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 Using a large-scale synthesis of peer-reviewed and unpublished data, we examine variability in abundance and biomass of herbivorous reef fishes and explore evidence for fishing impacts globally and within regions.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.

Janzen-Connell effects in a broadcast-spawning Caribbean coral: distance-dependent survival of larvae and settlers.

The Janzen-Connell hypothesis states that host-specific biotic enemies (pathogens and predators) promote the coexistence of tree species in tropical forests by causing distance- or density-dependent mortality of seeds and seedlings. Although coral reefs are the aquatic analogues of tropical forests, the Janzen-Connell model has never been proposed as an explanation for high diversity in these ecosystems. We tested the central predictions of the Janzen-Connell model in a coral reef, using swimming larvae and settled polyps of the common Caribbean coral Montastraea faveolata. In a field experiment to test for distance- or density-dependent mortality, coral settler mortality was higher and more strongly density dependent in locations down-current from adult corals. Survival did not increase monotoilically with distance, however, revealing the influence of fluid dynamics around adult corals in structuring spatial patterns of mortality. Complementary microbial profiles around adult coral heads revealed that one potential cause of settler mortality, marine microbial communities, are structured at the same spatial scale. In a field experiment to test whether factors causing juvenile mortality are host specific, settler mortality was 2.3-3.0 times higher near conspecific adults vs. near adult corals of other genera or in open reef areas. In four laboratory experiments to test for distance-dependent, host-specific mortality, swimming coral larvae were exposed to water collected near conspecific adult corals, near other coral genera, and in open areas of the reef. Microbial abundance in these water samples was manipulated with filters and antibiotics to test whether the cause of mortality was biotic (i.e., microbial). Juvenile survivorship was lowest in unfiltered water collected near conspecifics, and survivorship increased when this water was filter sterilized, collected farther away, or collected near other adult coral genera. Together these results demonstrate for the first time that the diversity-promoting mechanisms embodied in the Janzen-Connell model can operate in a marine ecosystem and in an animal. The distribution of adult corals across a reef will thus influence the spatial pattern of juvenile survival. When rare coral species have a survival advantage, coral species diversity per se becomes increasingly important for the persistence and recovery of coral cover on tropical reefs.

Fishing top predators indirectly affects condition and reproduction in a reef-fish community.

To examine the indirect effects of fishing on energy allocation in non-target prey species, condition and reproductive potential were measured for five representative species (two-spot red snapper Lutjanus bohar, arc-eye hawkfish Paracirrhites arcatus, blackbar devil Plectroglyphidodon dickii, bicolour chromis Chromis margaritifer and whitecheek surgeonfish Acanthurus nigricans) from three reef-fish communities with different levels of fishing and predator abundance in the northern Line Islands, central Pacific Ocean. Predator abundance differed by five to seven-fold among islands, and despite no clear differences in prey abundance, differences in prey condition and reproductive potential among islands were found. Body condition (mean body mass adjusted for length) was consistently lower at sites with higher predator abundance for three of the four prey species. Mean liver mass (adjusted for total body mass), an indicator of energy reserves, was also lower at sites with higher predator abundance for three of the prey species and the predator. Trends in reproductive potential were less clear. Mean gonad mass (adjusted for total body mass) was high where predator abundance was high for only one of the three species in which it was measured. Evidence of consistently low prey body condition and energy reserves in a diverse suite of species at reefs with high predator abundance suggests that fishing may indirectly affect non-target prey-fish populations through changes in predation and predation risk.

Survival and settlement success of coral planulae: independent and synergistic effects of macroalgae and microbes.

Restoration of degraded coral reef communities is dependent on successful recruitment and survival of new coral planulae. Degraded reefs are often characterized by high cover of fleshy algae and high microbial densities, complemented by low abundance of coral and coral recruits. Here, we investigated how the presence and abundance of macroalgae and microbes affected recruitment success of a common Hawaiian coral. We found that the presence of algae reduced survivorship and settlement success of planulae. With the addition of the broad-spectrum antibiotic, ampicillin, these negative effects were reversed, suggesting that algae indirectly cause planular mortality by enhancing microbial concentrations or by weakening the coral's resistance to microbial infections. Algae further reduced recruitment success of corals as planulae preferentially settled on algal surfaces, but later suffered 100% mortality. In contrast to survival, settlement was unsuccessful in treatments containing antibiotics, suggesting that benthic microbes may be necessary to induce settlement. These experiments highlight potential complex interactions that govern the relationships between microbes, algae and corals and emphasize the importance of microbial dynamics in coral reef ecology and restoration.