Coral reef fish larvae settle close to home

Population connectivity through larval dispersal is an essential parameter in models of marine population dynamics and the optimal size and spacing of marine reserves. However, there are remarkably few direct estimates of larval dispersal for marine organisms, and the actual birth sites of successful recruits have never been located. Here, we solve the mystery of the natal origin of clownfish (Amphiprion polymnus) juveniles by mass-marking via tetracycline immersion all larvae produced in a population. In addition, we established parentage by DNA genotyping all potential adults and all new recruits arriving in the population. Although no individuals settled into the same anemone as their parents, many settled remarkably close to home. Even though this species has a 9-12 day larval duration, one-third of settled juveniles had returned to a 2 hectare natal area, with many settling <100 m from their birth site. This represents the smallest scale of dispersal known for any marine fish species with a pelagic larval phase. The degree of local retention indicates that marine reserves can provide recruitment benefits not only beyond but also within their boundaries.     

Coral reef fish smell leaves to find island homes

Recent studies have shown that some coral reef fish larvae return to natal reefs, while others disperse to distant reefs. However, the sensory mechanisms used to find settlement sites are poorly understood. One hypothesis is that larvae use olfactory cues to navigate home or find other suitable reef habitats. Here we show a strong association between the clownfish Amphiprion percula and coral reefs surrounding offshore islands in Papua New Guinea. Host anemones and A. percula are particularly abundant in shallow water beneath overhanging rainforest vegetation. A series of experiments were carried out using paired-choice flumes to evaluate the potential role of water-borne olfactory cues in finding islands. Recently settled A. percula exhibited strong preferences for: (i) water from reefs with islands over water from reefs without islands; (ii) water collected near islands over water collected offshore; and (iii) water treated with either anemones or leaves from rainforest vegetation. Laboratory reared-juveniles exhibited the same positive response to anemones and rainforest vegetation, suggesting that olfactory preferences are innate rather than learned. We hypothesize that A. percula use a suite of olfactory stimuli to locate vegetated islands, which may explain the high levels of self-recruitment on island reefs. This previously unrecognized link between coral reefs and island vegetation argues for the integrated management of these pristine tropical habitats.     

Free—living fathead minnows rapidly learn to recognize pike as predators

Individuals from a natural population of approximately 20 000 fathead minnows from a pike–free pond did not respond with appropriate anti–predator behaviour upon encountering pike odour in laboratory tests. However, 14 days after 10 pike were stocked into the pond, minnows had acquired recognition of pike odour. Laboratory studies have indicated several possible mechanisms for acquiring predator recognition in fathead minnows. This study indicates that these, or similar processes, can produce major changes in predator recognition in the wild.     

Coral reef fish communities: a compromise view

Synopsis The fishes associated with coral reefs offer excellent opportunities for the study of the factors that determine the species composition of complex, highly interactive communities. Amenable to sampling and direct observations many patch reefs are small enough to be studied as entire units and yet diverse enough to include a wide range of interactions.Coral reef fishes appear to be highly specialized in morphology, color, behavior and life cycles and yet colonization experiments and repeated censusing have shown a surprisingly high variation in the fishes that are associated in similar habitats or in the same habitat at different times. This has led to two different views: (i) the chaos view that holds that the species composition is due to random factors and chance colonization, and (ii) the order view that resource sharing adaptations determine which species can live together.This paper reviews some of the obvious adaptations of reef fishes. An examination of the maximum sizes reached by infaunal reef dwellers shows that the largest individual of each species differs by a constant proportion from the next larger and next smaller species. This suggests growth limitations by interspecies competition. A hypothetical model showing how this might work is offered and it is shown that as long as there are more species than can be accommodated in the community at any one time the number of combinations is great enough to give the appearance of randomness even though the individual species may have precise environmental requirements.This paper forms 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. Hellman, ed.).     

Coral reef fish populations can persist without immigration

Determining the conditions under which populations may persist requires accurate estimates of demographic parameters, including immigration, local reproductive success, and mortality rates. In marine populations, empirical estimates of these parameters are rare, due at least in part to the pelagic dispersal stage common to most marine organisms. Here, we evaluate population persistence and turnover for a population of orange clownfish, Amphiprion percula, at Kimbe Island in Papua New Guinea. All fish in the population were sampled and genotyped on five occasions at 2-year intervals spanning eight years. The genetic data enabled estimates of reproductive success retained in the same population (reproductive success to self-recruitment), reproductive success exported to other subpopulations (reproductive success to local connectivity), and immigration and mortality rates of sub-adults and adults. Approximately 50% of the recruits were assigned to parents from the Kimbe Island population and this was stable through the sampling period. Stability in the proportion of local and immigrant settlers is likely due to: low annual mortality rates and stable egg production rates, and the short larval stages and sensory capacities of reef fish larvae. Biannual mortality rates ranged from 0.09 to 0.55 and varied significantly spatially. We used these data to parametrize a model that estimated the probability of the Kimbe Island population persisting in the absence of immigration. The Kimbe Island population was found to persist without significant immigration. Model results suggest the island population persists because the largest of the subpopulations are maintained due to having low mortality and high self-recruitment rates. Our results enable managers to appropriately target and scale actions to maximize persistence likelihood as disturbance frequencies increase.     

Density dependent recruitment in the reef fish Chaetodon miliaris

Synopsis Recruitment of reef fish has generally been found to be unrelated to or positively related to adult densities. This paper reports an inverse relationship between the number of recruits of Chaetodon miliaris and the density of conspecific adults. C. miliaris are non-territorial, planktivorous butterflyfish. The study populations occurred on the flanks of patch reefs of about 30 m in diameter. At their peak densities populations consisted of 300–400 C. miliaris. Recruitment (the appearance in the stock of fish less than 3.5cm in total length) occurred primarily from April to June and corresponded to the new moon period in these months. Populations generally showed a steady decline in numbers during months of the year when recruitment was not occurring: the finite rate of decline was approximately 30% per month. Populations varied in the density of adults at the start of the spring-summer period of recruitment, and the magnitude of recruitment to a reef was inversely related to the density of these conspecific adults. Reduction of the densities of adult C. miliaris through trap fishing, resulted in increased recruitment to the fished stock. These results provide evidence that in some. species of reef fish, benthic processes may play an important role in determining the magnitude of recruitment to an adult stock, in contrast to the widely held view that recruitment is in large part a result of chance events in the plankton or that recruitment is directly related to adult densities.     

Coral reef management and conservation in light of rapidly evolving ecological paradigms

The decline of many coral reef ecosystems in recent decades surprised experienced managers and researchers. It shattered old paradigms that these diverse ecosystems are spatially uniform and temporally stable on the scale of millennia. We now see reefs as heterogeneous, fragile, globally stressed ecosystems structured by strong positive or negative feedback processes. We review the causes and consequences of reef decline and ask whether management practices are addressing the problem at appropriate scales. We conclude that both science and management are currently failing to address the comanagement of extractive activities and ecological processes that drive ecosystems (e.g. productivity and herbivory). Most reef conservation efforts are directed toward reserve implementation, but new approaches are needed to sustain ecosystem function in exploited areas.     

Animal cognition: how archer fish learn to down rapidly moving targets

In extremely rapid maneuvers, animals including man can launch ballistic motor patterns that cannot immediately be corrected. Such patterns are difficult to direct at targets that move in three-dimensional space, and it is presently unknown how animals learn to acquire the precision required. Archer fish live in groups and are renowned for their ballistic hunting technique in which they knock down stationary aerial insect prey with a precisely aimed shot of water. Here we report that these fish can learn to release their shots so as to hit prey that moves rapidly at great height, a remarkable accomplishment in which the shooter must take both the target's three-dimensional motion as well as that of its rising shot into account. To successfully perform in the three-dimensional task, training with horizontal motion suffices. Moreover, all archer fish of a group were able to learn the complex sensomotor skill from watching a performing group member, without having to practice. This instance of social learning in a fish is most remarkable as it could imply that observers can "change their viewpoint," mapping the perceived shooting characteristics of a distant team member into angles and target distances that they later must use to hit.     

Variability in relative importance of determinants of reef fish recruitment

Multiple processes can act together to determine abundance of organisms and structure of communities. Recently, appreciation of this fact has motivated development of conceptual and statistical frameworks that quantitatively assess the relative importance of multiple causal factors. However, little consideration has been given to variability in the "importance" of processes through space and time (i.e. robustness), which represents another facet of a process's importance. Here, I focused on populations of a coral reef fish ( Thalassoma hardwicke ) and used an existing analytical method to assess the relative importance of initial population inputs (larval supply) and subsequent juvenile mortality in determining the average abundances of juvenile fish populations in different locations and times. The relative importance of processes varied significantly both temporally and spatially across a range of scales, and indicate a need for future assessments of relative importance to incorporate this variability.     

Coral reef fish assemblages: intra- and interspecific competition for shelter sites

Synopsis Observations were made on intra- and interspecific aggressive interactions among the fishes living in the rubble/sand coral reef habitat in St. Croix, U. S. Virgin Islands. Four species (beaugregory — Stegastes leucostictus; ocean surgeonfish — Acanthurus bahianus; doctor fish — A. chirurgus; common squirrelfish — Holocentrus rufus) which sheltered in holes on the reef all actively defended one to several shelter sites at dusk. Short-term shelter side fidelity was observed in three of these four species. Agonistic interactions over both food and shelter occurred during the daytime but much less frequently than agonistic interactions over shelter at dusk. Dominance in intraspecific aggression was determined almost completely by the relative sizes of the individuals involved, with the larger individuals dominating in 95–98% of all encounters. A similar, but less strong, relationship between size and dominance existed for interactions between closely related species. For aggressive encounters between unrelated species, however, both relative sizes and species identity determined the outcome. Species, both diurnal and nocturnal, which strongly defend several shelter sites may have a strong and disproportionate impact on the sheltering behavior of other fishes. Intraspecific and interspecific defense of shelter sites may determine the patterns of mortality that result from predation, thereby influencing population abundances and assemblage composition.     

Plants as reef fish: fitting the functional form of seedling recruitment

The life histories of many species depend first on dispersal to local sites and then on establishment. After dispersal, density-independent and density-dependent mortalities modify propagule supply, determining the number of individuals that establish. Because multiple factors influence recruitment, the dichotomy of propagule versus establishment limitation is best viewed as a continuum along which the strength of propagule or establishment limitation changes with propagule input. To evaluate the relative importance of seed and establishment limitation for plants, we (1) describe the shape of the recruitment function and (2) use limitation and elasticity analyses to quantify the sensitivity of recruitment to perturbations in seed limitation and density-independent and density-dependent mortality. Using 36 seed augmentation studies for 18 species, we tested four recruitment functions against one another. Although the linear model (accounting for seed limitation and density-independent mortality) fitted the largest number of studies, the nonlinear Beverton-Holt model (accounting for density-dependent mortality) performed better at high densities of seed augmentation. For the 18 species, seed limitation constrained population size more than other sources of limitation at ambient conditions. Seedling density reached saturation with increasing seed density in many studies, but at such high densities that seedling density was primarily limited by seed availability rather than microsite availability or density dependence.     

Coral bleaching, reef fish community phase shifts and the resilience of coral reefs

The 1998 global coral bleaching event was the largest recorded historical disturbance of coral reefs and resulted in extensive habitat loss. Annual censuses of reef fish community structure over a 12-year period spanning the bleaching event revealed a marked phase shift from a prebleach to postbleach assemblage. Surprisingly, we found that the bleaching event had no detectable effect on the abundance, diversity or species richness of a local cryptobenthic reef fish community. Furthermore, there is no evidence of regeneration even after 5–35 generations of these short-lived species. These results have significant implications for our understanding of the response of coral reef ecosystems to global warming and highlight the importance of selecting appropriate criteria for evaluating reef resilience.     

Resource limitation and recruitment patterns in a coral reef fish assemblage

The potential effects of food and shelter availability on the recruitment and early survivorship of coral reef fishes were studied on St. Croix, U.S. Virgin Islands. The faunal assemblage studied included diurnally active fishes found in the “rubble/sand” habitat. The most abundant members were: beaugregory, Stegastes leucostictus (Muller & Troschel), goldspotted goby, Gnatholepis thompsoni Jordan, bridled goby, Coryphopterus glaucofraenum Gill, surgeonfishes, Acanthurus bahianus Castelnau and A. chirurgus (Bloch), and French grunt, Haemulon flavolineatum (Desmarest). Comparisons of recruitment to reefs constructed from substrata that varied in morphological characteristics showed that there were differences in the relative abundances of recruits attracted to and/or surviving on the different reef types. Juveniles of most species appeared to prefer the branching coral Porites porites (Pallas), which provided a large number of small crevices between the branches.Manipulations of the availability of shelter sites for fishes demonstrated that recruitment and/or early survivorship were strongly limited by the number of refuges. This result was found in six separate carried out during different years and in different seasons. Shelter site availability presumably limits fish populations through its effects on prédation rates.Experimental manipulations of food availability indicated that food does not directly influence settlement or early survivorship of coral reef fishes. However, it is probable that correlations between habitat characteristics and food availability have influenced the evolution of settling preferences.     

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.     

The influence of multiple factors upon reef fish abundance and species richness in a tropical coral complex

The present study was conducted on Tamandaré reefs, northeast Brazil and aimed to analyse the importance of different factors (e.g. tourism activity, fishing activity, coral abundance and algal abundance) on reef fish abundance and species richness. Two distinct reef areas (A ver o mar and Caieiras) with different levels of influence were studied. A total of 8239 reef fish individuals were registered, including 59 species. Site 1 (A ver o mar) presented higher reef fish abundance and richness, with dominance of roving herbivores (29.9 %) and mobile invertebrate feeders (28.7 %). In contrast, at Site 2 (Caieiras) territorial herbivores (40.9 %) predominated, followed by mobile invertebrate feeders (24.6 %). Concerning the benthic community, at Site 1 macroalgae were recorded as the main category (49.3 %); however, Site 2 was dominated by calcareous algae (36.0 %). The most important variable explaining more than 90 % of variance on reef fish abundance and species richness was macroalgae abundance, followed by fishing activity. Phase shifts on coral reefs are evident, resulting in the replacement of coral by macroalgae and greatly influencing reef fish communities. In this context, it is important to understand the burden of the factors that affect reef fish communities and, therefore, influence the extinction vulnerability of coral reef fishes.     

Spatio-temporal variability in fish recruitment to a coral reef (Moorea,French Polynesia)

Spatial and temporal patterns of recruitment of juvenile coral reef fishes were studied on the reefs of the high island of Moorea (Society Archipelago, French Polynesia) during the wet season (October 1988 to April 1989). Two size-classes of fishes (new-recruits and juveniles) were censused by visual counts within 24 quadrats along a transect across the lagoon. Correspondence analysis was used to calculate the hypothetical movements of the two size classes. Spatial variability was far more important than temporal variability; six main spatial communities were revealed, which could result from differences among habitats. Temporal variability in recuitment occurred only at nearshore stations; stations on the outer fringing reef and inner barrier reef displayed stable recruitment patterns.