Food web structure and habitat loss

In this paper we explore simple food web models to study how metacommunity structure affects species response to habitat loss. We find that patch abundances and extinction thresholds vary according to the kind of food web. Second, for intermediate species, a slight decrease in the exploration cost of the better competitor has a strong effect on the extinction threshold of the poorer competitor. When predicting extinction risk one should consider not only the amount of habitat destroyed, but also the structure of the food web in which species are embedded. Both direct and indirect interactions are critical for predicting the consequences of habitat destruction.     

Metapopulation models for extinction threshold in spatially correlated landscapes

Simple analytical models assuming homogeneous space have been used to examine the effects of habitat loss and fragmentation on metapopulation size. The models predict an extinction threshold, a critical amount of suitable habitat below which the metapopulation goes deterministically extinct. The consequences of non-random loss of habitat for species with localized dispersal have been studied mainly numerically. In this paper, we present two analytical approaches to the study of habitat loss and its metapopulation dynamic consequences incorporating spatial correlation in both metapopulation dynamics as well as in the pattern of habitat destruction. One approach is based on a measure called metapopulation capacity, given by the dominant eigenvalue of a "landscape" matrix, which encapsulates the effects of landscape structure on population extinctions and colonizations. The other approach is based on pair approximation. These models allow us to examine analytically the effects of spatial structure in habitat loss on the equilibrium metapopulation size and the threshold condition for persistence. In contrast to the pair approximation based approaches, the metapopulation capacity based approach allows us to consider species with long as well as short dispersal range and landscapes with spatial correlation at different scales. The two methods make dissimilar assumptions, but the broad conclusions concerning the consequences of spatial correlation in the landscape structure are the same. Our results show that increasing correlation in the spatial arrangement of the remaining habitat increases patch occupancy, that this increase is more evident for species with short-range than long-range dispersal, and that to be most beneficial for metapopulation size, the range of spatial correlation in landscape structure should be at least a few times greater than the dispersal range of the species.     

Habitat destruction and extinction in competitive and mutualistic metacommunities

Because habitat loss is a leading cause of extinction, it is important to identify what kind of species is most vulnerable. Here, I use algebraic and graphical techniques to study metacommunity models of weak competition or locally facultative mutualism in which species may coexist within patches. Because a competition–colonization trade‐off is not required for regional coexistence of competitors, poor competitors are often regionally rare and most prone to extinction, in contrast to results from previous models of strongly competitive metapopulations. Metacommunities of mutualists can suffer the abrupt extinction of both species as habitat destruction is increased. These highlight the importance of identifying the mechanisms by which species coexist to predict their response to habitat loss.     

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.     

An integrative framework of coexistence mechanisms in competitive metacommunities

Species distribution in a metacommunity varies according to their traits, the distribution of environmental conditions and connectivity among localities. These ingredients contribute to coexistence across spatial scales via species sorting, patch dynamics, mass effects and neutral dynamics. These mechanisms however seldom act in isolation and the impact of landscape configuration on their relative importance remains poorly understood. We present a new model of metacommunity dynamics that simultaneously considers these four possible mechanisms over spatially explicit landscapes and propose a statistical approach to partition their contribution to species distribution. We find that landscape configuration can induce dispersal limitations that have negative consequences for local species richness. This result was more pronounced with neutral dynamics and mass effect than with species sorting or patch dynamics. We also find that the relative importance of the four mechanisms varies not only among landscape configurations, but also among species, with some species being mostly constrained by dispersal and/or drift and others by sorting. Changes in landscape properties might lead to a shift in coexistence mechanisms and, by extension, to a change in community composition. This confirms the importance of considering landscape properties for conservation and management. Our results illustrate the idea that ecological communities are the results of multiple mechanisms acting at the same time and complete our understanding of spatial processes in competitive metacommunities.     

物种演化对人类活动作用下不同性质栖息地毁坏的响应

栖息地毁坏是物种多样性减少的首要因素之一, 因此研究物种演化对栖息地毁坏的响应是非常必要的。而栖息地的毁坏又有瞬间毁坏和持续毁坏两种, 以往对栖息地毁坏的研究集中在瞬间毁坏上, 而该文则是通过N_物种 竞争共存模型分析对比了物种演化对栖息地瞬间毁坏和持续毁坏的响应特征。研究发现 :不同性质的栖息地毁坏都会导致物种强弱关系的变化, 并非如通常所认为的强物种将免于遭受物种灭绝的威胁, 也不是强物种首先灭绝, 而是因集合种群结构的不同而异。在热带雨林群落, 瞬间毁坏下物种演化一般经历了强迫适应和恢复上升阶段, 而持续毁坏下物种得不到恢复, 只能持续衰退, 在较长一段时间内持续毁坏比瞬间毁坏更有利于物种的续存 ;而在温带森林群落, 瞬间毁坏下物种演化一般经历强迫适应, 恢复上升和准周期振荡, 最后平衡, 而持续毁坏下物种只能持续衰退, 出现了在栖息地持续毁坏率小于瞬间毁坏率时, 物种的栖息地占有率却小于瞬间毁坏时的占有率。     

高阶作用对植物-传粉者-植食动物集合群落续存的影响

高阶作用通常指一个物种对另外两个物种之间相互作用强度的影响,对物种共存、群落构建及生物多样性具有重要影响。在集合种群水平上考虑了植食动物对动植物传粉关系造成的高阶作用,以及植食动物对传粉者的间接作用。通过分析基本生态过程,建立植物-传粉者-植食动物的集合群落模型,模型清楚地展示高阶作用和间接作用,可以用来研究它们对集合群落稳定性和续存的影响。结果表明：(1)互惠关系在集合群落尺度上会引起双稳态现象,说明了群落动态对初始条件的依赖性;(2)正高阶作用能够扩大集合群落双稳态的参数范围,负高阶作用和间接作用缩小它的参数范围,但都不会从本质上改变双稳态现象;(3)正高阶作用能够降低集合群落的灭绝阈值,增加集合群落稳定时的占有率,有利于集合群落续存,而负高阶作用和间接作用不利于续存。研究结果说明高阶和间接作用对调节多物种系统动态和物种共存具有重要作用。     

The competitive dynamics of metapopulations subject to the Allee-like effect

It is well recognized that individuals of many species can benefit from the presence of conspecifics, a concept broadly referred to as the Allee effect. At the metapopulation level, there is an analogous but essentially different phenomenon called the Allee-like effect that leads to metapopulation extinction thresholds at low habitat occupancy. But so far not adequate attention has been paid to this phenomenon. In this paper, the Allee-like effect is introduced into a metapopulation model of one species and also that of a three-state two-species competitive system. Phase plane analysis is used to investigate the dynamics of these models. We demonstrate that the Allee-like effect alone could lead to multiple stable states in three-state two-species competitive systems at the metapopulation level, and the number of stable states decrease as the Allee-like effect becomes more severe. Severe Allee-like effects may make coexistence impossible and may even lead to the extinction of both species even if their initial habitat occupancies are high and suitable habitats are enough. It is especially noticeable that depending on their initial conditions one species may exclude the other one that subjects to a weaker Allee-like effect than the former, while the second species always excludes the first one when both species are assumed to be in the absence of the Allee-like effect. We also investigate the habitat destructive effect on the Allee-like system mentioned above. Research indicates that the existence of the Allee-like effect makes a metapopulation more susceptible to habitat destruction. All in all, the Allee-like effect is probably a destabilizing factor that, together with habitat destruction, would affect the continuous existence of species. These conclusions may have important implications for conservation and metacommunity organization.     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

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.     

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.     

Coexistence in metacommunities: a tree-species model

Simple patch-occupancy models of competitive metacommunities have shown that coexistence is possible as long as there is a competition-colonization tradeoff such as that of superior competitors and dispersers. In this paper, we present a model of competition between three species in a dynamic landscape, where patches are being created and destroyed at a different rate. In our model, species interact according to a linear non-transitive hierarchy, such that species Y(3) outcompetes and can invade patches occupied by species Y(2) and this species in turn can outcompete and invade patches occupied by the inferior competitor Y(1). In this hierarchy, inferior competitors cannot invade patches of species with higher competitive ability. Analytical results show that there are regions in the parameter space where coexistence can occur, as well as regions where each of the species exists in isolation depending on species' life-history traits associated with their colonization abilities and extinction proneness as well as with the dynamics of habitat patches. In our model, the condition for coexistence depends explicitly on patch dynamics, which in turn modulate the limiting similarity for species coexistence. Coexistence in metacommunities inhabiting dynamic landscapes although possible is harder to attain than in static ones.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

Metapopulation Persistence in Random Fragmented Landscapes

Habitat destruction and land use change are making the world in which natural populations live increasingly fragmented, often leading to local extinctions. Although local populations might undergo extinction, a metapopulation may still be viable as long as patches of suitable habitat are connected by dispersal, so that empty patches can be recolonized. Thus far, metapopulations models have either taken a mean-field approach, or have modeled empirically-based, realistic landscapes. Here we show that an intermediate level of complexity between these two extremes is to consider random landscapes, in which the patches of suitable habitat are randomly arranged in an area (or volume). Using methods borrowed from the mathematics of Random Geometric Graphs and Euclidean Random Matrices, we derive a simple, analytic criterion for the persistence of the metapopulation in random fragmented landscapes. Our results show how the density of patches, the variability in their value, the shape of the dispersal kernel, and the dimensionality of the landscape all contribute to determining the fate of the metapopulation. Using this framework, we derive sufficient conditions for the population to be spatially localized, such that spatially confined clusters of patches act as a source of dispersal for the whole landscape. Finally, we show that a regular arrangement of the patches is always detrimental for persistence, compared to the random arrangement of the patches. Given the strong parallel between metapopulation models and contact processes, our results are also applicable to models of disease spread on spatial networks.     

Climate change and elevated extinction rates of reptiles from Mediterranean Islands

Recent climate change has caused the distributions of many species to shift poleward, yet few empirical studies have addressed which species will be vulnerable to longer-term climate changes. To investigate past consequences of climate change, we calculated the population extinction rates of 35 reptile species from 87 Greek land-bridge islands in the Mediterranean that occurred over the past 16,000 years. Population extinction rates were higher for those species that today have more northern distributions. We further found that northern species requiring cool, mesic habitats had less available suitable habitat among islands, implicating loss of suitable habitat in their elevated extinction rates. These extinctions occurred in the context of increasing habitat fragmentation, with islands shrinking and separating as sea levels rose. Thus, the circumstances faced by reptiles on the islands are similar to challenges for numerous species today that must cope with a changing climate while living in an increasingly human-fragmented landscape. Our island-biogeographical approach to investigating historical population extinctions gives insight into the long-term patterns of species responses to climate change.     

Habitat specialization predicts genetic response to fragmentation in tropical birds

Habitat fragmentation is one of the most severe threats to biodiversity as it may lead to changes in population genetic structure, with ultimate modifications of species evolutionary potential and local extinctions. Nonetheless, fragmentation does not equally affect all species and identifying which ecological traits are related to species sensitivity to habitat fragmentation could help prioritization of conservation efforts. Despite the theoretical link between species ecology and extinction proneness, comparative studies explicitly testing the hypothesis that particular ecological traits underlies species‐specific population structure are rare. Here, we used a comparative approach on eight bird species, co‐occurring across the same fragmented landscape. For each species, we quantified relative levels of forest specialization and genetic differentiation among populations. To test the link between forest specialization and susceptibility to forest fragmentation, we assessed species responses to fragmentation by comparing levels of genetic differentiation between continuous and fragmented forest landscapes. Our results revealed a significant and substantial population structure at a very small spatial scale for mobile organisms such as birds. More importantly, we found that specialist species are more affected by forest fragmentation than generalist ones. Finally, our results suggest that even a simple habitat specialization index can be a satisfying predictor of genetic and demographic consequences of habitat fragmentation, providing a reliable practical and quantitative tool for conservation biology.     

物种灭绝对不同时间尺度人类活动的响应机制研究

通过修改Tilman的多物种共存的经典模式中栖息地毁坏率（D）,使D随时间的推移呈线性增长情况下,本文模拟了百万年、万年和百年尺度人类活动对栖息地的破坏下,物种灭绝对栖息地毁坏的响应特征。结果表明,大时间尺度人类活动对栖息地毁坏导致物种的强弱关系发生变化,并且强物种先灭绝,而小时间尺度人类活动对栖息地破坏是弱物种先灭绝;在百万年和万年尺度上,物种对栖息地毁坏的响应是减幅振荡衰退直至灭绝,并且最强物种对栖息地的占有率（q）越大,振幅越大,而在百年尺度上,物种的演化几乎是直线衰退;在大时间尺度的栖息地毁坏情况下,q越大,则物种灭绝起始时间和所有物种灭绝的时间越长;而在较小的时间尺度的栖息地毁坏情况下,q越大,灭绝起始时间和所有物种最终灭绝的时间则越短。     

Metacommunity diversity depends on connectivity and patch arrangement in heterogeneous habitat networks

Connectivity is critical to the maintenance of biodiversity in fragmented landscapes, but its effects differ depending on the arrangement of linkages within a habitat network. Additionally, heterogeneity in habitat quality within the habitat network can alter patterns of diversity at local and regional scales in the metacommunity. Using a controlled experiment we examined the interactive effects of habitat connectivity, network form (linear vs square), and habitat patch quality on a moss‐inhabiting microarthropod community. We fragmented moss habitat while controlling for habitat loss, and altered habitat patch quality by regulating moisture conditions in landscapes differing in patch arrangement. Habitat patch quality had a significant effect on patterns of species richness, extinction, abundance and biomass. The effects of network form on diversity were strongest in heterogeneous landscapes. Gamma and beta diversity were greatest in continuous and linear landscapes. However, linear habitat networks showed marked patch specific edge effects that were detrimental to diversity under heterogeneous conditions. We provide direct evidence that habitat network structure impacts species community properties through mass effects, that are most evident when heterogeneity in habitat patch quality is present within the network. We conclude that habitat quality at the individual patch level and the distribution of high‐quality habitat within the network are important factors affecting biodiversity in metacommunities.     

集合种群强物种种群的演化特性

大量的数值模拟研究表明;(1)集合种群里最强物种种群对栖息地占有率相对较少时,较小的栖息地毁坏率就可以导致该最强物种种群与其它奇数物种种群一起退化;(2)而最强物种种群对栖息地占有率相对较多则是导致集合种群里弱物种种群集体灭绝的内因;(3)当栖息地的毁坏率大于最强物种种群对栖息地的占有率时,最强物种种群将先灭绝而使得弱物种种群进化为强物种种群或新的更强的强物种种群。