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。     

栖息地毁坏与动物物种灭绝关系的模拟研究

利用多个物种共存模式模拟了不同情况下的不同动物种群演化的动力学特性,研究结果表明：（1）由于栖息地的毁坏所导致的动手的种灭绝是依赖于对物种死亡率和有关平衡态的假设的,不同的假设下,既使栖息地的破坏率相同,灭绝的物种可能是竞争能力最强的若干物种,也可能是竞争能力相对较弱的若干物种,既不象传统的物种进化理论所认为的必是弱的物种先灭绝,也不象Tilman等人所认为的一定是最强的若干物种先灭绝;（2）如果弱的物种具有较高的平均死亡率,则当栖息地受到一定的毁坏时,将有较多强的物种灭绝,而且物种灭绝时间将大大缩短;（3）在物种死亡率不变的情形下,物种在未受毁坏栖息地上的平衡态和大占有率pl^0,将有利于物种的生存。     

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。     

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。     

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.     

集聚效应下的集合种群动力模式

引入种群集聚度和集聚效应的概念,通过建立物种的集聚效应模型,结合经典的Tilman集合种群模式,创建了集聚效应下的集合种群动力模式。通过大量的数值模拟分析栖息地未毁坏下的集合种群演化规律与集聚效应的关系,得到:(1)即使生境没有毁坏,种群的集聚效应也会影响种群的演化。(2)集合种群系统中不同种群对集聚效应反应有异同,相同点是各种群都要经历一段准周期波动才达到平衡态。不同点是不同种群对集聚效应反应的强度不一致,竞争能力越强的种群准周期波动的振幅越大,频率越低。竞争能力越弱的种群准周期波动的振幅越小,频率越高。(3)不同的群落对集聚效应的响应也不一致。优势种群相对明显的群落对集聚效应的响应幅度相对较小。(4)在优势种群明显的群落中,集聚效应对弱物种非常不利,弱物种很有可能由于集聚效应而灭绝。(5)群落或n集合种群里的各种群的集聚效应和建群种(或最优势种)的强弱是决定景观生态序的最为重要的2个因素。(6)每个物种对不同的不适集聚程度的响应不一致。不适集聚程度越大,物种演化波动幅度越大,频率越高。     

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。     

似Allee效应对2-物种集合种群竞争动态的影响

集合种群具有与局域种群Allee效应相似的现象被称为似Allee效应.将似Allee效应引入2-竞争物种集合种群系统,建立了具有似Allee效应的2-物种集合种群演化动态模型.大量的数值模拟表明:(1)似Allee效应导致集合种群水平上两竞争物种构成的系统具有多个平衡态;(2)似Allee效应使竞争共存物种无法续存甚至全部灭绝,即使种群具有很高的初始斑块占有率,并且最终平衡态随初始斑块占有率变化而改变;(3)似Allee效应可能使竞争排斥物种共同灭绝,且效应越强,物种存活时间越短;但似Allee效应不会增强强物种对弱物种的排斥强度,反而可能使强物种变为弱物种,弱物种变为强物种,其具有与栖息地毁坏类似的影响种群竞争等级排序的作用;(4)似Allee效应对竞争集合种群续存是一个不稳定的干扰因素,微小的变化都将引起系统平衡态的剧变.但对于已经达到平衡态的集合种群系统,似Allee效应对强弱种群多度起到调节与制约的作用,有助于平衡态集合种群的稳定与共存,这一结论更完整的揭示了似Allee效应在竞争集合种群系统发展的不同阶段所起的不同作用.以上这些结论对物种保护及集合群落的管理具有重要的指导意义.     

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。     

不同栖息地状态下外来种入侵及对本地种生存影响的模拟

外来种入侵引起的生态及经济问题日益严重,人类活动导致大规模栖息地持续变化的背景下,外来种的入侵过程及本地物种多样性的演化更为复杂.在Tilman多物种共存模型基础上,建立了栖息地持续变化条件下的外来种入侵干扰模式,分别模拟了栖息地不变、持续毁坏及持续恢复条件下的外来种入侵及对本地物种多样性的生态影响.模拟结果表明:(1)拓殖率小(入侵性弱)的外来种不能成功定殖,栖息地的持续变化对其没有任何影响.(2)拓殖率增大的外来种入侵过程复杂,栖息地持续毁坏导致其灭绝时间相比于栖息地不变时明显推迟,栖息地持续恢复使其灭绝提前;一定时间内持续增加栖息地可以减小此类外来种入侵危害.(3)拓殖率足够大的外来种能够成功定殖,快速定殖-扩散的入侵过程不会因栖息地毁坏而迅速改变,其响应具有时间滞后性;栖息地持续恢复也有利于其迅速蔓延,占有率呈线性疯狂增长.(4)在具有外来种入侵的本地生态系统中,栖息地持续改变(增加或减少)对本地物种多样性的发展均不利.     

Habitat destruction and the extinction debt revisited: The Allee effect

Habitat destruction, often caused by anthropogenic disturbance, can lead to the extinction of species at an unprecedented rate. It is important, therefore, to consider habitat destruction when assessing population viability. Another factor often ignored in population viability analysis, is the Allee effect that adds to the risk of populations already on the verge of extinction. Understanding the Allee effect on species dynamics and response to habitat destruction has intrinsic value in conservation prioritization. Here, the Allee effect was considered in a multi-species hierarchical competition model. Results showed that species persistence declines dramatically due to the Allee effect, and certain species become more susceptible to habitat destruction than others. Two extinction orders emerged under habitat destruction: either the best competitor becomes extinct first or the best colonizer first. The extinction debt and order, as well as the time lag between habitat destruction and species extinction, were found to be determined by species abundance and the intensity of the Allee effect.     

Habitat fragmentation and population extinction of birds

It has not been established that a major cause of extinction in birds or any other taxa is failure of metapopulation dynamics: the collapse of a network of ephemeral but discrete populations as movement between them becomes increasingly infrequent. The few data on who goes where and who mates with whom suggest that most species are structured as either a single large population or a small set of source populations and a larger set of sinks. The extinction of the latter is irrelevant to the persistence of the species. However, regional decline of a species in the face of habitat destruction and fragmentation can mimic a failure of metapopulation dynamics, because distinct aggregations of individuals will disappear much as they would if populations in an interacting network were eliminated one by one. Any species with highly restricted range is at great risk of extinction from spatially localized forces, such as cyclones or deforestation. Restricted range rather than inherent weakness is the main reason that so many island species have gone extinct or are endangered. Species with small populations in contact with much larger heterospecific ones with which they are interfertile are threatened with extinction by hybridization. Finally, the disappearance of a species from a site may be due to subtle habitat change, even if this observation seems superficially consistent with some general population theory, such as the dynamic equilibrium theory of island biogeography. Current theory is an inadequate substitute for intensive field studies as a means to address the conservation problems of individual species.     

生境破坏的模式对集合种群动态和续存的影响

构建了空间关联的集合种群模型,该模型不但包含了种群的空间结构信息,而且引入了破坏生境的全局密度和局部密度两个指标,它们描述了破坏生境的模式.模型揭示了破坏生境的空间分布格局复杂地影响了集合种群的动态和续存,破坏和未破坏生境斑块的均匀混合不利于集合种群的增长和续存,而生境类型聚集分布可以促进集合种群的快速增长和长期续存;对于两种斑块类型相对均匀混合的生境来说,均匀场假设可能会高估集合种群的续存,对于相对斑块类型高度聚集的生境,均匀场假设可能会低估集合种群的续存;物种的迁移范围也会影响集合种群的续存,迁移范围越大的物种越容易抵御生境的破坏而免遭灭绝.这意味着在生物保护中不能仅仅考虑生境的恢复和斑块质量的改善,生境结构的构建也是很重要的,加强生境斑块之间的连通性也有利于物种的长期续存.     

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。     

生境丧失对具有似Allee效应集合种群的影响及对策——以江苏盐城为例

似Allee效应对物种续存是潜在的扰动因素,稀有物种更易受其影响,可能增加生存于破碎化栖息地中的珍稀物种的死亡风险;但似Allee效应对多物种集合种群续存的影响及其在珍稀物种保护中的应用未能引起足够重视。将似Allee效应引入集合种群动力模式,建立了生境丧失下具有似Allee效应的n-珍稀物种的集合种群模式,并以江苏盐城滩涂湿地中的29种珍稀物种为研究实例。研究结果表明:(1)似Allee效应导致n-物种集合种群多度作长期变周期振荡,原本竞争共存物种可能无法继续共存,甚至灭绝。(2)似Allee效应增强对次强种及劣势种的生存极为不利,导致次强物种由强至弱灭绝,劣势物种由弱至强依次灭绝。(3)盐城天然湿地丧失29%后,11种劣势物种的集合种群由弱到强将最终依次灭绝,灭绝迟豫时间为304～890a,这些物种即Hanski所指的"活死者"。(4)适度增加栖息地面积是保护珍稀物种多样性的有效方法之一,在盐城现存3200km2的天然湿地基础上适度增加1801～2064km2左右栖息地面积,可以有效保护29种濒危物种的多样性,同时应注意结合针对具体物种的保护措施来提高濒危物种多度。研究结果对物种多样性保护及自然保护区建设具有重要的理论指导意义。     

Landscape Ecology of Eastern Spotted Skunks in Habitats Restored for Red‐Cockaded Woodpeckers

Although examples are rare, conflicts between species of conservation concern can result from habitat restoration that modifies habitat to benefit a single taxon. A forest restoration program designed to enhance habitat for endangered red‐cockaded woodpeckers (Picoides borealis) may be reducing available habitat for the eastern spotted skunk (Spilogale putorius), a forest‐adapted sympatric species of conservation concern that occurs in the Ouachita National Forest, Arkansas, U.S.A. At small scales, eastern spotted skunks select early successional forest with structural diversity, whereas red‐cockaded woodpeckers prefer mature pine (Pinus spp.) habitat. We surveyed for eastern spotted skunks at 50 managed forest stands, modeled occupancy as a function of landscape‐level habitat factors to examine how features of restoration practices influenced occurrence, and compared known occupied forest stands to those where active red‐cockaded woodpecker nesting clusters were located. The most‐supported occupancy models contained forest stand age (negatively associated) and size (positively associated); suggesting eastern spotted skunks primarily occupy large patches of habitat with dense understory and overhead cover. Red‐cockaded woodpecker nesting clusters were located in smaller and older forest stands. These results suggest that increased overhead cover, which can reduce risk of avian predation, enhances occupancy by small forest carnivores such as eastern spotted skunks. Management activities that increase forest stand rotation length reduce the availability of young dense forest. The practice may enhance the value of habitat for red‐cockaded woodpeckers, but may reduce the occurrence of eastern spotted skunks. Implementing plans that consider critical habitat and extinction risks for multiple species may reduce such conservation conflict.     

Are spatially correlated or uncorrelated disturbance regimes better for the survival of species?

The survival of species in dynamic landscapes (characterised by patch destruction and subsequent regeneration) depends on both the species' attributes and the disturbance pattern. Using a spatially explicit model we explored how the mean time to extinction of a metapopulation depends on the spatial correlation of patch destruction in relation to the population growth and dispersal abilities of species. Two contrasting answers are possible. On the one hand, increasing spatial correlation of patch destruction increases the spatial correlation of population growth and this is known to decrease metapopulation persistence. On the other hand, spatially correlated patch destruction and regeneration can lead to clustered habitat patches and this is known to increase metapopulation persistence. Therefore, we hypothesised that some species are better off under spatially correlated and alternatively uncorrelated disturbance regimes. However, contrary to this hypothesis, in all kinds of cases spatial correlation reduced metapopulation persistence. We found this to be due to the fact that the spatial correlation of patch destruction causes increasing temporal fluctuations in the regional carrying capacity of the metapopulation and is hence generally disadvantageous for long-term persistence. The main consequence for conservation biology is that reducing spatial correlation in disturbances is likely to be a reliable strategy in a dynamic landscape that will benefit practically all species with a low risk of adverse side effects .     

Fish species richness in spring‐fed ponds: effects of habitat size versus isolation in temporally variable environments

1. Species richness in a habitat patch is determined by immigration (regional) and extinction (local) processes, and understanding their relative importance is crucial for conservation of biodiversity. In this study, we applied the Island Biogeography concept to spring ponds connected to a river in southwestern Japan to examine how immigration and extinction processes interact to determine fish species richness in temporally variable environments. 2. Fish censuses were conducted 15 times in 13 study ponds at 1–4 month intervals from August 1998 through October 2000. Effects of habitat size (pond area), isolation (distance from the river) and temporal environmental variability (water level fluctuation) on (i) species richness, (ii) immigration and extinction rates and (iii) population size and persistence of each fish species were assessed. 3. The results revealed predominant effects of distance on species richness, immigration/extinction rates and population size and persistence. Species richness decreased with increasing distance but was not related to either pond area or water level fluctuation. A negative effect of distance on immigration rate was detected, while neither pond area nor water level fluctuation had significant effects on extinction rate. Further, population size and persistence of four species increased with decreasing distance, suggesting that, in ponds close to the river, immigrants from the river reduce the probability of extinction (i.e. provide a rescue effect), contributing to the maintenance of high species richness. 4. Overall results emphasise the importance of immigration processes, rather than extinction, in shaping patterns of species richness in our system. The predominant importance of immigration was probably because of (i) high temporal variability that negates habitat‐size effects and (ii) continuous immigration that easily compensates for local extinctions. Our results suggest that consideration of regional factors (e.g. connectivity, locations of source populations and barriers to colonisation) is crucial for conservation and restoration of local habitats.