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The amount of ocean protected from fishing and other human impacts has often been used as a metric of conservation progress. However, protection efforts have highly variable outcomes that depend on local conditions, which makes it difficult to quantify what coral reef protection efforts to date have actually achieved at a global scale. Here, we develop a predictive model of how local conditions influence conservation outcomes on ~2,600 coral reef sites across 44 ecoregions, which we used to quantify how much more fish biomass there is on coral reefs compared to a modeled scenario with no protection. Under the assumptions of our model, our study reveals that without existing protection efforts there would be ~10% less fish biomass on coral reefs. Thus, we estimate that coral reef protection efforts have led to approximately 1 in every 10 kg of existing fish biomass., Competing Interests: Competing interests statement:The authors declare no competing interest.

Scientific advances in environmental data coverage and machine learning algorithms have improved the ability to make large-scale predictions where data are missing. These advances allowed us to develop a spatially resolved proxy for predicting numbers of tropical nearshore marine taxa. A diverse marine environmental spatial database was used to model numbers of taxa from ∼1000 field sites, and the predictions were applied to all 7039 6.25-km 2 reef cells in 9 ecoregions and 11 nations of the western Indian Ocean. Our proxy for total numbers of taxa was based on the positive correlation (r 2  = 0.24) of numbers of taxa of hard corals and 5 highly diverse reef fish families. Environmental relationships indicated that the number of fish species was largely influenced by biomass, nearness to people, governance, connectivity, and productivity and that coral taxa were influenced mostly by physicochemical environmental variability. At spatial delineations of province, ecoregion, nation, and strength of spatial clustering, we compared areas of conservation priority based on our total species proxy with those identified in 3 previous priority-setting reports and with the protected area database. Our method identified 119 locations that fit 3 numbers of taxa (hard coral, fish, and their combination) and 4 spatial delineations (nation, ecoregion, province, and reef clustering) criteria. Previous publications on priority setting identified 91 priority locations of which 6 were identified by all reports. We identified 12 locations that fit our 12 criteria and corresponded with 3 previously identified locations, 65 that aligned with at least 1 past report, and 28 that were new locations. Only 34% of the 208 marine protected areas in this province overlapped with identified locations with high numbers of predicted taxa. Differences occurred because past priorities were frequently based on unquantified perceptions of remoteness and preselected priority taxa. Our environment-species proxy and modeling approach can be considered among other important criteria for making conservation decisions., (© 2024 The Authors. Conservation Biology published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.)

Marine reserves are known to impact the biomass, biodiversity, and functions of coral reef fish communities, but the effect of protective management on fish traits is less explored. We used a time-series modelling approach to simultaneously evaluate the abundance, biomass, and traits of eight fish families over a chronosequence spanning 44 years of protection. We constructed a multivariate functional space based on six traits known to respond to management or disturbance and affect ecosystem processes: size, diet, position in the water column, gregariousness, reef association, and length at maturity. We show that biomass increased with a log-linear trend over the time-series, but abundance only increased after 20 years of closure, and with more variation among reserves. This difference is attributed to recovery rates being dependent on body sizes. Abundance-weighted traits and the associated multivariate space of the community change is driven by increased proportions over time of the trait categories: 7-15 cm body size; planktivorous; species low in the water column; medium-large schools; and species with high levels of reef association. These findings suggest that the trait compositions emerging after the cessation of fishing are novel and dynamic., (© 2021. The Author(s).)

Identifying the species that are at risk of local extinction in highly diverse ecosystems is a big challenge for conservation science. Assessments of species status are costly and difficult to implement in developing countries with diverse ecosystems due to a lack of species-specific surveys, species-specific data, and other resources. Numerous techniques are devised to determine the threat status of species based on the availability of data and budgetary limits. On this basis, we developed a framework that compared occurrence data of historically exploited reef species in Kenya from existing disparate data sources. Occurrence data from archaeological remains (750-1500CE) was compared with occurrence data of these species catch assessments, and underwater surveys (1991-2014CE). This comparison indicated that only 67 species were exploited over a 750 year period, 750-1500CE, whereas 185 species were landed between 1995 and 2014CE. The first step of our framework identified 23 reef species as threatened with local extinction. The second step of the framework further evaluated the possibility of local extinction with Bayesian extinction analyses using occurrence data from naturalists' species list with the existing occurrence data sources. The Bayesian extinction analysis reduced the number of reef species threatened with local extinction from 23 to 15. We compared our findings with three methods used for assessing extinction risk. Commonly used extinction risk methods varied in their ability to identify reef species that we identified as threatened with local extinction by our comparative and Bayesian method. For example, 12 of the 15 threatened species that we identified using our framework were listed as either least concern, unevaluated, or data deficient in the International Union for the Conservation of Nature red list. Piscivores and macro-invertivores were the only functional groups found to be locally extinct. Comparing occurrence data from disparate sources revealed a large number of historically exploited reef species that are possibly locally extinct. Our framework addressed biases such as uncertainty in priors, sightings and survey effort, when estimating the probability of local extinction. Our inexpensive method showed the value and potential for disparate data to fill knowledge gaps that exist in species extinction assessments., Competing Interests: The authors have declared that no competing interests exist.

Coral reefs provide ecosystem goods and services for millions of people in the tropics, but reef conditions are declining worldwide. Effective solutions to the crisis facing coral reefs depend in part on understanding the context under which different types of conservation benefits can be maximized. Our global analysis of nearly 1,800 tropical reefs reveals how the intensity of human impacts in the surrounding seascape, measured as a function of human population size and accessibility to reefs ("gravity"), diminishes the effectiveness of marine reserves at sustaining reef fish biomass and the presence of top predators, even where compliance with reserve rules is high. Critically, fish biomass in high-compliance marine reserves located where human impacts were intensive tended to be less than a quarter that of reserves where human impacts were low. Similarly, the probability of encountering top predators on reefs with high human impacts was close to zero, even in high-compliance marine reserves. However, we find that the relative difference between openly fished sites and reserves (what we refer to as conservation gains) are highest for fish biomass (excluding predators) where human impacts are moderate and for top predators where human impacts are low. Our results illustrate critical ecological trade-offs in meeting key conservation objectives: reserves placed where there are moderate-to-high human impacts can provide substantial conservation gains for fish biomass, yet they are unlikely to support key ecosystem functions like higher-order predation, which is more prevalent in reserve locations with low human impacts., Competing Interests: The authors declare no conflict of interest., (Copyright © 2018 the Author(s). Published by PNAS.)

Government-managed marine protected areas (MPAs) can restore small fish stocks, but have been heavily criticized for excluding resource users and creating conflicts. A promising but less studied alternative are community-managed MPAs, where resource users are more involved in MPA design, implementation and enforcement. Here we evaluated effects of government- and community-managed MPAs on the density, size and biomass of seagrass- and coral reef-associated fish, using field surveys in Kenyan coastal lagoons. We also assessed protection effects on the potential monetary value of fish; a variable that increases non-linearly with fish body mass and is particularly important from a fishery perspective. We found that two recently established community MPAs (< 1 km2 in size, ≤ 5 years of protection) harbored larger fish and greater total fish biomass than two fished (open access) areas, in both seagrass beds and coral reefs. As expected, protection effects were considerably stronger in the older and larger government MPAs. Importantly, across management and habitat types, the protection effect on the potential monetary value of the fish was much stronger than the effects on fish biomass and size (6.7 vs. 2.6 and 1.3 times higher value in community MPAs than in fished areas, respectively). This strong effect on potential value was partly explained by presence of larger (and therefore more valuable) individual fish, and partly by higher densities of high-value taxa (e.g. rabbitfish). In summary, we show that i) small and recently established community-managed MPAs can, just like larger and older government-managed MPAs, play an important role for local conservation of high-value fish, and that ii) these effects are equally strong in coral reefs as in seagrass beds; an important habitat too rarely included in formal management. Consequently, community-managed MPAs could benefit both coral reef and seagrass ecosystems and provide spillover of valuable fish to nearby fisheries.

Coral reefs are among the most species-rich and threatened ecosystems on Earth, yet the extent to which human stressors determine species occurrences, compared with biogeography or environmental conditions, remains largely unknown. With ever-increasing human-mediated disturbances on these ecosystems, an important question is not only how many species can inhabit local communities, but also which biological traits determine species that can persist (or not) above particular disturbance thresholds. Here we show that human pressure and seasonal climate variability are disproportionately and negatively associated with the occurrence of large-bodied and geographically small-ranging fishes within local coral reef communities. These species are 67% less likely to occur where human impact and temperature seasonality exceed critical thresholds, such as in the marine biodiversity hotspot: the Coral Triangle. Our results identify the most sensitive species and critical thresholds of human and climatic stressors, providing opportunity for targeted conservation intervention to prevent local extinctions.

Ecological baselines are disappearing and it is uncertain how marine reserves, here called fisheries closures, simulate pristine communities. We tested the influence of fisheries closure age, size and compliance on recovery of community biomass and life-history metrics towards a baseline. We used census data from 324 coral reefs, including 41 protected areas ranging between 1 and 45 years of age and 0.28 and 1430 km(2), and 36 sites in a remote baseline, the Chagos Archipelago. Fish community-level life histories changed towards larger and later maturing fauna with increasing closure age, size and compliance. In high compliance closures, community biomass levelled at approximately 20 years and 10 km(2) but was still only at approximately 30% of the baseline and community growth rates were projected to slowly decline for more than 100 years. In low compliance and young closures, biomass levelled at half the value and time as high compliance closures and life-history metrics were not predicted to reach the baseline. Biomass does not adequately reflect the long-time scales for full recovery of life-history characteristics, with implications for coral reef management., (© 2015 The Author(s).)

Continuing degradation of coral reef ecosystems has generated substantial interest in how management can support reef resilience. Fishing is the primary source of diminished reef function globally, leading to widespread calls for additional marine reserves to recover fish biomass and restore key ecosystem functions. Yet there are no established baselines for determining when these conservation objectives have been met or whether alternative management strategies provide similar ecosystem benefits. Here we establish empirical conservation benchmarks and fish biomass recovery timelines against which coral reefs can be assessed and managed by studying the recovery potential of more than 800 coral reefs along an exploitation gradient. We show that resident reef fish biomass in the absence of fishing (B0) averages ∼1,000 kg ha(-1), and that the vast majority (83%) of fished reefs are missing more than half their expected biomass, with severe consequences for key ecosystem functions such as predation. Given protection from fishing, reef fish biomass has the potential to recover within 35 years on average and less than 60 years when heavily depleted. Notably, alternative fisheries restrictions are largely (64%) successful at maintaining biomass above 50% of B0, sustaining key functions such as herbivory. Our results demonstrate that crucial ecosystem functions can be maintained through a range of fisheries restrictions, allowing coral reef managers to develop recovery plans that meet conservation and livelihood objectives in areas where marine reserves are not socially or politically feasible solutions.

The failure of fisheries management among multispecies coral reef fisheries is well documented and has dire implications for the 100 million people engaged in these small-scale operations. Weak or missing management institutions, a lack of research capacity, and the complex nature of these ecosystems have heralded a call for ecosystem-based management approaches. However, ecosystem-based management of coral reef fisheries has proved challenging due to the multispecies nature of catches and the diversity of fish functional roles. We used data on fish communities collected from 233 individual sites in 9 western Indian Ocean countries to evaluate changes in the site's functional composition and associated life-history characteristics along a large range of fish biomass. As biomass increased along this range, fish were larger and grew and matured more slowly while the abundance of scraping and predatory species increased. The greatest changes in functional composition occurred below relatively low standing stock biomass (<600 kg/ha); abundances of piscivores, apex predators, and scraping herbivores were low at very light levels of fishing. This suggests potential trade-offs in ecosystem function and estimated yields for different management systems. Current fishing gear and area restrictions are not achieving conservation targets (proposed here as standing stock biomass of 1150 kg/ha) and result in losses of life history and ecological functions. Fish in reefs where destructive gears were restricted typically had very similar biomass and functions to young and low compliance closures. This indicates the potentially important role of fisheries restrictions in providing some gains in biomass and associated ecological functions when fully protected area enforcement potential is limited and likely to fail. Our results indicate that biomass alone can provide broad ecosystem-based fisheries management targets that can be easily applied even where research capacity and information is limited. Of particular value, is our finding that current management tools may be used to reach key ecosystem-based management targets, enabling ecosystem-based management in many socioeconomic contexts., (© 2014 Society for Conservation Biology.)

The impact of anthropogenic activity on ecosystems has highlighted the need to move beyond the biogeographical delineation of species richness patterns to understanding the vulnerability of species assemblages, including the functional components that are linked to the processes they support. We developed a decision theory framework to quantitatively assess the global taxonomic and functional vulnerability of fish assemblages on tropical reefs using a combination of sensitivity to species loss, exposure to threats and extent of protection. Fish assemblages with high taxonomic and functional sensitivity are often exposed to threats but are largely missed by the global network of marine protected areas. We found that areas of high species richness spatially mismatch areas of high taxonomic and functional vulnerability. Nevertheless, there is strong spatial match between taxonomic and functional vulnerabilities suggesting a potential win-win conservation-ecosystem service strategy if more protection is set in these locations., (© 2014 John Wiley & Sons Ltd/CNRS.)

There is an increasing need to evaluate the links between the social and ecological dimensions of human vulnerability to climate change. We use an empirical case study of 12 coastal communities and associated coral reefs in Kenya to assess and compare five key ecological and social components of the vulnerability of coastal social-ecological systems to temperature induced coral mortality [specifically: 1) environmental exposure; 2) ecological sensitivity; 3) ecological recovery potential; 4) social sensitivity; and 5) social adaptive capacity]. We examined whether ecological components of vulnerability varied between government operated no-take marine reserves, community-based reserves, and openly fished areas. Overall, fished sites were marginally more vulnerable than community-based and government marine reserves. Social sensitivity was indicated by the occupational composition of each community, including the importance of fishing relative to other occupations, as well as the susceptibility of different fishing gears to the effects of coral bleaching on target fish species. Key components of social adaptive capacity varied considerably between the communities. Together, these results show that different communities have relative strengths and weaknesses in terms of social-ecological vulnerability to climate change.

Globally, fisheries are challenged by the combined impacts of overfishing, degradation of ecosystems and impacts of climate change, while fisheries livelihoods are further pressured by conservation policy imperatives. Fishers' adaptive responses to these pressures, such as exiting from a fishery to pursue alternative livelihoods, determine their own vulnerability, as well as the potential for reducing fishing effort and sustaining fisheries. The willingness and ability to make particular adaptations in response to change, such as exiting from a declining fishery, is influenced by economic, cultural and institutional factors operating at scales from individual fishers to national economies. Previous studies of exit from fisheries at single or few sites, offer limited insight into the relative importance of individual and larger-scale social and economic factors. We asked 599 fishers how they would respond to hypothetical scenarios of catch declines in 28 sites in five western Indian Ocean countries. We investigated how socioeconomic variables at the individual-, household- and site-scale affected whether they would exit fisheries. Site-level factors had the greatest influence on readiness to exit, but these relationships were contrary to common predictions. Specifically, higher levels of infrastructure development and economic vitality - expected to promote exit from fisheries - were associated with less readiness to exit. This may be due to site level histories of exit from fisheries, greater specialisation of fishing households, or higher rewards from fishing in more economically developed sites due to technology, market access, catch value and government subsidies. At the individual and household scale, fishers from households with more livelihood activities, and fishers with lower catch value were more willing to exit. These results demonstrate empirically how adaptive responses to change are influenced by factors at multiple scales, and highlight the importance of understanding natural resource-based livelihoods in the context of the wider economy and society.

We tested the unsustainable fishing hypothesis that species in assemblages of fish differ in relative abundance as a function of their size, growth rates, vagility, trophic level, and diet by comparing species composition in historical bone middens, modern fisheries, and areas closed to fishing. Historical data came from one of the earliest and most enduring Swahili coastal settlements (approximately AD 750-1400). Modern data came from fisheries near the archeological site and intensively harvested fishing grounds in southern Kenya. The areas we sampled that were closed to fishing (closures) were small (<28 km(2) ) and permanent. The midden data indicated changes in the fish assemblage that are consistent with a weak expansion of fishing intensity and the unsustainable fishing hypothesis. Fishes represented in the early midden assemblages from AD 750 to 950 had longer life spans, older age at maturity, and longer generation times than fish assemblages after AD 950, when the abundance of species with longer maximum body lengths increased. Changes in fish life histories during the historical period were, however, one-third smaller than differences between the historical and modern assemblages. Fishes in the modern assemblage had smaller mean body sizes, higher growth and mortality rates, a higher proportion of microinvertivores, omnivores, and herbivores, and higher rates of food consumption, whereas the historical assemblage had a greater proportion of piscivores and macroinvertivores. Differences in fish life histories between modern closures and modern fishing grounds were also small, but the life histories of fishes in modern closures were more similar to those in the midden before AD 950 because they had longer life spans, older age at maturity, and a higher proportion of piscivores and macroinvertivores than the modern fisheries. Modern closures and historical fish assemblages were considerably different, although both contained species with longer life spans., (©2011 Society for Conservation Biology.)

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.)

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.

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.

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.

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.

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.

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.