基于功能性状的外来植物入侵预测模型框架构建

预测外来植物的潜在入侵性已成为生物多样性保护研究的重要内容,外来植物与乡土物种间的亲缘关系是预测外来植物能否成功入侵的一个重要途径。然而,达尔文归化难题却预测了两种截然不同的结果(即达尔文归化假说和预适应假说)。该研究解析了达尔文归化难题的内涵,提出了基于功能性状的外来植物与乡土群落间的相似性关系应该是进行外来植物入侵预测的重要切入点,而功能性状的种间分化与种内变异可能是外来植物成功入侵的两种不同生态策略。在此基础上,该研究还通过物种功能性状的多维超体积构建了外来植物与乡土群落间的相似性,提出了基于这种相似性的外来植物入侵预测的研究框架和基本流程。该模型框架的建立有助于理解外来植物的入侵机制,对外来植物的潜在入侵性预测提供了理论依据。当然,要实现外来植物能否成功入侵的准确预测,不仅依赖于功能性状的选择,还要考虑入侵的生境依赖性、空间尺度的重要性以及乡土群落的可入侵性等,未来的研究重点是通过控制实验对该模型进行验证和进一步完善。     

基于现代物种共存理论的入侵生态学概念框架

在入侵生态学60多年的发展历程中, 生态学家提出了多种多样的假说来解释生物入侵的机制。这些纷繁复杂的假说在丰富我们对生物入侵认知的同时, 也给入侵生态学概念的整合带来了困难。其中, 外来种和土著种是否存在生态学差异, 以及这种差异如何影响生物入侵, 是入侵生态学研究和争论的焦点问题。现代物种共存理论通过将外来种和土著种的生态学差异划分为生态位差异和适合度差异, 为入侵生态学概念的整合提供了新的视角。依据该理论, 外来种可以通过两种策略实现成功入侵: 一是扩大与土著种的生态位差异, 二是提高自身相较于土著种的适合度优势。因此, 外来种-土著种的生态位差异和适合度差异共同决定了入侵的成败与危害程度。通过对经典入侵假说进行梳理, 我们发现大部分假说都可以在该理论框架下进行解读, 不同假说的主要差别在于强调不同生态学过程对生态位和适合度差异的影响。同时, 这一理论框架很好地解释了为什么外来种-土著种的亲缘关系和性状差异会对生物入侵产生复杂的影响, 为达尔文归化谜团的和解以及外来种-土著种功能性状的比较研究提供了新的思路。目前, 现代物种共存理论还处于快速发展的阶段, 依旧存在很多不足, 但将其运用到生物入侵的实证研究中将是入侵生态学今后一个重要的发展方向。     

外来种-本地种亲缘关系对外来植物归化和入侵的影响

达尔文归化难题描述了外来种-本地种亲缘关系促进(预适应假说)或阻止(达尔文归化假说)外来种成功入侵的悖论。目前, 在中国仍缺少针对达尔文归化难题的研究。为系统研究外来种-本地种亲缘关系对中国外来植物入侵的影响, 该文利用线性混合效应模型从省级、市级和群落3个空间尺度以及归化、扩散和入侵3个阶段探究了外来种-本地种谱系距离和外来植物表现的关系。结果表明: 在省级和市级(区域)尺度上, 与本地种亲缘关系较近的外来植物更有可能在当地归化和扩散, 符合预适应假说的预期; 而在群落(局域)尺度上, 外来种-本地种亲缘距离与外来种是否在群落中成功定居及其入侵程度无关。该研究结果表明与本地区系亲缘较近的外来种和本地种的竞争并不强烈, 却能较好地适应本地气候环境而具有更强的归化和入侵潜力。因此, 在今后的外来植物管理和治理中需要尤其重视与本地区系亲缘关系较近的外来植物。     

Effects of exotic-native species relationship on naturalization and invasion of exotic plant species

达尔文归化难题描述了外来种-本地种亲缘关系促进(预适应假说)或阻止(达尔文归化假说)外来种成功入侵的悖论。目前, 在中国仍缺少针对达尔文归化难题的研究。为系统研究外来种-本地种亲缘关系对中国外来植物入侵的影响, 该文利用线性混合效应模型从省级、市级和群落3个空间尺度以及归化、扩散和入侵3个阶段探究了外来种-本地种谱系距离和外来植物表现的关系。结果表明: 在省级和市级(区域)尺度上, 与本地种亲缘关系较近的外来植物更有可能在当地归化和扩散, 符合预适应假说的预期; 而在群落(局域)尺度上, 外来种-本地种亲缘距离与外来种是否在群落中成功定居及其入侵程度无关。该研究结果表明与本地区系亲缘较近的外来种和本地种的竞争并不强烈, 却能较好地适应本地气候环境而具有更强的归化和入侵潜力。因此, 在今后的外来植物管理和治理中需要尤其重视与本地区系亲缘关系较近的外来植物。     

外来生物入侵中的达尔文归化谜团

针对外来生物入侵问题,达尔文在《物种起源》中提出了相反的两个假说,分别主张和本地物种亲缘关系近和远的外来物种更容易归化和入侵.这两个假说综合在一起现在被称为达尔文归化谜团.在沉寂了百余年后,随着系统发育和功能性状分析在生态学中的广泛运用,该谜团在近30年内迅速成为生物入侵领域研究的热点和前沿问题.但由于不同的研究往往得出矛盾的结论,错综复杂的研究结果使得该谜团的和解仍面临很大困难.本文梳理了达尔文归化谜团的内涵和外延,回顾了该谜团研究的发展历程和脉络,总结了近年来从空间尺度、入侵阶段、差异度量方式和控制性实验等角度尝试和解这一谜团的探索及其进展.本文认为,充分考虑环境梯度和人类活动的影响,综合分析入侵进程中的本地群落特征和动态,选择恰当的统计方法和零模型,并将该谜团的研究思路和理念引入到疾病生态学等诸多相关领域,是未来达尔文归化谜团研究的重点方向.对这一长达160余年谜团的探索,不仅有助于发展和完善入侵生态学理论,也将为外来生物入侵的防控提供重要科学指导.     

Scale dependence in the phylogenetic relatedness of alien and native taxa

达尔文的归化假说提出,由于生态位的不同,成功建群的外来物种与本地物种的关系不太密切。先前的研究对这一假设有支持也有反对,其中一个原因是外来物种和本地物种在大的空间尺度上有系统发育聚类的倾向,而在细微尺度上存在过度分散的倾向。然而,对于外来物种的系统发育关系如何改变其入侵群落的系统发育结构,以及在何种空间尺度上可能表现出这种影响,人们知之甚少。在本研究中,我们调查被入侵的森林下层植物群落在系统发育上是聚集的还是或过度分散的,亲缘关系如何随空间尺度变化,以及外来物种如何影响下层群落的系统发育模式。在澳大利亚东南部干旱森林的下层群落进行了5个空间尺度(1, 20, 500, 1500和4500 m²)的实地调查。使用两个指标的标准化效应量[(i)平均成对距离和(ii)平均最近分类单元距离]来量化群落与其外来和本地亚群落之间的系统发育关系,并研究系统发育模 式如何随空间尺度变化。研究结果表明,外来物种之间的亲缘关系非常密切,而且这种亲缘关系会随着尺度的增加而增加。在中等空间尺 度下(20–500 m²), 整个群落呈随机分布趋势,而本地物种高度分散,外来亚群落高度聚集。这说明亲缘关系密切的外来物种入侵使群落系统发育结构由过度分散向随机分布转变。外来物种和本地物种在空间尺度上是远亲,这支持了达尔文的归化假说,但只是在系统发育距离被量化为平均最近分类单元距离时成立。外来物种和本地物种的系统发育差异随着空间尺度的增加而增加,这与预期的模式相反。我们的研究结果表明,外来物种强大的系统发育聚类是由人类干预的引入驱动的,牵涉能够成功建群和传播的密切相关的类群。系统发育相关性的尺度依赖模式可能是由火灾和散布等随机过程引起的,这表明竞争和生境过滤并不是分别在小和大尺度上控制系统发育关系的唯一因素。区分不同进化深度的指标很重要,因为不同的指标可以显示不同的尺度依赖模式。     

群落组成及物种间相互作用对外来植物入侵的影响

外来植物入侵已成为严重的环境和社会问题,了解外来植物的入侵机制是有效控制其入侵的前提。生物阻抗假说认为,入侵地本地植物群落中的许多生物因子及生物过程能够抵御外来植物入侵。但关于群落抵抗外来种入侵的主要机制,目前还没有确定的结论。本文综述了群落中物种功能特征的多样性以及与外来种功能特征的相似度、植物与动物、植物与植物以及植物与土壤微生物间的相互作用等因素对外来植物入侵的影响,以及以前研究存在的不足。未来研究应该注重不同条件下植物与植物间的相互作用;不同竞争强度下,植物与食草动物的相互作用;植物、动物及土壤微生物三者之间的相互作用对外来植物入侵的影响。这些研究不仅能够丰富和完善入侵生态学理论,而且对于预测外来植物未来的扩散范围,合理有效地管理生态系统,防止外来植物入侵,保护本地生物多样性具有重要的实际意义。     

Contrasting responses of native and alien plant species to soil properties shed new light on the invasion of dune systems

在陆地生态系统中，沿海沙丘特别容易受到外来植物的入侵。许多研究将沙丘生境的入侵归因于人为因素，但对土壤性质和植物性状在植物 入侵中的作用却知之甚少。本研究考察了沙丘系统土壤特征与外来植物入侵的关系，重点研究了土壤养分、土壤盐分和植物功能特征之间的相互作 用。研究地点是马拉诺和格拉多泻湖(北亚得里亚海)的沙质堰洲岛。根据沙丘系统生态梯度上的主要生境(前沙丘、后沙丘和盐沼)，在10 个区域内选择100个地块(4 m × 4 m)。在每个地块中记录所有植物物种的发生和丰度，并收集一个土壤核。对每个土壤样品进行土壤质地、电导率(代表土壤盐 分)、有机碳和氮含量分析，并与本地和外来植物的种类数和覆盖度相关联。分析外来和本地物种主要的生殖功能和营养功能性状在生境中的变化。研究结果表明，土壤性质对外来物种库和本地物种库的影响不同，与植物整体多样性密切相关。在后沙丘(最易入侵)的生境中，高的土壤导电性限制了外来物种的数量，但土壤有机碳含量随着外来植物丰度的增加而增加，这也表明植物入侵与土壤之间存在潜在的反馈过程。只有在后沙丘生境中，随着环境条件的改善和植物竞争的加剧，本地植物和外来植物的功能性状谱有显著的趋同。本研究结果表明，在恶劣条件下，只有本地特有植物才能茁壮成长，而在中等条件下，土壤特性梯度与植物特性协同作用，抑制/促进外来植物丰富度。     

化感——外来入侵植物的“Novel Weapons”

综述了与外来植物入侵相关的理论假说。并主要介绍了从化感角度提出的“N ove lW eapons”假说(NW假说)。某些外来入侵植物能产生一些化学物质。这些物质可以发挥较强的化感作用,或者成为植物和土壤微生物之间相互作用的调节者。由于不同区域的植物群落共同进化的轨迹不同,被入侵群落的植物对这些化学物质缺乏适应性,因此,这些物质成为了外来种入侵的利器。而对外来种发源地的群落来说,由于长期的共存关系,这些物质对其伴生植物的影响相对较小。NW假说为化感作为外来植物入侵的一种机制奠定了理论基础,为外来入侵植物的防治提供了理论依据,但作为一种较新的理论假说,还需要更多的实验研究加以支持。     

可利用资源波动对外来种入侵的抵抗性

生物入侵带来的生态和经济危害引起了人们的广泛关注。在入侵生态学研究方面,生物多样性与生物入侵之间的关系长久以来成为群落可入侵性探讨的焦点。Elton经典假说认为,物种多样性越高对外来种入侵的抵抗能力越强,许多模型或野外试验都支持这一假说。但现在越来越多的试验对此提出了异议,各种假说纷纷出现。究竟生物多样性会不会影响外来种的入侵?假设两种不同的群落结构(功能群),设计6种外来种入侵土著群落的情景分析不同多样性及相同多样性下外来种的入侵状况。结果发现,在多样性相同的情况下,两种群落对外来种入侵的抵抗力不同。外来种成功入侵等比群落,却被倍数群落排斥在系统之外。进一步分析表明这主要是由于可利用资源的波动引起的,即Davis提出的"资源机遇假说"。在相同的物种多样性下,由于倍数群落的特殊结构,整个群落所占有资源远远大于等比群落资源比率。因此,外来种在等比群落中更易找到合适的入侵机会。而在物种多样性不同的情况下,由于物种多样性与已占有资源的变化是成正比的,因此,混淆了多样性与剩余资源可利用性对外来种入侵的影响。     

A phylogenetic approach towards understanding the drivers of plant invasiveness on Robben Island,South Africa

Invasive plant species are a considerable threat to ecosystems globally and on islands in particular where species diversity can be relatively low. In this study, we examined the phylogenetic basis of invasion success on Robben Island in South Africa. The flora of the island was sampled extensively and the phylogeny of the local community was reconstructed using the two core DNA barcode regions, rbcLa and matK. By analysing the phylogenetic patterns of native and invasive floras at two different scales, we found that invasive alien species are more distantly related to native species, a confirmation of Darwin's naturalization hypothesis. However, this pattern also holds even for randomly generated communities, therefore discounting the explanatory power of Darwin's naturalization hypothesis as the unique driver of invasion success on the island. These findings suggest that the drivers of invasion success on the island may be linked to species traits rather than their evolutionary history alone, or to the combination thereof. This result also has implications for the invasion management programmes currently being implemented to rehabilitate the native diversity on Robben Island. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 142–152.     

A phylogenetic analysis of the British flora sheds light on the evolutionary and ecological factors driving plant invasions

Darwin's naturalization hypothesis predicts that invasive species should perform better in their novel range in the absence of close relatives in the native flora due to reduced competition. Evidence from recent taxonomic and phylogenetic‐based studies, however, is equivocal. We test Darwin's naturalization hypothesis at two different spatial scales using a fossil‐dated molecular phylogenetic tree of the British native and alien flora (ca. 1600 species) and extensive, fine‐scale survey data from the 1998 Countryside Survey. At both landscape and local scales, invasive species were neither significantly more nor less related to the native flora than their non‐invasive alien counterparts. Species invasiveness was instead correlated with higher nitrogen and moisture preference, but not other life history traits such as life‐form and height. We argue that invasive species spread in Britain is hence more likely determined by changes in land use and other anthropogenic factors, rather than evolutionary history. Synthesis. The transition from non‐invasive to invasive is not related to phylogenetic distinctiveness to the native community, but instead to their environmental preferences. Therefore, combating biological invasions in the Britain and other industrialized countries need entirely different strategies than in more natural environments.     

Dissecting spatiotemporal patterns of functional diversity through the lens of Darwin's naturalization conundrum

Darwin's naturalization conundrum describes the paradigm that community assembly is regulated by two opposing processes, environmental filtering and competitive interactions, which predict both similarity and distinctiveness of species to be important for establishment. Our goal is to use long‐term, large‐scale, and high‐resolution temporal data to examine diversity patterns over time and assess whether environmental filtering or competition plays a larger role in regulating community assembly processes. We evaluated Darwin's naturalization conundrum and how functional diversity has changed in the Laurentian Great Lakes fish community from 1870 to 2010, which has experienced frequent introductions of non‐native species and extirpations of native species. We analyzed how functional diversity has changed over time by decade from 1870 to 2010 at three spatial scales (regional, lake, and habitat) to account for potential noninteractions between species at the regional and lake level. We also determined which process, environmental filtering or competitive interactions, is more important in regulating community assembly and maintenance by comparing observed patterns to what we would expect in the absence of an ecological mechanism. With the exception of one community, all analyses show that functional diversity and species richness has increased over time and that environmental filtering regulates community assembly at the regional level. When examining functional diversity at the lake and habitat level, the regulating processes become more context dependent. This study is the first to examine diversity patterns and Darwin's conundrum by integrating long‐term, large‐scale, and high‐resolution temporal data at multiple spatial scales. Our results confirm that Darwin's conundrum is highly context dependent.     

The effects of phylogenetic relatedness on invasion success and impact: deconstructing Darwin's naturalisation conundrum

Darwin's naturalisation conundrum describes the paradox that the relatedness of exotic species to native residents could either promote or hinder their success through opposing mechanisms: niche pre‐adaptation or competitive interactions. Previous studies focusing on single snapshots of invasion patterns have provided support to both sides of the conundrum. Here, by examining invasion dynamics of 480 plots over 40 years, we show that exotic species more closely related to native species were more likely to enter, establish and dominate the resident communities, and that native residents more closely related to these successful exotics were more likely to go locally extinct. Therefore, non‐random displacement of natives during invasion could weaken or even reverse the negative effects of exotic–native phylogenetic distances on invasion success. The scenario that exotics more closely related to native residents are more successful, but tend to eliminate their closely related natives, may help to reconcile the 150‐year‐old conundrum.     

Similarities between invaders and native species: Moving past Darwin's naturalization conundrum

Darwin's naturalization conundrum states that successful invaders must be closely related to native species to possess the traits to tolerate that environment, but distantly related enough to possess traits allowing exploitation of underutilized niches, thereby minimizing competition. Although influential, this hypothesis is based on several simplistic assumptions. In particular, the relationship among phylogenetic relatedness, similarity, and competition is more complex than assumed and changes with spatial and phylogenetic scale. Competitive interactions are determined by limiting similarity and trait hierarchies associated with separate traits. Successful invaders thus need to be similar to native species in some respects, but different in others. This combination of similarities and differences is unlikely to be conserved. Further, many invasive species are represented in their novel range by genotypes with extreme trait values or plasticity relative to the species mean. Selection for these genotypes may alter the similarity between invasive and native species, thus obscuring the relationship between competition and phylogenetic relatedness. As environmental filtering and competition often act on different spatial scales, approaches assessing how individual traits relate to invasion at these scales (species pools vs local community) may improve our understanding of the relationship between similarity and invasion.     

It takes one to know one: Similarity to resident alien species increases establishment success of new invaders

Aim

Darwin's naturalization hypothesis states that dissimilarity to native species may benefit alien species establishment due to empty niches and reduced competition. We here add a new dimension to large‐scale tests of community invasibility, investigating the role that previously established alien species play in facilitating or hindering new invasions in plant communities.

Location

Permanent grasslands across France (including mainland and Corsica), as a receding ecosystem of great conservation importance.

Methods

Focusing on 121 alien plant species occurring in 7,215 vegetation plots, we quantified biotic similarity between new invaders and resident alien species (i.e., alien species with longer residence times) based on phylogenetic and trait distances. Additionally, we calculated distances to native species for each alien species and plot. Using multispecies distribution models, we analysed the influence of these biotic similarity measures and additional covariates on establishment success (presence/absence) of new invaders.

Results

We found that biotic similarity to resident alien species consistently increased establishment success of more recently introduced species. Phylogenetic relatedness to previous invaders had an equally strong positive effect as relatedness to native species. Conversely, trait similarity to natives hindered alien establishment as predicted by Darwin's naturalization hypothesis. These results highlight that various mechanisms may act simultaneously to determine alien establishment success.

Main conclusions

Our results suggest that, with greater similarity among alien species, invasion success increases. Such a pattern may arise either due to actual facilitation among invaders or as a result of weaker competitive interactions among invaders than between native and alien species, leading to an indirect facilitative effect. Alternatively, recent environmental changes (e.g., eutrophication, climate change) may have added new environmental filters. Determining how initial invasions might pave the road for subsequent invasions is crucial for effective multispecies management decisions and contributes a new aspect to our understanding of community assembly.

     

Ecological and morphological determinants of evolutionary diversification in Darwin's finches and their relatives

Darwin''s finches are a classic example of adaptive radiation, a process by which multiple ecologically distinct species rapidly evolve from a single ancestor. Such evolutionary diversification is typically explained by adaptation to new ecological opportunities. However, the ecological diversification of Darwin''s finches following their dispersal to Galápagos was not matched on the same archipelago by other lineages of colonizing land birds, which diversified very little in terms of both species number and morphology. To better understand the causes underlying the extraordinary variation in Darwin''s finches, we analyze the evolutionary dynamics of speciation and trait diversification in Thraupidae, including Coerebinae (Darwin''s finches and relatives) and, their closely related clade, Sporophilinae. For all traits, we observe an early pulse of speciation and morphological diversification followed by prolonged periods of slower steady‐state rates of change. The primary exception is the apparent recent increase in diversification rate in Darwin''s finches coupled with highly variable beak morphology, a potential key factor explaining this adaptive radiation. Our observations illustrate how the exploitation of ecological opportunity by contrasting means can produce clades with similarly high diversification rate yet strikingly different degrees of ecological and morphological differentiation.     

Limiting similarity and Darwin’s naturalization hypothesis: understanding the drivers of biotic resistance against invasive plant species

Several hypotheses have been proposed to explain biotic resistance of a recipient plant community based on reduced niche opportunities for invasive alien plant species. The limiting similarity hypothesis predicts that invasive species are less likely to establish in communities of species holding similar functional traits. Likewise, Darwin’s naturalization hypothesis states that invasive species closely related to the native community would be less successful. We tested both using the invasive alien Ambrosia artemisiifolia L. and Solidago gigantea Aiton, and grassland species used for ecological restoration in central Europe. We classified all plant species into groups based on functional traits obtained from trait databases and calculated the phylogenetic distance among them. In a greenhouse experiment, we submitted the two invasive species at two propagule pressures to competition with communities of ten native species from the same functional group. In another experiment, they were submitted to pairwise competition with native species selected from each functional group. At the community level, highest suppression for both invasive species was observed at low propagule pressure and not explained by similarity in functional traits. Moreover, suppression decreased asymptotically with increasing phylogenetic distance to species of the native community. When submitted to pairwise competition, suppression for both invasive species was also better explained by phylogenetic distance. Overall, our results support Darwin’s naturalization hypothesis but not the limiting similarity hypothesis based on the selected traits. Biotic resistance of native communities against invasive species at an early stage of establishment is enhanced by competitive traits and phylogenetic relatedness.