Habitat choice,recruitment and the response of coral reef fishes to coral degradation

The global degradation of coral reefs is having profound effects on the structure and species richness of associated reef fish assemblages. Historically, variation in the composition of fish communities has largely been attributed to factors affecting settlement of reef fish larvae. However, the mechanisms that determine how fish settlers respond to different stages of coral stress and the extent of coral loss on fish settlement are poorly understood. Here, we examined the effects of habitat degradation on fish settlement using a two-stage experimental approach. First, we employed laboratory choice experiments to test how settlers responded to early and terminal stages of coral degradation. We then quantified the settlement response of the whole reef fish assemblage in a field perturbation experiment. The laboratory choice experiments tested how juveniles from nine common Indo-Pacific fishes chose among live colonies, partially degraded colonies, and dead colonies with recent algal growth. Many species did not distinguish between live and partially degraded colonies, suggesting settlement patterns are resilient to the early stages of declining coral health. Several species preferred live or degraded corals, and none preferred to associate with dead, algal-covered colonies. In the field experiment, fish recruitment to coral colonies was monitored before and after the introduction of a coral predator (the crown-of-thorns starfish) and compared with undisturbed control colonies. Starfish reduced live coral cover by 95–100%, causing persistent negative effects on the recruitment of coral-associated fishes. Rapid reductions in new recruit abundance, greater numbers of unoccupied colonies and a shift in the recruit community structure from one dominated by coral-associated fishes before degradation to one predominantly composed of algal-associated fish species were observed. Our results suggest that while resistant to coral stress, coral death alters the process of replenishment of coral reef fish communities.     

SMURFs: standard monitoring units for the recruitment of temperate reef fishes

I evaluated a standard monitoring unit for the recruitment of reef fishes (SMURF) as a tool for ascertaining spatial and temporal patterns of reef fish recruitment in central California, USA. SMURFs consisted of a 1.0×0.35 m dia. cylinder of fine mesh plastic grid that contained a folded section of larger mesh plastic grid. SMURFs collected new recruits of 20 species of fish with 92% of the individuals collected from 10 species, mostly rockfish (genus Sebastes). An experiment varying depth of SMURFs in the water column (surface, mid-depth, or bottom) showed that surface SMURFs collected the greatest diversity of species and significantly greater abundance for eight species, with two species having significantly greater abundance on mid-depth SMURFs and three species having significantly greater abundance on bottom SMURFs. A comparison of cumulated recruitment from SMURFs that varied in sampling frequency (removal of new recruits every 1-3, 7, or 28 days) suggested that increasing the time between sampling caused a significant decrease in recruitment estimates for some species but not for others. To determine how well temporal patterns of recruitment to SMURFs reflected patterns to nearby reefs, I compared within season temporal patterns of recruitment to SMURFs with that at nearby reefs, estimated by visual transect surveys conducted on scuba. Temporal patterns of recruitment to SMURFs were significantly and positively related to early recruitment on reefs for one group of benthic-algal associated rockfish species when diver surveys were lagged by 30 days (r=0.87) and for another group of canopy-algal associated rockfish species when lagged by 5 days (r=0.72). SMURFs appeared to be an effective and efficient method for indexing relative rates of delivery of competent juveniles for many temperate nearshore reef fishes.     

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     

Ecological structure of assemblages of coral reef fishes on isolated patch reefs

A 9-year study of the structure of assemblages of fish on 20 coral patch reefs, based on 20 non-manipulative censuses, revealed a total of 141 species from 34 families, although 40 species accounted for over 95% of sightings of fish. The average patch reef was 8.5 m² in surface area, and supported 125 fish of 20 species at a census. All reefs showed at least a two-fold variation among censuses in total numbers of fish present, and 12 showed ten-fold variations. There was also substantial variation in the composition and relative abundances of species present on each patch reef, such that censuses of a single patch reef were on average about 50% different from each other in percent similarity of species composition (Czekanowski's index). Species differed substantially in the degree to which their numbers varied from census to census, and in the degree to which their dispersion among patch reefs was modified from census to census. We characterize the 40 most common species with respect to these attributes. The variations in assemblage structure cannot be attributed to responses of fish to a changing physical structure of patch reefs, nor to the comings and goings of numerous rare species. Our results support and extend earlier reports on this study, which have stressed the lack of persistant structure for assemblages on these patch reefs. While reef fishes clearly have microhabitat preferences which are expressed at settlement, the variations in microhabitat offered by the patch reefs are insufficient to segregate many species of fish by patch reef. Instead, at the scale of single patch reefs, and, to a degree, at the larger scale of the 20 patch reefs, most of the 141 species of fish are distributed without regard to differences in habitat structure among reefs, and patterns of distribution change over time. Implications for general understanding of assemblage dynamics for fish over more extensive patches of reef habitat are considered.     

Growth of reef fishes in response to live coral cover

Although the global decline in coral reef health is likely to have profound effects on reef associated fishes, these effects are poorly understood. While declining coral cover can reduce the abundance of reef fishes through direct effects on recruitment and/or mortality, recent evidence suggests that individuals may survive in disturbed habitats, but may experience sublethal reductions in their condition. This study examined the response of 2 coral associated damselfishes (Pomacentridae), Chrysiptera parasema and Dascyllus melanurus, to varying levels of live coral cover. Growth, persistence, and the condition of individuals were quantified on replicate coral colonies in 3 coral treatments: 100% live coral (control), 50% live coral (partial) and 0% live coral (dead). The growth rates of both species were directly related to the percentage live coral cover, with individuals associated with dead corals exhibiting the slowest growth, and highest growth on control corals. Such differences in individual growth between treatments were apparent after 29 d. There was no significant difference in the numbers of fishes persisting or the physiological condition of individuals between different treatments on this time-scale. Slower growth in disturbed habitats will delay the onset of maturity, reduce lifetime fecundity and increase individual's vulnerability to gape-limited predation. Hence, immediate effects on recruitment and survival may underestimate the longer-term impacts of declining coral on the structure and diversity of coral-associated reef fish communities.     

Nocturnal relocation of adult and juvenile coral reef fishes in response to reef noise

Juvenile and adult reef fishes often undergo migration, ontogenic habitat shifts, and nocturnal foraging movements. The orientation cues used for these behaviours are largely unknown. In this study, the use of sound as an orientation cue guiding the nocturnal movements of adult and juvenile reef fishes at Lizard Island, Great Barrier Reef was examined. The first experiment compared the movements of fishes to small patch reefs where reef noise was broadcast, with those to silent reefs. No significant responses were found in the 79 adults that were collected, but the 166 juveniles collected showed an increased diversity each morning on the reefs with broadcast noise, and significantly greater numbers of juveniles from three taxa (Apogonidae, Gobiidae and Pinguipedidae) were collected from reefs with broadcast noise. The second experiment compared the movement of adult and juvenile fishes to reefs broadcasting high (>570 Hz), or low (<570 Hz) frequency reef noise, or to silent reefs. Of the 122 adults collected, the highest diversity was seen at the low frequency reefs; and adults from two families (Gobiidae and Blenniidae) preferred these reefs. A similar trend was observed in the 372 juveniles collected, with higher diversity at the reefs with low frequency noises. This preference was seen in the juvenile apogonids; however, juvenile gobiids were attracted to both high and low sound treatments equally, and juvenile stage Acanthuridae preferred the high frequency noises. This evidence that juvenile and adult reef fishes orientate with respect to the soundscape raises important issues for management, conservation and the protection of sound cues used in natural behaviour.     

Interpreting coral reef monitoring data: A guide for improved management decisions

Coral reef monitoring programmes exist in all regions of the world, recording reef attributes such as coral cover, fish biomass and macroalgal cover. Given the cost of such monitoring programs, and the degraded state of many of the world’s reefs, understanding how reef monitoring data can be used to shape management decisions for coral reefs is a high priority. However, there is no general guide to understanding the ecological implications of the data in a format that can trigger a management response. We attempt to provide such a guide for interpreting the temporal trends in 41 coral reef monitoring attributes, recorded by seven of the largest reef monitoring programmes. We show that only a small subset of these attributes is required to identify the stressors that have impacted a reef (i.e. provide a diagnosis), as well as to estimate the likely recovery potential (prognosis). Two of the most useful indicators, turf algal canopy height and coral colony growth rate are not commonly measured, and we strongly recommend their inclusion in reef monitoring. The diagnosis and prognosis system that we have developed may help guide management actions and provides a foundation for further development as biological and ecological insights continue to grow.     

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.     

How does shoreline development impact the recruitment patterns of coral reef fish juveniles (Moorea Island,French Polynesia)?

The study examined the effects of coastal embankment building on fish recruitment in three habitat types (beach-rock, white sand and muddy sand) in the near shore and fringing reef habitats of Moorea lagoon (French Polynesia). The results showed a positive relationship between the presence of embankments and the density and species richness of juvenile fish along the shoreline (whatever the habitat types). However, embankments deteriorated adjacent fringing reefs (decrease of live coral), which led to a decrease of fish density on beach-rock and white sand sites, and a decrease of fish species richness on muddy sand sites.     

Extended breeding and recruitment periods of fishes on a low latitude coral reef

The temporal dynamics of fish recruitment to equatorial Indo-Pacific coral reefs are not well known. This paper documents fish recruitment over a 2.5-year period in Kimbe Bay (PNG) and shows that it is much less seasonal than is typically described for higher latitude coral reefs. Two families, wrasses (Labridae) and damselfishes (Pomacentridae), which accounted for 90% of all non-cryptic reef fish settlers, exhibited contrasting patterns. Most wrasse species had year-round recruitment with irregular peaks in abundance between November and May. Damselfish species showed a wider range of recruitment patterns, but most had negligible recruitment during the wet season (December–February), followed by one or two recruitment peaks between May and November. Species with longer seasonal recruitment periods exhibited higher cumulative levels of recruitment. For three focal damselfish species, reproductive output was reduced during the wet season, but this alone did not account for the low recruitment at this time. The lack of damselfish recruitment during the wet season is hypothesised to be due to a combination of reduced reproductive output and increased larval mortality associated with monsoonal conditions. The results indicate that there are consistent family-wide recruitment strategies that may play a significant role in the dynamics of populations in equatorial waters.     

An appraisal of methods used in coral recruitment studies

 A new method for attaching individual artificial settlement plates directly to the reef surface using small stainless steel base plates is described. Recruitment of corals to settlement plates attached to the reef substratum and to steel mesh racks is compared. The effects of differences in depth, settlement plate angle, and local topography on recruitment of corals were also investigated. No significant difference in mean recruit density was found between settlement plates deployed using the two attachment methods. Small differences in depth and plate angle among replicate plates explained less than 6% of the variability in coral recruitment on replicate settlement plates. The direct-attachment method is less obtrusive, more cost and time efficient, and settlement plates can be deployed at precise locations. Additionally, because settlement plates are deployed individually rather than grouped on racks or frames, the direct-attachment method avoids complications associated with assumptions of independence implicit in most statistical procedures. Accepted: 24 November 1999     

Evidence of density-independent mortality in a settling coral reef damselfish, Chromis viridis

To know if the variation in the number of settling fish larvae can be dampened by density-dependent postsettlement mortality, we investigated the relationship between settler density and predator-induced mortality of a coral reef damselfish, Chromis viridis. Totals of 2, 3, 5, 8, 10, 12, 14, 16, 18, and 20 fish of 10 or 20 mm total length were released in experimental cages enclosing a coral head of Porites rus (to provide settlement habitat) and five predators. The results showed that the mortality rate of both 10- and 20-mm fish was density independent.     

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.     

Impact of micropredatory gnathiid isopods on young coral reef fishes

The ecological role of parasites in the early life-history stages of coral reef fish, and whether this varies between fish with and without a pelagic phase, was investigated. The susceptibility to, and effect of reef-based micropredatory gnathiid isopods on larval, recently settled, and juvenile fishes was tested using two damselfishes (Pomacentridae): Neopomacentrus azysron, which has pelagic larvae, and Acanthochromis polyacanthus, which does not. When larval and recently settled stages of N. azysron and very young A. polyacanthus juveniles (smaller than larval N. azysron) were exposed to one or three gnathiids, the proportion of infections did not vary significantly among the three host types or between the number of gnathiids to which the fish were exposed. The overall infection was 35%. Mortality, however, differed among the three gnathiid-exposed host types with most deaths occurring in larval N. azysron; no mortalities occurred for recently settled N. azysron exposed to one or three gnathiids, and A. polyacanthus exposed to one gnathiid. Mortality did not differ significantly between larval N. azysron and A. polyacanthus juveniles, failing to provide support for the hypothesis that reef-based A. polyacanthus juveniles are better adapted to gnathiid attack than fish with a pelagic phase. The study suggests that settling on the reef exposes young fish to potentially deadly micropredators. This supports the idea that the pelagic phase may allow young fish to avoid reef-based parasites.     

Diel variability in counts of reef fishes and its implications for monitoring

Studies of reef fish assemblages in space rarely consider the effects of temporal variability on spatial comparisons, and when they do, usually examine timescales of months to years. The nature of fish monitoring surveys is such that particular locations may be surveyed at one time of day, and surveys designed to establish the degree of spatial variability in assemblages may be confounded if the order of sampling within treatments is not randomised with respect to time of day. In this study, we tested the degree of temporal variability in temperate reef fish counts at the same sites in New Zealand and Italy, within and between days. Repeated counts separated by months returned quite different assemblages, whereas counts separated by days did not. Although overall assemblage structure did not generally differ significantly with time of day, at some locations there was evidence of changes in the relative density of certain species in the afternoon relative to the morning. Care must be taken not to introduce systematic bias to spatial comparisons due to behavioural variability in fishes at differing times of day.     

Effects of microhabitat characteristics on the settlement and recruitment of a coral reef fish at two spatial scales

Populations of fishes on coral reefs are replenished by the settlement of pelagic larvae to demersal populations. Recruitment varies spatially and temporally and can exert strong effects on the dynamics of reef fish populations. This study examined the effect of microhabitat characteristics on small-scale and large-scale recruitment variation in the three-spot damselfish, Stegastes planifrons (Cuvier). Comparison of 0.25-m² quadrats occupied by three-spots with randomly sampled null quadrats showed that three-spots quadrats contained a higher percent cover of the coral Montastrea annularis than would be expected at random. Manipulative experiments on three types of 1.0-m² patch reefs (living M. annularis, dead Porites Porites and dead Acropora palmata) patch reefs on showed that this non-random distribution was established by microhabitat choice during settlement and not by differential post-settlement survival. The presence of conspecific juveniles did not affect settlement. Recruitment was monitored at nine sites on three islands over 3 years. Recruitment showed no consistent pattern in the relative levels of recruitment among sites. Similarly, no consistent relationship emerged between recruitment levels and microhabitat characteristics at the nine sites. For example, at this large scale, the percent cover of M. annularis explained variation in recruitment in only 1 out of 3 years. These results suggest that small-scale recruitment patterns are influenced by microhabitat choice during settlement, but that these habitat effects do not scale up to influence large-scale variation in recruitment.     

Indices for assessing coral reef fish biodiversity: the need for a change in habits

We present the first representative and quantified overview of the indices used worldwide for assessing the biodiversity of coral reef fishes. On this basis, we discuss the suitability and drawbacks of the indices most widely used in the assessment of coral fish biodiversity. An extensive and systematic survey of the literature focused on coral reef fish biodiversity was conducted from 1990 up to the present. We found that the multicomponent aspect of biodiversity, which is considered as a key feature of biodiversity for numerous terrestrial and marine ecosystems, has been poorly taken into account in coral reef fish studies. Species richness is still strongly dominant while other diversity components, such as functional diversity, are underestimated even when functional information is available. We also demonstrate that the reason for choosing particular indices is often unclear, mainly based on empirical rationales and/or the reproduction of widespread habits, but generally with no clear relevance with regard to the aims of the studies. As a result, the most widely used indices (species richness, Shannon, etc.) would appear to be poorly suited to meeting the main challenges facing the monitoring of coral reef fish biodiversity in the future. Our results clearly show that coral reef scientists should rather take advantage of the multicomponent aspect of biodiversity. To facilitate this approach, we propose general guidelines to serve as a basis for the selection of indices that provide complementary and relevant information for monitoring the response of coral reef fish biodiversity in the face of structuring factors (natural or anthropic). The aim of these guidelines was to achieve a better match between the properties of the selected indices and the context of each study (e.g. expected effect of the main structuring factors, nature of data available).     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

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     

A key phase in the recruitment dynamics of coral reef fishes: post-settlement transition

Synopsis Recent studies of recruitment dynamics in demersal fishes have placed major emphasis on presettlement mortality, and little on events bridging late larval and early juvenile periods. Observations on 68 taxa of Caribbean coral reef fishes before and during settlement revealed the existence of a distinct post-settlement life phase called the transition juvenile, associated with the act of recruitment. Transition juveniles were found as solitary individuals, in conspecific groups, or in heterospecific groups. The groups were either uniform or heterogenous in appearance. The complexity of the transition phase and its apparently widespread occurrence in coral reef fishes suggests that important aspects of population structure may be determined between settlement and first appearance as a full-fledged juvenile.