种间互作网络的结构、生态系统功能及稳定性机制研究

生态群落中不同物种间发生多样化的相互作用, 形成了复杂的种间互作网络。复杂生态网络的结构如何影响群落的生态系统功能及稳定性是群落生态学的核心问题之一。种间互作直接影响到物质和能量在生态系统不同组分之间的流动和循环以及群落构建过程, 使得网络结构与生态系统功能和群落稳定性密切相关。在群落及生态系统水平上开展种间互作网络研究将为群落的构建机制、生物多样性维持、生态系统稳定性、物种协同进化和性状分化等领域提供新的视野。当前生物多样性及生态系统功能受到全球变化的极大影响, 研究种间互作网络的拓扑结构、构建机制、稳定性和生态功能也可为生物多样性的保护和管理提供依据。该文从网络结构、构建机制、网络结构和稳定性关系、种间互作对生态系统功能的影响等4个方面综述当前种间网络研究进展, 并提出在今后的研究中利用机器学习和多层网络等来探究环境变化对种间互作网络结构和功能的影响, 并实现理论和实证研究的有效整合。     

羌塘高寒草地物种多样性与生态系统多功能关系格局

传统的生物多样性-生态系统功能研究大多侧重于单一生态系统功能与物种多样性的关系,忽略了生态系统的重要价值在于其能够同时提供多种功能或服务,即生态系统的多功能性。基于藏北羌塘高寒草地样带调查数据,选取植被地上生物量、地下生物量、土壤全氮、硝态氮及铵态氮含量、土壤全磷含量、土壤有机碳储量等7个与植物生长、养分循环、土壤有机碳蓄积相关的参数来表征生态系统多功能性。采用上述参数转换为Z分数后的平均值计算多功能性指数(M)。分析了不同生物多样性指数与生态系统多功能指数的关系以及年降水量和年均温度对物种多样性和生态系统多功能性指数的影响。结果表明,物种丰富度指数与生态系统多功能性之间呈极显著的正相关关系,Shannon-wiener和Simpson物种多样性指数也与多功能性指数间呈显著的正相关,但多功能性指数与Pielou均匀度指数没有表现出明显的相关关系。物种丰富度与表征植物生长、养分循环以及土壤有机碳蓄积的生态系统功能指数间也均呈极显著的正相关关系。降水格局显著影响羌塘高原物种丰富度和生态系统多功能指数,二者均随年降雨量的增加而显著增加,但物种多样性指数并未与年降水量呈现显著相关关系。研究强调了群落物种丰富度即群落物种数量对维持生态系统多功能性的重要意义,这意味着由于人类活动导致的物种丧失可能会给藏北高寒草地生态系统多功能和生态服务带来更为严重的后果。就退化草地恢复或草地可持续管理而言,在藏北羌塘地区,本地植物种的物种丰富度恢复和维持应作为重要目标之一。     

长期施肥对黄土高原典型草原群落稳定性的影响及机制研究

施肥是当前草地生态系统最常见的人为干扰方式之一,可导致草地生物多样性和生态系统稳定性发生显著变化。该研究以黄土高原典型草原为研究对象,通过连续8年的氮肥(尿素)野外添加试验,分析不同氮肥处理(分别为0、5、10、20、40和80 g·m^-2)对草地群落稳定性的影响,并分析检验可能影响群落稳定性的四个潜在机制(物种多样性、物种同步性、投资组合效应和优势度),以明确施肥对群落稳定性的影响及其潜在机制,为黄土高原恢复草地的合理利用提供理论依据。结果表明:(1)随着施肥水平的增加,群落稳定性(S)降低,并在施肥处理为20 g·m^-2时开始极显著下降(P<0.01)。(2)施肥极显著降低了物种的丰富度(R)和优势度指数(D)(P<0.01),但物种同步性(B)和均值-方差的比例指数(z)并没有发生显著变化(P>0.05)。(3)物种丰富度(R)与群落稳定性(S)之间呈正相关关系,而优势度指数(D)与群落稳定性(S)不相关。研究认为,在人为施肥干扰下,黄土高原典型草原的物种丰富度下降导致群落稳定性下降,表明物种多样性对维持草地群落稳定具有重要作用。     

生态学的多样性指数:功能与系统发育

生物多样性是生态学的核心问题。传统的多样性指数仅包含物种数和相对多度的信息,这类基于分类学的多样性指数并不能很好地帮助理解群落构建和生态系统功能。不同物种对群落构建和生态系统功能所起到的作用类型和贡献也不完全相同,且物种在生态过程中的作用和贡献往往与性状密切相关,因此功能多样性已经成为反映物种群落构建、干扰以及环境因素对群落影响的重要指标。同时,由于亲缘关系相近的物种往往具有相似的性状,系统发育多样性也可以作为功能多样性的一个替代。功能多样性和系统发育多样性各自具有优缺点,但二者均比分类多样性更能揭示群落和生态系统的构建、维持与功能。     

基于支付意愿法的黄海海域物种多样性维持服务价值评估(2002-2012)

物种多样性维持是支撑生态系统正常运转的基础性支持服务功能,为生态系统向人类提供服务和生态福利发挥着关键作用。正确认识与评估物种多样性维持服务价值,是开展生态保护决策的基础性工作。基于支付意愿法(也称为条件价值法),开展了2002—2012年黄海物种多样性维持服务价值评估。结果显示,黄海海域沿岸城市居民对于17种黄海珍稀濒危物种生物多样性维持的支付意愿,呈现逐年上升的趋势。截至2012年,总支付意愿高达97.06亿元,其中山东省比例最高(45.75%),相对2002年增长了247%。沿岸9座城市居民对于黄海珍稀濒危物种生物多样性维持的人均支付意愿在121至259元之间,青岛市最高,其次为烟台市和南通市。17种珍稀濒危物种中,以黄岛长吻虫、白枕鹤、丹顶鹤的多样性维持服务价值最高,均超过了8亿元。为黄海生物多样性价值发现、价值展示、价值实现提供科学基础和理论支持。     

内蒙古典型草原建群种羊草基因型多样性抑制群落物种多样性的生态功能

植物群落的物种多样性以及群落建群种的基因型多样性对群落生态功能是否存在交互影响已成为群落生态学研究的热点内容。以内蒙古典型草原群落内常见物种为研究对象,研究了群落物种多样性与建群种羊草(Leymus chinensis)基因型多样性及其交互作用对群落生物量生态功能特性的影响。结果表明:(1)羊草基因型多样性、物种多样性及其交互作用对群落地上、地下和总生物量无显著影响(P0.05);(2)羊草基因型多样性、物种多样性及其交互作用对多样性效应(净多样性效应、互补效应和选择效应)有显著影响(P0.05)。羊草基因型多样性抑制多样性净效应的发挥,且主要抑制互补效应;而物种多样性则促进多样性净效应的发挥,主要表现为选择效应对地上生物量的正效应;(3)互补效应对群落生物量多样性净效应起主要贡献。实验所得结果不仅为探讨多样性效应在物种水平以及群落水平上对群落生物量的影响因素提供了重要启示,而且为内蒙古草原种质资源的保护及合理利用,乃至生态系统的恢复和重建提供理论指导。     

群落构建的中性理论和生态位理论

物种共存和生物多样性维持一直是生态学研究的中心论题。基于物种生态位分化的群落构建理论已经发展了近一个世纪, 但我们对群落构建和生物多样性维持的机理仍然不清楚。近年来, 群落中性理论以其简约性和预测能力成为群落生态学研究的焦点, 但由于其“物种在生态功能上等价”的假设与大量研究结果相悖, 同时对自然群落结构的准确预测也只限于少数的生态系统, 因而饱受质疑。如今, 越来越多的生态学家认为群落构建的生态位理论与中性理论之争的最终归宿应该是二者的整合。 在本文中, 我们在简要回顾生态位理论和群落中性理论发展的基础上, 分析二者之间的主要分歧和互补性, 试图梳理二者整合的途径。我们认为, 尽管中性理论的发展极大地丰富了群落构建理论, 但二者的整合尚处于初级阶段; 群落构建零模型假说、中性—生态位连续体假说、随机生态位假说等都不失为有价值的尝试, 今后需要在其他类型的生态系统中进行实验验证, 以更好地理解确定性过程和随机过程在决定群落构建和生物多样性维持中的作用。     

化学除草剂对农田生态系统野生植物多样性的影响

农田生物多样性是全球生物多样性的重要组成部分, 除草剂的大量施用对其产生了严重影响。本文综述了化学除草剂对农田生态系统中野生植物多样性的影响, 并分析归纳了其影响机制。除草剂的施用会使敏感植物减少, 抗药性植物增多, 从而改变农田生态系统中的野生植物物种组成, 并使其趋同化, 降低遗传多样性和物种多样性, 以致植物功能群单一化, 群落稳定性下降。除草剂的主要影响机制是杀死植物或改变其生长代谢、抗性、繁殖等, 改变生境, 并与人为因素、环境因素等产生协同影响。不同种类的除草剂影响程度不同, 且不同物种间、不同群落间的响应也存在差异。我国化学除草剂使用量持续增长, 应加强除草剂对野生植物多样性的影响及其机制研究, 重视除草剂使用历史记录和野生植物的长期监测, 以及除草剂使用规范和相关政策法律研究, 更好地保护我国农田生态系统中的生物多样性。     

生物多样性与森林生态系统健康的几个关键问题

生物多样性和生态系统健康问的关系,是生态学领域的一个重大科学问题,近年来已成为关注的热点.首先,生物多样性在复杂的时空尺度上维持着生态系统过程的运行,是生态系统功能得以维持的生物基础;其次,生物多样性是生态系统抗干扰能力、恢复能力及适应环境变化能力的物质基础.生态系统健康的维持取决于系统的生物多样性、可再生能力和生产力,维护生物多样性是生态系统管理计划中不可缺少的部分;第三,对生态系统健康程度的评价,很大程度上依赖于对生态过程的认识,而生物多样性及其生态系统功能又是认识和了解生态过程的基础.本文通过对森林生态系统异质性、种间关系及关键种、外来物种入侵、"绿色沙漠"等问题的论述,阐述了生物多样性与森林生态系统健康之间的关系,提出了我国所面临的问题和相应的对策.     

应用中性理论分析局域群落中的物种多样性及稳定性

如何解释群落中物种的丰富与稀少,并对物种多样性和群落稳定性进行合理的量化评价是群落生态学研究中的一个热点问题。20世纪中期,MacArthur将影响自然群落稳定性的因素归结为物种数量多少以及物种间相互作用的大小,20世纪末Doak等学者提出群落的容纳能力和物种间的维持机制是决定群落稳定性的关键因素。同时对群落结构及物种间维持机制的研究也有了新的突破,Hubbell提出"生物多样性与生物地理学统一的中性理论(Unified Neutral Theory of Biodiversity and Biogeography)"为群落生态学研究提供了新的思路和方法。从群落中性理论的基本假设出发,对Hubbell中性理论中局域群落的物种多度动态模型进行分析,归纳得出群落中性理论中物种多样性与群落稳定性之间的量化关系。封闭的局域群落中,出现物种灭绝或单物种独占的时间与群落大小及物种相对多度成正比,物种多样性程度的增加可延长物种灭绝或独占的时间;开放的局域群落中,物种多度期望值与局域群落大小、物种在集合群落中的物种相对多度成正比,周围群落中物种的灭绝会引起局域群落中相应物种的灭绝,最终导致整个生态群落物种多样性的降低;群落中物种多度的方差与局域群落大小、迁移率、物种在集合群落中的物种相对多度相关,局域群落物种多度的波动幅度随着群落间生态隔离的减弱或物种多样性程度的增加而减小。由此,集合群落物种多样性是影响局域群落物种多样性的重要因素,生态隔离程度的减弱及物种多样性的增加都将增强群落的稳定性。     

Relationships between functional diversity and ecosystem functioning: A review

Functional diversity, which is the value, variation and distribution of traits in a community assembly, is an important component of biodiversity. Functional diversity is generally viewed as a key to understand ecosystem and community functioning. There are three components of functional diversity, i.e. functional richness, evenness and divergence. Functional diversity and species diversity can be either positively or negatively correlated, or uncorrelated, depending on the environmental conditions and disturbance intensity. Ecosystem functioning includes ecosystem processes, ecosystem properties and ecosystem stability. The diversity hypothesis and the mass ratio hypothesis are the two major hypotheses of explaining the effect of functional diversity on ecosystem functioning, diversity hypothesis reflects that organisms and their functional traits in a assemblage effect on ecosystem functioning by the complementarity of using resources, and mass ratio hypothesis emphasises the identify of the dominant species in a assemblage. These two hypotheses do not contradict each other and instead they reflect the two different sides of functional diversity and functional composition. The effect of functional diversity on ecosystem functioning also depends on abiotic factors, perturbation, management actions, etc. Function diversity potentially influences ecosystem service and management by effecting on ecosystem functioning. Ecosystem management groups should include functional diversity in their scheme and not just species richness.     

Biodiversity influences ecosystem functioning in aquatic angiosperm communities

Knowledge of the connection between aquatic plant diversity and ecosystem processes is still limited. To examine how plant species diversity affects primary productivity, plant nutrient use, functional diversity of secondary producers and population/community stability, we manipulated submerged angiosperm species diversity in a field experiment lasting 15 weeks. Plant richness increased the shoot density for three of four species. Polyculture biomass production was enhanced by increasing richness, with positive complementarity and selection effects causing positive biodiversity effects. Species richness enhanced the community stability for biomass production and shoot density. Sediment ammonium availability decreased with plant diversity, suggesting improved nutrient usage with increasing plant richness. Interestingly, positive multitrophic effects of plant species richness on structural and functional diversity of macrobenthic secondary producers were recorded. The results suggest that mixed seagrass meadows play an important role for ecosystem functioning and thus contribute to the provision of goods and services in coastal areas.     

物种多样性和功能群多样性与生态系统生产力的关系——以内蒙古短花针茅草原为例

生物多样性与生态系统生产力之间的关系是当前生态学领域的热点问题。短花针茅(Stipa breviflora)草原是内蒙古荒漠草原的主要类型, 生态系统脆弱, 气候波动剧烈, 研究内蒙古短花针茅草原生物多样性与生产力的关系具有十分重要的意义。该研究在内蒙古短花针茅草原区设置了202个样地进行群落调查, 在干旱区及半干旱区两种资源供给下, 分析了物种丰富度、功能群丰富度与生产力的关系, 旨在解决两个科学问题: 1)物种多样性和功能群多样性中, 哪一种与生产力关系更为密切?2)资源供给对多样性和生产力关系的影响。结果表明: 1)物种丰富度、群落生产力与年降水量呈正相关关系, 而功能群丰富度与年降水量之间不存在显著相关性; 2)群落生产力随物种丰富度的增加而增加, 且两者间呈正线性关系, 功能群丰富度与生产力之间不存在显著相关关系; 3)资源供给会影响多样性与生产力之间的关系, 资源供给低时, 多样性对生产力贡献较低, 资源供给高时, 多样性对生产力的贡献较高。该研究丰富了多样性与生产力关系的研究, 同时, 考虑到植物功能性状的研究在近几年受到生态学家的重视, 且多数研究集中于小尺度的人工控制实验, 因此, 在大尺度自然生态系统中开展功能性状多样性与生态系统功能关系的研究将十分必要。     

Linking species functional roles to their network roles

Species roles in ecological networks combine to generate their architecture, which contributes to their stability. Species trait diversity also affects ecosystem functioning and resilience, yet it remains unknown whether species’ contributions to functional diversity relate to their network roles. Here, we use 21 empirical pollen transport networks to characterise this relationship. We found that, apart from a few abundant species, pollinators with original traits either had few interaction partners or interacted most frequently with a subset of these partners. This suggests that narrowing of interactions to a subset of the plant community accompanies pollinator niche specialisation, congruent with our hypothesised trade‐off between having unique traits vs. being able to interact with many mutualist partners. Conversely, these effects were not detected in plants, potentially because key aspects of their flowering traits are conserved at a family level. Relating functional and network roles can provide further insight into mechanisms underlying ecosystem functioning.     

Nitrogen enrichment weakens ecosystem stability through decreased species asynchrony and population stability in a temperate grassland

Biodiversity generally promotes ecosystem stability. To assess whether the diversity–stability relationship observed under ambient nitrogen (N) conditions still holds under N enriched conditions, we designed a 6‐year field experiment to test whether the magnitude and frequency of N enrichment affects ecosystem stability and its relationship with species diversity in a temperate grassland. Results of this experiment showed that the frequency of N addition had no effect on either the temporal stability of ecosystem and population or the relationship between diversity and stability. Nitrogen addition decreased ecosystem stability significantly through decreases in species asynchrony and population stability. Species richness was positively associated with ecosystem stability, but no significant relationship between diversity and the residuals of ecosystem stability was detected after controlling for the effects of the magnitude of N addition, suggesting collinearity between the effects of N addition and species richness on ecosystem stability, with the former prevailing over the latter. Both population stability and the residuals of population stability after controlling for the effects of the magnitude of N addition were positively associated with ecosystem stability, indicating that the stabilizing effects of component populations were still present after N enrichment. Our study supports the theory predicting that the effects of environmental factors on ecosystem functioning are stronger than those of biodiversity. Understanding such mechanisms is important and urgent to protect biodiversity in mediating ecosystem functioning and services in the face of global changes.     

How biological diversity influences ecosystem function: a test with a tropical stream detritivore guild

We investigated the relationship between diversity and ecosystem function, which is controversial and has rarely been examined for consumer assemblages, for the process of leaf breakdown by the shredder guild in a tropical stream. We manipulated species richness, evenness and identity of four macroinvertebrate shredder species (three caddisflies and one mayfly) in microcosms and tested their effect on leaf breakdown rates measured as leaf mass loss per capita and per milligram of animal. Species richness, evenness and species identity all affected leaf breakdown rates. Breakdown rates tended to increase with higher richness, but only for the three caddisflies, probably through a release of intraspecific interference, although other mechanisms such as niche complementarity or facilitation cannot be discarded. Leaf breakdown by the caddisflies was reduced in the presence of the mayfly, possibly because of its mode of movement by swimming instead of crawling and its similarity to some predators that are common in leaf litter. Species identity was more important than species richness in determining leaf breakdown rates, indicating that some species within the shredder guild are not redundant, and suggesting important consequences of particular species loss for the functioning of the ecosystem.     

Functional diversity predicts overyielding effect of species combination on primary productivity

The effect of biodiversity on ecosystem functioning has proven variable both within and among manipulative studies. Species richness is the most commonly used measure of biodiversity in such studies, but the range of species’ functional traits (functional diversity), not the number of species per se, likely underpins a key mechanistic link between species richness and ecosystem functioning. However, the majority of experiments that have examined the effect of functional diversity have manipulated functional group richness, an approach recognised to suffer numerous limitations. Continuous measures of functional diversity avoid many of these limitations, but the relationship between continuous functional diversity and the magnitude of ecosystem processes has been largely untested. Using one vs two‐species mixtures of rock pool macroalgae as a model, we conducted a field experiment to determine the effect of a continuous measure of functional diversity (functional attribute diversity, FAD, the degree of functional differentiation based on four functional traits) on the magnitude of net primary productivity and overyielding, based upon two alternative null‐models. The total magnitude of productivity was largely determined by the identity of species present, not FAD. However, FAD proved to be a good predictor of overyielding (variation in productivity after the dominant effects of species identity had been accounted for). Furthermore, despite differences in the mean magnitude of the effect of combining species, the positive relationship between FAD and overyielding was consistent according to both additive and substitutive null‐models. Our findings imply that whilst knowledge of species’ independent contributions remains indispensable in the prediction of biotic effects on ecosystem functioning within a trophic level, continuous measures of functional diversity should be used as a supplementary tool to predict the magnitude of overyielding, thereby refining predictions.     

Population density of North American elk: effects on plant diversity

Large, herbivorous mammals have profound effects on ecosystem structure and function and often act as keystone species in ecosystems they inhabit. Density-dependent processes associated with population structure of large mammals may interact with ecosystem functioning to increase or decrease biodiversity, depending on the relationship of herbivore populations relative to the carrying capacity (K) of the ecosystem. We tested for indirect effects of population density of large herbivores on plant species richness and diversity in a montane ecosystem, where increased net aboveground primary productivity (NAPP) in response to low levels of herbivory has been reported. We documented a positive, linear relationship between plant-species diversity and richness with NAPP. Structural equation modeling revealed significant indirect relationships between population density of herbivores, NAPP, and species diversity. We observed an indirect effect of density-dependent processes in large, herbivorous mammals and species diversity of plants through changes in NAPP in this montane ecosystem. Changes in species diversity of plants in response to herbivory may be more indirect in ecosystems with long histories of herbivory. Those subtle or indirect effects of herbivory may have strong effects on ecosystem functioning, but may be overlooked in plant communities that are relatively resilient to herbivory.     

Quantifying effects of biodiversity on ecosystem functioning across times and places

Biodiversity loss decreases ecosystem functioning at the local scales at which species interact, but it remains unclear how biodiversity loss affects ecosystem functioning at the larger scales of space and time that are most relevant to biodiversity conservation and policy. Theory predicts that additional insurance effects of biodiversity on ecosystem functioning could emerge across time and space if species respond asynchronously to environmental variation and if species become increasingly dominant when and where they are most productive. Even if only a few dominant species maintain ecosystem functioning within a particular time and place, ecosystem functioning may be enhanced by many different species across many times and places (β‐diversity). Here, we develop and apply a new approach to estimate these previously unquantified insurance effects of biodiversity on ecosystem functioning that arise due to species turnover across times and places. In a long‐term (18‐year) grassland plant diversity experiment, we find that total insurance effects are positive in sign and substantial in magnitude, amounting to 19% of the net biodiversity effect, mostly due to temporal insurance effects. Species loss can therefore reduce ecosystem functioning both locally and by eliminating species that would otherwise enhance ecosystem functioning across temporally fluctuating and spatially heterogeneous environments.     

Predicting productivity in tropical reservoirs: The roles of phytoplankton taxonomic and functional diversity

Primary productivity is intimately linked with biodiversity and ecosystem functioning. Much of what is known today about such relationship has been based on the manipulation of species richness. Other facets of biodiversity, such as functional diversity, have been neglected within this framework, particularly in freshwater systems. We assess the adequacy of different diversity measures, from species richness and evenness, to functional groups richness and functional diversity indices, to predict primary productivity in 19 tropical reservoirs of central Brazil, built to generate hydroelectric energy. We applied linear mixed models (and model selection based on the Akaike’s information criterion) to achieve our goal, using chlorophyll-a concentration as a surrogate for primary productivity. A total of 412 species were collected in this study. Overall we found a positive relation between productivity and diversity, with functional evenness representing the only exception. The most parsimonious models never included functional group classifications, with at least one continuous measure of functional diversity being present in many models. The best model included only species richness and explained 24.1% of variability in productivity. We therefore advise the use of species richness as an indicator of productivity in tropical freshwater environments. However, since the productivity–diversity relationship is known to be scale dependent, we recommend the use of continuous measures of functional diversity in future biodiversity and ecosystem functioning studies, in order to be certain that all functional differences between communities are being accounted for.