破碎栖息地中物种灭绝机制

栖息地毁坏既会直接降低物种多度,又会间接地降低物种迁移繁殖力,同时还会改变原有的种间平衡.尽管已有研究表明栖息地毁坏是物种灭绝的主要原因之一,但是尚未揭示破碎的栖息地中物种灭绝的驱动机制.通过元胞自动机模拟了物种灭绝对栖息地毁坏空间异质性响应的基础上,进一步研究了栖息地毁坏和种间竞争对物种灭绝的影响.结果发现:强物种的灭绝主要来自栖息地毁坏,而弱物种的灭绝,在随机毁坏下,主要由栖息地毁坏与种间竞争共同决定,而在边缘毁坏下则主要由种间竞争所引起的.栖息地毁坏与种间竞争共同引起的物种灭绝的时间非常短,而栖息地毁坏或种间竞争所引起的物种灭绝时间则较长.     

Isodars unveil asymmetric effects on habitat use caused by competition between two endangered species

In order for competing species to coexist, segregation on some ecological niche component is required and is often mediated by differential habitat use. When unequal competitors are involved, the dominant species tends to displace the subordinate one to its less preferred habitat. Here, we use habitat isodars, an approach which reflects evolutionary stable strategies of habitat selection, to evaluate whether interspecific competition between two competing species with distinct habitat preferences, the little bustard Tetrax tetrax and the great bustard Otis tarda, modulates their habitat use. Field data on these endangered species demonstrate that unequal competitors can coexist without completely segregating on their preferred habitats. The negatively sloped isodar of the subordinate little bustard unveils its competition with the dominant great bustard. Interference from great bustards in secondary cereal habitats reinforces use of preferred natural habitat by little bustards. Studies of density‐dependent habitat selection by a single‐species can thus aid in identifying the effects of competition on community composition, and guide the conservation of at‐risk species. Isodars, in particular, represent a promising method to gain clear knowledge on interspecific competition for species in which experimental manipulations are not feasible.     

Competition and coexistence in regional habitats

Habitat heterogeneity plays a key role in the dynamics and structures of communities. In this article, a two-species metapopulation model that includes local competitive dynamics is analyzed to study the population dynamics of two competing species in spatially structured habitats. When local stochastic extinction can be ignored, there are, as in Lotka-Volterra equations, four outcomes of interspecific competition in this model. The outcomes of competition depend on the competitive intensity between the competing pairs. An inferior competitor and a superior competitor, or two strongly competing species, can never stably coexist, whereas two weak competitors (even if they are very similar species) may coexist over the long term in such environments. Local stochastic extinction may greatly affect the outcomes of interspecific competition. Two competing species can or cannot stably coexist depending not only on the competitive intensity between the competing pairs but also on their precompetitive distributions. Two weak competitors that have similar precompetitive distributions can always regionally coexist. Two strongly competing species that competitively exclude each other in more stable habitats may be able to stably coexist in highly heterogenous environments if they have similar precompetitive distributions. There is also a chance for an inferior competitor to coexist regionally or even to exclude a superior competitor when the superior competitor has a narrow precompetitive distribution and the inferior competitor has a wide precompetitive distribution.     

不同生境毁坏速度下的物种灭绝机制

已有似Levins的多物种模型,在研究生境毁坏的影响时,一方面主要集中在对瞬间毁坏影响的研究,另一方面主要研究生境毁坏对强物种影响的研究。在Tilman的多物种竞争共存模型的基础上,同时考虑了生境毁坏直接效应和生境毁坏时间异质性,提出了全新的普适的多物种竞争共存的非自治动力模式。通过模拟物种灭绝对不同速度的生境毁坏时间异质性的响应发现：（1）物种灭绝既存在强物种由强到弱的灭绝,也存在弱物种由弱到强的灭绝。同时,弱物种灭绝机制进一步分为弱物种瞬间集体灭绝,以及较长时间由弱到强的灭绝。（2）生境毁坏速度越快,物种灭绝的时间越短,弱物种灭绝的越多,因此,生境毁坏速度越慢,越有利于弱物种的长期续存。（3）最强物种的多度越大,强-强物种抵御生境毁坏的能力越强,而弱-弱物种抵御生境毁坏的能力越弱,集体灭绝的弱-弱物种就越多。最强物种的多度大的群落（如温带森林）,主要发生的是弱-弱物种灭绝,而最强物种多度小的群落（如热带雨林）同时发生强-强和弱-弱物种的灭绝。因此,争对不同结构的集合种群,不同的保护对象,应采取不同的管理策略。     

Determinism in a transient assemblage: the roles of dispersal and local competition

Both dispersal and local competitive ability may determine the outcome of competition among species that cannot coexist locally. I develop a spatially implicit model of two-species competition at a small spatial scale. The model predicts the relative fitness of two competitors based on local reproductive rates and regional dispersal rates in the context of the number, size, and extinction probability of habitat patches in the landscape. I test the predictions of this model experimentally using two genotypes of the bacteriophagous soil nematode Caenorhabditis elegans in patchy microcosms. One genotype has higher fecundity while the other is a better disperser. With such a fecundity-dispersal trade-off between competitors, the model predicts that relative fitness will be affected most by local population size when patches do not go extinct and by the number of patches when there is a high probability of patch extinction. The microcosm experiments support the model predictions. Both approaches suggest that competitive dominance in a patchily distributed transient assemblage will depend upon the architecture and predictability of the environment. These mechanisms, operating at a small scale with high spatial admixture, may be embedded in a larger metacommunity process.     

集合种群动态对生境毁坏空间异质性的响应

首次将分形几何(Fractal geometry)与元胞自动机(Cellular automata)相结合,研究了破碎化生境中集合种群的空间分布格局动态,以及集合种群动态对生境毁坏空间异质性的响应。研究发现:(1)各个物种种群在生境中的分布具有很好的分形特征,物种的计盒维数(Box dimension)不仅可以很好地反映种群的空间分布结构,也能很好地反映种群动态。(2)如果将空间因素考虑进来的话,生境毁坏的灭绝债务(Time debt)将大于空间隐含模式所模拟的结果。(3)物种灭绝同时存在强物种灭绝和弱物种灭绝。并且只有在生境随机毁坏下,才与空间隐含的模拟结果比较接近,即强物种中将是最强物种率先灭绝。而在边缘毁坏这种比较集中成块的开发方式下,将是较强的物种灭绝。(4)边缘毁坏相对随机毁坏有利于物种,尤其是弱物种的长期续存。     

Interspecific competition in perennial sedentary organisms: An individual-based model

We investigated a mathematical model of the dynamics of the ecological system consisting of two competing perennial species, each of which leads a sedentary life. It is an individual-based model, in which the growth of each individual is described. The rate of this growth is weakened by competition from neighboring individuals. The strength of the competitors' influence depends on their size and distance to them. The conditions, in which the competitive exclusion of one of the competitors and the coexistence of both competitors take place are provided. The influence of the parameters responsible for the strength of competition, the degree of competitive asymmetry, and consideration of the importance of specific elements of the spatial structure of this ecological system on the results of the competition were analyzed. Both species co-exist when they are equal competitors. Permanent coexistence is possible only when interspecific competition is weaker than intraspecific. When interspecific competition is stronger, the coexistence of equal interspecific competitors is random. Both species have equal probability of extinction. If species are not equal competitors, the stronger one wins. This result can be modified by different strengths of intraspecific competition. The weaker interspecific competitor can permanently coexist with stronger one, when its individuals suffer stronger intraspecific competition.     

集合种群动态对栖息地毁坏时空异质性的响应

栖息地毁坏既有时间异质性,也有空间异质性,而以往的研究往往只关注其中的一种。将两种不同的异质性共同引入到元胞自动机中,模拟了集合种群动态对栖息地毁坏时空异质性的响应。发现,在随机离散的栖息地毁坏下,由于物种的迁移繁殖力受栖息地毁坏的影响很大,迁移繁殖力弱而竞争力强的物种先灭绝。在连续的栖息地毁坏下,物种的迁移繁殖力受栖息地毁坏的影响较小,物种的灭绝由竞争力和迁移繁殖力共同决定:在有绝对优势种的群落里,种间竞争显著,弱物种先灭绝,而在没有绝对优势种的群落里,种间竞争较小,则以强物种先灭绝。因此,随机毁坏不利于强物种续存,而连续毁坏则不利于具有绝对优势种群的群落里的弱物种续存。在实际开发某一栖息地时,根据集合种群结构和被保护的对象采取相应的开发模式。     

How species diversity responds to different kinds of human-caused habitat destruction

Human-caused habitat destruction, the major cause of species diversity losses, can be classified into two basic types, instantaneous destruction and continuous destruction. Thus, a universal model should be established to simulate and forecast the effects of different kinds of habitat destruction on species diversity during different historical periods. In this paper, we explore a multi-time-scale n-species model to study and compare species responses to instantaneous and continuous destruction. We find that (1) under instantaneous destruction, there are two different mechanisms of species extinction: one is a time-delayed deterministic extinction of superior competitors in order from the best to the poorest; the other is the extinction in a short time of inferior competitors. The survivors will experience three phases: decline, adjustment, and equilibrium. (2) When the total amounts of habitat destruction for both instantaneous and continuous cases are equal, the oscillation amplitudes of species abundances under instantaneous destruction are much greater than under continuous destruction, especially for inferior competitors, which make inferior competitors under instantaneous destruction more prone to stochastic extinction. Therefore instantaneous destruction is more detrimental to the survival of inferior competitors. (3) Under continuous destruction with habitat eventually being destroyed completely, there also are two types of species extinction mechanisms: the first is extinction in order from the best competitors to the poorest before complete destruction; the second is collective extinction due to complete destruction.     

Rock-scissors-paper game in a chaotic flow: the effect of dispersion on the cyclic competition of microorganisms

Laboratory experiments and numerical simulations have shown that the outcome of cyclic competition is significantly affected by the spatial distribution of the competitors. Short-range interaction and limited dispersion allows for coexistence of competing species that cannot coexist in a well-mixed environment. In order to elucidate the mechanisms that destroy species diversity we study the intermediate situation of imperfect mixing, typical in aquatic media, in a model of cyclic competition between toxin producing, sensitive and resistant phenotypes. It is found, that chaotic mixing, by changing the character of the spatial distribution, induces coherent oscillations in the populations. The magnitude of the oscillations increases with the strength of mixing, leading to the extinction of some species beyond a critical mixing rate. When mixing is non-uniform in space, coexistence can be sustained at much stronger mixing by the formation of partially isolated regions, that prevent global extinction. The heterogeneity of mixing may enable toxin producing and sensitive strains to coexist for very long time at strong mixing.     

Spatial patterns of coexistence of competing species in patchy habitat

The single-species spatially realistic patch occupancy metapopulation model is, in this study, extended to a metacommunity of many competing species. Competition is assumed to reduce the local carrying capacity (effective patch area), which in turn increases local extinction rates and reduces colonization rates because of smaller population sizes. Each species is described by three parameters: pre-competitive abundance (equilibrium incidence of patch occupancy, which reflects the rate of colonization in relation to extinction rate), the spatial range of migration, and competitive ability. The model ignores spatio–temporal correlations caused by interspecific interactions, because in metacommunities of unequal competitors inhabiting heterogeneous landscapes, correlations in the occurrence of species are driven more by patch heterogeneity than by competition. The model allows the calculation of multispecies equilibria in patchy habitats without simulations. In general, the number of coexisting species in the metacommunity increases with decreasing strength of competition, increasing rate of colonization, and decreasing range of migration. Habitat heterogeneity in the form of spatial variation in patch areas tends to facilitate coexistence. Poor competitors may coexist with superior competitors in the patch network if the former have higher colonization rates (competition–colonization trade-off). When migration distances are short, competition leads to spatial pattern formation: Species tend to have restricted spatial distributions in the network, but contrary to intuitive expectations, often the distributions of many species are nested. Having more dispersive species enhances both local and global diversity, whereas more local migration decreases local but increases global diversity.     

不同栖息地状态下物种竞争模式及模拟研究与应用

物种竞争是影响生态系统演化的重要生态过程之一.而物种在受人类影响出现不同程度毁坏的栖息地上的演化又是非常复杂的,因此研究物种演化对栖息地毁坏的响应是非常必要的.在Tilman研究工作的基础上,将竞争系数引入集合种群动力模式,建立了多物种集合种群竞争共存的数学模型,并对5-物种集合种群在不同栖息地状态下的竞争动态进行了计算机模拟研究.结果表明：（1）不同结构的群落（q值不同）,物种之间的竞争排斥作用强度不同,优势物种明显的群落,物种之间的排斥强度大;（2）随着栖息地毁坏程度的增加,对优势物种的负面影响逐渐减小,而对弱势物种的负面影响逐渐增加;（3）随着栖息地恢复幅度的增加,优势物种和弱势物种之间的竞争越强烈,优势物种受到的竞争排斥加大,而弱势物种逐渐变强,出现了强者变弱、弱者变强的格局;（4）物种竞争排斥与共存受迁移扩散能力和竞争能力影响很大,竞争共存的条件是其竞争能力与扩散能力呈非线性负相关关系;（5）竞争共存的物种的强弱序列发生了变化.     

Population size dependence, competitive coexistence and habitat destruction

1. Spatial dynamics can lead to coexistence of competing species even with strong asymmetric competition under the assumption that the inferior competitor is a better colonizer given equal rates of extinction. Patterns of habitat fragmentation may alter competitive coexistence under this assumption.
2. Numerical models were developed to test for the previously ignored effect of population size on competitive exclusion and on extinction rates for coexistence of competing species. These models neglect spatial arrangement.
3. Cellular automata were developed to test the effect of population size on competitive coexistence of two species, given that the inferior competitor is a better colonizer. The cellular automata in the present study were stochastic in that they were based upon colonization and extinction probabilities rather than deterministic rules.
4. The effect of population size on competitive exclusion at the local scale was found to have little consequence for the coexistence of competitors at the metapopulation (or landscape) scale. In contrast, population size effects on extinction at the local scale led to much reduced landscape scale coexistence compared to simulations not including localized population size effects on extinction, especially in the cellular automata models. Spatially explicit dynamics of the cellular automata vs. deterministic rates of the numerical model resulted in decreased survival of both species. One important finding is that superior competitors that are widespread can become extinct before less common inferior competitors because of limited colonization.
5. These results suggest that population size–extinction relationships may play a large role in competitive coexistence. These results and differences are used in a model structure to help reconcile previous spatially explicit studies which provided apparently different results concerning coexistence of competing species.     

人类周期性活动对物种多样性的影响及其预测

人类活动通过作用于栖息地而影响着物种的种群动态,从而影响着物种多样性的变化。首次提出了不同时间尺度人类周期性活动干扰下的多物种竞争动力模式,模拟了千年时间尺度,物种多样性对人类周期性活动的响应过程,开展了人类周期性活动所导致的物种多样性的量的变化的预测研究。有关模拟和预测结果表明:在人类周期性活动的作用下,物种多度变化对栖息地变化的响应也做准周期振荡;同时人类活动强度越大,物种多度振荡的幅度也越大;并且在栖息地减少过程中,人类周期性活动对物种多样性的影响幅度要小于栖息地扩充过程中的。在一个完整的周期内,并没有物种的灭绝,只是物种的多度和强弱关系变化很大。当人类周期性活动仅持续1/4个周期时,最强的几个物种将灭绝,而其它物种做准周期振荡,但振幅相对较小。     

Habitat complexity modifies ant–parasitoid interactions: implications for community dynamics and the role of disturbance

Species must balance effective competition with avoidance of mortality imposed by predators or parasites to coexist within a local ecological community. Attributes of the habitat in which species interact, such as structural complexity, have the potential to affect how species balance competition and mortality by providing refuge from predators or parasites. Disturbance events such as fire can drastically alter habitat complexity and may be important modifiers of species interactions in communities. This study investigates whether the presence of habitat complexity in the form of leaf litter can alter interactions between the behaviorally dominant host ants Pheidole diversipilosa and Pheidole bicarinata, their respective specialist dipteran parasitoids (Phoridae: Apocephalus sp. 8 and Apocephalus sp. 25) and a single species of ant competitor (Dorymyrmex insanus). We used a factorial design to manipulate competition (presence/absence of competitors), mortality risk (presence/absence of parasitoids) and habitat complexity (presence/absence of leaf litter). Parasitoid presence reduced soldier caste foraging, but refuge from habitat complexity allowed increased soldier foraging in comparison to treatments in which no refuge was available. Variation in soldier foraging behavior correlated strongly with foraging success, a proxy for colony fitness. Habitat complexity allowed both host species to balance competitive success with mortality avoidance. The effect of fire on habitat complexity was also studied, and demonstrated that the immediate negative impact of fire on habitat complexity can persist for multiple years. Our findings indicate that habitat complexity can increase dominant host competitive success even in the presence of parasitoids, which may have consequences for coexistence of subordinate competitors and community diversity in general.     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

Effects of predation pressure on species packing on a resource gradient: insights from nonlinear dynamics

The classical case of three competitors arranged on a resource gradient such that the central competitor will be excluded due to competition from the other two is studied from the point of view of the effects of added predators. The basic formulation is motivated by a desire to understand the effects of asymmetries in multidimensional Lotka-Volterra systems. We first study the effects of perfectly specialist predators and find a rich collection of possible behaviors of the system including (1) extinction of all predators and subsequent extinction of the subordinate competitor, (2) dominant competitors and their predators coexist but the subdominant competitor goes extinct, (3) all species except the predator of the subordinate competitor coexist in coordinated phase-reversed chaos, (4) exclusion of one or more species occurs through an expanding heteroclinic cycle, and (5) all species coexist in an uncoordinated chaos. We then study the effects of five qualitatively distinct forms of polyphagy. In one case, corresponding to the well-known vulnerability to predation versus competitive ability trade-off, it is possible to have the subordinate competitor be the only survivor in the system. The other three cases of polyphagy lead to distortions in the basic pattern seen in the previously analyzed specialist case. Studying this case of ecologically motivated asymmetries in the basic Lotka-Volterra formulation is a step in the direction of fully understanding interacting populations.     

The form of direct interspecific competition modifies secondary extinction patterns in multi‐trophic food webs

Forcibly removing species from ecosystems has important consequences for the remaining assemblage, leading to changes in community structure, ecosystem functioning and secondary (cascading) extinctions. One key question that has arisen from single‐ and multi‐trophic ecosystem models is whether the secondary extinctions that occur within competitive communities (guilds) are also important in multi‐trophic ecosystems? The loss of consumer–resource links obviously causes secondary extinction of specialist consumers (topological extinctions), but the importance of secondary extinctions in multi‐trophic food webs driven by direct competitive exclusion remains unknown. Here I disentangle the effects of extinctions driven by basal competitive exclusion from those caused by trophic interactions in a multi‐trophic ecosystem (basal producers, intermediate and top consumers). I compared food webs where basal species either show diffuse (all species compete with each other identically: no within guild extinctions following primary extinction) or asymmetric competition (unequal interspecific competition: within guild extinctions are possible). Basal competitive exclusion drives extra extinction cascades across all trophic levels, with the effect amplified in larger ecosystems, though varying connectance has little impact on results. Secondary extinction patterns based on the relative abundance of the species lost in the primary extinction differ qualitatively between diffuse and asymmetric competition. Removing asymmetric basal species with low (high) abundance triggers fewer (more) secondary extinctions throughout the whole food web than removing diffuse basal species. Rare asymmetric competitors experience less pressure from consumers compared to rare diffuse competitors. Simulations revealed that diffuse basal species are never involved in extinction cascades, regardless of the trophic level of a primary extinction, while asymmetric competitors were. This work highlights important qualitative differences in extinction patterns that arise when different assumptions are made about the form of direct competition in multi‐trophic food webs.     

Ecological Predictors of Extinction Risk in the Flora of Lowland England, UK

We used historical and contemporary records to determine the scale of plant extinction in Bedfordshire and Northamptonshire, and to assess whether extinct species share a range of ecological and phytogeographical traits. Since 1700 both counties have lost 94 species (11% of their native floras) with the rate of extinction increasing from 3.8 to 4.8 species per decade in the 19th century to 6–8 species per decade after 1950. The most important predictors of extinction risk were English range size and traits associated with habitat specialisation and competitive ability: poor competitors (i.e. short stress-tolerators) associated with open habitats with very low or high pH and soil moisture (e.g. lowland bogs, dwarf-shrub heath and acid and calcareous grassland) were much more likely to have become extinct in the study region than would have been expected by chance alone. Many of these species have very localised distributions and/or occur at the northern, southern or eastern edges of their range in southern England (i.e. Northern and Oceanic). In contrast, there was no clear or significant relationship between extinction and dispersal ability or reproductive mode. These findings, which parallel national trends, indicate that habitat loss and eutrophication have been the main causes of population extinction in lowland England over the last 300 years. However, more fine-scaled studies are required to assess whether ‘low-level’ stresses, such as habitat fragmentation, climate change and atmospheric pollution, are having additional impacts on populations already severely depleted by habitat loss, as well as to quantify changes in the abundance of more widespread species which are known to have declined over the same period.     

Habitat loss alters effects of intransitive higher-order competition on biodiversity: a new metapopulation framework

Recent studies have suggested that intransitive competition, as opposed to hierarchical competition, allows more species to coexist. Furthermore, it is recognized that the prevalent paradigm, which assumes that species interactions are exclusively pairwise, may be insufficient. More importantly, whether and how habitat loss, a key driver of biodiversity loss, can alter these complex competition structures (and therefore species coexistence) remain unclear. We thus present a new, simple yet comprehensive metapopulation framework that can account for any competition pattern and more complex higher-order interactions (HOIs) among species. We find that competitive intransitivity increases community diversity and that HOIs generally enhance this effect. Essentially, intransitivity promotes species richness by preventing the dominance of a few species, unlike the hierarchical competition, while HOIs facilitate species coexistence through stabilizing community fluctuations. However, variation in species’ vital rates and habitat loss can weaken or even reverse such higher-order effects, as their interaction can lead to a more rapid decline in competitive intransitivity under HOIs. Thus, it is essential to correctly identify the most appropriate interaction model for a given system before models are used to inform conservation efforts. Overall, our simple model framework provides a more parsimonious explanation for biodiversity maintenance than the existing theory.