High population density enhances recruitment and survival of a harvested coral reef fish.

A negative relationship between population growth and population density (direct density dependence) is necessary for population regulation and is assumed in most models of harvested populations. Experimental tests for density dependence are lacking for large-bodied, harvested fish because of the difficulty of manipulating population density over large areas. We studied a harvested coral reef fish, Lutjanus apodus (schoolmaster snapper), using eight large, isolated natural reefs (0.4-1.6 ha) in the Bahamas as replicates. An initial observational test for density dependence was followed by a manipulation of population density. The manipulation weakened an association between density and shelter-providing habitat features and revealed a positive effect of population density on recruitment and survival (inverse density dependence), but no effect of density on somatic growth. The snappers on an individual reef were organized into a few shoals, and we hypothesize that large shoals on high-density reefs were less vulnerable to large piscivores (groupers and barracudas) than the small shoals on low-density reefs. Reductions in predation risk for individuals in large social groups are well documented, but because snapper shoals occupied reefs the size of small marine reserves, these ecological interactions may influence the outcome of management actions.

Synthesizing mechanisms of density dependence in reef fishes: behavior, habitat configuration, and observational scale.

Coral and rocky reef fish populations are widely used as model systems for the experimental exploration of density-dependent vital rates, but patterns of density-dependent mortality in these systems are not yet fully understood. In particular, the paradigm for strong, directly density-dependent (DDD) postsettlement mortality stands in contrast to recent evidence for inversely density-dependent (IDD) mortality. We review the processes responsible for DDD and IDD per capita mortality in reef fishes, noting that the pattern observed depends on predator and prey behavior, the spatial configuration of the reef habitat, and the spatial and temporal scales of observation. Specifically, predators tend to produce DDD prey mortality at their characteristic spatial scale of foraging, but prey mortality is IDD at smaller spatial scales due to attack-abatement effects (e.g., risk dilution). As a result, DDD mortality may be more common than IDD mortality on patch reefs, which tend to constrain predator foraging to the same scale as prey aggregation, eliminating attack-abatement effects. Additionally, adjacent groups of prey on continuous reefs may share a subset of refuges, increasing per capita refuge availability and relaxing DDD mortality relative to prey on patch reefs, where the patch edge could prevent such refuge sharing. These hypotheses lead to a synthetic framework to predict expected mortality patterns for a variety of scenarios. For nonsocial, nonaggregating species and species that aggregate in order to take advantage of spatially clumped refuges, IDD mortality is possible but likely superseded by DDD refuge competition, especially on patch reefs. By contrast, for species that aggregate socially, mortality should be IDD at the scale of individual aggregations but DDD at larger scales. The results of nearly all prior reef fish studies fit within this framework, although additional work is needed to test many of the predicted outcomes. This synthesis reconciles some apparent contradictions in the recent reef fish literature and suggests the importance of accounting for the scale-sensitive details of predator and prey behavior in any study system.

Recent region-wide declines in Caribbean reef fish abundance.

Profound ecological changes are occurring on coral reefs throughout the tropics, with marked coral cover losses and concomitant algal increases, particularly in the Caribbean region. Historical declines in the abundance of large Caribbean reef fishes likely reflect centuries of overexploitation. However, effects of drastic recent degradation of reef habitats on reef fish assemblages have yet to be established. By using meta-analysis, we analyzed time series of reef fish density obtained from 48 studies that include 318 reefs across the Caribbean and span the time period 1955-2007. Our analyses show that overall reef fish density has been declining significantly for more than a decade, at rates that are consistent across all subregions of the Caribbean basin (2.7% to 6.0% loss per year) and in three of six trophic groups. Changes in fish density over the past half-century are modest relative to concurrent changes in benthic cover on Caribbean reefs. However, the recent significant decline in overall fish abundance and its consistency across several trophic groups and among both fished and nonfished species indicate that Caribbean fishes have begun to respond negatively to habitat degradation.