Density dependence explains tree species abundance and diversity in tropical forests

The recurrent patterns in the commonness and rarity of species in ecological communities--the relative species abundance--have puzzled ecologists for more than half a century. Here we show that the framework of the current neutral theory in ecology can easily be generalized to incorporate symmetric density dependence. We can calculate precisely the strength of the rare-species advantage that is needed to explain a given RSA distribution. Previously, we demonstrated that a mechanism of dispersal limitation also fits RSA data well. Here we compare fits of the dispersal and density-dependence mechanisms for empirical RSA data on tree species in six New and Old World tropical forests and show that both mechanisms offer sufficient and independent explanations. We suggest that RSA data cannot by themselves be used to discriminate among these explanations of RSA patterns--empirical studies will be required to determine whether RSA patterns are due to one or the other mechanism, or to some combination of both.     

Coral reef diversity refutes the neutral theory of biodiversity

The global decline of coral reefs highlights the need to understand the mechanisms that regulate community structure and sustain biodiversity in these systems. The neutral theory, which assumes that individuals are demographically identical regardless of species, seeks to explain ubiquitous features of community structure and biodiversity patterns. Here we present a test of neutral-theory predictions with the use of an extensive species-level data set of Indo-Pacific coral communities. We show that coral assemblages differ markedly from neutral-model predictions for patterns of community similarity and the relative abundance of species. Within local communities, neutral models do not fit relative abundance distributions as well as the classical log-normal distribution. Relative abundances of species across local communities also differ markedly from neutral-theory predictions: coral communities exhibit community similarity values that are far more variable, and lower on average, than the neutral theory can produce. Empirical community similarities deviate from the neutral model in a direction opposite to that predicted in previous critiques of the neutral theory. Instead, our results support spatio-temporal environmental stochasticity as a major driver of diversity patterns on coral reefs.     

Mangroves enhance the biomass of coral reef fish communities in the Caribbean

Mangrove forests are one of the world's most threatened tropical ecosystems with global loss exceeding 35% (ref. 1). Juvenile coral reef fish often inhabit mangroves, but the importance of these nurseries to reef fish population dynamics has not been quantified. Indeed, mangroves might be expected to have negligible influence on reef fish communities: juvenile fish can inhabit alternative habitats and fish populations may be regulated by other limiting factors such as larval supply or fishing. Here we show that mangroves are unexpectedly important, serving as an intermediate nursery habitat that may increase the survivorship of young fish. Mangroves in the Caribbean strongly influence the community structure of fish on neighbouring coral reefs. In addition, the biomass of several commercially important species is more than doubled when adult habitat is connected to mangroves. The largest herbivorous fish in the Atlantic, Scarus guacamaia, has a functional dependency on mangroves and has suffered local extinction after mangrove removal. Current rates of mangrove deforestation are likely to have severe deleterious consequences for the ecosystem function, fisheries productivity and resilience of reefs. Conservation efforts should protect connected corridors of mangroves, seagrass beds and coral reefs.     

Ecological consequences of major hydrodynamic disturbances on coral reefs

A recent tsunami and an apparent increase in the frequency of severe tropical storms underscore the need to understand and predict the ecological consequences of major hydrodynamic disturbances. Reef corals provide the habitat structure that sustains the high biodiversity of tropical reefs, and thus provide the foundation for the ecosystem goods and services that are critical to many tropical societies. Here we integrate predictions from oceanographic models with engineering theory, to predict the dislodgement of benthic reef corals during hydrodynamic disturbances. This generalizes earlier work, by incorporating colonies of any shape and by explicitly examining the effects of hydrodynamic gradients on coral assemblage structure. A field test shows that this model accurately predicts changes in the mechanical vulnerability of coral colonies, and thus their size and shape, with distance from the reef crest. This work provides a general framework for understanding and predicting the effects of hydrodynamic disturbances on coral reef communities; such disturbances have a major role in determining species zonation and coexistence on coral reefs, and are critical determinants of how coral assemblages will respond to changes in the frequency and intensity of tropical storms associated with a changing climate.     

Coral communities are regionally enriched along an oceanic biodiversity gradient

Ecological communities are influenced by processes operating at multiple scales. Thus, a better understanding of how broad- as well as local-scale processes affect species diversity and richness is increasingly becoming a central focus in modern community ecology. Here, in a study of unprecedented geographical scope, we show significant regional and local variation in the species richness of coral assemblages across an oceanic biodiversity gradient. The gradient that we sampled extends 10,000 km eastwards from the world's richest coral biodiversity hotspot in the central Indo-Pacific. Local richness and the size of regional species pools decline significantly across 15 islands spanning the gradient. In addition, richness declines across three adjacent habitats (reef slopes, crests and flats). In each habitat, a highly consistent linear relationship between local and regional species richness indicates strong regional enrichment. Thus, even on the most diverse coral reefs in the world, local coral assemblages are profoundly affected by regional-scale processes. Understanding these historical and biogeographical influences is essential for the effective management and preservation of these endangered communities.     

Host specificity of Lepidoptera in tropical and temperate forests

For numerous taxa, species richness is much higher in tropical than in temperate zone habitats. A major challenge in community ecology and evolutionary biogeography is to reveal the mechanisms underlying these differences. For herbivorous insects, one such mechanism leading to an increased number of species in a given locale could be increased ecological specialization, resulting in a greater proportion of insect species occupying narrow niches within a community. We tested this hypothesis by comparing host specialization in larval Lepidoptera (moths and butterflies) at eight different New World forest sites ranging in latitude from 15 degrees S to 55 degrees N. Here we show that larval diets of tropical Lepidoptera are more specialized than those of their temperate forest counterparts: tropical species on average feed on fewer plant species, genera and families than do temperate caterpillars. This result holds true whether calculated per lepidopteran family or for a caterpillar assemblage as a whole. As a result, there is greater turnover in caterpillar species composition (greater beta diversity) between tree species in tropical faunas than in temperate faunas. We suggest that greater specialization in tropical faunas is the result of differences in trophic interactions; for example, there are more distinct plant secondary chemical profiles from one tree species to the next in tropical forests than in temperate forests as well as more diverse and chronic pressures from natural enemy communities.     

Low host specificity of herbivorous insects in a tropical forest

Two decades of research have not established whether tropical insect herbivores are dominated by specialists or generalists. This impedes our understanding of species coexistence in diverse rainforest communities. Host specificity and species richness of tropical insects are also key parameters in mapping global patterns of biodiversity. Here we analyse data for over 900 herbivorous species feeding on 51 plant species in New Guinea and show that most herbivorous species feed on several closely related plant species. Because species-rich genera are dominant in tropical floras, monophagous herbivores are probably rare in tropical forests. Furthermore, even between phylogenetically distant hosts, herbivore communities typically shared a third of their species. These results do not support the classical view that the coexistence of herbivorous species in the tropics is a consequence of finely divided plant resources; non-equilibrium models of tropical diversity should instead be considered. Low host specificity of tropical herbivores reduces global estimates of arthropod diversity from 31 million (ref. 1) to 4 6 million species. This finding agrees with estimates based on taxonomic collections, reconciling an order of magnitude discrepancy between extrapolations of global diversity based on ecological samples of tropical communities with those based on sampling regional faunas.     

Thresholds and the resilience of Caribbean coral reefs

The deteriorating health of the world's coral reefs threatens global biodiversity, ecosystem function, and the livelihoods of millions of people living in tropical coastal regions. Reefs in the Caribbean are among the most heavily affected, having experienced mass disease-induced mortality of the herbivorous urchin Diadema antillarum in 1983 and two framework-building species of coral. Declining reef health is characterized by increases in macroalgae. A critical question is whether the observed macroalgal bloom on Caribbean reefs is easily reversible. To answer this question, we must resolve whether algal-dominated reefs are an alternative stable state of the ecosystem or simply the readily reversible result of a phase change along a gradient of some environmental or ecological parameter. Here, using a fully parameterized simulation model in combination with a simple analytical model, we show that Caribbean reefs became susceptible to alternative stable states once the urchin mortality event of 1983 confined the majority of grazing to parrotfishes. We reveal dramatic hysteresis in a natural system and define critical thresholds of grazing and coral cover beyond which resilience is lost. Most grazing thresholds lie near the upper level observed for parrotfishes in nature, suggesting that reefs are highly sensitive to parrotfish exploitation. Ecosystem thresholds can be combined with stochastic models of disturbance to identify targets for the restoration of ecosystem processes. We illustrate this principle by estimating the relationship between current reef state (coral cover and grazing) and the probability that the reef will withstand moderate hurricane intensity for two decades without becoming entrained in a shift towards a stable macroalgal-dominated state. Such targets may help reef managers face the challenge of addressing global disturbance at local scales.     

涠洲岛珊瑚礁特点、演变及保护与修复对策的思考

涠洲岛珊瑚礁最早形成于约7 000年前的末次海侵,是中国大陆沿海成礁珊瑚分布最北缘的典型边缘珊瑚礁（Marginal Coral Reefs）,与区域内其他珊瑚礁缺少生态关联,其特征不同于热带珊瑚礁,即块状珊瑚占优势。20多年来,涠洲岛珊瑚礁持续退化,枝状珊瑚大量死亡,石珊瑚盖度急剧降低,反映珊瑚礁良好状况的指标生物如鹦嘴鱼、蝴蝶鱼等的数量大幅减少。近10年来,珊瑚虽未出现异常死亡现象,但珊瑚自然修复不明显,表明珊瑚补充不足。功能性植食性鱼类的缺乏,显示珊瑚礁恢复力不足。涠洲岛珊瑚礁国家级海洋公园的建立,体现了保护涠洲岛珊瑚礁的政治意愿。只有恢复鹦嘴鱼等功能性植食性鱼类的数量,才能恢复和增强珊瑚礁固有恢复力,进而促进涠洲岛珊瑚礁的自然恢复。人为修复可采取移植珊瑚断枝的方式,并以实现生态、经济和社会增益为目标,构建能应对人为干预且能持续演化的珊瑚礁。     

Coral reefs: corals' adaptive response to climate change

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

Twenty-five years of change in scleractinian coral communities of Daya Bay （northern South China Sea） and its response to the 2008 AD extreme cold climate event

Coral reefs worldwide are becoming increasingly and detrimentally impacted upon by a variety of factors including significant climate changes, such as global warming and increased El Nino-Southern Oscillation activity. Generally, the persistence of coral reefs, especially at low-latitudes, is governed, in part, by sea surface temperatures not exceeding the critical limit （-30℃） at which mass mortality can occur. Thus, it is thought that corals living at high-latitudes （i.e., currently cooler sea surface temperatures） will likely respond more favourably to hypothesized future temperature increases than corals living at low-latitudes （i.e., currently warmer sea surface temperatures）. Consequently, high-latitude coral communities may have the potential to act as regions of refugia for many coral species in the face of potential future global warming. The Daya Bay （22°31′--22°50′N）, northern South China Sea, contains several high-latitude non-reefal coral communities and represents one of the most northerly distributions of scleractinian corals within the region. Significantly, Daya Bay has experienced dramatic warming in both air and sea surface temperatures throughout the past 50 years. In this paper, we analyze 25 years of change in the Daya Bay coral communities, based both on historic surveys and our latest 2006--2008 regional ecological surveys. Our results suggest that, contrary to predictions, there have been significant declines in coral cover within the Daya Bay during the past 25 years （i.e., 76.6% coral cover in 1983/1984 to only 15.3% coral cover by 2008）. Such changes also reflect a significant shift in the most abundant coral species, from Acropora pruinosa to Favites abdita. Most of the modern coral communities became established between 15 and 30 years ago, corresponding to a period of increased winter sea surface temperature. However, very few colonies have become established within the last 15 years, despite a more intense period of warming. By taking into account additional factors, we hypothesize that direct anthropogenic impacts, rather than climatic events, have both restricted the development, and drove the decline, of Daya Bay coral communities in the last 15 years. The Daya Bay has also been subjected to occasional extreme cold events during the past 50 years, with the most recent occurring in early 2008 （13 January-13 February）. During the 2008 cold event, the lowest air temperature reaches only 6.6℃, and the mean sea surface temperature for February fall to 〈 14℃, including six continuous days at 12.3℃. Significantly, the sea surface temperatures fall below the hypothesized critical lower temperature threshold （-13℃） that commonly leads to mass mortality in scleractinian coral communities. Surprisingly, our coral community surveys, conducted both before （August 2007） and after （late February 2008） the extreme 2008 cold event, demonstrate that the Daya Bay coral ecosystems are barely impacted upon during the cold period. Those observations suggest that the Daya Bay scleractinian coral communities have developed adaptations to low sea surface temperatures. Overall, our data support the hypothesis that high-latitude coral communities, such as Daya Bay, have the potential to act as areas of refugia for scleractinian corals in the advent of potential future global warming.     

Experimental evidence for apparent competition in a tropical forest food web

The herbivorous insects of tropical forests constitute some of the most diverse communities of living organisms. For this reason it has been difficult to discover the degree to which these communities are structured, and by what processes. Interspecific competition for resources does occur, but its contemporary importance is limited because most pairs of potentially competing insects feed on different host plants. An alternative way in which species can interact is through shared natural enemies, a process called apparent competition. Despite extensive theoretical discussion there are few field demonstrations of apparent competition, and none in hyper-diverse tropical communities. Here, we experimentally removed two species of herbivore from a community of leaf-mining insects in a tropical forest. We predicted that other species that share natural enemies with the two removed species would experience lower parasitism and have higher population densities in treatment compared with control sites. In both cases (on removal of a dipteran and a coleopteran leaf-miner species) we found significantly lower parasitism, and in one case (removal of the dipteran) we found significantly higher abundance a year after the manipulation. Our results suggest that apparent competition may be important in structuring tropical insect communities.     

珊瑚礁相关病毒的研究进展

珊瑚礁被称为海洋中的“热带雨林”,是海洋生态系统中的核心组成部分之一。珊瑚礁病毒包含珊瑚本身感染的病毒和珊瑚共生生物（如藻类、细菌等）感染的病毒,对于珊瑚礁健康与疾病、生长与死亡起着独特而重要的作用,在维持珊瑚礁生态系统的物质循环和能量流动等生态学功能方面起着潜在作用。由于技术的局限以及关注度较低等原因,有关珊瑚礁病毒的研究较少。近年来,随着高通量测序技术的发展,宏转录组和宏基因组逐渐成为研究珊瑚礁病毒多样性及其生态学功能的重要方法之一。本文对国内外相关研究进行总结和归纳,拟为珊瑚礁病毒后续研究提供新的方向和思路,同时指导我国珊瑚礁的保护以及修复。     

热带岛礁型海洋牧场中珊瑚礁生境与资源的修复

我国南海的离岸岛礁以珊瑚礁生境为主。珊瑚礁在提升海洋生态资源和维护我国海洋国土权益上发挥了至关重要的作用。近年来,在全球气候变化以及人类活动的影响下,南海珊瑚礁生境和资源严重退化,威胁着渔业资源、海洋环境以及国土安全。现代化的海洋牧场被认为是实现南海资源可持续发展战略的新型海洋生物资源开发模式。珊瑚礁生境与资源的修复则是热带岛礁型海洋牧场建设的基础和关键。基于此,本文整理了珊瑚礁生境与资源修复的技术方法,详细介绍了珊瑚礁生境构建框架生物造礁石珊瑚的有性繁殖、断枝培育、底播移植等技术方法以及珊瑚礁其他特色生物资源人工放流技术,并提出了在珊瑚礁生境与资源修复中所面临的问题及相应对策。     

黔南石炭纪珊瑚礁与南海现代珊瑚礁的对比研究

根据沉积环境和珊瑚礁的特征,分别对两种礁体进行了沉积相带的划分和建造过程的对比研究.分析表明:扁平村珊瑚礁中充填了大量的蜓、海百合茎和腕足碎屑等附礁生物组分,而现在珊瑚礁中的充填物大都以珊瑚碎屑为主;两种礁体虽形成于不同的时代,但具有类似的沉积相带分布和礁体形成过程,这与礁群落的盛衰状况密不可分.与晚石炭世的四射珊瑚相比,现代六射珊瑚对环境的适应能力更强,现代珊瑚礁的形成过程也更加曲折.     

Heterotrophic plasticity and resilience in bleached corals

Mass coral bleaching events caused by elevated seawater temperatures have resulted in extensive coral mortality throughout the tropics over the past few decades. With continued global warming, bleaching events are predicted to increase in frequency and severity, causing up to 60% coral mortality globally within the next few decades. Although some corals are able to recover and to survive bleaching, the mechanisms underlying such resilience are poorly understood. Here we show that the coral host has a significant role in recovery and resilience. Bleached and recovering Montipora capitata (branching) corals met more than 100% of their daily metabolic energy requirements by markedly increasing their feeding rates and CHAR (per cent contribution of heterotrophically acquired carbon to daily animal respiration), whereas Porites compressa (branching) and Porites lobata (mounding) corals did not. These findings suggest that coral species with high-CHAR capability during bleaching and recovery, irrespective of morphology, will be more resilient to bleaching events over the long term, could become the dominant coral species on reefs, and may help to safeguard affected reefs from potential local and global extinction.     

Soil pathogens and spatial patterns of seedling mortality in a temperate tree

The Janzen-Connell hypothesis proposes that host-specific, distance- and/or density-dependent predators and herbivores maintain high tree diversity in tropical forests. Negative feedback between plant and soil communities could be a more effective mechanism promoting species coexistence because soil pathogens can increase rapidly in the presence of their host, causing conditions unfavourable for local conspecific recruitment. Here we show that a soil pathogen leads to patterns of seedling mortality in a temperate tree (Prunus serotina) as predicted by the Janzen-Connell hypothesis. In the field, the mean distance to parent of seedling cohorts shifted away from maternal trees over a period of 3 years. Seedlings were grown in soil collected 0-5 m or 25-30 m from Prunus trees. Sterilization of soil collected beneath trees improved seedling survival relative to unsterilized soil, whereas sterilization of distant soil did not affect survival. Pythium spp., isolated from roots of dying seedlings and used to inoculate healthy seedlings, decreased survival by 65% relative to controls. Our results provide the most complete evidence that native pathogens influence tree distributions, as predicted by the Janzen-Connell hypothesis, and suggest that similar ecological mechanisms operate in tropical and temperate forests.     

Density-dependent mortality and the latitudinal gradient in species diversity

Ecologists have long postulated that density-dependent mortality maintains high tree diversity in the tropics. If species experience greater mortality when abundant, then more rare species can persist. Agents of density-dependent mortality (such as host-specific predators, and pathogens) may be more prevalent or have stronger effects in tropical forests, because they are not limited by climatic factors. If so, decreasing density-dependent mortality with increasing latitude could partially explain the observed latitudinal gradient in tree diversity. This hypothesis has never been tested with latitudinal data. Here we show that several temperate tree species experience density-dependent mortality between seed dispersal and seedling establishment. The proportion of species affected is equivalent to that in tropical forests, failing to support the hypothesis that this mechanism is more prevalent at tropical latitudes. We further show that density-dependent mortality is misinterpreted in previous studies. Our results and evidence from other studies suggest that density-dependent mortality is important in many forests. Thus, unless the strength of density-dependent mortality varies with latitude, this mechanism is not likely to explain the high diversity of tropical forests.     

Theoretical biology: comparing models of species abundance

Ecologists are struggling to explain how so many tropical tree species can coexist in tropical forests, and several empirical studies have demonstrated that negative density dependence is an important mechanism of tree-species coexistence. Volkov et al. compare a model incorporating negative density dependence with a dispersal-limited neutral model and claim that each predicts six empirical species-abundance distributions of tropical-tree communities equally well. However, we show here that their main conclusion is premature: when the two models are compared in an improved analysis, we find that the dispersal-limited model outcompetes the density-dependent model in all six cases. Hence, although density dependence is certainly an important diversity-maintaining mechanism, our improved approach indicates that the dispersal-limited model provides a more parsimonious explanation of empirical species-abundance distributions.     

Catastrophic shifts in ecosystems

All ecosystems are exposed to gradual changes in climate, nutrient loading, habitat fragmentation or biotic exploitation. Nature is usually assumed to respond to gradual change in a smooth way. However, studies on lakes, coral reefs, oceans, forests and arid lands have shown that smooth change can be interrupted by sudden drastic switches to a contrasting state. Although diverse events can trigger such shifts, recent studies show that a loss of resilience usually paves the way for a switch to an alternative state. This suggests that strategies for sustainable management of such ecosystems should focus on maintaining resilience.