The majority of marine populations are demographically open; their replenishment is largely or exclusively dependent on a supply of juveniles from the plankton. In spite of much recent research, no consensus has yet been reached regarding the importance of recruitment relative to other demographic processes in determining local population densities. We argue 1. that demographic theory suggests that, except under restrictive and unlikely conditions, recruitment must influence local population density to some extent. Therefore, 2. the question as to whether the size of a particular population is limited by recruitment is misguided. Finally, 3. the effect of recruitment on population size can be difficult to detect but is nonetheless real. A major weakness of most existing studies is a lack of attention to the survival of recruits over appropriate scales of time and space. Acknowledgment of the multifactorial determination of population density should guide the design of future experimental studies of the demography of open populations.

The worldwide decline in coral cover has serious implications for the health of coral reefs. But what is the future of reef fish assemblages? Marine reserves can protect fish from exploitation, but do they protect fish biodiversity in degrading environments? The answer appears to be no, as indicated by our 8-year study in Papua New Guinea. A devastating decline in coral cover caused a parallel decline in fish biodiversity, both in marine reserves and in areas open to fishing. Over 75% of reef fish species declined in abundance, and 50% declined to less than half of their original numbers. The greater the dependence species have on living coral as juvenile recruitment sites, the greater the observed decline in abundance. Several rare coral-specialists became locally extinct. We suggest that fish biodiversity is threatened wherever permanent reef degradation occurs and warn that marine reserves will not always be sufficient to ensure their survival.M any ecologists have expressed concern over the worldwide decline in coral cover due to global warming and associated coral bleaching, overfishing, and coastal pollution (1-5). Coral reefs support a high diversity of fishes that may ultimately depend on corals for their survival; however, the impact of long-term reef degradation on fish populations is unknown. Most attention to the protection of marine fish populations has focused on the benefits of controlling exploitation by establishing ''no-take'' marine reserves (6-8). However, comprehensive strategies for protecting marine biodiversity also require an understanding of how species respond to degradation of their habitats.In the past, there has been a dichotomy of opinion over how closely fish communities are linked to their habitat, with some information indicating a high degree of variability that is independent of habitat change (9-14) and other data showing that coral-specialists clearly suffer when coral cover is reduced (13-17). Here we ask the following questions. If coral reefs continue along a path of degradation, what will be the fate of fish communities as a whole? Will marine reserves provide fish communities with any resilience to the effects of habitat loss? MethodsIn 1996, we observed the beginning of what progressed into a long-term decline in coral cover in four marine reserves in the Tamane Puli Conservation Area, Kimbe Bay, Papua New Guinea (150°06ЈE, 5°25ЈS). To predict the potential response of fish assemblages to declining coral in this area, we began by estimating the proportion of reef fish species that only fed on coral tissue or those that only lived in association with branching corals. We surveyed all species in 20 different families of fishes associated with coral reefs in the region (18). Those species dependent on live coral as food or living space were distinguished from the rest, based both on our own observations and published accounts of diet and habitat associations (19,20).The cover of branching scleractinian corals was estimated from annual surveys of eight reefs between 1996 a...

Increased frequency of disturbances and anthropogenic activities are predicted to have a devastating impact on coral reefs that will ultimately change the composition of reef associated fish communities. We reviewed and analysed studies that document the effects of disturbance-mediated coral loss on coral reef fishes. Meta-analysis of 17 independent studies revealed that 62% of fish species declined in abundance within 3 years of disturbances that resulted in 410% decline in coral cover. Abundances of species reliant on live coral for food and shelter consistently declined during this time frame, while abundance of some species that feed on invertebrates, algae and/or detritus increased. The response of species, particularly those expected to benefit from the immediate loss of coral, is, however, variable and is attributed to erratic replenishment of stocks, ecological versatility of species and sublethal responses, such as changes in growth, body condition and feeding rates. The diversity of fish communities was found to be negatively and linearly correlated to disturbance-mediated coral loss. Coral loss 420% typically resulted in a decline in species richness of fish communities, although diversity may initially increase following small declines in coral cover from high coverage. Disturbances that result in an immediate loss of habitat complexity (e.g. severe tropical storms), have a greater impact on fishes from all trophic levels, compared with disturbances that kill corals, but leave the reef framework intact (e.g. coral bleaching and outbreaks of Acanthaster planci). This is most evident among small bodied species and suggests the long-term consequences of coral loss through coral bleaching and crown-ofthorn starfish outbreaks may be much more substantial than the short-term effects currently documented.

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The persistence of most coastal marine species depends on larvae finding suitable adult habitat at the end of an offshore dispersive stage that can last weeks or months. We tested the effects that ocean acidification from elevated levels of atmospheric carbon dioxide (CO2) could have on the ability of larvae to detect olfactory cues from adult habitats. Larval clownfish reared in control seawater (pH 8.15) discriminated between a range of cues that could help them locate reef habitat and suitable settlement sites. This discriminatory ability was disrupted when larvae were reared in conditions simulating CO2-induced ocean acidification. Larvae became strongly attracted to olfactory stimuli they normally avoided when reared at levels of ocean pH that could occur ca. 2100 (pH 7.8) and they no longer responded to any olfactory cues when reared at pH levels (pH 7.6) that might be attained later next century on a business-as-usual carbon-dioxide emissions trajectory. If acidification continues unabated, the impairment of sensory ability will reduce population sustainability of many marine species, with potentially profound consequences for marine diversity.climate change ͉ larval sensory mechanisms ͉ population connectivity ͉ population replenishment O cean acidification caused by the uptake of additional carbon dioxide (CO 2 ) at the ocean surface is now recognized as a serious threat to marine ecosystems (1-4). At least 30% of the anthropogenic CO 2 released into the atmosphere in the past 200 years has been absorbed by the oceans, causing ocean pH to decline at a rate Ϸ100 times faster than at any time in the past 650,000 years (1, 4). Global ocean pH is estimated to have dropped by 0.1 units since preindustrial times and is projected to fall another 0.3-0.4 units by 2100 because of existing and future CO 2 emissions (1, 5-6). Considerable research effort has focused on predicting the impact that reduced carbonate-ion saturation states that accompany ocean acidification will have on calcifying marine organisms, particularly corals and other invertebrates that precipitate aragonite skeletons (2-3, 6). However, the effects that ocean acidification will have on other marine organisms, including fishes, remain almost completely unknown, especially for conditions of atmospheric carbon dioxide and seawater pH that could occur in the near future (4, 7-9).The persistence of most coastal marine species depends on the ability of larvae to locate suitable settlement habitat at the end of a pelagic stage that can last weeks or months. Accumulating evidence for reef fishes suggests that both reef sounds (10) and olfactory cues (11-13) are used by larvae to locate reefs. The olfactory organs of many reef fishes are well-developed by the end of the larval phase (14-15), and it has recently been shown that larvae of some species can discriminate the smell of water from their natal reef compared with water from other reefs (13), which provides a mechanism to explain high levels of selfrecruitment in some reef fish populations (16)(...

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.

Marine reserves, areas closed to all forms of fishing, continue to be advocated and implemented to supplement fisheries and conserve populations. However, although the reproductive potential of important fishery species can dramatically increase inside reserves, the extent to which larval offspring are exported and the relative contribution of reserves to recruitment in fished and protected populations are unknown. Using genetic parentage analyses, we resolve patterns of larval dispersal for two species of exploited coral reef fish within a network of marine reserves on the Great Barrier Reef. In a 1,000 km(2) study area, populations resident in three reserves exported 83% (coral trout, Plectropomus maculatus) and 55% (stripey snapper, Lutjanus carponotatus) of assigned offspring to fished reefs, with the remainder having recruited to natal reserves or other reserves in the region. We estimate that reserves, which account for just 28% of the local reef area, produced approximately half of all juvenile recruitment to both reserve and fished reefs within 30 km. Our results provide compelling evidence that adequately protected reserve networks can make a significant contribution to the replenishment of populations on both reserve and fished reefs at a scale that benefits local stakeholders.