Differences in the structures of fish assemblages inThalassia testudinum beds in Guadeloupe,French West Indies,and their ecological significance

The structures of fish assemblages in twoThalassia testudinum beds in Guadeloupe, French West Indies, one adjacent to mangroves and the other adjacent to coral reefs, were compared between January 1983 and May 1984. The aim of the study was to compare the influences of mangroves and coral reefs on the utilization of seagrass beds by fishes through examination of species composition, catch rate, size of fishes and temporal changes. The two fish assemblages were similar in terms of the number of species they had in common (nearly 44% of the total number of species collected) and the great abundance of juveniles. They both comprised species that usually inhabit other habitats, i.e., estuaries, open waters or coral reefs. Estuary-associated species (e.g. Gerreidae) were the most abundant species in the seagrass bed near the mangroves, while small pelagic species (e.g. Clupeidae) were the most abundant species in the seagrass bed near the coral reefs. The seagrass bed near the mangroves was preferentially utilized as a nursery area by small juveniles of various species (e.g. Clupeidae, Sparidae, Gerreidae, and at least one coral reef species,Ocyurus chrysurus). The abundance of these species varied frequently, suggesting successive arrivals and departures of juveniles over time. The seagrass bed near the coral reefs was characteristically utilized by fishes that are more able to avoid predation, i.e., fishes that forage over seagrass beds at night and shelter in or near the coral reefs during the day (large juveniles of coral reef species and adults of schooling pelagic species, respectively). The constant migrations of these fishes between the coral reefs and seagrass beds explained the relative stability of the structure of the fish assemblage in the seagrass bed over time. Thus, the two seagrass beds were not equivalent habitats for fishes. The distinct ecological influences of the mangroves (as a nursery for small juveniles) and coral reefs (as a shelter for larger fishes) on the nearby seagrass beds was clearly reflected by the distinct utilizations of these seagrass beds by fishes.     

Potential of Marine Reserves to Cause Community-Wide Changes beyond Their Boundaries

Abstract:  Fishing and other human activities can alter the abundances, size structure, and behavior of species playing key roles in shaping marine communities (e.g., keystone predators), which may in turn cause ecosystem shifts. Despite extensive evidence that cascading trophic interactions can underlie community-wide recovery inside no-take marine reserves by protecting high-level predators, the spatial extent of these effects into adjacent fished areas is unknown. I examined the potential for community-wide changes (i.e., the transition from overgrazed coralline barrens to macroalgal beds) in temperate rocky reefs within and around a no-take marine reserve. For this purpose I assessed distribution patterns of predatory fishes, sea urchins, and barrens across the reserve boundaries. Predatory fishes were significantly more abundant within the reserve than in adjacent locations, with moderate spillover across the reserve edges. In contrast, community-wide changes of benthic assemblages were apparent well beyond the reserve boundaries, which is consistent with temporary movements of predatory fishes (e.g., foraging migration) from the reserve to surrounding areas. My results suggest that no-take marine reserves can promote community-wide changes beyond their boundaries.     

Effects of Artisanal Fishing on Caribbean Coral Reefs

Abstract:  Although the impacts of industrial fishing are widely recognized, marine ecosystems are generally considered less threatened by artisanal fisheries. To determine how coral reef fish assemblages and benthic communities are affected by artisanal fishing, we studied six Caribbean islands on which fishing pressure ranged from virtually none in Bonaire, increasing through Saba, Puerto Rico, St Lucia, and Dominica, and reaching very high intensities in Jamaica. Using stationary-point fish counts at 5 m and 15 m depth, we counted and estimated the lengths of all noncryptic, diurnal fish species within replicate 10-m-diameter areas. We estimated percent cover of coral and algae and determined reef structural complexity. From fish numbers and lengths we calculated mean fish biomass per count for the five most commercially important families. Groupers (Serranidae), snappers (Lutjanidae), parrotfish (Scaridae), and surgeonfish (Acanthuridae) showed order-of-magnitude differences in biomass among islands. Biomass fell as fishing pressure increased. Only grunts (Haemulidae) did not follow this pattern. Within families, larger-bodied species decreased as fishing intensified. Coral cover and structural complexity were highest on little-fished islands and lowest on those most fished. By contrast, algal cover was an order of magnitude higher in Jamaica than in Bonaire. These results suggest that following the Caribbean-wide mass mortality of herbivorous sea urchins in 1983–1984 and consequent declines in grazing pressure on reefs, herbivorous fishes have not controlled algae overgrowing corals in heavily fished areas but have restricted growth in lightly fished areas. In summary, differences among islands in the structure of fish and benthic assemblages suggest that intensive artisanal fishing has transformed Caribbean reefs.     

The ecology of Indo-Pacific grouper (Serranidae) species and the effects of a small scale no take area on grouper assemblage,abundance and size frequency distribution

This paper used the case study of the Wakatobi Marine National Park, Indonesia to examine changes in the diversity, density and maturity of grouper species over a 5-year period following the establishment of a small-scale no-take area (NTA). This work was carried out to investigate whether “small” NTAs could be effective management strategies over a time scale that is relevant to local fishery communities and their perception of management success. Our research also documents the ecology of these species, information essential if we are to understand how management practises are to affect coral reef fish species. Designation of this “small NTA” increased the density of groupers by 30% over a 5-year period of protected status. After 5 years of protection, grouper populations within this NTA were more mature and double the density of those within the adjacent lightly fished sites and nearly five times those of a heavily fished site. During this time all other nearby fished sites underwent large declines in grouper density. The nearby lightly fished Kaledupa site decreased by up to 50% year⁻¹. Such drastic declines are considered the impact of the exponential development of ever efficient and unsustainable methods of fishing within the study region. This NTA was not of benefit to all grouper species; the reasons for which are not clear. Such questions require further detailed research about the life history, population and behavioural ecology of Indo-Pacific grouper species. Such information is critical for urgently needed fisheries management. The present study found that a small scale NTA of 500 m length was large enough to increase the population of top predatory fish. In conjunction with other socially acceptable small scale NTAs it could help maintain and increase important fish stocks over a larger area. The use of “small” NTAs within networks of reserves should become a useful tool in the management of the locally exploited coral reefs.     

Relative influence of environmental factors and fishing on coral reef fish assemblages

Understanding whether assemblages of species respond more strongly to bottom-up (availability of trophic resources or habitats) or top-down (predation pressure) processes is important for effective management of resources and ecosystems. We determined the relative influence of environmental factors and predation by humans in shaping the density, biomass, and species richness of 4 medium-bodied (10–40 cm total length [TL]) coral reef fish groups targeted by fishers (mesopredators, planktivores, grazer and detritivores, and scrapers) and the density of 2 groups not targeted by fishers (invertivores, small fish ≤10 cm TL) in the central Philippines. Boosted regression trees were used to model the response of each fish group to 21 predictor variables: 13 habitat variables, 5 island variables, and 3 fishing variables (no-take marine reserve [NTMR] presence or absence, NTMR size, and NTMR age). Targeted and nontargeted fish groups responded most strongly to habitat variables, then island variables. Fishing (NTMR) variables generally had less influence on fish groups. Of the habitat variables, live hard coral cover, structural complexity or habitat complexity index, and depth had the greatest effects on density, biomass, and species richness of targeted fish groups and on the density of nontargeted fishes. Of the island variables, proximity to the nearest river and island elevation had the most influence on fish groups. The NTMRs affected only fishes targeted by fishers; NTMR size positively correlated with density, biomass, and species richness of targeted fishes, particularly mesopredatory, and grazing and detritivorous fishes. Importantly, NTMRs as small as 15 ha positively affected medium-bodied fishes. This finding provides reassurance for regions that have invested in small-scale community-managed NTMRs. However, management strategies that integrate sound coastal land-use practices to conserve adjacent reef fish habitat, strategic NTMR placement, and establishment of larger NTMRs will be crucial for maintaining biodiversity and fisheries.     

Community-wide ramifications of an associational refuge on shallow rocky reefs

Little attention has been given to associational refuges in ecology, despite their potential for maintaining species diversity and supporting higher trophic levels. Here I show how the colonial anemone, Corynactis californica, creates a refuge for benthic macroalgae and invertebrate fish prey on intensively grazed shallow rocky reefs in the Santa Barbara Channel off southern California, U.S.A. On reefs heavily grazed by sea urchins, benthic macroalgae and invertebrate fish prey were relatively more abundant among Corynactis colonies than adjacent areas lacking the anemone. Results from field experiments showed that Corynactis facilitated the recruitment of macroalgae and tubicolous amphipods in "urchin-barren" areas subjected to intensive grazing. In areas forested by giant kelp (Macrocystis pyrifera), where grazing intensity from urchins was low, Corynactis suppressed algal recruitment but facilitated tubicolous amphipods. A manipulation of fish and sea urchins suggested that grazing by urchins, as opposed to predation from fish (primarily surfperch Embiotocidae), suppressed tubicolous amphipods, and this activity was hindered by the presence of Corynactis. In systems where human activity has intensified herbivory, associational refuges may maintain species diversity and support higher trophic levels.     

Associations between the abundance of piscivorous fishes and their prey on coral reefs: implications for prey-fish mortality

Few studies have examined predator-prey relationships in diverse communities such as those found on coral reefs. Here we examined patterns of spatial and temporal association between the local abundance of predator and prey fishes at Lizard Island on the Great Barrier Reef, Australia. We predicted that the nature of this association would have implications for patterns of prey-fish mortality. Strong positive relationships between prey and piscivore abundance were found throughout the study. Greater densities of predators and of prey were found on patch-reef habitats, compared with contiguous reef-slope habitats. Declines in prey-fish abundance on patch reefs were density-dependent and correlated with the densities of predators. The relative roles of recruitment and piscivore movement in determining patterns of predator and prey abundance were assessed from surveys of recruit densities and an intensive programme of tagging two species of rock-cod, Cephalopholis cyanostigma and C. boenak (Serranidae), over 2 years. Patterns of recruitment explained little of the variation in the abundance and distribution of piscivorous fish. If movement explains large-scale patterns of distribution, this was not evident from the tagging study. The two rock-cod species were highly sedentary, with individuals on patch reefs seldom moving among reefs. Individuals on reef slopes were also highly site-attached, although they moved greater distances than those on patch reefs. Although the mechanisms responsible remain to be determined, this study demonstrated strong associations between the abundance of piscivorous fish and their prey on coral reefs. This relationship appeared to be an important factor in producing density-dependent declines in the abundance of prey. Received: 30 April 2000 / Accepted: 22 September 2000     

Patterns in fish response to seagrass bed loss at the southern Ryukyu Islands, Japan

An extensive seagrass bed on a fringing coral reef at Amitori Bay (southern Ryukyu Islands) disappeared completely in 2009 after a typhoon. Seagrass bed loss had a significant negative influence on not only seagrass bed residents but also commercially important coral reef fishes that utilize seagrass beds as nurseries or feeding grounds. With seagrass bed loss, mean species’ richness and densities of overall seagrass bed fishes per transect decreased by more than 75 and 85%, respectively. Most of the affected fishes were benthivores, piscivores, detritivores, and herbivores. Of 21 dominant species, 13 disappeared completely and 4 showed severe reductions in densities following seagrass bed loss, whereas the densities of 4 bottom-dwelling gobies did not change significantly. Thus, this study demonstrated that most seagrass bed fishes lack the ability to adapt to seagrass habitat loss, suggesting that increasing global seagrass loss will cause serious reductions in seagrass-associated fishes and fishery resources.     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

Trophic cascades result in large-scale coralline algae loss through differential grazer effects

Removal of predators can have strong indirect effects on primary producers through trophic cascades. Crustose coralline algae (CCA) are major primary producers worldwide that may be influenced by predator removal through changes in grazer composition and biomass. CCA have been most widely studied in Caribbean and temperate reefs, where cover increases with increasing grazer biomass due to removal of competitive fleshy algae. However, each of these systems has one dominant grazer type, herbivorous fishes or sea urchins, which may not be functionally equivalent. Where fishes and sea urchins co-occur, fishing can result in a phase shift in the grazing community with subsequent effects on CCA and other substrata. Kenyan reefs have herbivorous fishes and sea urchins, providing an opportunity to determine the relative impacts of each grazer type and evaluate potential human-induced trophic cascades. We hypothesized that fish benefit CCA, abundant sea urchins erode CCA, and that fishing indirectly reduces CCA cover by removing sea urchin predators. We used closures and fished reefs as a large-scale, long-term natural experiment to assess how fishing and resultant changes in communities affect CCA abundance. We used a short-term caging experiment to directly test the effects of grazing on CCA. CCA cover declined with increasing fish and sea urchin abundance, but the negative impact of sea urchin grazing was much stronger than that of fishes. Abundant sea urchins reduced the CCA growth rate to almost zero and prevented CCA accumulation. A warming event (El Ni?o Southern Oscillation, ENSO) occurred during the 18-year study and had a strong but short-term positive effect on CCA cover. However, the effect of the ENSO on CCA was lower in magnitude than the effect of sea urchin grazing. We compare our results with worldwide literature on bioerosion by fishes and sea urchins. Grazer influence depends on whether benefits of fleshy algae removal outweigh costs of grazer-induced bioerosion. However, the cost-benefit ratio for CCA appears to change with grazer type, grazer abundance, and environment. In Kenya, predator removal leads to a trophic cascade that is expected to reduce net calcification of reefs and therefore reduce reef stability, growth, and resilience.     

Lag Effects in the Impacts of Mass Coral Bleaching on Coral Reef Fish, Fisheries, and Ecosystems

Abstract:  Recent episodes of coral bleaching have led to wide-scale loss of reef corals and raised concerns over the effectiveness of existing conservation and management efforts. The 1998 bleaching event was most severe in the western Indian Ocean, where coral declined by up to 90% in some locations. Using fisheries-independent data, we assessed the long-term impacts of this event on fishery target species in the Seychelles, the overall size structure of the fish assemblage, and the effectiveness of two marine protected areas (MPAs) in protecting fish communities. The biomass of fished species above the size retained in fish traps changed little between 1994 and 2005, indicating no current effect on fishery yields. Biomass remained higher in MPAs, indicating they were effective in protecting fish stocks. Nevertheless, the size structure of the fish communities, as described with size-spectra analysis, changed in both fished areas and MPAs, with a decline in smaller fish (<30 cm) and an increase in larger fish (>45 cm). We believe this represents a time-lag response to a reduction in reef structural complexity brought about because fishes are being lost through natural mortality and fishing, and are not being replaced by juveniles. This effect is expected to be greater in terms of fisheries productivity and, because congruent patterns are observed for herbivores, suggests that MPAs do not offer coral reefs long-term resilience to bleaching events. Corallivores and planktivores declined strikingly in abundance, particularly in MPAs, and this decline was associated with a similar pattern of decline in their preferred corals. We suggest that climate-mediated disturbances, such as coral bleaching, be at the fore of conservation planning for coral reefs.     

Stable C and N isotope analysis elucidated the importance of zooplankton in a tropical seagrass bed of Santiago Island,Northwestern Philippines

The role of zooplankton in a tropical seagrass ecosystem was investigated in milkfish farms pollution-impacted and -unimpacted seagrass beds in Santiago Island coral reefs, Northwestern Philippines. The aim was to compare between the two sites: (1) abiotic factors and zooplankton community parameters, and (2) the trophic structure using C and N stable isotopes. Low water (98–119?mV) and sediment (–121 to ?138?mV) Oxidation Reduction Potential values indicated a reducing environment in the impacted site. Zooplankton in the impacted site showed the typical community response to eutrophication (low diversity, but high total abundance due to the dominance of the cyclopoid copepod Oithona oculata), generally few elevated δ¹⁵N values, but a significant shift towards depleted ¹³C due to the organic enrichment of fish-farm feeds. Apart from suggesting a highly complex food web with POM and zooplankton as main food sources in the unimpacted site, the Bayesian mixing model simulation generated reduced complexity in feeding interactions between basal sources, zooplankton, and fish including adults of a key fish species, Siganus fuscescens, in the impacted sites. In this study, C and N stable isotope analysis has clarified the importance of zooplankton as fish prey in a seagrass bed food web.     

Impact of conservation areas on trophic interactions between apex predators and herbivores on coral reefs

Apex predators are declining at alarming rates due to exploitation by humans, but we have yet to fully discern the impacts of apex predator loss on ecosystem function. In a management context, it is critically important to clarify the role apex predators play in structuring populations of lower trophic levels. Thus, we examined the top‐down influence of reef sharks (an apex predator on coral reefs) and mesopredators on large‐bodied herbivores. We measured the abundance, size structure, and biomass of apex predators, mesopredators, and herbivores across fished, no‐take, and no‐entry management zones in the Great Barrier Reef Marine Park, Australia. Shark abundance and mesopredator size and biomass were higher in no‐entry zones than in fished and no‐take zones, which indicates the viability of strictly enforced human exclusion areas as tools for the conservation of predator communities. Changes in predator populations due to protection in no‐entry zones did not have a discernible influence on the density, size, or biomass of different functional groups of herbivorous fishes. The lack of a relationship between predators and herbivores suggests that top‐down forces may not play a strong role in regulating large‐bodied herbivorous coral reef fish populations. Given this inconsistency with traditional ecological theories of trophic cascades, trophic structures on coral reefs may need to be reassessed to enable the establishment of appropriate and effective management regimes. El Impacto de las Áreas de Conservación sobre las Interacciones Tróficas entre los Depredadores Dominantes y los Herbívoros en los Arrecifes de Coral     

A new apparatus for force measurement in marine bioadhesion

In an experimental study on the effect of parrotfish (probably Scarus taeniurus) grazing on the structure of benthic reef communities, fishes in densities of 0.6 to 1.5 parrotfish per m² or 9 to 17 g wet weight of fish per m² of feeding surface were found to have an optimum effect, resulting in the greatest benthic species richness and biomass on 2-dimensional surfaces. The presence of refuges (3-dimensional habitats), however, has a greater impact on bemthic community structure (number of species and biomass) than does just the density of parrotfish in such an experimental system. Coral recruitment is enhanced by the presence of refuges and, like coralline algae, is more successful under increased grazing pressure. These optimum densities of parrotfishes relate well to observed field densities where, in a collection from a Hawaiian patch reef, there were 1.1 fish or 10.8 g wet weight of parrotfish per square meter of collection area. The success of coralline algae and corals under high grazing pressure may have important consequences for the stability and structure of modern coral reefs.     

Importance of macroalgal fields as coral reef fish nursery habitat in north-west Australia

Macroalgal fields are a feature of the shallow tropical benthos, yet their importance for coral reef fish population dynamics remains poorly understood. The abundance of fish recruits was recorded using underwater visual census at six macroalgal and 11 coral reef sites in the Montebello and Barrow Islands. Surveys identified 6,935 individual recruit fish from 105 species, 54 genera and 20 families. Of these, 1,401 recruits from 48 species, 31 genera and 14 families were observed in macroalgal sites. Sixteen of the 105 recruit species (15.2 %) were observed exclusively at macroalgal sites. Forty-two (87.5 %) of these species have been observed as adults on adjacent coral reefs. Species composition of fish recruits differed significantly between the two habitats. Corallivore, small omnivore and zooplanktivore recruits had significantly higher numbers in the coral sites, while the results clearly demonstrate that juveniles, within the genera Lethrinus and Choerodon, as well as large algal croppers, are predominantly found at macroalgal (74–100 %) rather than coral-dominated sites. High-canopy macroalgae cover was positively correlated with abundance of these taxa, particularly Lethrinids (r ² = 0.40). This study is the first to highlight the important attributes of tropical macroalgal fields and suggests that they have a similar role to seagrass meadows as essential juvenile habitat, thus warranting greater attention in conservation planning and ecological studies.     

Body condition of predatory fishes linked to the availability of sandeels

Lesser sandeels Ammodytes marinus are eaten by a range of predatory fishes including commercially fished species, but are also exploited at large scale by industrial fisheries. Is availability of sandeels, as key prey source, linked to the body condition of predatory fishes? In the North Sea, the largest sandeel biomass is concentrated in the Dogger Bank region. Here we studied predator–sandeel interactions at two sites differing widely in sandeel abundance and local sandeel fishing effort. Surveys took place in 2004, 2005, and 2006, years when local sandeel densities observed at these sites were low, intermediate, and high, respectively. Five predator species––whiting, lesser weever, grey gurnard, plaice, and haddock––showed better body condition indices in either the years or study area (or both) characterised by higher local sandeel densities, when compared to sandeel-poorer conditions. Moreover, whiting, weever, and gurnard condition was better for those individuals actually observed to have eaten sandeels (based on stomach contents) than for those that had not. As body condition relates to growth, reproduction, and survival, predators in sandeel-rich conditions may be inferred to have a higher fitness. These links between sandeel availability, sandeel consumption, and predator condition hint that, if large-scale localised depletions of sandeels were to occur, negative indirect effects on predatory fish might become apparent, underlining the importance of considering the sandeel fishery in an ecosystem context.     

Ichthyofauna of seagrass meadows along the Caribbean coast of Panamá and in the Gulf of Mexico: Composition,structure and community ecology

Collections of fishes from seagrass meadows along the coast of the Republic of Panamá and the Canal Zone during 1974–1975 revealed that juveniles of reefassociated predators are common in this habitat. There are also important diurnal changes in species composition: members of the families Pomadasyidae, Lutjanidae, Sciaenidae, Apogonidae and Muraenidae were noticeably more abundant in the seagrass meadows at night. Although there were significant differences in species composition and abundance among sampling sites, seasonal differences within sites appeared to be insignificant. The ichthyofauna of the Panamanian seagrass beds is significantly richer than that of similar habitats which have been studied in the Gulf of Mexico. This difference in species richness appears to be due in part to the influence of nearby coral reefs, which add an additional structural dimension to the habitat.     

Predicting the Vulnerability of Tropical Reef Fishes to Exploitation with Phylogenies and Life Histories

Abstract: Fishing has led to local extirpations of reef fishes. For conservation and management purposes, it is important to identify all those species that are vulnerable to fishing, but this cannot be done using a priori assessments or by describing trends in abundance because the necessary scientific resources are not available. Thus the predictions of vulnerability that provide the basis for conservation action will have to be made with existing data or data that can be acquired rapidly before further extirpations occur. The life histories of species may determine their responses to exploitation, and we describe how an easily measured parameter, maximum observed size, is related to population trends of exploited fishes on coral reefs. Using a phylogenetic comparative approach, we demonstrated that species of grouper (Epinephelinae), snapper (Lutjanidae), and parrotfish (Scaridae) that decreased in abundance more than their nearest phylogenetic relative had greater maximum size. Our results suggest that one can predict the vulnerability of reef fishes to exploitation based on responses of their relatives. The quality of the prediction was good for the intensively fished groupers and snappers but poor for the lightly fished parrotfishes. Our approach may help proactive conservationists and fishery managers identify and conserve vulnerable species in new, developing, or lightly exploited fisheries, thereby reducing their reliance on reactive management methods.     

Indirect effects of an exploited predator on recruitment of coral-reef fishes

The more ecologists examine the role of trait-mediated indirect interactions (TMIIs), especially in regulating predator-prey interactions, the more we recognize their fundamental role in structuring food webs. However, most empirical evidence for TMIIs comes from studies that are either conducted in laboratory or mesocosm venues or are restricted to simple food webs involving lower trophic-level animals. Here, I quantified the direct and indirect effects of interactions between high-level vertebrate predators on their vertebrate prey using a field experiment. Specifically, I tested how varying densities of a large-bodied, top predator (Nassau grouper; Epinephelus striatus) affected persistence, growth, and behavior of two smaller-bodied, intermediate predators (coney and graysby groupers; Cephalopholis fulva and C. cruentata) on 20 isolated patch reefs in the Bahamas. Large-bodied groupers are capable of consuming their smaller-bodied counterparts, and previous observational studies have indicated that local abundances of these groupers are negatively correlated. I measured the effects of interactions among groupers on lower trophic-level prey by quantifying recruitment of coral-reef fishes to the reefs. The field experiment demonstrated a strong trophic cascade that was entirely mediated by modified behavior of the intermediate predators. These results indicate that indirect, nonlethal interactions in natural systems can have strong cascading effects even at high trophic levels and in high-diversity food webs. Incorporating the complexity of such indirect effects into fisheries management may improve the sustainability of fished populations and strengthen marine conservation efforts; however these results also indicate that the effects of fishing are complex and difficult to predict.     

Variability in nursery function of tropical seagrass beds during fish ontogeny: timing of ontogenetic habitat shift

Seagrass beds are often considered to be important nurseries for coral reef fish, yet the effectiveness of these nursery functions (refuge and food availability) at different juvenile stages is poorly understood. To understand how the demands of juvenile fish on seagrass nursery functions determines the timing of ontogenetic habitat shifts from seagrass beds to coral reefs, we conducted visual transect survey and field tethering and caging experiments on three different sizes of the coral reef fish Pacific yellowtail emperor (Lethrinus atkinsoni) during its juvenile tenure in seagrass beds at Ishigaki Island, southern Japan. The study showed that although the number of individual L. atkinsoni juveniles decreased by >90 % during their stay in the seagrass nursery, the shelter and/or food availability functions of the nursery, at least for a juvenile size of approximately 5 cm total length (TL), provided the best survival and growth option. The timing of ontogenetic migration to coral reefs of larger fish (>8 cm TL) was attributed to foraging efficiency for larger food items in different habitats. Overall, the function of the seagrass bed nursery changed with juvenile body size, with marginally higher survival and significantly greater growth rates during early juvenile stages in seagrass beds compared to coral reefs. This would contribute to the enhancement in the number of individuals eventually recruited to adult populations.