生物多样性 ›› 2020, Vol. 28 ›› Issue (11): 1362-1375. DOI: 10.17520/biods.2020243
于文波1, 黎绍鹏1,2,*(
)
收稿日期:2020-06-15
接受日期:2020-07-24
出版日期:2020-11-20
发布日期:2020-08-16
通讯作者:
黎绍鹏
作者简介:* E-mail: spli@des.ecnu.edu.cn基金资助:
Wenbo Yu1, Shaopeng Li1,2,*()
Received:2020-06-15
Accepted:2020-07-24
Online:2020-11-20
Published:2020-08-16
Contact:
Shaopeng Li
摘要：
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。
于文波, 黎绍鹏 (2020) 基于现代物种共存理论的入侵生态学概念框架. 生物多样性, 28, 1362-1375. DOI: 10.17520/biods.2020243.
Wenbo Yu, Shaopeng Li (2020) Modern coexistence theory as a framework for invasion ecology. Biodiversity Science, 28, 1362-1375. DOI: 10.17520/biods.2020243.
图1 基于现代物种共存理论的入侵生态学概念框架。该框架认为入侵的成功和危害取决于外来种-土著种的生态位差异和适合度差异。其中外来种-土著种的生态位差异促进生物入侵但会降低入侵危害, 而外来种相对于土著种的适合度优势会同时促进生物入侵和加剧危害程度。修改自MacDougall等(2009)、Cadotte等(2018)和Li等(2019)。
Fig. 1 A conceptual diagram using modern coexistence theory as a framework for invasive ecology. According to the framework, the invasion outcome depends on both exotic-native niche and fitness differences. On one hand, increasing exotics-native niche difference promotes the success of the exotics but hinders their impacts on natives. On the other hand, fitness difference, here measured as the fitness advantage of invaders over the natives, could enhance both invasion success and their impacts on native species. Adapted from MacDougall et al (2009), Cadotte et al (2018), and Li et al (2019).
| 假说 Hypothesis | 定义 Definition | 强调的差异 Ecological difference | 在现代物种共存理论框架下的解读 Interpretation under the framework of modern coexistence theory | 参考文献 References | ||
|---|---|---|---|---|---|---|
| 空余生态位假说 Empty niche hypothesis | 外来种通过占领入侵地未被土著种占据的空余生态位, 进而实现成功入侵。Exotics successfully invade when occupying niche space unused by the native communities. | 生态位差异 Niche difference | 外来种和土著种的生态位差异导致空余生态位的存在, 促进入侵。Exotic-native niche differences create empty niches and promote invasion. | |||
| 生物抵抗假说 Biotic resistance hypothesis | 多样性高的群落能够更有效地利用资源, 空余的生态位更少, 因此更难以被入侵。More diverse communities have less available resources and vacant niches, preventing invasion. | 生态位差异 Niche difference | 多样性高的群落土著种和外来种生态位重叠的概率更高, 抑制入侵。Higher probability of exotic-native niche overlap in more diverse communities inhibits invasion. | |||
| 资源机遇假说 Resource opportunity hypothesis | 入侵地具备外来种必需的生物和非生物资源, 且这些资源大多未被土著种有效利用, 促进外来种入侵。The resident communities are more susceptible to invasions when they contain high availability of resources on which a potential invader depends. | 生态位差异 Niche difference | 外来种和土著种的生态位差异导致入侵地中外来种的必需资源未被利用, 促进入侵。 Exotic-native niche differences making the essential resources of exotics available, promoting invasion. | |||
| 环境异质性假说 Environmental heterogeneity hypothesis | 异质性高的生境中具有更多的生态位, 外来种可以占据其中可用的生态位实现成功入侵。 Heterogeneous environment contain a diverse array of niches, and invaders would be successful by filling the available niches. | 生态位差异 Niche difference | 在异质性高的环境中, 外来种和土著种的生态位差异大, 促进入侵。Exotic-native niche differences would be larger in heterogeneous environment, promoting invasion. | |||
| 干扰假说 Disturbance hypothesis | 干扰在群落中形成空余生态位, 降低土著群落的抵抗力, 促进外来种成功定居。Disturbance events increase vacant niches and decrease the resistance of native communities, promoting invasion. | 生态位差异 Niche difference | 干扰导致空余生态位的存在, 促进了和土著种生态位差异大的外来种的入侵。Empty niches created by disturbance events benefit the invasion of exotics with large niche differences to natives. | |||
| 波动资源假说 Fluctuating resources hypothesis | 资源在时间上的波动打破了资源供给和获取的平衡, 导致外来种可利用资源的增加, 提高了群落的可入侵性。Fluctuations in resource availability change resource input and uptake rates, and promote invasion. | 生态位差异 Niche difference | 资源的波动提供了空余生态位, 促进了和土著种生态位差异大的外来种的入侵。 Fluctuating resources offer empty niches, which benefit the invasion of exotics with large niche differences to natives. | |||
| 达尔文归化假说 Darwin’s naturalization hypothesis | 达尔文认为和本土物种亲缘关系远的外来种更容易归化, 因为它们和本土物种差异大, 生态位重叠小, 种间竞争弱。Exotics distantly related to natives are more likely to naturalize, as they occupy distinct niches and complete less stronger with natives. | 生态位差异 Niche difference | 远缘物种生态位差异大, 因此与土著种亲缘关系远的外来种更容易入侵。Distantly related species have large niche differences, which promote the invasion of exotics that distantly related to natives. | |||
| 内禀优势假说 Inherent superiority hypothesis | 成功入侵的外来种在形态、生理、生态、遗传和行为等特征上具备独特的内禀优势。Successfully invaded exotics have intrinsic characteristics superior to native species. | 适合度差异 Fitness difference | 相比土著种, 成功入侵的外来种本身具有更高适合度。Successfully invaded exotics are at a fitness advantage relative to the resident communities. | |||
| 理想杂草特征假说 Ideal weed characteristics hypothesis | 入侵种具备类似理想杂草的生物特性, 这些特征使其很好地适应环境, 并在竞争中占据优势。Invasive species share similar characteristics to ideal weeds, making them well adapt to the environment and exhibit competitive advantages. | 适合度差异 Fitness difference | 相比土著种, 成功入侵的外来种具有的杂草特征使其具有更高的适合度。Exotic species with ideal weed characteristics show higher fitness than the native species in the resident communities. | |||
| 繁殖体压力假说 Propagule pressure hypothesis | 繁殖体引入的数量、频率和质量在入侵初期起决定性作用, 繁殖体压力越大, 成功入侵概率越高。 High quantity, frequency and quality of propagule introductions increase chance of successful invasion. | 适合度差异 Fitness difference | 繁殖体压力越大, 外来种适合度越高, 越容易入侵成功。Exotic species with high propagule pressure show high fitness advantage, and are more likely to invade. | |||
| 竞争能力增强进化假说 Evolution of increased competitive ability | 外来入侵植物到达入侵地后, 会把原来用于防御天敌的物质和能量转移到生长系统, 增强自身竞争能力, 实现成功入侵。After releasing from native specialist predators, exotic species evolve with less investment in anti-herbivore defenses and allocate more resources to vegetative and reproductive growth. This strategy promotes invasion. | 适合度差异 Fitness difference | 竞争能力的增强使外来种具有更高的适合度, 进而在和土著种竞争时占据优势, 成功入侵。The increased competitive ability makes the exotics show higher fitness advantage than the natives, promoting invasion. | |||
| 假说 Hypothesis | 定义 Definition | 强调的差异 Ecological difference | 在现代物种共存理论框架下的解读 Interpretation under the framework of modern coexistence theory | 参考文献 References | ||
| 氮分配进化假说 Hypothesis of the evolution of nitrogen allocation | 相比原产地, 外来种在入侵地降低体内氮元素向防御作用的分配, 增加了氮向光合机构的转移, 提高了自身光合能力。Comparing to native ranges, exotics evolve to decrease nitrogen allocation to defenses and increase nitrogen allocation to photosynthesis in introduced ranges. | 适合度差异 Fitness difference | 氮分配的改变提高了外来种在入侵地的适合度。The evolution of nitrogen allocation increases the fitness of exotics in the introduced habitats. | |||
| 新武器假说 New weapon hypothesis | 入侵植物能够分泌特异的化感物质抑制土著种。土著种对这些化感物质缺乏适应性, 导致外来种在竞争中占据上风。Exotics release allopatric chemicals that inhibit and repress the native competitors in new range, enhancing the exotics’ competitive ability and success. | 适合度差异 Fitness difference | 化感物质提高了外来种的适合度, 降低了土著种的适合度, 促进入侵。Allopatric chemicals increase the fitness of exotics and decrease the fitness of natives, promoting invasion. | |||
| 天敌逃逸假说 Enemy release hypothesis | 当外来种被引入入侵地后, 脱离了原产地天敌的控制, 进而可以快速生长繁殖扩散。After introduction into new areas, exotics escape from their old enemies, benefiting their growth and spread. | 适合度差异 Fitness difference | 脱离天敌控制后, 外来种适合度增加, 提高了相对于土著种的适合度优势, 有利于成功入侵。After escaping from the natural enemies, exotics increase their fitness, promoting invasion. | |||
| 预适应假说 Preadaptation hypothesis | 达尔文认为近缘种偏好相似的生境, 因此土著种占据的生境更适合与其近缘的外来种定居。Close relatives favor similar environments, the habitats occupied by native species were more suitable for the establishment of their closely related exotics. | 适合度差异 Fitness difference | 在被土著种占据的生境中, 与土著种亲缘关系近的外来种具有更高适合度, 因此更容易入侵。In the habitats occupied by native species, the exotics that closely related to these natives would have high fitness and are more likely to invade. | |||
| 新生态位假说 Novel niche hypothesis | 环境变化和人类活动形成了新的生态位, 这些新生态位更适合外来种生存。Anthropogenic changes create novel niches, and these niches are better exploited by exotic species. | 生态位差异和适合度差异 Niche and fitness difference | (1)由于外来种和土著种的生态位差异, 新生态位只能被外来种占据。The novel niches could only be occupied by exotics due to the large exotic-native niche differences. (2)外来种比土著种在新生态位中具有更高的适合度。Exotics exhibit higher fitness than natives in exploiting the new niches. | |||
表1 主要生物入侵假说及其在现代物种共存理论框架下的解读
Table 1 Leading invasion hypotheses and their relationship to modern coexistence theory
| 假说 Hypothesis | 定义 Definition | 强调的差异 Ecological difference | 在现代物种共存理论框架下的解读 Interpretation under the framework of modern coexistence theory | 参考文献 References | ||
|---|---|---|---|---|---|---|
| 空余生态位假说 Empty niche hypothesis | 外来种通过占领入侵地未被土著种占据的空余生态位, 进而实现成功入侵。Exotics successfully invade when occupying niche space unused by the native communities. | 生态位差异 Niche difference | 外来种和土著种的生态位差异导致空余生态位的存在, 促进入侵。Exotic-native niche differences create empty niches and promote invasion. | |||
| 生物抵抗假说 Biotic resistance hypothesis | 多样性高的群落能够更有效地利用资源, 空余的生态位更少, 因此更难以被入侵。More diverse communities have less available resources and vacant niches, preventing invasion. | 生态位差异 Niche difference | 多样性高的群落土著种和外来种生态位重叠的概率更高, 抑制入侵。Higher probability of exotic-native niche overlap in more diverse communities inhibits invasion. | |||
| 资源机遇假说 Resource opportunity hypothesis | 入侵地具备外来种必需的生物和非生物资源, 且这些资源大多未被土著种有效利用, 促进外来种入侵。The resident communities are more susceptible to invasions when they contain high availability of resources on which a potential invader depends. | 生态位差异 Niche difference | 外来种和土著种的生态位差异导致入侵地中外来种的必需资源未被利用, 促进入侵。 Exotic-native niche differences making the essential resources of exotics available, promoting invasion. | |||
| 环境异质性假说 Environmental heterogeneity hypothesis | 异质性高的生境中具有更多的生态位, 外来种可以占据其中可用的生态位实现成功入侵。 Heterogeneous environment contain a diverse array of niches, and invaders would be successful by filling the available niches. | 生态位差异 Niche difference | 在异质性高的环境中, 外来种和土著种的生态位差异大, 促进入侵。Exotic-native niche differences would be larger in heterogeneous environment, promoting invasion. | |||
| 干扰假说 Disturbance hypothesis | 干扰在群落中形成空余生态位, 降低土著群落的抵抗力, 促进外来种成功定居。Disturbance events increase vacant niches and decrease the resistance of native communities, promoting invasion. | 生态位差异 Niche difference | 干扰导致空余生态位的存在, 促进了和土著种生态位差异大的外来种的入侵。Empty niches created by disturbance events benefit the invasion of exotics with large niche differences to natives. | |||
| 波动资源假说 Fluctuating resources hypothesis | 资源在时间上的波动打破了资源供给和获取的平衡, 导致外来种可利用资源的增加, 提高了群落的可入侵性。Fluctuations in resource availability change resource input and uptake rates, and promote invasion. | 生态位差异 Niche difference | 资源的波动提供了空余生态位, 促进了和土著种生态位差异大的外来种的入侵。 Fluctuating resources offer empty niches, which benefit the invasion of exotics with large niche differences to natives. | |||
| 达尔文归化假说 Darwin’s naturalization hypothesis | 达尔文认为和本土物种亲缘关系远的外来种更容易归化, 因为它们和本土物种差异大, 生态位重叠小, 种间竞争弱。Exotics distantly related to natives are more likely to naturalize, as they occupy distinct niches and complete less stronger with natives. | 生态位差异 Niche difference | 远缘物种生态位差异大, 因此与土著种亲缘关系远的外来种更容易入侵。Distantly related species have large niche differences, which promote the invasion of exotics that distantly related to natives. | |||
| 内禀优势假说 Inherent superiority hypothesis | 成功入侵的外来种在形态、生理、生态、遗传和行为等特征上具备独特的内禀优势。Successfully invaded exotics have intrinsic characteristics superior to native species. | 适合度差异 Fitness difference | 相比土著种, 成功入侵的外来种本身具有更高适合度。Successfully invaded exotics are at a fitness advantage relative to the resident communities. | |||
| 理想杂草特征假说 Ideal weed characteristics hypothesis | 入侵种具备类似理想杂草的生物特性, 这些特征使其很好地适应环境, 并在竞争中占据优势。Invasive species share similar characteristics to ideal weeds, making them well adapt to the environment and exhibit competitive advantages. | 适合度差异 Fitness difference | 相比土著种, 成功入侵的外来种具有的杂草特征使其具有更高的适合度。Exotic species with ideal weed characteristics show higher fitness than the native species in the resident communities. | |||
| 繁殖体压力假说 Propagule pressure hypothesis | 繁殖体引入的数量、频率和质量在入侵初期起决定性作用, 繁殖体压力越大, 成功入侵概率越高。 High quantity, frequency and quality of propagule introductions increase chance of successful invasion. | 适合度差异 Fitness difference | 繁殖体压力越大, 外来种适合度越高, 越容易入侵成功。Exotic species with high propagule pressure show high fitness advantage, and are more likely to invade. | |||
| 竞争能力增强进化假说 Evolution of increased competitive ability | 外来入侵植物到达入侵地后, 会把原来用于防御天敌的物质和能量转移到生长系统, 增强自身竞争能力, 实现成功入侵。After releasing from native specialist predators, exotic species evolve with less investment in anti-herbivore defenses and allocate more resources to vegetative and reproductive growth. This strategy promotes invasion. | 适合度差异 Fitness difference | 竞争能力的增强使外来种具有更高的适合度, 进而在和土著种竞争时占据优势, 成功入侵。The increased competitive ability makes the exotics show higher fitness advantage than the natives, promoting invasion. | |||
| 假说 Hypothesis | 定义 Definition | 强调的差异 Ecological difference | 在现代物种共存理论框架下的解读 Interpretation under the framework of modern coexistence theory | 参考文献 References | ||
| 氮分配进化假说 Hypothesis of the evolution of nitrogen allocation | 相比原产地, 外来种在入侵地降低体内氮元素向防御作用的分配, 增加了氮向光合机构的转移, 提高了自身光合能力。Comparing to native ranges, exotics evolve to decrease nitrogen allocation to defenses and increase nitrogen allocation to photosynthesis in introduced ranges. | 适合度差异 Fitness difference | 氮分配的改变提高了外来种在入侵地的适合度。The evolution of nitrogen allocation increases the fitness of exotics in the introduced habitats. | |||
| 新武器假说 New weapon hypothesis | 入侵植物能够分泌特异的化感物质抑制土著种。土著种对这些化感物质缺乏适应性, 导致外来种在竞争中占据上风。Exotics release allopatric chemicals that inhibit and repress the native competitors in new range, enhancing the exotics’ competitive ability and success. | 适合度差异 Fitness difference | 化感物质提高了外来种的适合度, 降低了土著种的适合度, 促进入侵。Allopatric chemicals increase the fitness of exotics and decrease the fitness of natives, promoting invasion. | |||
| 天敌逃逸假说 Enemy release hypothesis | 当外来种被引入入侵地后, 脱离了原产地天敌的控制, 进而可以快速生长繁殖扩散。After introduction into new areas, exotics escape from their old enemies, benefiting their growth and spread. | 适合度差异 Fitness difference | 脱离天敌控制后, 外来种适合度增加, 提高了相对于土著种的适合度优势, 有利于成功入侵。After escaping from the natural enemies, exotics increase their fitness, promoting invasion. | |||
| 预适应假说 Preadaptation hypothesis | 达尔文认为近缘种偏好相似的生境, 因此土著种占据的生境更适合与其近缘的外来种定居。Close relatives favor similar environments, the habitats occupied by native species were more suitable for the establishment of their closely related exotics. | 适合度差异 Fitness difference | 在被土著种占据的生境中, 与土著种亲缘关系近的外来种具有更高适合度, 因此更容易入侵。In the habitats occupied by native species, the exotics that closely related to these natives would have high fitness and are more likely to invade. | |||
| 新生态位假说 Novel niche hypothesis | 环境变化和人类活动形成了新的生态位, 这些新生态位更适合外来种生存。Anthropogenic changes create novel niches, and these niches are better exploited by exotic species. | 生态位差异和适合度差异 Niche and fitness difference | (1)由于外来种和土著种的生态位差异, 新生态位只能被外来种占据。The novel niches could only be occupied by exotics due to the large exotic-native niche differences. (2)外来种比土著种在新生态位中具有更高的适合度。Exotics exhibit higher fitness than natives in exploiting the new niches. | |||
图2 外来种和土著种不同维度的生态学差异对生物入侵的影响。外来种和土著种的谱系距离会影响性状差异, 二者共同影响外来种和土著种的生态位差异和适合度差异, 进而决定生物入侵的成功和危害。
Fig. 2 The effects of the ecological differences between exotics and natives on invasion outcome. Exotic-native phylogenetic distances influence their trait differences, which translate into exotic-native niche and/or fitness differences that combine to determine invasion success and impact.
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