外来植物物种的入侵机制及其对生境特征的响应
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摘要
随着经济、贸易、旅游和其他人类活动的日益加剧，生物入侵已经成为一个全球化问题，不仅导致了生物多样性的丧失，而且带来了巨大的经济和生态危害。尽管生物入侵研究已经有50多年的历史，但至今还没有一个统一的理论和预测框架。
人们对生物入侵的理解和研究主要从两个方面着手，一方面围绕入侵的三个焦点问题，即物种的入侵性，群落的可入侵性和造成的危害。另一方面围绕外来种的入侵过程，即引入、建群、扩散和稳定(或危害)四个阶段。然而这些研究多集中于成功入侵的外来种而没有考虑从生态系统中迅速消失的外来种。而且，只关注了产生巨大可见生态和经济影响的入侵存活者而忽视了没有造成危害的归化种，这一趋势在入侵数据中将造成很大的偏差。因此本研究尝试在研究外来种入侵不确定性的基础上区分两类植物对象，一类是通过根茎(匍匐茎)传播的克隆植物，一类是通过产生轻而小的种子传播的植物，两种植物类型的空间扩散方式不同，通过空间显含的元胞自动机模型一方面可以体现不同的生活史特征和空间表现形式，另一方面可以将入侵生物学的“三个焦点”和“四个阶段”融合在一起，总体把握入侵的整个过程，在“逐类分析，总体把握”思想的指导下分析外来种的入侵机制。最后通过室内和野外调查以及资料分析以江苏沿海外来种互花米草的种群动态为例进行了验证，取得了很好的一致性。
本文建立了具有创新意义的非自治生物入侵干扰模型和生物入侵元胞自动机模型，通过大量的数值模拟实验、理论分析和野外调查的研究，基本揭示了外来植物物种的入侵机制及其对生境特征的响应机制：1)通过对不同群落结构的研究，外来种在不同的迁移扩散能力下可以表现出入侵失败、共存、先成功入侵后失败或成功入侵4种可能的入侵结果；2)克隆植物在6方向上传播最快；3)竞争能力强的外来种不一定能够成功入侵土著生态系统，除了与迁移能力有关外，还与物种本身的生物学特征和生境特征有关。在外来种竞争力弱的情况下。外来种可以通过克隆优势优先占据生境实现成功入侵(优先占据优先发展原则)。同时，一定程度的生境增加和生境破坏会减弱种间竞争对入侵的影响；4)生境破坏影响了外来植物的入侵，而且因不同的植物类型而异。对于克隆植物来说，生境破坏减弱了外来种的入侵性，这是因为克隆植物的迁移能力有限，它只能在临域6个方向上传播。生境破坏严重时，初始多度较小的外来种在传播过程中死亡率增加，因此，强的生境破坏减弱了克隆植物的入侵力。对种子植物而言，生境破坏超过一定的阈值，外来种的入侵性增强，这是因为一定程度的生境破坏减弱了竞争在种群间的作用；5)生境增加有利于边缘效应的产生。竞争力弱的物种在边缘地区可以逃避强物种的干扰，逃避的能力取决于生境增长的速度和物种的传播速度；6)生活史特征对外来种的入侵产生很大的影响。外来种从幼苗到成体阶段对物种入侵的影响最大，成体存活率的的影响稍弱，外来种多度与迁移距离之间呈多项式拟合关系，而与两龄后的繁殖年龄呈对数关系。
With increasing economic, trading, traveling and other human activities, invasions of exotic species has become one of the most important global-scale problems, which not noly alters ecological functioning and threatens biodiversity, but also causes huge economic losses and ecological threats. More than 50 years have past, however, biological invasions are still scarce of uniform theoretical and predictive frame.
Researches on biological invasions are mainly discussed by two angles, one focuses on three key problems on biological invasions, i. e. species invasiveness, invisibility of community and pest (impact); the other emphasizes the invasion process by exotic species, i. e. four stages of introduction, establishment, spread and stabilization (impact/pest), which makes the research on invasion mechanism much easier. However, these studies on species invasions have tended to focus on species that are more likely to be successful colonizers than those that quickly disappear from the invaded communites. Additionally, surviors producing highly visible ecological and economic impacts are more likely to be studied than inconspicuous or innocuous species. Given this tendency, failed or non-visible colonists are underrecorded, and this creates a significant bias in the data on invasive species. So in this paper, invasion mechanism by exotic species was investigated by differtiating exotic clonal plant species (rhizomatic plant and stoloniferous plant) and exotic sexual plant species producing slight and small seeds. The study not only can exhibit various life history traits and spatial patterns of two plant types, but also can involve the "three focus" and "four stages" involved in biological invasions. Also, the model results are tested by population dynamics of exotic species Spartina alterniflora through indoor experiment, field investigations and data analysis, which accords well.
Innovative models, biological invasion disturbance model and cellular automata model, were set up in the paper, which principly disclosed the mechanisms of biological invasions and response to various habitat traits. Six conclusions can be drawn by a great deal of numerical simulations, theoretical analysis and field investigations: 1) Exotic species invasions can result in one of the following four possibilities according to various dispersal abilities: a) rejection failure, b) coexistence, c) indirect failure and d) rejection; 2) The optimal spread of exotic clonal plant is in 6 directions; 3) Superior exotic species can not necessarilly invade native ecosystem, which is associated with biology characteristics and habitat conditions besides dispersal abilities. In virtue of clonal advantage, inferior exotic species can precolonize available habitat (First colonize, first develop rule). And a certain habitat increase and habitat destruction can clear off ability of competition; 4) Effect of habitat destruction on exotic species invasions is determined by dispersal and competitive ability and variated by plant types as well. For exotic clonal plant, habitat destruction lowers invasiveness by exotic plant species and clonal plant spreads in 6 neighboring directions, which limits their survivals, as serious habitat destructions increase species mortality and propagule pressure in the transportation. Severe habitat destruction slacks invasibility by exotic clonal plant species. For seed plant, competition between native species and colonizers can affect colonizer success and estabilishment. When superior exotic species are introduced, slight destruction has no obvious effect on population dynamics, but median habitat destruction can protect native species 1 to some certain extent. When the introduced exotic speces is inferior, serious destruction decreases the abundance of native species 1 and 2 while increases the colonization of exotic species with high dispersal abilities; 5) Habitat increases in favor of the occurance of edge effect. Inferior species (native or exotic) can escape superior ones in the edges, which is decided by the comparison between the speed of habitat increases and species spread; 6) Exotic invasion is sensitive to its main life history traits. Results from cellular automata model showed that growth stages from seedlings to adults made the greatest contributions to population growth of exotic species, while the adult survival probability can not significantly affect it. The variation of dispersal distance and the age of reproduction maturity show that exotic species abundance grew logarithmitically with dispersal distance and decreased logarithmically with reproductive maturity age for those species than two years old.
人们对生物入侵的理解和研究主要从两个方面着手，一方面围绕入侵的三个焦点问题，即物种的入侵性，群落的可入侵性和造成的危害。另一方面围绕外来种的入侵过程，即引入、建群、扩散和稳定(或危害)四个阶段。然而这些研究多集中于成功入侵的外来种而没有考虑从生态系统中迅速消失的外来种。而且，只关注了产生巨大可见生态和经济影响的入侵存活者而忽视了没有造成危害的归化种，这一趋势在入侵数据中将造成很大的偏差。因此本研究尝试在研究外来种入侵不确定性的基础上区分两类植物对象，一类是通过根茎(匍匐茎)传播的克隆植物，一类是通过产生轻而小的种子传播的植物，两种植物类型的空间扩散方式不同，通过空间显含的元胞自动机模型一方面可以体现不同的生活史特征和空间表现形式，另一方面可以将入侵生物学的“三个焦点”和“四个阶段”融合在一起，总体把握入侵的整个过程，在“逐类分析，总体把握”思想的指导下分析外来种的入侵机制。最后通过室内和野外调查以及资料分析以江苏沿海外来种互花米草的种群动态为例进行了验证，取得了很好的一致性。
本文建立了具有创新意义的非自治生物入侵干扰模型和生物入侵元胞自动机模型，通过大量的数值模拟实验、理论分析和野外调查的研究，基本揭示了外来植物物种的入侵机制及其对生境特征的响应机制：1)通过对不同群落结构的研究，外来种在不同的迁移扩散能力下可以表现出入侵失败、共存、先成功入侵后失败或成功入侵4种可能的入侵结果；2)克隆植物在6方向上传播最快；3)竞争能力强的外来种不一定能够成功入侵土著生态系统，除了与迁移能力有关外，还与物种本身的生物学特征和生境特征有关。在外来种竞争力弱的情况下。外来种可以通过克隆优势优先占据生境实现成功入侵(优先占据优先发展原则)。同时，一定程度的生境增加和生境破坏会减弱种间竞争对入侵的影响；4)生境破坏影响了外来植物的入侵，而且因不同的植物类型而异。对于克隆植物来说，生境破坏减弱了外来种的入侵性，这是因为克隆植物的迁移能力有限，它只能在临域6个方向上传播。生境破坏严重时，初始多度较小的外来种在传播过程中死亡率增加，因此，强的生境破坏减弱了克隆植物的入侵力。对种子植物而言，生境破坏超过一定的阈值，外来种的入侵性增强，这是因为一定程度的生境破坏减弱了竞争在种群间的作用；5)生境增加有利于边缘效应的产生。竞争力弱的物种在边缘地区可以逃避强物种的干扰，逃避的能力取决于生境增长的速度和物种的传播速度；6)生活史特征对外来种的入侵产生很大的影响。外来种从幼苗到成体阶段对物种入侵的影响最大，成体存活率的的影响稍弱，外来种多度与迁移距离之间呈多项式拟合关系，而与两龄后的繁殖年龄呈对数关系。
With increasing economic, trading, traveling and other human activities, invasions of exotic species has become one of the most important global-scale problems, which not noly alters ecological functioning and threatens biodiversity, but also causes huge economic losses and ecological threats. More than 50 years have past, however, biological invasions are still scarce of uniform theoretical and predictive frame.
Researches on biological invasions are mainly discussed by two angles, one focuses on three key problems on biological invasions, i. e. species invasiveness, invisibility of community and pest (impact); the other emphasizes the invasion process by exotic species, i. e. four stages of introduction, establishment, spread and stabilization (impact/pest), which makes the research on invasion mechanism much easier. However, these studies on species invasions have tended to focus on species that are more likely to be successful colonizers than those that quickly disappear from the invaded communites. Additionally, surviors producing highly visible ecological and economic impacts are more likely to be studied than inconspicuous or innocuous species. Given this tendency, failed or non-visible colonists are underrecorded, and this creates a significant bias in the data on invasive species. So in this paper, invasion mechanism by exotic species was investigated by differtiating exotic clonal plant species (rhizomatic plant and stoloniferous plant) and exotic sexual plant species producing slight and small seeds. The study not only can exhibit various life history traits and spatial patterns of two plant types, but also can involve the "three focus" and "four stages" involved in biological invasions. Also, the model results are tested by population dynamics of exotic species Spartina alterniflora through indoor experiment, field investigations and data analysis, which accords well.
Innovative models, biological invasion disturbance model and cellular automata model, were set up in the paper, which principly disclosed the mechanisms of biological invasions and response to various habitat traits. Six conclusions can be drawn by a great deal of numerical simulations, theoretical analysis and field investigations: 1) Exotic species invasions can result in one of the following four possibilities according to various dispersal abilities: a) rejection failure, b) coexistence, c) indirect failure and d) rejection; 2) The optimal spread of exotic clonal plant is in 6 directions; 3) Superior exotic species can not necessarilly invade native ecosystem, which is associated with biology characteristics and habitat conditions besides dispersal abilities. In virtue of clonal advantage, inferior exotic species can precolonize available habitat (First colonize, first develop rule). And a certain habitat increase and habitat destruction can clear off ability of competition; 4) Effect of habitat destruction on exotic species invasions is determined by dispersal and competitive ability and variated by plant types as well. For exotic clonal plant, habitat destruction lowers invasiveness by exotic plant species and clonal plant spreads in 6 neighboring directions, which limits their survivals, as serious habitat destructions increase species mortality and propagule pressure in the transportation. Severe habitat destruction slacks invasibility by exotic clonal plant species. For seed plant, competition between native species and colonizers can affect colonizer success and estabilishment. When superior exotic species are introduced, slight destruction has no obvious effect on population dynamics, but median habitat destruction can protect native species 1 to some certain extent. When the introduced exotic speces is inferior, serious destruction decreases the abundance of native species 1 and 2 while increases the colonization of exotic species with high dispersal abilities; 5) Habitat increases in favor of the occurance of edge effect. Inferior species (native or exotic) can escape superior ones in the edges, which is decided by the comparison between the speed of habitat increases and species spread; 6) Exotic invasion is sensitive to its main life history traits. Results from cellular automata model showed that growth stages from seedlings to adults made the greatest contributions to population growth of exotic species, while the adult survival probability can not significantly affect it. The variation of dispersal distance and the age of reproduction maturity show that exotic species abundance grew logarithmitically with dispersal distance and decreased logarithmically with reproductive maturity age for those species than two years old.
引文
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