珊瑚礁生态脆弱性评价--以泰国思仓岛为例

珊瑚礁生态系统受到环境变化、人类活动等各种因素的严重威胁,保护珊瑚礁生态系统是目前全球海洋生态保护的热点,对珊瑚礁开展定量的生态脆弱性评估能够为保护管理对策的制定提供重要科学依据。本研究选取泰国思仓岛作为研究区域,结合空间分析技术建立了具有通用性的珊瑚礁生态脆弱性评估方法。基于ESA模型构建了珊瑚礁生态脆弱性综合指数和评价指标体系,系统分析了思仓岛珊瑚礁脆弱性的来源、构成,并直观展现了脆弱性的区域空间分布。结果表明:思仓岛研究区东北侧的珊瑚礁生态脆弱性大于西南侧,当地珊瑚礁的关键影响因子分别为驳船排污、港口码头、水体透明度等。根据脆弱性评价的结果,提出了当地珊瑚礁保护与修复的空间分区管理对策。本研究为印度-太平洋区系珊瑚礁生态脆弱性评价提供了可行的示例,也为中国的珊瑚礁可持续管理研究提供了借鉴和参照。     

Operationalizing resilience for adaptive coral reef management under global environmental change

Cumulative pressures from global climate and ocean change combined with multiple regional and local‐scale stressors pose fundamental challenges to coral reef managers worldwide. Understanding how cumulative stressors affect coral reef vulnerability is critical for successful reef conservation now and in the future. In this review, we present the case that strategically managing for increased ecological resilience (capacity for stress resistance and recovery) can reduce coral reef vulnerability (risk of net decline) up to a point. Specifically, we propose an operational framework for identifying effective management levers to enhance resilience and support management decisions that reduce reef vulnerability. Building on a system understanding of biological and ecological processes that drive resilience of coral reefs in different environmental and socio‐economic settings, we present an Adaptive Resilience‐Based management (ARBM) framework and suggest a set of guidelines for how and where resilience can be enhanced via management interventions. We argue that press‐type stressors (pollution, sedimentation, overfishing, ocean warming and acidification) are key threats to coral reef resilience by affecting processes underpinning resistance and recovery, while pulse‐type (acute) stressors (e.g. storms, bleaching events, crown‐of‐thorns starfish outbreaks) increase the demand for resilience. We apply the framework to a set of example problems for Caribbean and Indo‐Pacific reefs. A combined strategy of active risk reduction and resilience support is needed, informed by key management objectives, knowledge of reef ecosystem processes and consideration of environmental and social drivers. As climate change and ocean acidification erode the resilience and increase the vulnerability of coral reefs globally, successful adaptive management of coral reefs will become increasingly difficult. Given limited resources, on‐the‐ground solutions are likely to focus increasingly on actions that support resilience at finer spatial scales, and that are tightly linked to ecosystem goods and services.     

An Integrated Coral Reef Ecosystem Model to Support Resource Management under a Changing Climate

Millions of people rely on the ecosystem services provided by coral reefs, but sustaining these benefits requires an understanding of how reefs and their biotic communities are affected by local human-induced disturbances and global climate change. Ecosystem-based management that explicitly considers the indirect and cumulative effects of multiple disturbances has been recommended and adopted in policies in many places around the globe. Ecosystem models give insight into complex reef dynamics and their responses to multiple disturbances and are useful tools to support planning and implementation of ecosystem-based management. We adapted the Atlantis Ecosystem Model to incorporate key dynamics for a coral reef ecosystem around Guam in the tropical western Pacific. We used this model to quantify the effects of predicted climate and ocean changes and current levels of current land-based sources of pollution (LBSP) and fishing. We used the following six ecosystem metrics as indicators of ecosystem state, resilience and harvest potential: 1) ratio of calcifying to non-calcifying benthic groups, 2) trophic level of the community, 3) biomass of apex predators, 4) biomass of herbivorous fishes, 5) total biomass of living groups and 6) the end-to-start ratio of exploited fish groups. Simulation tests of the effects of each of the three drivers separately suggest that by mid-century climate change will have the largest overall effect on this suite of ecosystem metrics due to substantial negative effects on coral cover. The effects of fishing were also important, negatively influencing five out of the six metrics. Moreover, LBSP exacerbates this effect for all metrics but not quite as badly as would be expected under additive assumptions, although the magnitude of the effects of LBSP are sensitive to uncertainty associated with primary productivity. Over longer time spans (i.e., 65 year simulations), climate change impacts have a slight positive interaction with other drivers, generally meaning that declines in ecosystem metrics are not as steep as the sum of individual effects of the drivers. These analyses offer one way to quantify impacts and interactions of particular stressors in an ecosystem context and so provide guidance to managers. For example, the model showed that improving water quality, rather than prohibiting fishing, extended the timescales over which corals can maintain high abundance by at least 5–8 years. This result, in turn, provides more scope for corals to adapt or for resilient species to become established and for local and global management efforts to reduce or reverse stressors.     

The Reef Environment Centralized InFormation System (RECIFS): An integrated geo-environmental database for coral reef research and conservation

Motivation

Host to intricate networks of marine species, coral reefs are among the most biologically diverse ecosystems on Earth. Over the past few decades, major degradations of coral reefs have been observed worldwide, which is largely attributed to the effects of climate change and local stressors related to human activities. Now more than ever, characterizing how the environment shapes the dynamics of the reef ecosystem (e.g., shifts in species abundance, community changes, emergence of locally adapted populations) is key to uncovering the environmental drivers of reef degradation, and developing efficient conservation strategies in response. To achieve these objectives, it is pivotal that environmental data describing the processes driving such ecosystem dynamics, which occur across specific spatial and temporal scales, are easily accessible to coral reef researchers and conservation stakeholders alike.

Main types of variable contained

Multiple environmental variables characterizing various facets of the reef environment, including water chemistry and physics (e.g., temperature, pH, chlorophyll concentration), local anthropogenic pressures (e.g., boat traffic, distance from agricultural or urban areas) and sea currents patterns.

Spatial location and grain

Worldwide reef cells of 5 by 5 km.

Time period and grain

Last 3–4 decades, monthly and yearly resolution.

Major taxa and level of measurement

Environmental data important for coral reefs and associated biodiversity.

Software format

Interactive web application available at https://recifs.epfl.ch .     

Climate warming, marine protected areas and the ocean-scale integrity of coral reef ecosystems

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.     

Coral reef conservation in Bali in light of international best practice,a literature review

Bali, Indonesia sits within the coral triangle and is internationally recognised for its high coral reef diversity. The health of Bali’s marine ecosystems has declined in recent decades, and this is thought to be due to threats from climate change, destructive fishing practices, pollution, outbreaks coral eating invertebrates, coral disease and unsustainable tourism. As a response, multiple conservation strategies have been introduced by the island’s communities, non-government organisations and governments, with the aim of preventing further decline, as well as restoring already degraded coral reefs. This literature review provides an in-depth analysis of the tools used to conserve Bali’s coral reefs, and compares them to those used in other countries. In light of international ‘best practice’ in coral reef conservation, this review makes suggestions on how Bali could better conserve its coral reef ecosystems. These include (1) increasing its designation of official Marine Protected Areas (MPAS) and strengthening management of existing ones, (2) creating an MPA network, (3) substantially reducing marine plastic pollution, (4) continuing artificial reef construction in degraded habitats, (5) continuing to develop Bali as an ecotourism destination, (6) increasing engagement in global science to inform marine conservation decision-making, and (7) developing more marine monitoring programmes.     

How much time can herbivore protection buy for coral reefs under realistic regimes of hurricanes and coral bleaching?

Coral reefs have been more severely impacted by recent climate instability than any other ecosystem on Earth. Corals tolerate a narrow range of physical environmental stress, and increases in sea temperature of just 1 °C over several weeks can result in mass coral mortality, often exceeding 95% of individuals over hundreds of square kilometres. Even conservative climate models predict that mass coral bleaching events could occur annually by 2050. Unfortunately, managers of coral‐reef resources have few options available to meet this challenge. Here, we investigate the role that fisheries conservation tools, including the designation of marine reserves, can play in altering future trajectories of Caribbean coral reefs. We use an individual‐based model of the ecological dynamics to test the influence of spatially realistic regimes of disturbance on coral populations. Two major sources of disturbance, hurricanes and coral bleaching, are simulated in contrasting regions of the Caribbean: Belize, Bonaire, and the Bahamas. Simulations are extended to 2099 using the HadGEM1 climate model. We find that coral populations can maintain themselves under all levels of hurricane disturbance providing that grazing levels are high. Regional differences in hurricane frequency are found to cause strikingly different spatial patterns of reef health with greater patchiness occurring in Belize, which has less frequent disturbance, than the Bahamas. The addition of coral bleaching led to a much more homogenous reef state over the seascape. Moreover, in the presence of bleaching, all reefs exhibited a decline in health over time, though with substantial variation among regions. Although the protection of herbivores does not prevent reef degradation it does delay rates of coral loss even under the most severe thermal and hurricane regimes. Thus, we can estimate the degree to which local conservation can help buy time for reefs with values ranging between 18 years in the Bahamas and over 50 years in Bonaire, compared with heavily fished systems. Ultimately, we demonstrate that local conservation measures can benefit reef ecosystem services but that their impact will vary spatially and temporally. Recognizing where such management interventions will either help or fail is an important step towards both achieving sustainable use of coral‐reef resources and maximizing resource management investments.     

珊瑚礁生态系统病毒研究进展

病毒对珊瑚礁生态系统中的生物进化、生物地球化学循环、珊瑚疾病等方面具有重要的生态影响。随着珊瑚礁的全球性退化,病毒在珊瑚礁生态系统中的功能与危害日益显现。综述了珊瑚礁生态系统中病毒的研究现状与进展,包括：（1）珊瑚礁病毒的多样性与分布特征（水体、宿主、核心病毒组）;（2）珊瑚礁病毒的生态功能（感染方式、促进生物进化、生物地球化学循环）;（3）珊瑚礁病毒对全球气候变化的响应（热压力、珊瑚疾病）。总体而言,珊瑚礁生态系统具有极高的病毒多样性,所发现的60个科占已知所有病毒科数量的58%。珊瑚的核心病毒组主要由双链DNA病毒、单链DNA病毒、单链逆转录病毒所组成,珊瑚黏液层对病毒具有富集作用。"Piggyback-the-Winner"（依附-胜利）是病毒在珊瑚礁中主要的生物动力学模式,其可通过水平基因迁移的方式促进礁区生物进化。病毒可通过裂解细菌与浮游藻类的途径参与珊瑚礁的生物地球化学循环,尤其是碳循环与氮循环过程。此外,病毒还具有介导珊瑚热白化与直接引发珊瑚疾病的能力,这会影响珊瑚礁生态系统应对气候变化的适应性与恢复力。基于国际上的研究进展综述,结合南海珊瑚礁生态现状提出以下研究方向,以期促进我国珊瑚礁病毒学的发展：（1）开展南海珊瑚礁中病毒多样性的识别及其时-空分布特征研究;（2）探索病毒对南海珊瑚热白化、珊瑚疾病的介导作用及其与气候变化的关系;（3）揭示病毒对南海珊瑚礁生物地球化学循环的贡献。     

Anticipative management for coral reef ecosystem services in the 21st century

Under projections of global climate change and other stressors, significant changes in the ecology, structure and function of coral reefs are predicted. Current management strategies tend to look to the past to set goals, focusing on halting declines and restoring baseline conditions. Here, we explore a complementary approach to decision making that is based on the anticipation of future changes in ecosystem state, function and services. Reviewing the existing literature and utilizing a scenario planning approach, we explore how the structure of coral reef communities might change in the future in response to global climate change and overfishing. We incorporate uncertainties in our predictions by considering heterogeneity in reef types in relation to structural complexity and primary productivity. We examine 14 ecosystem services provided by reefs, and rate their sensitivity to a range of future scenarios and management options. Our predictions suggest that the efficacy of management is highly dependent on biophysical characteristics and reef state. Reserves are currently widely used and are predicted to remain effective for reefs with high structural complexity. However, when complexity is lost, maximizing service provision requires a broader portfolio of management approaches, including the provision of artificial complexity, coral restoration, fish aggregation devices and herbivore management. Increased use of such management tools will require capacity building and technique refinement and we therefore conclude that diversification of our management toolbox should be considered urgently to prepare for the challenges of managing reefs into the 21st century.     

Shifting paradigms in restoration of the world's coral reefs

Many ecosystems around the world are rapidly deteriorating due to both local and global pressures, and perhaps none so precipitously as coral reefs. Management of coral reefs through maintenance (e.g., marine‐protected areas, catchment management to improve water quality), restoration, as well as global and national governmental agreements to reduce greenhouse gas emissions (e.g., the 2015 Paris Agreement) is critical for the persistence of coral reefs. Despite these initiatives, the health and abundance of corals reefs are rapidly declining and other solutions will soon be required. We have recently discussed options for using assisted evolution (i.e., selective breeding, assisted gene flow, conditioning or epigenetic programming, and the manipulation of the coral microbiome) as a means to enhance environmental stress tolerance of corals and the success of coral reef restoration efforts. The 2014–2016 global coral bleaching event has sharpened the focus on such interventionist approaches. We highlight the necessity for consideration of alternative (e.g., hybrid) ecosystem states, discuss traits of resilient corals and coral reef ecosystems, and propose a decision tree for incorporating assisted evolution into restoration initiatives to enhance climate resilience of coral reefs.     

Simulations of Long-Term Community Dynamics in Coral Reefs - How Perturbations Shape Trajectories

Tropical coral reefs feature extraordinary biodiversity and high productivity rates in oligotrophic waters. Due to increasing frequencies of perturbations – anthropogenic and natural – many reefs are under threat. Such perturbations often have devastating effects on these unique ecosystems and especially if they occur simultaneously and amplify each other''s impact, they might trigger a phase shift and create irreversible conditions.We developed a generic, spatially explicit, individual-based model in which competition drives the dynamics of a virtual benthic reef community – comprised of scleractinian corals and algae – under different environmental settings. Higher system properties, like population dynamics or community composition arise through self-organization as emergent properties. The model was parameterized for a typical coral reef site at Zanzibar, Tanzania and features coral bleaching and physical disturbance regimes as major sources of perturbations. Our results show that various types and modes (intensities and frequencies) of perturbations create diverse outcomes and that the switch from high diversity to single species dominance can be evoked by small changes in a key parameter.Here we extend the understanding of coral reef resilience and the identification of key processes, drivers and respective thresholds, responsible for changes in local situations. One future goal is to provide a tool which may aid decision making processes in management of coral reefs.     

New directions in coral reef microbial ecology

Microbial processes largely control the health and resilience of coral reef ecosystems, and new technologies have led to an exciting wave of discovery regarding the mechanisms by which microbial communities support the functioning of these incredibly diverse and valuable systems. There are three questions at the forefront of discovery: What mechanisms underlie coral reef health and resilience? How do environmental and anthropogenic pressures affect ecosystem function? What is the ecology of microbial diseases of corals? The goal is to understand the functioning of coral reefs as integrated systems from microbes and molecules to regional and ocean‐basin scale ecosystems to enable accurate predictions of resilience and responses to perturbations such as climate change and eutrophication. This review outlines recent discoveries regarding the microbial ecology of different microenvironments within coral ecosystems, and highlights research directions that take advantage of new technologies to build a quantitative and mechanistic understanding of how coral health is connected through microbial processes to its surrounding environment. The time is ripe for natural resource managers and microbial ecologists to work together to create an integrated understanding of coral reef functioning. In the context of long‐term survival and conservation of reefs, the need for this work is immediate.     

Modelling Coral Reef Futures to Inform Management: Can Reducing Local-Scale Stressors Conserve Reefs under Climate Change?

Climate change has emerged as a principal threat to coral reefs, and is expected to exacerbate coral reef degradation caused by more localised stressors. Management of local stressors is widely advocated to bolster coral reef resilience, but the extent to which management of local stressors might affect future trajectories of reef state remains unclear. This is in part because of limited understanding of the cumulative impact of multiple stressors. Models are ideal tools to aid understanding of future reef state under alternative management and climatic scenarios, but to date few have been sufficiently developed to be useful as decision support tools for local management of coral reefs subject to multiple stressors. We used a simulation model of coral reefs to investigate the extent to which the management of local stressors (namely poor water quality and fishing) might influence future reef state under varying climatic scenarios relating to coral bleaching. We parameterised the model for Bolinao, the Philippines, and explored how simulation modelling can be used to provide decision support for local management. We found that management of water quality, and to a lesser extent fishing, can have a significant impact on future reef state, including coral recovery following bleaching-induced mortality. The stressors we examined interacted antagonistically to affect reef state, highlighting the importance of considering the combined impact of multiple stressors rather than considering them individually. Further, by providing explicit guidance for management of Bolinao''s reef system, such as which course of management action will most likely to be effective over what time scales and at which sites, we demonstrated the utility of simulation models for supporting management. Aside from providing explicit guidance for management of Bolinao''s reef system, our study offers insights which could inform reef management more broadly, as well as general understanding of reef systems.     

Community involvement in management for maintaining coral reef resilience and biodiversity in southern Caribbean marine protected areas

Climate change is posing new challenges to conservation because management policies on protected coral reefs are less effective than they were before the current ecosystem degradation. Coral reefs, the most diverse and complex marine ecosystem provide economic services for millions, but are seriously threatened worldwide because reef-building corals are experiencing bleaching phenomena and a steady decline in abundance. The resources of a Marine Protected Area (MPA) in Cartagena, Colombia, are in constant decline, despite a current management plan and on-site staff, urging new conservation actions. A multidisciplinary team gathered to evaluate management effectiveness including biophysical, socioeconomic and governance indicators. Coral cover and fish diversity and abundance were low both inside and outside the MPA, which suggests a limited efficiency of management. Currently, the MPA is a reef with low coral cover and high algae cover as well as large dead coral areas, which are generally signs of highly degraded reef habitats. Live coral cover in the MPA was represented by pioneer coral species such as Agaricia tenuifolia and Porites astreoides. Nonetheless, 35% of the scleractinian species sampled in the area harbored more than one zooxanthellae symbiont, which suggests potential resistance and resilience against coral bleaching. Maintenance of trophic structure and functional diversity is an important endeavor that should be a priority for management in order to allow ecosystem resilience. Social and governance indicators showed low-income levels and few opportunities for communities living in and around the park, low governability, weak communication among stakeholders and with authorities at different levels. As a result, problems related to over exploitation of resources were commonplace in the MPA. These results reflect low adaptive capacity of communities to comply with restrictive conservation rules, showing that establishment of a protected area is a necessary but insufficient condition to guarantee conservation goals. Ignoring the role of local communities only will exacerbate the problems associated with natural resources. Involvement of communities in strategic ecosystems management appears to be a requisite to improve effectiveness of protected areas, and participatory strategies, such as co-management, offer opportunities to improve governability while letting communities adapt to MPA needs.     

Rehabilitation of coral reefs through removal of macroalgae: state of knowledge and considerations for management and implementation

Coral reef ecosystems are under increasing pressure by multiple stressors that degrade reef condition and function. Although improved management systems have yielded benefits in many regions, broad‐scale declines continue and additional practical and effective solutions for reef conservation and management are urgently needed. Ecological interventions to assist or enhance ecosystem recovery are standard practice in many terrestrial management regimes, and they are now increasingly being implemented in the marine environment. Intervention activities in coral reef systems include the control of coral predators (e.g. crown‐of‐thorns starfish), substrate modification, the creation of artificial habitats and the cultivation, transplantation, and assisted recruitment of corals. On many coastal reefs, corals face competition and overgrowth by fleshy macroalgae whose abundance may be elevated due to acute disturbance events, chronic nutrient enrichment, and reduced herbivory. Active macroalgae removal has been proposed and trialed as a management tool to reduce competition between algae and corals and provide space for coral recruitment, in the hope of restoring the spatial dominance of habitat‐forming corals. However, macroalgae removal has received little formal attention as a method of reef restoration. This review synthesizes available knowledge of the ecological role of macroalgae on coral reefs and the potential benefits and risks associated with their active removal.     

Region‐wide temporal and spatial variation in Caribbean reef architecture: is coral cover the whole story?

The architectural complexity of coral reefs is largely generated by reef‐building corals, yet the effects of current regional‐scale declines in coral cover on reef complexity are poorly understood. In particular, both the extent to which declines in coral cover lead to declines in complexity and the length of time it takes for reefs to collapse following coral mortality are unknown. Here we assess the extent of temporal and spatial covariation between coral cover and reef architectural complexity using a Caribbean‐wide dataset of temporally replicated estimates spanning four decades. Both coral cover and architectural complexity have declined rapidly over time, with little evidence of a time‐lag. However, annual rates of change in coral cover and complexity do not covary, and levels of complexity vary greatly among reefs with similar coral cover. These findings suggest that the stressors influencing Caribbean reefs are sufficiently severe and widespread to produce similar regional‐scale declines in coral cover and reef complexity, even though reef architectural complexity is not a direct function of coral cover at local scales. Given that architectural complexity is not a simple function of coral cover, it is important that conservation monitoring and restoration give due consideration to both architecture and coral cover. This will help ensure that the ecosystem services supported by architectural complexity, such as nutrient recycling, dissipation of wave energy, fish production and diversity, are maintained and enhanced.     

Baselines and degradation of coral reefs in the Northern Line Islands

Effective conservation requires rigorous baselines of pristine conditions to assess the impacts of human activities and to evaluate the efficacy of management. Most coral reefs are moderately to severely degraded by local human activities such as fishing and pollution as well as global change, hence it is difficult to separate local from global effects. To this end, we surveyed coral reefs on uninhabited atolls in the northern Line Islands to provide a baseline of reef community structure, and on increasingly populated atolls to document changes associated with human activities. We found that top predators and reef-building organisms dominated unpopulated Kingman and Palmyra, while small planktivorous fishes and fleshy algae dominated the populated atolls of Tabuaeran and Kiritimati. Sharks and other top predators overwhelmed the fish assemblages on Kingman and Palmyra so that the biomass pyramid was inverted (top-heavy). In contrast, the biomass pyramid at Tabuaeran and Kiritimati exhibited the typical bottom-heavy pattern. Reefs without people exhibited less coral disease and greater coral recruitment relative to more inhabited reefs. Thus, protection from overfishing and pollution appears to increase the resilience of reef ecosystems to the effects of global warming.     

PROCESSES OF ORGANIC PRODUCTION ON CORAL REEFS

1. The first quantitative studies of production on coral reefs were those of Sargent & Austin who showed that productivity on reefs was considerably higher than in surrounding waters. This high production occurred in spite of nutrient limitation and low productivity of offshore waters. Their conclusions have since been confirmed by numerous other workers in both the Atlantic and the Pacific. 2. Primary production on reefs has been studied by flow respirometry, measuring changes in oxygen or carbon dioxide concentrations in water flowing over reefs. Production of benthic organisms has also been measured in situ by light and dark bottle methods and by radioactive tracer techniques. Production values obtained by the various methods are not identical but their use in combination is to be recommended. 3. Rates of gross primary production on reefs vary between 300–5000 gC/m²/yr. These rates are higher than general oceanic values and as high as those of the most productive marine communities. 4. Sources of primary production include fleshy macrophytes, calcareous algae, filamentous algae on the coral skeletons or calcareous rock, marine grasses and the zooxanthellae within coral tissue. Production values from the various sources fall within the range of production of reefs as a whole. 5. Concentrations of nitrogen and phosphorus in waters flowing over reefs are consistently low. There is evidence to suggest that both these nutrients are recycled rapidly on the reef and that nitrogen is fixed by bacteria and primary producers. 6. In many instances the mass of detritus over coral reefs exceeds the biomass of zooplankton. While the quantitative significance of detritus as food for corals and other benthic organisms has not been evaluated, there is a growing body of evidence to show that this may be the key to understanding secondary production. 7. Opinions differ on the adequacy of zooplankton in satisfying the food requirements of corals and other benthic invertebrates on reefs. The weight of evidence suggests that while there is a removal of zooplankton by benthic organisms, the total biomass carried over the reef is too small to support the energy needs of secondary production. 8. Bacteria are a potential source of energy for secondary production on reefs and are implicated in nitrogen fixation, decomposition and biogeochemical cycling. 9. There is an abundance of sessile invertebrates other than corals on reefs but there are few quantitative data on their importance in secondary production. 10. The biomass of fish on reefs may be very high but the quantitative significance of grazing and predation is not fully established. 11. Studies on the growth of corals themselves have been based on measurements of skeletal accretion. These methods do not lead directly to estimates of reef organic production. Growth rates of corals vary considerably between and within species. 12. Estimates of reef growth have been made from measurements of coral growth and from the flux of calcium carbonate. There is less quantitative information on erosion caused by mechanical damage, by boring organisms and by human pollution. 13. Hydrographic factors influence growth and form of reefs and there is some evidence to show that production is enhanced by conservation of water in lagoonal areas.     

西沙珊瑚礁生态系统完整性评价与动态仿真模拟研究

以中国西沙珊瑚礁生态监控区为例,从生态系统完整性视角出发,结合珊瑚礁功能群组与环境变化的多重反馈效应,运用系统动力学方法构建了西沙珊瑚礁生态系统动态诊断模型,并通过基础情景和典型干扰情景（捕捞活动、陆源沉积、长棘海星（Acanthaster planci）暴发）的模拟过程,分析该生态监控区2010-2050年生态系统完整性的发展趋势及演变机理。结果显示：（1）系统动力学与生态系统完整性评价方法的整合,为珊瑚礁生态系统完整性的定量研究提供了一种可行性的方法;（2）珊瑚礁生态系统各效应指数中累积效应的变化最为明显,2010-2050年指数最大增长17.2,年均增长率约为1.2%,综合完整性指数在波动变化中稳步上升,珊瑚礁受损程度由中受损向中、低受损转变;（3）不同状态的珊瑚礁生态系统完整性,在其演化的不同阶段各有差异性,并且这种差异通常随时间推移才逐渐放大;（4）无论何种情景下,珊瑚礁生态系统都具有一定的恢复能力,尤其处于系统演化的释放重组阶段（2035-2040年）,生态系统的不稳定将创造了一定的可恢复性。研究结果对于分析西沙珊瑚礁生态系统的整体演化过程提供了理论借鉴,也为我国后续的珊瑚礁生态系统适应性修复理论模式研究提供了重要的基础。     

Millennial-scale change in the structure of a Caribbean reef ecosystem and the role of human and natural disturbance

Caribbean coral reefs have transformed into algal-dominated habitats over the past half-century, but the role of specific anthropogenic drivers is unresolved due to the lack of ecosystem-level data predating human disturbance. To better understand the extent and causes of long-term Caribbean reef declines, we produced a continuous 3000-yr record of the ecosystem state of three reefs in Bocas del Toro, Caribbean Panama. From fossils and sediments obtained from reef matrix cores, we tracked changes in reef accretion rates and the taxonomic and functional group composition of fish, coral, urchin, bivalve and benthic foraminifera. This dataset provided a comprehensive picture of reef community and environmental change. At all sites, reefs shifted from systems with greater relative abundance of herbivorous fish, epifaunal suspension feeding bivalves and Diadema urchins to systems with greater relative abundance of micropredator fish, infaunal bivalves and Echinometra urchins. These transitions were initiated a millennium ago at two less-degraded reefs fringing offshore islands and ~250 yr ago at a degraded patch reef near the continental coast. Ecosystem shifts were accompanied by a decline in reef accretion rates, and at the patch reef, a decline in water quality since the 18th century. Within all cores, synchronous increases in infaunal bivalves and declines in herbivorous fish regardless of water quality suggest a loss of hard substrate and increasingly hypoxic sediment conditions related to herbivore loss. While the early timing of ecosystem transitions at the fringing reefs implicates large-scale hydrological change, the more recent timing of change and loss of water quality at the patch reef implicates terrigenous runoff from land-clearing. Our whole-ecosystem reconstruction reveals that reef ecosystem deterioration appears to follow a predictable trajectory whether driven by natural or anthropogenic disturbances and that historical local human activities have quickly unraveled reefs at a scale similar to longer-term natural environmental change.