Peter Doherty

An Empirical Test of Recruitment Limitation in a Coral Reef Fish

A long-term, large-scale empirical test of the recruitment limitation hypothesis was done by sampling fish populations from the southern Great Barrier Reef after having monitored their recruitment histories for 9 years. After adjustment for demographic differences, recruitment patterns explained over 90 percent of the spatial variation in abundance of a common damselfish among seven coral reefs. The age structures from individual reefs also preserved major temporal variations in the recruitment signal over at least 10 years. Abundance and demography of this small fish at these spatial and temporal scales can be explained almost entirely as variable recruitment interacting with density-independent mortality.

High mortality during settlement is a population bottleneck for a tropical surgeonfish

Replenishment of benthic marine populations typically involves “settlement” from pelagic larval to benthic juvenile habitats. Mortality during this transition has been unknown because of the difficulty of measuring propagule supply in open water. For three weeks, we compared the nocturnal passage of presettlement fishes across the barrier reef encircling Moorea Island (French Polynesia) with the abundance of benthic recruits in the back-reef lagoon on the following morning. During this time, >40,000 presettlement unicornfish, Naso unicornis entered our study area of ∼1 km2 with half arriving on just two nights. Using coupled Beverton-Holt functions to describe the decay of each cohort, we were able to predict the daily abundance of recruits and their final age structure from the presettlement inputs. The best model estimated that ∼61% of the potential settlers were lost between their nocturnal arrival and the following morning, independent of cohort size. Postsettlement mortality was density dependent, varying between 9% and 20% per day. We attribute all mortality to predation and suggest that high risk associated with settlement has shaped colonization strategies. Because fishing targets the survivors of this population bottleneck, aquarium fisheries may be more sustainable when sourced from pelagic juveniles.

Chapter 15 – Variable Replenishment and the Dynamics of Reef Fish Populations

Publisher Summary This chapter focuses on the relationship between variable replenishment and the dynamics of reef fish populations. The populations of coral reef fishes can experience variable replenishment and these variations may have lasting impacts on patterns of demography and abundance. Populations of siganids in Guam, acanthurids in Minicoy Atoll and Johnston Atoll, balistids in Panama, and serranids in Moorea and La Reunion have all shown conspicuous mass mortalities following exceptional replenishment. Modern theoretical models of reef fish populations tend to be based upon bounded stochastic dynamics, with variable replenishment of populations that provide a strong source of change when populations are not near their upper limits. Under this density-vague scenario, population dynamics feature both density-dependent and density-independent fluctuations, and the relative importance of each varies with context. When replenishment is drawn from strongly skewed probability distributions, density dependence is most important at the extremes of density, where its primary effect is to dampen exceptional cohorts. For sedentary prey that fall between these extremes, the foraging behavior of predators interacting with the spatial dispersion of settlers has the potential to produce density-dependent outcomes that are linked only indirectly to absolute abundance. Fish populations are very tolerant of crowding, with numerical responses detected only at very high and very low levels of abundance. Between these extremes, social behavior can have a big effect on individual access to essential resources, including food, with impacts on growth and maturation. Some populations may be able to store fecundity in the form of stunted subadults that could be recruited into the breeding pool as required to buffer egg production from the impacts of variable settlement. Thus, it is likely that numerical fluctuations in fish populations are a poor proxy for changes in egg production.

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.

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 sites 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.

Large scale spatial and temporal variation in recruitment to fish populations on coral reefs

SummaryVisual census was used to sample young of the year of fish species recruited to each of two habitats on seven lagoonal platform reefs of the Capricorn-Bunker Group, Great Barrier Reef. The reefs sampled span an area 70 km in extent. In 1983, 62 species from 13 families were detected as recruits on reef slope sites. The total number of cruits, and the number of each of 6 of 16 species tested, differed significantly among reefs, despite the fact that differences among sites within reefs did not exist, and that sampled slopes were chosen to be hydrographically, and physiographically as similar as possible. Lagoonal patch reefs were sampled in two years. In 1982, 76 species of 11 families occurred as recruits. In 1983, 86 species of 12 families were recorded. All of 22 species common enough to test showed some significant variation in abundance among reefs, years, or both. For 9 species, significant year x reef interactions occurred, demonstrating that relative recruitment success among reefs varied between years. Reasons for the substantial levels of variability are discussed, and implications for the organisation of reef fish communities are considered.

Some effects of density on the juveniles of two species of tropical, territorial damselfish

Abstract Small juveniles of Pomacentrus wardi Whitley and P. flavicauda Whitley were transplanted onto replicate reefs, at various densities, to assess the importance of intraspecific and interspecific competition among fishes of the same size, age, and experience. Although P. flavicauda disappeared from the reefs faster than did P. wardi , survival of neither species was affected by the density of conspecifics or the presence of its congener (during a 1-yr period). Thus densities at the end of the experiment reflected the initial stocking densities which simulated a range of recruitment densities exceeding those observed in wild populations. Crowding caused both species to change their intraspecific behaviours from territorialism to participation in dominance hierarchies. For P. wardi , this was correlated with unequal growth among the individuals on each patch, which partially disguised a trend for the average size of all fish to decrease with increasing density. Over the range of crowding and the time scales explored, total biomass ( = production) did not reach an asymptotic value which might have indicated an absolute shortage of food for these small fishes. Variations in the survival and biomass of P. wardi among replicate reefs were not correlated with the distribution and abundance of any other species also resident on these reefs. The highest densities of damselfishes used in this experiment were more than 50 times the average year-class strength observed at this site. Yet proportional losses from the experimental and natural cohorts were not distinguishable. The lack of compensatory mortality over this range of densities suggests that competition within a year-class does not control the density of young fish in wild populations. This result is consistent with other evidence that recruit densities are extrinsically controlled by a limiting supply of pelagic larvae.

Monitoring of Benthic Reference Sites: Using an Autonomous Underwater Vehicle

We have established an Australia-wide observation program that exhibits recent developments in autonomous underwater vehicle (AUV) systems to deliver precisely navigated time series benthic imagery at selected reference stations on Australias continental shelf. These observations are designed to help characterize changes in benthic assemblage composition and cover derived from precisely registered maps collected at regular intervals. This information will provide researchers with the baseline ecological data necessary to make quantitative inferences about the long-term effects of climate change and human activities on the benthos. Incorporating a suite of observations that capitalize on the unique capabilities of AUVs into Australias integrated marine observation system (IMOS) [1] is providing a critical link between oceanographic and benthic processes. IMOS is a nationally coordinated program designed to establish and maintain the research infrastructure required to support Australias marine science research. It has, and will maintain, a strategic focus on the impact of major boundary currents on continental shelf environments, ecosystems, and biodiversity. The IMOS AUV facility observation program is designed to generate physical and biological observations of benthic variables that cannot be cost effectively obtained by other means.

Directional Swimming of Fish Larvae Determines Connectivity of Fish Populations on the Great Barrier Reef

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The adaptive significance of larval dispersal in coral reef fishes

SynopsisCoral reef fishes almost universally disperse over relatively great distances during a pelagic larval phase. Barlow (1981) suggested that this dispersal is adaptive because adult fishes inhabit a patchy, uncertain environment. This reiterated an older idea that the random extinction of local populations necessarily favours dispersal, since ultimately all populations of non-dispersers will disappear. Whereas this view is based on adult survival, we emphasize a less frequent view that substantial larval dispersal may be adaptive when offspring experience patchy and unpredictable survival in the pelagic habitat. We do not address the question of why these animals ‘broadcast’ rather than ‘brood’, but suggest that species committed to pelagic offspring will be under selection to disperse siblings to spread the risk of failure among members of a cohort. Our arguments are supported by a heuristic computer simulation.