Predicting which tropical tree species are vulnerable to forest disturbances

Tropical forest management often focuses on a few high‐value timber species because they are thought to be the most vulnerable in logged forests. However, other tree species may be vulnerable to secondary effects of logging, like loss of vertebrate dispersers. We examined vulnerability of tree species to loss of vertebrate dispersers in Mabira, a heavily disturbed tropical rainforest in Uganda. Fruit characteristics and shade tolerance regimes of 269 tree species were compiled. Stem densities of tree species producing fruits of various sizes and having different shade tolerance regimes were computed for Mabira and compared with densities of conspecifics in Budongo, a less disturbed forest with similar floral composition. Seventy per cent of tree species in Mabira are animal‐dispersed, of which 10% are large‐fruited light demanders. These species are the most vulnerable because they rarely recruit beneath adult conspecifics and are exclusively dispersed by large vertebrates, also vulnerable in heavily disturbed forests. Comparison of densities between Mabira and Budongo showed that large‐fruited light demanders had a lower density in Mabira. Other categories of tree species had similar densities in both forests. It is plausible that the low density of large‐fruited light demanders is due to limited recruitment caused by dispersal limitations.     

Mapping tree species vulnerability to multiple threats as a guide to restoration and conservation of tropical dry forests

Understanding the vulnerability of tree species to anthropogenic threats is important for the efficient planning of restoration and conservation efforts. We quantified and compared the effects of future climate change and four current threats (fire, habitat conversion, overgrazing and overexploitation) on the 50 most common tree species of the tropical dry forests of northwestern Peru and southern Ecuador. We used an ensemble modelling approach to predict species distribution ranges, employed freely accessible spatial datasets to map threat exposures, and developed a trait‐based scoring approach to estimate species‐specific sensitivities, using differentiated trait weights in accordance with their expected importance in determining species sensitivities to specific threats. Species‐specific vulnerability maps were constructed from the product of the exposure maps and the sensitivity estimates. We found that all 50 species face considerable threats, with an average of 46% of species’ distribution ranges displaying high or very high vulnerability to at least one of the five threats. Our results suggest that current levels of habitat conversion, overexploitation and overgrazing pose larger threats to most of the studied species than climate change. We present a spatially explicit planning strategy for species‐specific restoration and conservation actions, proposing management interventions to focus on (a) in situ conservation of tree populations and seed collection for tree planting activities in areas with low vulnerability to climate change and current threats; (b) ex situ conservation or translocation of populations in areas with high climate change vulnerability; and (c) active planting or assisted regeneration in areas under high current threat vulnerability but low climate change vulnerability, provided that interventions are in place to lower threat pressure. We provide an online, user‐friendly tool to visualize both the vulnerability maps and the maps indicating priority restoration and conservation actions.     

Variation in population structure and dynamics of montane forest tree species in Ethiopia guide priorities for conservation and research

The greatest extent of Afromontane environments in the world is found in Ethiopia. These areas support exceptional biodiversity, but forest cover and ecological integrity have declined sharply in recent decades. Conservation and management efforts are hampered in part by an inadequate understanding of the basic ecology of major tree species. We investigated population structure and inferred population dynamics from size frequency distributions of 22 forest tree species encountered in montane forests of Ethiopia. We collected new empirical data from four sites in the Bale Mountains, where some of the country's most extensive and least disturbed forests remain, and conducted a systematic review and analysis of all such studies that reported population structure for one or more of these species in Ethiopia. Thirteen widespread montane tree species showed a reverse‐J size distribution, indicating a relatively stable population structure. Six other species had size‐frequency distributions that indicate episodic recruitment and/or removal of certain size classes. Specific causes of these patterns are uncertain: they may involve timber harvesting, herbivory, fire, or natural disturbances, but patterns were inconsistent and locality dependent. For three other tree species, existing data are inadequate for any interpretation of population structure and dynamics. A species of particular conservation concern that emerged from this analysis was Hagenia abyssinica, which was found in all areas to consist only of larger individuals with no recent recruitment. For management and conservation purposes, the species in most urgent need of new research are those with inadequate or inconsistent data, and H. abyssinica.     

Population extinctions in the Italian diurnal lepidoptera: an analysis of possible causes

In depth studies of patterns of extinction are fundamental to understand species vulnerability, in particular when population extinctions are not driven by habitat loss, but related to subtle changes in habitat quality and are due to ‘unknown causes’. We used a dataset containing over 160,000 non-duplicate individual records of occurrence (referred to 280 butterflies and 43 zygenid moths), and their relative extinction data, to carry out a twofold analysis. We identified ecological preferences that influence extinction probability, and we analysed if all species were equally vulnerable to the same factors. Our analyses revealed that extinctions were non-randomly distributed in space and time, as well as across species. Most of the extinctions were recorded in 1901–1950 and, as expected, populations at their range edges were more prone to become extinct for non-habitat-related causes. Ecological traits were not only unequally distributed between extinction and non-extinction events, but also not all ecological features had the same importance in driving population vulnerability. Hygrophilous and nemoral species were the most likely to experience population losses and the most prone to disappear even when their habitat remained apparently unchanged. Species vulnerability depends on both ecological requirements and threat type: in fact, each species showed a distinct pattern of vulnerability, depending on threats. We concluded that the analysis may be an important step to prevent butterfly declines: species that are strongly suffering due to ‘unknown changes’ are in clear and urgent need of more detailed auto-ecological studies.     

Correlates of vulnerability among arthropod species threatened by invasive ants

Invasive species are causing population declines and extinctions of native species worldwide. Correlates of species vulnerability, which help identify at-risk taxa, are not well developed for arthropods, particularly with respect to threats from invasive species. At five sites undergoing invasion by ants in the Hawaiian Islands, we assessed body size, population density, trophic role and provenance (introduced or endemic to the Hawaiian Islands) as potential correlates of vulnerability for 300 arthropod species. Among rare species, provenance was the most important factor associated with absence from invaded plots, with endemic species much more commonly absent. Trophic role was also important, but only when interacting with provenance: endemic carnivores were by far the most vulnerable group, followed by endemic detritivores. For non-rare species, Hawaii endemics were significantly more reduced in invaded plots compared to introduced species. In addition, species that occurred at lower population densities were more vulnerable than those occurring at higher densities. Body size did not correlate with vulnerability for either rare or non-rare species. Despite these trends, there was relatively high variability in responses to invasion among species in many taxonomic orders, as well as among populations of particular species at different sites. While the consideration of additional intrinsic traits might increase predictive ability to some degree (e.g., intrinsic traits only explained 21% of the variation in impact among non-rare species), community-specific extrinsic factors appear to play a large role in influencing outcomes for many species, making prediction substantially more difficult.     

Vulnerability of eastern US tree species to climate change

Climate change is expected to alter the distribution of tree species because of critical environmental tolerances related to growth, mortality, reproduction, disturbances, and biotic interactions. How this is realized in 21st century remains uncertain, in large part due to limitations on plant migration and the impacts of landscape fragmentation. Understanding these changes is of particular concern for forest management, which requires information at an appropriately fine spatial resolution. Here we provide a framework and application for tree species vulnerability to climate change in the eastern United States that accounts for influential drivers of future distributions. We used species distribution models to project changes in habitat suitability at 800 m for 40 tree species that vary in physiology, range, and environmental niche. We then developed layers of adaptive capacity based on migration potential, forest fragmentation, and propagule pressure. These were combined into metrics of vulnerability, including an overall index and spatially explicit categories designed to inform management. Despite overall favorable changes in suitability, the majority of species and the landscape were considered vulnerable to climate change. Vulnerability was significantly exacerbated by projections of pests and pathogens for some species. Northern and high‐elevation species tended to be the most vulnerable. There were, however, some notable areas of particular resilience, including most of West Virginia. Our approach combines some of the most important considerations for species vulnerability in a straightforward framework, and can be used as a tool for managers to prioritize species, areas, and actions.     

Frugivorous butterfly species in tropical forest fragments: correlates of vulnerability to extinction

Insects exhibit a variety of population-level responses to forest fragmentation, ranging from population increase to extinction. However, the biological attributes that underlie differences in extinction vulnerability among insects have been little-studied. Using the frugivorous butterfly community of tropical dry forest in Venezuela, we studied body size, population density and colonization ability as attributes that might underlie the range of responses of insects to forest fragmentation. The study was carried out in a set of forest fragments in the reservoir Lago Guri, formed by the damming of the Caroni River in eastern Venezuela. Results show that larger butterfly species were more vulnerable to extinction from habitat fragments than smaller ones. Rarer species were not more vulnerable to extinction, showing that rarity may not be an important correlate of vulnerability to extinction amongst insects. Contrary to expectation, faster-flying species were more and not less vulnerable to extinction from small habitat fragments. We speculate on the possible reasons for the observed patterns in extinction vulnerability using additional observations on behavioural patterns and larval host plant distributions of some of the butterfly species.     

Geographical adaptation prevails over species‐specific determinism in trees’ vulnerability to climate change at Mediterranean rear‐edge forests

Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short‐term extreme events and gradual environmental changes are quite uncertain across the species’ limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short‐ and long‐term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species’ vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species‐specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species‐specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine‐scale limitations of the climate data generated by global climate models.     

Comparing environmental vulnerability in the montane cloud forest of eastern Mexico: A vulnerability index

The montane cloud forest (MCF) is one of the most threatened ecosystems, in spite of its high strategic value for sustainable development, the role it plays in the hydrological cycle maintenance, and as reservoir of endemic biodiversity. For Mexico, this forest is considered the most threatened terrestrial ecosystem at national level because of land-use changes and the effects of global climate change. To compare and assess the environmental vulnerability in the MCF we measured two physiological traits (stomatal conductance and leaf water potential), four climate variables (air temperature, photosynthetically active radiation, vapor pressure deficit, water availability) and the potential geographic distribution of eleven tree species from this forest. We evaluated stomatal conductance responses using the envelope function method (EFM), and after analyzing these responses we developed a vulnerability index that allowed us to compare the environmental vulnerability among species. We proposed the EFM as a useful tool to assess regional environmental vulnerability by comparing species. Our results showed differential species responses to all the studied variables; however, the vulnerability index allowed us to conclude that the most vulnerable species was Liquidambar styraciflua, and the least vulnerable Persea longipes. We also found that temperatures above 34 °C, and vapor pressure deficit above 2.9 kPa with relative humidity below 30% jeopardized the stomatal conductance performance of all species. We also found leaf water potential as the most influential variable over the studied species followed by vapor pressure deficit, showing that even in the MCF water is a determinant factor for species’ development.     

The effect of logging on fission-fusion behaviour of tree-dwelling bats explored by an agent-based model

Logging is one of the greatest threats to global biodiversity, while forests are one of the most important habitats for bats. Bats that roost in tree cavities require a large number of potential roosts due to their frequent roost switching. However, the density of tree cavities and hollows sufficient to sustain large populations of bat species in forests is unknown. The fission-fusion dynamics of bat groups in forest environment is associated with ritualised dawn swarming behaviour at potential tree cavities that serves to exchange information in a non-centralised decision-making process. We used a computer model based on the swarm algorithm, SkyBat, that resembles this complex process and aimed to determine how population size changes over time when cavity trees are removed from roosting territory of the local population of Leisler's bats (Nyctalus leisleri), which inhabit a forest habitat in Central Europe. Simulations revealed that social bonds between bats, maintained by frequent switching among groups, play an important role in this highly dynamic system. When strong social contact was not considered, reducing the original number of trees with cavities (20 cavities × ha⁻¹) to 50% was still acceptable to bats, but further interventions and/or increased demand for social contact would have led to local extinction of the species. Results suggest that potential bat roosts in mature forest stands should be preserved as much as possible and that non-intensive logging and management can be beneficial to tree-dwelling bats.     

The bigger they come, the harder they fall: body size and prey abundance influence predator-prey ratios

Large carnivores are highly threatened, yet the processes underlying their population declines are still poorly understood and widely debated. We explored how body mass and prey abundance influence carnivore density using data on 199 populations obtained across multiple sites for 11 carnivore species. We found that relative decreases in prey abundance resulted in a five- to sixfold greater decrease in the largest carnivores compared with the smallest species. We discuss a number of possible causes for this inherent vulnerability, but also explore a possible mechanistic link between predator size, energetics and population processes. Our results have important implications for carnivore ecology and conservation, demonstrating that larger species are particularly vulnerable to anthropogenic threats to their environment, especially those which have an adverse affect on the abundance of their prey.     

Genomic signals reveal past evolutionary dynamics of Quercus schottkyana and its response to future climate change

Understanding the past and future evolutionary dynamics of dominant species in a forest is important for guiding decisions for biodiversity conservation, forest management, and vegetation restoration. This study used Quercus schottkyana, a dominant tree in subtropical evergreen broad-leaved forests in southwest China, to investigate the influences of past environmental fluctuations and future changes in climate on the dynamics of tree demographics. Genomic data were obtained for 133 samples of Q. schottkyana from 22 populations using double-digest genotyping by sequencing. The single-nucleotide polymorphisms in the genome showed a uniform distribution. Based on principal component analysis and Admixture analysis, two distinct lineages and one mixed group were identified that corresponded to their geographical distribution. Approximate Bayesian computation analyses provided evidence that the divergence among Q. schottkyana populations could be driven by the collision between the Asian and Indian plates during the Miocene, and that climatic fluctuations in the late Pleistocene led to the introgression. The analysis of genotype-environment relationships showed that annual precipitation and geographic distance were associated with spatial genetic variation. Populations of Q. schottkyana in the northern area of the Jinsha River basin were predicted to be the most vulnerable to future climate change. To increase genetic diversity in the northern Jinsha River basin and to buffer threats from future climate change, managers could use a mixture of local and alien seeds during forest restoration and management. This case study can promote further investigations into assessing how past and future climate change impacts genetic divergence and local adaptation of trees in forests.     

Threatened and Endangered Subspecies with Vulnerable Ecological Traits Also Have High Susceptibility to Sea Level Rise and Habitat Fragmentation

The presence of multiple interacting threats to biodiversity and the increasing rate of species extinction make it critical to prioritize management efforts on species and communities that maximize conservation success. We implemented a multi-step approach that coupled vulnerability assessments evaluating threats to Florida taxa such as climate change, sea-level rise, and habitat fragmentation with in-depth literature surveys of taxon-specific ecological traits. The vulnerability, adaptive capacity, and ecological traits of 12 threatened and endangered subspecies were compared to non-listed subspecies of the same parent species. Overall, the threatened and endangered subspecies showed high vulnerability and low adaptive capacity, in particular to sea level rise and habitat fragmentation. They also exhibited larger home ranges and greater dispersal limitation compared to non-endangered subspecies, which may inhibit their ability to track changing climate in fragmented landscapes. There was evidence for lower reproductive capacity in some of the threatened or endangered taxa, but not for most. Taxa located in the Florida Keys or in other low coastal areas were most vulnerable to sea level rise, and also showed low levels of adaptive capacity, indicating they may have a lower probability of conservation success. Our analysis of at-risk subspecies and closely related non-endangered subspecies demonstrates that ecological traits help to explain observed differences in vulnerability and adaptive capacity. This study points to the importance of assessing the relative contributions of multiple threats and evaluating conservation value at the species (or subspecies) level when resources are limited and several factors affect conservation success.     

Forest fragmentation, synergisms and the impoverishment of neotropical forests

It is well documented that the negative effects of habitat fragmentation are strong enough to promote local as well as regional extinction of canopy and emergent trees in neotropical forests. However, forest fragmentation does not occur alone, but is always associated with other human-induced threats to trees, such as logging, forest burning and hunting of key vertebrate seed dispersers within forest remnants. This association occurs because forest resources are, at least during a certain period, the main income source for local human populations. It is now possible to establish how these threats act in concert causing tree species impoverishment. Based on a predictive model, it is predicted that the most fragmented forest regions have lost or will lose an important part of their tree diversity. New integrated research must urgently test this prediction and investigate how human activities might be regulated in both old and new tropical forest frontiers to avoid species loss. If we fail to do this we will miss the opportunity of proposing sound and efficient guidelines to rescue neotropical forests from species impoverishment.     

Vulnerability of Australian tropical savanna birds to climate change

Assessments of species vulnerability to climate change should increase the effectiveness of interventions in the current decline in biodiversity. Species vulnerability to climate change is a consequence of their sensitivity and adaptive capacity, in combination with their exposure to climate change. We apply a vulnerability assessment framework to 243 bird species inhabiting the tropical savannas of northern Australia. We build on previous vulnerability studies by including detailed data for variables relating to species sensitivity to change (relative abundance, clutch size, sensitivity to fire and distribution area), species adaptive capacity (movement behaviour and dietary breadth) and proportional changes predicted for their geographic range (i.e. exposure to climate change). These are integrated to provide a ranking of vulnerability. Our analysis found that birds of Australian tropical savannas cluster together with high sensitivity, with a few wide‐ranging increasing species with very low sensitivity. Australian tropical savanna birds have a range of adaptive capacities, and the impact of climate change on these species is predicted to be substantial. Two already endangered species are among the most vulnerable. Species largely restricted to Cape York Peninsula (a geographically distinct region) had the greatest overall vulnerability; these species were, in general, sensitive due to small distributions, sensitivity to fire frequency and had a lower capacity for dispersal. It will be important for the future of Australian tropical savanna birds to mitigate ecological threats and maintain extensive areas of suitable habitat to facilitate species dispersal.     

An Amazonian rainforest and its fragments as a laboratory of global change

We synthesize findings from one of the world's largest and longest‐running experimental investigations, the Biological Dynamics of Forest Fragments Project (BDFFP). Spanning an area of ∼ 1000 km² in central Amazonia, the BDFFP was initially designed to evaluate the effects of fragment area on rainforest biodiversity and ecological processes. However, over its 38‐year history to date the project has far transcended its original mission, and now focuses more broadly on landscape dynamics, forest regeneration, regional‐ and global‐change phenomena, and their potential interactions and implications for Amazonian forest conservation. The project has yielded a wealth of insights into the ecological and environmental changes in fragmented forests. For instance, many rainforest species are naturally rare and hence are either missing entirely from many fragments or so sparsely represented as to have little chance of long‐term survival. Additionally, edge effects are a prominent driver of fragment dynamics, strongly affecting forest microclimate, tree mortality, carbon storage and a diversity of fauna. Even within our controlled study area, the landscape has been highly dynamic: for example, the matrix of vegetation surrounding fragments has changed markedly over time, succeeding from large cattle pastures or forest clearcuts to secondary regrowth forest. This, in turn, has influenced the dynamics of plant and animal communities and their trajectories of change over time. In general, fauna and flora have responded differently to fragmentation: the most locally extinction‐prone animal species are those that have both large area requirements and low tolerance of the modified habitats surrounding fragments, whereas the most vulnerable plants are those that respond poorly to edge effects or chronic forest disturbances, and that rely on vulnerable animals for seed dispersal or pollination. Relative to intact forests, most fragments are hyperdynamic, with unstable or fluctuating populations of species in response to a variety of external vicissitudes. Rare weather events such as droughts, windstorms and floods have had strong impacts on fragments and left lasting legacies of change. Both forest fragments and the intact forests in our study area appear to be influenced by larger‐scale environmental drivers operating at regional or global scales. These drivers are apparently increasing forest productivity and have led to concerted, widespread increases in forest dynamics and plant growth, shifts in tree‐community composition, and increases in liana (woody vine) abundance. Such large‐scale drivers are likely to interact synergistically with habitat fragmentation, exacerbating its effects for some species and ecological phenomena. Hence, the impacts of fragmentation on Amazonian biodiversity and ecosystem processes appear to be a consequence not only of local site features but also of broader changes occurring at landscape, regional and even global scales.     

西双版纳片断热带雨林30多年来植物种类组成及种群结构的变化

通过对西双版纳景洪曼仰广的“龙山”片断热带雨雨林30年前后植物区系组成和样方调查资料的对比研究,探讨了30年来该“龙山”片断热带雨林植物区系组成、生活型及群落生态成分和乔木树种的种群变化规律。在植物区系组成上,已确认有种子植物7科53属55种从该片断雨林中消失。在消失的这55种植物中,属于群落顶级成分的有18种,属于耐阴的林下和层间植物有30种,喜阳种类有2种,广生态幅的随遇种有5种。初步得出该龙山热带雨林随着30多年的片断化,8．8％的科,26．8％的属和22．4％的种类消失或被后来的成分替换了。在生活型及群落生态成分变化上,群落中小高位芽及一年生植物相对增加,阳性植物明显增加,阴生（耐阴）植物明显减少,亦即雨林固有成分减少,非雨林成分增加。在乔木树种的种群变化上,过去该片断雨林的优势成份,绝大多数现在仍存在,多数仍在优势种之列,少数优势种衰退了,少数变得更优势。首先消失的种类,或为先锋树种（短命）或为种群数量很少的树种。     

The vulnerability of species to range expansions by predators can be predicted using historical species associations and body size

Climate change threatens species directly through environmental changes and indirectly through its effects on species interactions. We need tools to predict which species are most vulnerable to these threats. Pairwise species associations and body size are simple but promising predictors of the relative impact of species introduced outside of their historical ranges. We examined the vulnerability of 30 fish species to the impacts of three centrarchid predators that are being introduced to lakes north of their historical range boundaries. Species that were negatively associated with each centrarchid in their historical range were more likely to be lost from lakes with centrarchid introductions. Total body length was most important in predicting impact for the most gape-limited predator. At the regional scale, our method identifies those species most vulnerable to introductions facilitated by climate change and can easily be applied to a range of taxa undergoing range expansions.     

Competition‐interaction landscapes for the joint response of forests to climate change

The recent global increase in forest mortality episodes could not have been predicted from current vegetation models that are calibrated to regional climate data. Physiological studies show that mortality results from interactions between climate and competition at the individual scale. Models of forest response to climate do not include interactions because they are hard to estimate and require long‐term observations on individual trees obtained at frequent (annual) intervals. Interactions involve multiple tree responses that can only be quantified if these responses are estimated as a joint distribution. A new approach provides estimates of climate–competition interactions in two critical ways, (i) among individuals, as a joint distribution of responses to combinations of inputs, such as resources and climate, and (ii) within individuals, due to allocation requirements that control outputs, such as demographic rates. Application to 20 years of data from climate and competition gradients shows that interactions control forest responses, and their omission from models leads to inaccurate predictions. Species most vulnerable to increasing aridity are not those that show the largest growth response to precipitation, but rather depend on interactions with the local resource environment. This first assessment of regional species vulnerability that is based on the scale at which climate operates, individual trees competing for carbon and water, supports predictions of potential savannification in the southeastern US.     

The intrinsic vulnerability to fishing of coral reef fishes and their differential recovery in fishery closures

Coral reef fishes differ in their intrinsic vulnerability to fishing and rates of population recovery after cessation of fishing. We reviewed life history-based predictions about the vulnerability of different groups of coral reef fish and examined the empirical evidence for different rates of population recovery inside no-take marine reserves to (1) determine if the empirical data agree with predictions about vulnerability and (2) show plausible scenarios of recovery within fully protected reserves and periodically-harvested fishery closures. In general, larger-bodied carnivorous reef fishes are predicted to be more vulnerable to fishing while smaller-bodied species lower in the food web (e.g., some herbivores) are predicted to be less vulnerable. However, this prediction does not always hold true because of the considerable diversity of life history strategies in reef fishes. Long-term trends in reef fish population recovery inside no-take reserves are consistent with broad predictions about vulnerability, suggesting that moderately to highly vulnerable species will require a significantly longer time (decades) to attain local carrying capacity than less vulnerable species. We recommend: (1) expanding age-based demographic studies of economically and ecologically important reef fishes to improve estimates of vulnerability; (2) long term (20–40 years), if not permanent, protection of no-take reserves to allow full population recovery and maximum biomass export; (3) strict compliance to no-take reserves to avoid considerable delays in recovery; (4) carefully controlling the timing and intensity of harvesting periodic closures to ensure long-term fishery benefits; (5) the use of periodically-harvested closures together with, rather than instead of, permanent no-take reserves.