net increase, despite an increase in parrotfish biomass and settlement space. Variation in the overall percent cover of an alga is not indicative of the underlying patch dynamics. The steady rise in the cover of Lobophora took place despite a high turnover of patches (12–60% of patches per year). The patch dynamics of Dictyota were slower (7–20%), but a greater patch density and threefold higher lateral growth rate led to greater fluctuations in total cover. The dynamics of algal patches are size-specific such that larger patches are less likely to become extinct during hurricanes.     

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.