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With rapidly increasing rates of contemporary extinction, predicting extinction vulnerability and identifying how multiple stressors drive non-random species loss have become key challenges in ecology. These assessments are crucial for avoiding the loss of key functional groups that sustain ecosystem processes and services. We developed a novel predictive framework of species extinction vulnerability and applied it to coral reef fishes. Although relatively few coral reef fishes are at risk of global extinction from climate disturbances, a negative convex relationship between fish species locally vulnerable to climate change vs. fisheries exploitation indicates that the entire community is vulnerable on the many reefs where both stressors co-occur. Fishes involved in maintaining key ecosystem functions are more at risk from fishing than climate disturbances. This finding is encouraging as local and regional commitment to fisheries management action can maintain reef ecosystem functions pending progress towards the more complex global problem of stabilizing the climate., (© 2011 Blackwell Publishing Ltd/CNRS.)

Background: The decline of coral reefs globally underscores the need for a spatial assessment of their exposure to multiple environmental stressors to estimate vulnerability and evaluate potential counter-measures., Methodology/principal Findings: This study combined global spatial gradients of coral exposure to radiation stress factors (temperature, UV light and doldrums), stress-reinforcing factors (sedimentation and eutrophication), and stress-reducing factors (temperature variability and tidal amplitude) to produce a global map of coral exposure and identify areas where exposure depends on factors that can be locally managed. A systems analytical approach was used to define interactions between radiation stress variables, stress reinforcing variables and stress reducing variables. Fuzzy logic and spatial ordinations were employed to quantify coral exposure to these stressors. Globally, corals are exposed to radiation and reinforcing stress, albeit with high spatial variability within regions. Based on ordination of exposure grades, regions group into two clusters. The first cluster was composed of severely exposed regions with high radiation and low reducing stress scores (South East Asia, Micronesia, Eastern Pacific and the central Indian Ocean) or alternatively high reinforcing stress scores (the Middle East and the Western Australia). The second cluster was composed of moderately to highly exposed regions with moderate to high scores in both radiation and reducing factors (Caribbean, Great Barrier Reef (GBR), Central Pacific, Polynesia and the western Indian Ocean) where the GBR was strongly associated with reinforcing stress., Conclusions/significance: Despite radiation stress being the most dominant stressor, the exposure of coral reefs could be reduced by locally managing chronic human impacts that act to reinforce radiation stress. Future research and management efforts should focus on incorporating the factors that mitigate the effect of coral stressors until long-term carbon reductions are achieved through global negotiations.

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 Niño 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.

The adoption of fisheries closures and gear restrictions in the conservation of coral reefs may be limited by poor understanding of the economic profitability of competing economic uses of marine resources. Over the past 12 years, I evaluated the effects of gear regulation and fisheries closures on per person and per area incomes from fishing in coral reefs of Kenya. In two of my study areas, the use of small-meshed beach seines was stopped after 6 years; one of these areas was next to a fishery closure. In my third study area, fishing was unregulated. Fishing yields on per capita daily wet weight basis were 20% higher after seine-net fishing was stopped. The per person daily fishing income adjacent to the closed areas was 14 and 22% higher than the fishing income at areas with only gear restrictions before and after the seine-net restriction, respectively. Incomes differed because larger fish were captured next to the closed area and the price per weight (kilograms) increased as fish size increased and because catches adjacent to the closure contained fish species of higher market value. Per capita incomes were 41 and 135% higher for those who fished in gear-restricted areas and near-closed areas, respectively, compared with those who fished areas with no restrictions. On a per unit area basis (square kilometers), differences in fishing income among the three areas were not large because fishing effort increased as the number of restrictions decreased. Changes in catch were, however, larger and often in the opposite direction expected from changes in effort alone. For example, effort declined 21% but nominal profits per square kilometer (not accounting for inflation) increased 29% near the area with gear restrictions. Gear restrictions also reduced the cost of fishing and increased the proportion of self-employed fishers., (© 2010 Society for Conservation Biology.)

Global climate change has the potential to substantially alter the production and community structure of marine fisheries and modify the ongoing impacts of fishing. Fish community composition is already changing in some tropical, temperate and polar ecosystems, where local combinations of warming trends and higher environmental variation anticipate the changes likely to occur more widely over coming decades. Using case studies from the Western Indian Ocean, the North Sea and the Bering Sea, we contextualize the direct and indirect effects of climate change on production and biodiversity and, in turn, on the social and economic aspects of marine fisheries. Climate warming is expected to lead to (i) yield and species losses in tropical reef fisheries, driven primarily by habitat loss; (ii) community turnover in temperate fisheries, owing to the arrival and increasing dominance of warm-water species as well as the reduced dominance and departure of cold-water species; and (iii) increased diversity and yield in Arctic fisheries, arising from invasions of southern species and increased primary production resulting from ice-free summer conditions. How societies deal with such changes will depend largely on their capacity to adapt--to plan and implement effective responses to change--a process heavily influenced by social, economic, political and cultural conditions.

Decadal-scale observations of marine reserves suggest that indirect effects on taxa that occur through cascading trophic interactions take longer to develop than direct effects on target species. Combining and analyzing a unique set of long-term time series of ecologic data in and out of fisheries closures from disparate regions, we found that the time to initial detection of direct effects on target species (±SE) was 5.13 ± 1.9 years, whereas initial detection of indirect effects on other taxa, which were often trait mediated, took significantly longer (13.1 ± 2.0 years). Most target species showed initial direct effects, but their trajectories over time were highly variable. Many target species continued to increase, some leveled off, and others decreased. Decreases were due to natural fluctuations, fishing impacts from outside reserves, or indirect effects from target species at higher trophic levels. The average duration of stable periods for direct effects was 6.2 ± 1.2 years, even in studies of more than 15 years. For indirect effects, stable periods averaged 9.1 ± 1.6 years, although this was not significantly different from direct effects. Populations of directly targeted species were more stable in reserves than in fished areas, suggesting increased ecologic resilience. This is an important benefit of marine reserves with respect to their function as a tool for conservation and restoration.

Marine reserves are increasingly recognized as having linked social and ecological dynamics. This study investigates how the ecological performance of 56 marine reserves throughout the Philippines, Caribbean, and Western Indian Ocean (WIO) is related to both reserve design features and the socioeconomic characteristics in associated coastal communities. Ecological performance was measured as fish biomass in the reserve relative to nearby areas. Of the socioeconomic variables considered, human population density and compliance with reserve rules had the strongest effects on fish biomass, but the effects of these variables were region specific. Relationships between population density and the reserve effect on fish biomass were negative in the Caribbean, positive in the WIO, and not detectable in the Philippines. Differing associations between population density and reserve effectiveness defy simple explanation but may depend on human migration to effective reserves, depletion of fish stocks outside reserves, or other social factors that change with population density. Higher levels of compliance reported by resource users was related to higher fish biomass in reserves compared with outside, but this relationship was only statistically significant in the Caribbean. A heuristic model based on correlations between social, cultural, political, economic, and other contextual conditions in 127 marine reserves showed that high levels of compliance with reserve rules were related to complex social interactions rather than simply to enforcement of reserve rules. Comparative research of this type is important for uncovering the complexities surrounding human dimensions of marine reserves and improving reserve management.

Many of the world's coral reefs suffered high coral mortality during the 1998 ENSO, with the highest mortality in the western Indian Ocean (WIO). A meta-analysis of field data on change in coral cover across the 1998 ENSO event was conducted for 36 major reef areas in the WIO, and relationship of the change with the historical sea-surface temperature (SST) variability investigated. WIO reefs were categorized into three major SST groups of differing coral cover change. Cover change was negatively associated with standard deviation (SD) SST until about SD 2.3, with increasing flatness of the SST frequency distributions. It increased with further increase in SD as the SST distributions became strongly bimodal in the Arabian/Persian Gulf area. The study indicates that environmental resistance/tolerance to extreme anomalous events could be predicted and management priorities directed accordingly for a warmer and more variable future climate., (Copyright 2010 Elsevier Ltd. All rights reserved.)

After a long history of overexploitation, increasing efforts to restore marine ecosystems and rebuild fisheries are under way. Here, we analyze current trends from a fisheries and conservation perspective. In 5 of 10 well-studied ecosystems, the average exploitation rate has recently declined and is now at or below the rate predicted to achieve maximum sustainable yield for seven systems. Yet 63% of assessed fish stocks worldwide still require rebuilding, and even lower exploitation rates are needed to reverse the collapse of vulnerable species. Combined fisheries and conservation objectives can be achieved by merging diverse management actions, including catch restrictions, gear modification, and closed areas, depending on local context. Impacts of international fleets and the lack of alternatives to fishing complicate prospects for rebuilding fisheries in many poorer regions, highlighting the need for a global perspective on rebuilding marine resources.

Priorities for conservation, management, and associated activities will differ based on the interplay between nearness of ecosystems to full recovery from a disturbance (pristineness), susceptibility to climate change (environmental susceptibility [ES]), and capacity of human communities to cope with and adapt to change (social adaptive capacity [AC]). We studied 24 human communities and adjacent coral reef ecosystems in 5 countries of the southwestern Indian Ocean. We used ecological measures of abundance and diversity of fishes and corals, estimated reef pristineness, and conducted socioeconomic household surveys to determine the AC of communities adjacent to selected coral reefs. We also used Web-based oceanographic and coral mortality data to predict each site's ES to climate warming. Coral reefs of Mauritius and eastern Madagascar had low ES and consequently were not predicted to be affected strongly by warm water, although these sites were differentiated by the AC of the human community. The higher AC in Mauritius may increase the chances for successful self-initiated recovery and protective management of reefs of this island. In contrast, Madagascar may require donor support to build AC as a prerequisite to preservation efforts. The Seychelles and Kenya had high ES, but their levels of AC and disturbance differed. The high AC in the Seychelles could be used to develop alternatives to dependence on coral reef resources and reduce the effects of climate change. Pristineness weighted toward measures of fish recovery was greatest for Kenya's marine protected areas; however, most protected areas in the region were far from pristine. Conservation priorities and actions with realistic chances for success require knowledge of where socioecological systems lie among the 3 axes of environment, ecology, and society., (©2009 Society for Conservation Biology.)

The ecosystem goods and services provided by coral reefs are critical to the social and economic welfare of hundreds of millions of people, overwhelmingly in developing countries [1]. Widespread reef degradation is severely eroding these goods and services, but the socioeconomic factors shaping the ways that societies use coral reefs are poorly understood [2]. We examine relationships between human population density, a multidimensional index of socioeconomic development, reef complexity, and the condition of coral reef fish populations in five countries across the Indian Ocean. In fished sites, fish biomass was negatively related to human population density, but it was best explained by reef complexity and a U-shaped relationship with socioeconomic development. The biomass of reef fishes was four times lower at locations with intermediate levels of economic development than at locations with both low and high development. In contrast, average biomass inside fishery closures was three times higher than in fished sites and was not associated with socioeconomic development. Sustaining coral reef fisheries requires an integrated approach that uses tools such as protected areas to quickly build reef resources while also building capacities and capital in societies over longer time frames to address the complex underlying causes of reef degradation.

The emerging world crisis created by declining fish stocks poses a challenge to resource users and managers. The problem is particularly acute in poor nations, such as those in East Africa, where fishing is an important subsistence activity but high fishing intensity and use of destructive gear have resulted in declining catches. In this context developing effective management strategies requires an understanding of how fishers may respond to declines in catch. We examined the readiness of 141 Kenyan fishers to stop fishing under hypothetical scenarios of declines in catch and how socioeconomic conditions influenced their decisions. As expected, the proportion of fishers that would exit the fishery increased with magnitude of decline in catch. Fishers were more likely to say they would stop fishing if they were from households that had a higher material style of life and a greater number of occupations. Variables such as capital investment in the fishery and the proportion of catch sold had weak, nonsignificant relationships. Our finding that fishers from poorer households would be less likely to exit a severely declining fishery is consistent with the literature on poverty traps, which suggests the poor are unable to mobilize the necessary resources to overcome either shocks or chronic low-income situations and consequently may remain in poverty. This finding supports the proposition that wealth generation and employment opportunities directed at the poorest fishers may help reduce fishing effort on overexploited fisheries, but successful interventions such as these will require an understanding of the socioeconomic context in which fishers operate.

This study examined trends along a gradient of fishing intensity in an artisanal coral reef fishery over a 10-year period along 75 km of Kenya's most populated coastline. As predicted by Malthusian scenarios, catch per unit effort (CPUE), mean trophic level, the functional diversity of fished taxa, and the diversity of gear declined, while total annual catch and catch variability increased along the fishing pressure gradient. The fishery was able to sustain high (approximately 16 Mg x km(-2) x yr(-1)) but variable yields at high fishing pressure due to the dominance of a few productive herbivorous fish species in the catch. The effect of two separate management strategies to overcome this Malthusian pattern was investigated: fisheries area closure and elimination of the dominant and most "competitive" gear. We found that sites within 5 km of the enforced closure showed significantly lower total catch and CPUE, but increased yield stability and trophic level of catch than predicted by regression models normalized for fishing effort. Sites that had excluded illegal beach seine use through active gear management exhibited increased total catch and CPUE. There was a strong interaction between closure and gear management, which indicates that, for closures to be effective at increasing catch, there must be simultaneous efforts at gear management around the periphery of the closures. We propose that Malthusian effects are responsible for the variation in gear and catch and that active management through reduced effort and reductions in the most competitive gear have the greatest potential to increase the functional and trophic diversity and per-person productivity.

Coral reefs have emerged as one of the ecosystems most vulnerable to climate variation and change. While the contribution of a warming climate to the loss of live coral cover has been well documented across large spatial and temporal scales, the associated effects on fish have not. Here, we respond to recent and repeated calls to assess the importance of local management in conserving coral reefs in the context of global climate change. Such information is important, as coral reef fish assemblages are the most species dense vertebrate communities on earth, contributing critical ecosystem functions and providing crucial ecosystem services to human societies in tropical countries. Our assessment of the impacts of the 1998 mass bleaching event on coral cover, reef structural complexity, and reef associated fishes spans 7 countries, 66 sites and 26 degrees of latitude in the Indian Ocean. Using Bayesian meta-analysis we show that changes in the size structure, diversity and trophic composition of the reef fish community have followed coral declines. Although the ocean scale integrity of these coral reef ecosystems has been lost, it is positive to see the effects are spatially variable at multiple scales, with impacts and vulnerability affected by geography but not management regime. Existing no-take marine protected areas still support high biomass of fish, however they had no positive affect on the ecosystem response to large-scale disturbance. This suggests a need for future conservation and management efforts to identify and protect regional refugia, which should be integrated into existing management frameworks and combined with policies to improve system-wide resilience to climate variation and change.

The hypothesis that herbivory is higher in areas without fishing and will increase the rate at which hard coral communities return to pre-disturbance conditions was tested in and out of the marine protected areas (MPA) of Kenya after the 1998 El Niño Southern Oscillation (ENSO). Herbivory was estimated by assay and biomass methods, and both methods indicated higher herbivory in fishery closures. Despite higher herbivory, the effect of the ENSO disturbance was larger within these closures, with reefs undergoing a temporary transition from dominance by hard and soft coral to a temporary dominance of turf and erect algae that ended in the dominance of calcifying algae, massive Porites, Pocillopora and a few faviids six years after the disturbance. The fished reefs changed the least but had a greater cover of turf and erect algae and sponge shortly after the disturbance. Higher herbivory in the fishery closures reduced the abundance and persistence of herbivore-susceptible erect algae and created space and appropriate substratum for recruiting corals. Nonetheless, other post-settlement processes may have had strong influences such that annual rates of coral recovery were low ( approximately 2%) and not different between the management regimes. Recovery, as defined as and measured by the return to pre-disturbance coral cover and the dominant taxa, was slower in fishery closures than unmanaged reefs.

Nitrogen and phosphorous fertilizers were used to determine their short-term summer effects on algal colonization, abundance, and species composition in moderate herbivory treatments. Secondary succession of algae on coral skeletons was examined in four treatments: an untreated control, a pure phosphate fertilizer, a pure nitrogen fertilizer, and an equal mix of the two fertilizers. Turf algae cover was the only measure of algae abundance to respond significantly to fertilization. Turf cover was three times higher in treatments with added nitrogen when compared with the pure phosphorus treatment. These turfs were dominated by green and cyanobacteria taxa, namely Enteromorpha prolifera, Lyngbya confervoides, and two species of Cladophora. The phosphate treatment was dominated by encrusting corallines and the cyanobacteria L. confervoides, while the controls had the highest cover of frondose brown algae, namely Padina sanctae-crucis and two species of Dictyota. Results indicate that turf algae were co-limited by nitrogen and phosphorus but enrichment appeared to inhibit brown frondose algae that currently dominate these reefs. Number of species was lowest on the pure phosphorus and nitrogen treatments, highest in the controls and intermediate in the mixed treatments, which suggests that diversity is reduced most by an imbalanced nutrient ratio.

Identifying the rates of recovery of fish in no-take areas is fundamental to designing protected area networks, managing fisheries, estimating yields, identifying ecological interactions, and informing stakeholders about the outcomes of this management. Here we study the recovery of coral reef fishes through 37 years of protection using a space-for-time chronosequence of four marine national parks in Kenya. Using AIC model selection techniques, we assessed recovery trends using five ecologically meaningful production models: asymptotic, Ricker, logistic, linear, and exponential. There were clear recovery trends with time for species richness, total and size class density, and wet masses at the level of the taxonomic family. Species richness recovered rapidly to an asymptote at 10 years. The two main herbivorous families displayed differing responses to protection, scarids recovering rapidly, but then exhibiting some decline while acanthurids recovered more slowly and steadily throughout the study. Recovery of the two invertebrate-eating groups suggested competitive interactions over resources, with the labrids recovering more rapidly before a decline and the balistids demonstrating a slower logistic recovery. Remaining families displayed differing trends with time, with a general pattern of decline in smaller size classes or small-bodied species after an initial recovery, which suggests that some species- and size-related competitive and predatory control occurs in older closures. There appears to be an ecological succession of dominance with an initial rapid rise in labrids and scarids, followed by a slower rise in balistids and acanthurids, an associated decline in sea urchins, and an ultimate dominance in calcifying algae. Our results indicate that the unfished "equilibrium" biomass of the fish assemblage > 10 cm is 1100-1200 kg/ha, but these small parks (< 10 km2) are likely to underestimate pre-human influence values due to edge effects and the rarity of taxa with large area requirement, such as apex predators, including sharks.

Marine protected areas (MPAs) have been widely adopted as the leading tool for coral-reef conservation, but resource users seldom accept them , and many have failed to produce tangible conservation benefits [3]. Few studies have objectively and simultaneously examined the types of MPAs that are most effective in conserving reef resources and the socioeconomic factors responsible for effective conservation [4-6]. We simultaneously explored measures of reef and socioeconomic conservation success at four national parks, four comanaged reserves, and three traditionally managed areas in Indonesia and Papua New Guinea. Underwater visual censuses of key ecological indicators [7, 8] revealed that the average size and biomass of fishes were higher in all areas under traditional management and at one comanaged reserve when compared to nearby unmanaged areas. Socioeconomic assessments [6, 9, 10] revealed that this "effective conservation" was positively related to compliance, visibility of the reserve, and length of time the management had been in place but negatively related to market integration, wealth, and village population size. We suggest that in cases where the resources for enforcement are lacking, management regimes that are designed to meet community goals can achieve greater compliance and subsequent conservation success than regimes designed primarily for biodiversity conservation.

We studied the responses of algae, corals, and small fish to elevated inorganic fertilizer, organic matter, and their combination over a 49-day summer period in cages that simulated the coral reef in the remote Glovers reef atoll, Belize. The addition of organic matter reduced while fertilization had no effect on the numbers of herbivorous damsel and parrotfishes. All measures of algal biomass were influenced by fertilization. The combined inorganic and organic enrichment produced the highest algal biomass, which is most likely due to the combined effect of higher nutrients and lower herbivory. The cover of turf and total algae were influenced by all treatments and their interactions and most strongly and positively influenced by fertilization followed by organic matter and the combination of organic matter and inorganic fertilizer. The inorganic and combined treatments were both dominated by two turf algae, Enteromorpha prolifera and Digenia simplex, while the nonfertilized treatments were dominated by brown frondose algae Lobophora variegata, Padina sanctae, and Dictyota cervicornis. The organic matter treatment had greater cover of P. sanctae and D. cervicornis than the untreated control, which was dominated by Lobophora variegata, also the dominant algae on the nearby patch reefs. Crustose corallines grew slowly ( approximately 2.5 mm/49 days) and were not influenced by the treatments when grown on vertical surfaces but decreased on horizontal coral plates in the combined organic matter and fertilization treatment. No mortality occurred for the two coral species that were added to the cages. Porites furcata darkened in the fertilized cages while there was a mix of paling and darkening for a small amount of the coral tissue of Diploria labyrinthiformes. Inorganic fertilization stimulates small filamentous turf algae and Symbiodinium living in coral but inhibits brown frondose algae. Organic matter inhibits small herbivorous fish, L. variegata, and encrusting coralline algae when growing on horizontal surfaces.

The long-term response of coral reefs to climate change depends on the ability of reef-building coral symbioses to adapt or acclimatize to warmer temperatures, but there has been no direct evidence that such a response can occur. Here we show that corals containing unusual algal symbionts that are thermally tolerant and commonly associated with high-temperature environments are much more abundant on reefs that have been severely affected by recent climate change. This adaptive shift in symbiont communities indicates that these devastated reefs could be more resistant to future thermal stress, resulting in significantly longer extinction times for surviving corals than had been previously assumed.

We compared the bleaching and mortality response (BMI) of 19 common scleractinian corals to an anomalous warm-water event in 1998 to determine the degree of variation between depths, sites, and regions. Mombasa corals experienced a greater temperature anomaly than those on the Great Barrier Reef (GBR) sites and this was reflected in the greater BMI response of most taxa. Comparing coral taxa in different sites at the same depth produced high correlation coefficients in the bleaching response in Kenya at 2 m (r=0.86) and GBR at 6 m depth sites (r=0.80) but less in the GBR for shallow 2 m sites (r=0.49). The pattern of taxa susceptibility was remarkably consistent between the regions. Coral taxa explained 52% of the variation in the response of colonies to bleaching between these two regions (Kenya BMI=0.90 GBR BMI+26; F(1,19) - 18.3; p < 0.001; r2 = 0.52). Stylophora and Pocillopora were consistently susceptible while Cyphastrea, Goniopora Galaxea and Pavona were resistant in both regions. Three taxa behaved differently between the two regions; Acropora, and branching Porites were both moderately affected on the GBR but were highly affected in Kenya while the opposite was true for Pavona. These results suggest that a colonies response to bleaching is phylogenetically constrained, emphasizing the importance of features of the host's physiology or morphology in determining the response to thermal stress.

The proliferation of algae on disturbed coral reefs has often been attributed to (1) a loss of large-bodied herbivorous fishes, (2) increases in sea water nutrient concentrations, particularly phosphorus, and (3) a loss of hard coral cover or a combination of these and other factors. We performed replicated small-scale caging experiments in the offshore lagoon of Glovers Reef atoll, Belize where three treatments had closed-top (no large-bodied herbivores) and one treatment had open-top cages (grazing by large-bodied herbivores). Closed-top treatments simulated a reduced-herbivory situation, excluding large fishes but including small herbivorous fishes such as damselfishes and small parrotfishes. Treatments in the closed-top cages included the addition of high phosphorus fertilizer, live branches of Acropora cervicornis and a third unmanipulated control treatment. Colonization, algal biomass and species composition on dead A. palmata "plates" were studied weekly for 50 days in each of the four treatments. Fertilization doubled the concentration of phosphorus from 0.35 to 0.77 microM. Closed-top cages, particularly the fertilizer and A. cervicornis additions, attracted more small-bodied parrotfish and damselfish than the open-top cages such that there was moderate levels of herbivory in closed-top cages. The open-top cages did, however, have a higher abundance of the chemically and morphologically defended erect algal species including Caulerpa cupressoides, Laurencia obtusa, Dictyota menstrualis and Lobophora variegata. The most herbivore-resistant calcareous green algae (i.e. Halimeda) were, however, uncommon in all treatments. Algal biomass increased and fluctuated simultaneously in all treatments over time, but algal biomass, as measured by wet, dry and decalcified weight, did not differ greatly between the treatments with only marginally higher biomass (p < 0.06) in the fertilized compared to open-top cages. Algal species composition was influenced by all treatments with a maximum between-treatment Bray-Curtis similarity of only 29%. The fertilized cages showed rapid colonization by a mixed turf community largely composed of the filamentous brown (Hincksia mitchelliae) and green (Enteromorpha prolifera) species. Algal cover in the fertilized cages leveled at 80% after 20 days compared to less than 50% in the other treatments. There was no evidence that A. cervicornis suppressed algal colonization compared to the unmanipulated controls. Instead, the herbivore susceptible Padina sanctae-crucis was the most abundant algae followed by Jania capillacea in this treatment in contrast to the more chemically defended Dictyota menstrualis that dominated the unmanipulated controls. We conclude that A. cervicornis was not suppressing algae as a group and its loss cannot account for the observed changes in algal abundance in most reefs except for creating space. In contrast, A. cervicornis appears to attract aggressive damselfish that may reduce herbivory by larger herbivores. Phosphorus enrichment can lead to rapid colonization of space by filamentous turf communities but not high biomass and dominance of erect frondose algae within 50 days. Moderate levels of herbivory by large-bodied herbivores promoted moderately herbivore-resistant erect brown and green algae that are commonly reported on disturbed reefs. Consequently, all the studied factors influenced algal communities but seldom as commonly predicted.

During feeding, echinoids remove a large proportion of calcium carbonate in addition to the algae growing on dead coral and are consequently of importance in estimating the turnover of organic and inorganic carbon in coral reefs. Rates of herbivory and the erosion of dead coral substratum, referred to as bioerosion, by the most abundant echinoid species in Kenyan reefs, Echinothrix diadema (Linnaeus), Diadema setosum (Leske), D. savignyi (Michelin) and Echinometra mathaei (de Blainville), were compared in three different reef categories with different histories of fishing and its exclusion. These were reefs: (i) protected within Marine National Parks, which exclude all forms of fishing, coral and shell collection for more than 25 years; (ii) one reef within a Marine Park, which has received protection from fishing activities for 8 years (referred to as 'newly protected' reef); and (iii) unprotected reefs, which experience heavy fishing and some coral collection. The aim was to investigate the grazing and bioerosion activity by the above echinoid species in these reef categories. We surveyed sea urchin population densities and determined their rates of bioerosion and herbivory per individual and square meter. Individual rates of bioerosion and herbivory, of the species D. setosum, D. savignyi and E. diadema were estimated from laboratory gut content analysis and gut evacuation experiments in the field, using elevated underwater cages. Individual rates of bioerosion and herbivory of E. mathaei were obtained from a previous field study [J. Exp. Mar. Biol. Ecol. 147 (1991) 121]. Sea urchin bioerosion was greater than herbivory for all studied species and proportional to the body size of the sea urchin species. The large-bodied E. diadema exhibited the highest bioerosion and herbivory rates (5.5+/-0.9 and 2.2+/-0.3 g individual(-1) day(-1), respectively) followed by D. setosum (1.8+/-0.3 and 1.1+/-0.2 g individual(-1) day(-1)) and D. savignyi (0.7+/-0.2 and 0.4+/-0.1 g individual(-1) day(-1)). Highest sea urchin densities were recorded at unprotected reefs (6.2+/-1.5 individual m(-2)), and therefore, bioerosion and herbivory by sea urchins were also highest in this reef category (1180+/-230 g CaCO(3) m(-2) year(-1) and 450+/-77 g algae m(-2) year(-1)). Protected reefs recorded 20 times lower sea urchin bioerosion and herbivory rates (50.3+/-25.8 g CaCO(3) m(-2) year(-1) and 20.7+/-10.4 g algae m(-2) year(-1)), due to the low sea urchin population densities in these reefs (0.06+/-0.01 individual m(-2)). The newly protected reef, with intermediate number of sea urchins (1.2+/-0.1 individual m(-2)), had intermediate rates of sea urchin bioerosion and herbivory (711+/-157 g CaCO(3) m(-2) year(-1) and 299+/-63 g algae m(-2) year(-1)). These findings suggest that echinoids are important in the carbon cycle and reef development, and that fishing can influence these ecological processes.

A 194-year annual record of skeletal delta(18)O from a coral growing at Malindi, Kenya, preserves a history of sea surface temperature (SST) change that is coherent with instrumental and proxy records of tropical Pacific climate variability over interannual to decadal periods. This variability is superimposed on a warming of as much as 1.3 degrees C since the early 1800s. These results suggest that the tropical Pacific imparts substantial decadal climate variability to the western Indian Ocean and, by implication, may force decadal variability in other regions with strong El Nino-Southern Oscillation teleconnections.

Large differences in community structure of sea urchins and finfish have been observed in Kenyan reef lagoons. Differences have been attributed to removal of finfish predators through human fishing activities. This study attempted to determine (i) the major sea urchin finfish predators, (ii) the effect of predation on sea-urchin community structure, and (iii) the possible effect of sea urchin increases and finfish decreases on the lagoonal substrate. Six reefs, two protected and four unprotected, were compared for differences in finfish abundance, sea urchin abundance and diversity and substrate cover, diversity and complexity. Comparisons between protected and unprotected reefs indicated that finfish populations were ca. 4 x denser in protected than unprotected reefs. Sea urchin populations were >100 x denser and predation rates on a sea urchin, Echinometra mathaei, were 4 x lower in unprotected than in protected reefs. The balistidae (triggerfish) was the single sea-urchin finfish predator family which had a higher population density in protected than in unprotected reefs. Balistid density was positively correlated with predation rates on tethered E. mathaei (r=0.88; p<0.025) and negatively correlated with total sea-urchin density (r=-0.89; p<0.025) on the six reefs. We conclude from observations that the balistids Balistaphus undulatus and Rhinecanthus aculeatus are the dominant sea-urchin predators. The sea-urchin assemblage had its greatest diversity and species richness at intermediate predation rates and low to intermediate sea-urchin densities. At low predation rates and high sea-urchin density E. mathaei dominated the assemblage's species composition. Preferential predation on the competitive dominant maintains the assemblage's diversity, supporting the compensatory mortality hypothesis (Connell 1978) of coral reef diversity. Protected reefs had greater cover of hard coral, calcareous and coralline algae, and greater substrate diversity and topographic complexity than unprotected reefs which had greater algal turf and sponge cover. Coral cover and topographic complexity were negatively correlated with total sea urchin density. Although experimentation is lacking, these substrate changes may be due to the switch from finfish to sea-urchins as consumers which results from overfishing of finfish. Removal of top invertebrate-eating carnivores appears to have cascading effects on the entire coral reef ecosystem.

Coexistence between the coral reef inhabiting sea urchins Echinometra mathaei, Diadema savignyi and D. setosum was studied by comparing differences in body morphology, distribution, diet, susceptibility to predators, intra- and interspecific competition and settlement. The three species share similar diets and broad within-habitat distributions but differ in their microspatial preferences. E. mathaei is the smallest species, has the highest settlement rates and lives territorially within small burrows or crevices. D. savignyi is intermediate in size and lives frequently in intermediate size crevices or occassionally in social groups. D. setosum is the largest species and occassionally lives in large crevices or more frequently in social groups. Both Diadema have similarily low settlement rates. Competition experiments showed that E. mathaei was consistently the top competitor for crevice space. Diadema species shared larger crevices but competition occured within smaller crevices and was frequently won by the largest individual, regardless of species. D. savignyi may be the top competitor for crevice space between the Diadema species due to a reduced spine length/test size ratio which gives it a larger test for the same crevice size requirement. Predation rates were high for E. mathaei and low for both Diadema species. Coexistence is mediated by predation on the competitive-dominant while predation coupled with different body morphologies and behavior allows spatial resource partitioning of the reef's variable topography. Consequently, the three variables of predation, topographic complexity and differing body shapes create the observed species diversity. A reduction in predators due to stochastic fluctuations or from fishing pressure can lead to E. mathaei population increases and competitive exclusion of Diadema.