Stability criteria for complex ecosystems

Forty years ago, May proved that sufficiently large or complex ecological networks have a probability of persisting that is close to zero, contrary to previous expectations. May analysed large networks in which species interact at random. However, in natural systems pairs of species have well-defined interactions (for example predator-prey, mutualistic or competitive). Here we extend May's results to these relationships and find remarkable differences between predator-prey interactions, which are stabilizing, and mutualistic and competitive interactions, which are destabilizing. We provide analytic stability criteria for all cases. We use the criteria to prove that, counterintuitively, the probability of stability for predator-prey networks decreases when a realistic food web structure is imposed or if there is a large preponderance of weak interactions. Similarly, stability is negatively affected by nestedness in bipartite mutualistic networks. These results are found by separating the contribution of network structure and interaction strengths to stability. Stable predator-prey networks can be arbitrarily large and complex, provided that predator-prey pairs are tightly coupled. The stability criteria are widely applicable, because they hold for any system of differential equations.     

An ecological and evolutionary perspective on human-microbe mutualism and disease

The microbial communities of humans are characteristic and complex mixtures of microorganisms that have co-evolved with their human hosts. The species that make up these communities vary between hosts as a result of restricted migration of microorganisms between hosts and strong ecological interactions within hosts, as well as host variability in terms of diet, genotype and colonization history. The shared evolutionary fate of humans and their symbiotic bacteria has selected for mutualistic interactions that are essential for human health, and ecological or genetic changes that uncouple this shared fate can result in disease. In this way, looking to ecological and evolutionary principles might provide new strategies for restoring and maintaining human health.     

Ecological networks and their fragility

Darwin used the metaphor of a 'tangled bank' to describe the complex interactions between species. Those interactions are varied: they can be antagonistic ones involving predation, herbivory and parasitism, or mutualistic ones, such as those involving the pollination of flowers by insects. Moreover, the metaphor hints that the interactions may be complex to the point of being impossible to understand. All interactions can be visualized as ecological networks, in which species are linked together, either directly or indirectly through intermediate species. Ecological networks, although complex, have well defined patterns that both illuminate the ecological mechanisms underlying them and promise a better understanding of the relationship between complexity and ecological stability.     

Non-random coextinctions in phylogenetically structured mutualistic networks

The interactions between plants and their animal pollinators and seed dispersers have moulded much of Earth's biodiversity. Recently, it has been shown that these mutually beneficial interactions form complex networks with a well-defined architecture that may contribute to biodiversity persistence. Little is known, however, about which ecological and evolutionary processes generate these network patterns. Here we use phylogenetic methods to show that the phylogenetic relationships of species predict the number of interactions they exhibit in more than one-third of the networks, and the identity of the species with which they interact in about half of the networks. As a consequence of the phylogenetic effects on interaction patterns, simulated extinction events tend to trigger coextinction cascades of related species. This results in a non-random pruning of the evolutionary tree and a more pronounced loss of taxonomic diversity than expected in the absence of a phylogenetic signal. Our results emphasize how the simultaneous consideration of phylogenetic information and network architecture can contribute to our understanding of the structure and fate of species-rich communities.     

Consequences of a biological invasion reveal the importance of mutualism for plant communities.

Seed-dispersal mutualisms have a fundamental role in regenerating natural communities. Interest in the importance of seed dispersal to plant communities has been heightened by worldwide declines in animal dispersers. One view, the 'keystone mutualist hypothesis', predicts that these human-caused losses will trigger a cascade of linked extinctions throughout the community. Implicitly, this view holds that mutualisms, such as seed dispersal, are crucial ecological interactions that maintain the structure and diversity of natural communities. Although many studies suggest the importance of mutualism, empirical evidence for community-level impacts of mutualists has remained anecdotal, and the central role of mutualism, relative to other species interactions, has long been debated in the theoretical literature. Here I report the community-level consequences of a biological invasion that disrupts important seed-dispersal mutualisms. I show that invasion of South African shrublands by the Argentine ant (Linepithema humile) leads to a shift in composition of the plant community, owing to a disproportionate reduction in the densities of large-seeded plants. This study suggests that the preservation of mutualistic interactions may be essential for maintaining natural communities.     

Patterns of relative species abundance in rainforests and coral reefs

A formidable many-body problem in ecology is to understand the complex of factors controlling patterns of relative species abundance (RSA) in communities of interacting species. Unlike many problems in physics, the nature of the interactions in ecological communities is not completely known. Although most contemporary theories in ecology start with the basic premise that species interact, here we show that a theory in which all interspecific interactions are turned off leads to analytical results that are in agreement with RSA data from tropical forests and coral reefs. The assumption of non-interacting species leads to a sampling theory for the RSA that yields a simple approximation at large scales to the exact theory. Our results show that one can make significant theoretical progress in ecology by assuming that the effective interactions among species are weak in the stationary states in species-rich communities such as tropical forests and coral reefs.     

Strong contributors to network persistence are the most vulnerable to extinction

The architecture of mutualistic networks facilitates coexistence of individual participants by minimizing competition relative to facilitation. However, it is not known whether this benefit is received by each participant node in proportion to its overall contribution to network persistence. This issue is critical to understanding the trade-offs faced by individual nodes in a network. We address this question by applying a suite of structural and dynamic methods to an ensemble of flowering plant/insect pollinator networks. Here we report two main results. First, nodes contribute heterogeneously to the overall nested architecture of the network. From simulations, we confirm that the removal of a strong contributor tends to decrease overall network persistence more than the removal of a weak contributor. Second, strong contributors to collective persistence do not gain individual survival benefits but are in fact the nodes most vulnerable to extinction. We explore the generality of these results to other cooperative networks by analysing a 15-year time series of the interactions between designer and contractor firms in the New York City garment industry. As with the ecological networks, a firm's survival probability decreases as its individual nestedness contribution increases. Our results, therefore, introduce a new paradox into the study of the persistence of cooperative networks, and potentially address questions about the impact of invasive species in ecological systems and new competitors in economic systems.     

Niche lability in the evolution of a Caribbean lizard community

Niche conservatism--the tendency for closely related species to be ecologically similar--is widespread. However, most studies compare closely related taxa that occur in allopatry; in sympatry, the stabilizing forces that promote niche conservatism, and thus inhibit niche shifts, may be countered by natural selection favouring ecological divergence to minimize the intensity of interspecific interactions. Consequently, the relative importance of niche conservatism versus niche divergence in determining community structure has received little attention. Here, we examine a tropical lizard community in which species have a long evolutionary history of ecological interaction. We find that evolutionary divergence overcomes niche conservatism: closely related species are no more ecologically similar than expected by random divergence and some distantly related species are ecologically similar, leading to a community in which the relationship between ecological similarity and phylogenetic relatedness is very weak. Despite this lack of niche conservatism, the ecological structuring of the community has a phylogenetic component: niche complementarity only occurs among distantly related species, which suggests that the strength of ecological interactions among species may be related to phylogeny, but it is not necessarily the most closely related species that interact most strongly.     

聚果榕小蜂群落在中国西双版纳和缅甸曼德勒之间的地理变异

榕树及其传粉榕小蜂之间的相互作用是互惠共生的经典例子，每种雌雄同株的榕树还庇护着多样的非传粉小蜂，并共生形成多营养级的榕小蜂群落，是研究群落生态学的有趣模型。区域的物种库、物种的生物地理特征及种间相互作用可能影响榕小蜂群落，但研究很少。本研究选择分布于中国西双版纳和缅甸曼德勒的雌雄同株聚果榕小蜂群落作为研究对象，比较地理分布差异对榕小蜂个体大小及共存网络的影响。结果显示：聚果榕小蜂群落在两地由相同的6个种组成，但分布于西双版纳的6种榕小蜂的个体均显著大于分布在曼德勒的同种榕小蜂。在两个分布地传粉榕小蜂均为优势种，然而在西双版纳传粉榕小蜂种群较大，而在曼德勒非传粉小蜂更丰富。在两地聚果榕有不同的榕小蜂群落结构和物种丰富度，且榕小蜂共存的网络也存在变异，在两地共存网络有相同的节点、边、连接度和全局聚类系数，但是在西双版纳的共存网络有较大的直径和较高的群落矩阵温度，这表明分布于西双版纳的聚果榕小蜂群落种间互作不太紧密，群落受到的干扰较大。由此，阐明了由地理分布所驱动的榕小蜂群落变异。     

Competition and phylogeny determine community structure in Müllerian co-mimics

Until recently, the study of negative and antagonistic interactions (for example, competition and predation) has dominated our understanding of community structure, maintenance and assembly. Nevertheless, a recent theoretical model suggests that positive interactions (for example, mutualisms) may counterbalance competition, facilitating long-term coexistence even among ecologically undifferentiated species. Müllerian mimics are mutualists that share the costs of predator education and are therefore ideally suited for the investigation of positive and negative interactions in community dynamics. The sole empirical test of this model in a Müllerian mimetic community supports the prediction that positive interactions outweigh the negative effects of spatial overlap (without quantifying resource acquisition). Understanding the role of trophic niche partitioning in facilitating the evolution and stability of Müllerian mimetic communities is now of critical importance, but has yet to be formally investigated. Here we show that resource partitioning and phylogeny determine community structure and outweigh the positive effects of Müllerian mimicry in a species-rich group of neotropical catfishes. From multiple, independent reproductively isolated allopatric communities displaying convergently evolved colour patterns, 92% consist of species that do not compete for resources. Significant differences in phylogenetically conserved traits (snout morphology and body size) were consistently linked to trait-specific resource acquisition. Thus, we report the first evidence, to our knowledge, that competition for trophic resources and phylogeny are pivotal factors in the stable evolution of Müllerian mimicry rings. More generally, our work demonstrates that competition for resources is likely to have a dominant role in the structuring of communities that are simultaneously subject to the effects of both positive and negative interactions.     

Invasion sequence affects predator-prey dynamics in a multi-species interaction

Ecologists seek to understand the rules that govern the assembly, coexistence and persistence of communities of interacting species. There is, however, a variety of sequences in which a multi-species community can be assembled--unlike more familiar one- and two-species systems. Ecological systems can exhibit contrasting dynamics depending on initial conditions, but studies have been focused on simple communities initiated at different densities, not on multi-species communities constructed in different sequences. Investigations of permanence and convergence in ecological communities have been concerned with the flux of whole species (presence or absence) but have not addressed the central issues concerning the dynamics exhibited by individual species in particular interactions. Here we examine data for replicated three-species systems and demonstrate that the dynamic trajectories of both a predator and its prey within the system are determined by the sequence in which it is constructed, and that for one construction-sequence alternative dynamic patterns are possible.     

General patterns of taxonomic and biomass partitioning in extant and fossil plant communities

A central goal of evolutionary ecology is to identify the general features maintaining the diversity of species assemblages. Understanding the taxonomic and ecological characteristics of ecological communities provides a means to develop and test theories about the processes that regulate species coexistence and diversity. Here, using data from woody plant communities from different biogeographic regions, continents and geologic time periods, we show that the number of higher taxa is a general power-function of species richness that is significantly different from randomized assemblages. In general, we find that local communities are characterized by fewer higher taxa than would be expected by chance. The degree of taxonomic diversity is influenced by modes of dispersal and potential biotic interactions. Further, changes in local diversity are accompanied by regular changes in the partitioning of community biomass between taxa that are also described by a power function. Our results indicate that local and regional processes have consistently regulated community diversity and biomass partitioning for millions of years.     

Field parameterization and experimental test of the neutral theory of biodiversity

Ecologists would like to explain general patterns observed across multi-species communities, such as species-area and abundance-frequency relationships, in terms of the fundamental processes of birth, death and migration underlying the dynamics of all constituent species. The unified neutral theory of biodiversity and related theories based on these fundamental population processes have successfully recreated general species-abundance patterns without accounting for either the variation among species and individuals or resource-releasing processes such as predation and disturbance, long emphasized in ecological theory. If ecological communities can be described adequately without estimating variation in species and their interactions, our understanding of ecological community organization and the predicted consequences of reduced biodiversity and environmental change would shift markedly. Here, I introduce a strong method to test the neutral theory that combines field parameterization of the underlying population dynamics with a field experiment, and apply it to a rocky intertidal community. Although the observed abundance-frequency distribution of the system follows that predicted by the neutral theory, the neutral theory predicts poorly the field experimental results, indicating an essential role for variation in species interactions.     

Positive interactions among alpine plants increase with stress

Plants can have positive effects on each other. For example, the accumulation of nutrients, provision of shade, amelioration of disturbance, or protection from herbivores by some species can enhance the performance of neighbouring species. Thus the notion that the distributions and abundances of plant species are independent of other species may be inadequate as a theoretical underpinning for understanding species coexistence and diversity. But there have been no large-scale experiments designed to examine the generality of positive interactions in plant communities and their importance relative to competition. Here we show that the biomass, growth and reproduction of alpine plant species are higher when other plants are nearby. In an experiment conducted in subalpine and alpine plant communities with 115 species in 11 different mountain ranges, we find that competition generally, but not exclusively, dominates interactions at lower elevations where conditions are less physically stressful. In contrast, at high elevations where abiotic stress is high the interactions among plants are predominantly positive. Furthermore, across all high and low sites positive interactions are more important at sites with low temperatures in the early summer, but competition prevails at warmer sites.     

Competition and mutualism among the gut helminths of a mammalian host

Most animal species are infected with multiple parasite species; however, the role of interspecific parasite interactions in influencing parasite dynamics and shaping parasite communities has been unclear. Although laboratory studies have found evidence of cross-immunity, immunosuppression and competition, analyses of hosts in the field have generally concluded that parasite communities are little more than random assemblages. Here we present evidence of consistent interspecific interactions in a natural mammalian system, revealed through the analysis of parasite intensity data collected from a free-ranging rabbit (Oryctolagus cuniculus) population, sampled monthly for a period of 23 yr. The wild rabbit plays host to a diverse gut helminth community that reflects the communities seen in other economically important domestic herbivores. These findings suggest that parasite interactions could have profound implications for the dynamics of parasite communities. The efficacy of parasite control programmes could be jeopardized if such interactions are not taken into account. In contrast, a clear understanding of such interactions may provide the basis for the development of more environmentally acceptable methods of parasite control.     

植食性昆虫、植物和共生微生物的功能关联

植食性昆虫和植物均能与微生物形成密切关系，它们各自的生态功能及相互关系也常被共生微生物所影响。近年来，随着分子水平研究方法的进展，植食性昆虫和植物中很多可遗传共生微生物（细菌、真菌等）被发现。共生微生物能够在营养、生殖、防御和解毒等方面给宿主带来显著影响，并与宿主形成竞争、互利或寄生等关系。植食性昆虫体内的含菌胞、肠道、血淋巴、唾液腺等常含有重要功能的共生微生物。新的分子生物学手段和高通量测序技术的应用使得我们能够增加对宿主和共生微生物（即使处于低丰度）之间关系的了解。本文尝试总结了植食性昆虫和植物共生微生物的多样性及其关系、昆虫和植物互作机制、昆虫共生菌解毒植物毒素等方面的近期证据，突出强调了应以系统观思维来理解共生微生物、植食性昆虫和植物间的功能关联，并就将来值得研究的问题提出了建议。     

The diversity-stability debate

There exists little doubt that the Earth's biodiversity is declining. The Nature Conservancy, for example, has documented that one-third of the plant and animal species in the United States are now at risk of extinction. The problem is a monumental one, and forces us to consider in depth how we expect ecosystems, which ultimately are our life-support systems, to respond to reductions in diversity. This issue--commonly referred to as the diversity-stability debate--is the subject of this review, which synthesizes historical ideas with recent advances. Both theory and empirical evidence agree that we should expect declines in diversity to accelerate the simplification of ecological communities.     

川西高原森林生物多样性及其生态功能的研究

对川西高原森林物种多样性及其生态功能进行了研究。结果表明，在采伐迹地上，人工林的物种多样性及其生态功能随着林龄的增长呈现增加和增强的趋势。天然林群落的物种多样性随着顺行演替的进行逐渐增加，至顶极阶段，群落的物种多样性有所减少。但是，群落的生态功能却始终随着演替的进行逐步增强。此外，在相似立地条件、同密度和同龄的天然林中，针阔叶混交林的物种多样性指数要高于针叶混交林；而在相似立地条件、同龄的天然林中，高密度林分的物种多样性低于低密度林分。     

Policing stabilizes construction of social niches in primates

All organisms interact with their environment, and in doing so shape it, modifying resource availability. Termed niche construction, this process has been studied primarily at the ecological level with an emphasis on the consequences of construction across generations. We focus on the behavioural process of construction within a single generation, identifying the role a robustness mechanism--conflict management--has in promoting interactions that build social resource networks or social niches. Using 'knockout' experiments on a large, captive group of pigtailed macaques (Macaca nemestrina), we show that a policing function, performed infrequently by a small subset of individuals, significantly contributes to maintaining stable resource networks in the face of chronic perturbations that arise through conflict. When policing is absent, social niches destabilize, with group members building smaller, less diverse, and less integrated grooming, play, proximity and contact-sitting networks. Instability is quantified in terms of reduced mean degree, increased clustering, reduced reach, and increased assortativity. Policing not only controls conflict, we find it significantly influences the structure of networks that constitute essential social resources in gregarious primate societies. The structure of such networks plays a critical role in infant survivorship, emergence and spread of cooperative behaviour, social learning and cultural traditions.