群落构建的中性理论和生态位理论

doi:10.3724/SP.J.1003.2009.09142

生物多样性 ›› 2009, Vol. 17 ›› Issue (6): 579-593. DOI: 10.3724/SP.J.1003.2009.09142

所属专题：群落中的物种多样性：格局与机制

群落构建的中性理论和生态位理论

牛克昌¹^,^*(), 刘怿宁¹, 沈泽昊¹, 何芳良², 方精云¹

1 北京大学城市与环境学院生态学系, 北京大学地表过程分析与模拟教育部重点实验室, 北京 100871
2 Department of Renewable Resources, University of Alberta, Edmonton, Alberta, Canada T6G 2H1

收稿日期:2009-06-03 接受日期:2009-12-06 出版日期:2009-11-20 发布日期:2009-11-20
通讯作者: 牛克昌
作者简介:*E-mail: kechangniu@pku.edu.cn
基金资助:
国家自然科学基金(40638039);国家科技支撑计划(2007BAC06B01)

Community assembly: the relative importance of neutral theory and niche theory

Kechang Niu¹^,^*(), Yining Liu¹, Zehao Shen¹, Fangliang He², Jingyun Fang¹

1 Department of Ecology, College of Urban and Environmental Sciences and Key Laboratory for Earth Surface Processes of the Ministry of Education, Peking University, Beijing 100871, China
2 Department of Renewable Resources, University of Alberta, Edmonton, Alberta, Canada T6G 2H1

Received:2009-06-03 Accepted:2009-12-06 Online:2009-11-20 Published:2009-11-20
Contact: Kechang Niu

摘要/Abstract

摘要：

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。

关键词: 物种特征, 生态等价, 随机, 物种相对多度, 物种多度分布

Abstract

Understanding species coexistence and the maintenance of biodiversity has long been the central interest of ecologists. The niche-based theory of community assembly has dominated community ecology for nearly a century, yet understanding of the mechanisms of species coexistence has remained elusive. The newly developed neutral theory of biodiversity has offered a promising alternative to the niche paradigm. The analytical elegance and simplicity of the neutral theory and its predictive power have made the theory widely popular. However, it is the very same simplicity of the theory (e.g. the symmetric assumption) that makes the theory vulnerable to stark criticisms. Widespread empirical evidence has shown that species in communities are not functionally symmetric; ecological equivalence is more a conceptual simplicity than a biological realism. Recognizing that niche and neutral processes do not have to diametrically oppose each other and a community is likely determined by the interplay of the two processes, ecologists currently are searching to reconcile the two theories by either incorporating drift into niche theory or niche into the neutral framework. However, this reconciliation process is still at its very early stage, we expect this direction will lead to a more complete understanding of community assembly mechanisms. In this paper, we provide a review on the brief histories of the niche and neutral theories, with the focus on comparing the distinct importance of the two theories in explaining community assembly. We discuss in details several integrated models that attempt to unify the niche and neutral theories. We argue that it is an essential step for any successful theory to withstand substantial experimental and field tests. The experimental tests of neutral theories are an important direction that has currently not received due attention.

Key words: species traits, ecological equivalence, stochasticity, species relative abundance, species abundance distribution

牛克昌, 刘怿宁, 沈泽昊, 何芳良, 方精云 (2009) 群落构建的中性理论和生态位理论. 生物多样性, 17, 579-593. DOI: 10.3724/SP.J.1003.2009.09142.

Kechang Niu, Yining Liu, Zehao Shen, Fangliang He, Jingyun Fang (2009) Community assembly: the relative importance of neutral theory and niche theory. Biodiversity Science, 17, 579-593. DOI: 10.3724/SP.J.1003.2009.09142.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2009.09142

https://www.biodiversity-science.net/CN/Y2009/V17/I6/579

图/表 4

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验证项	待检验假设	检验区域	主要结论	相关文献
物种等价性	检验群落物种的出生率、死亡率和拓殖能力是否相同	热带雨林乔木	大部分物种的出生率、死亡率和拓殖能力不符合中性假设	Chave, 2004; Hubbell, 2006
	自然群落中, 物种的功能属性与群落结构是否有关	高寒草地、温带草原、热带雨林	群落组分种对环境响应各异, 物种功能性状与群落结构变化有关, 不符合中性假设	Fargione et al., 2003; Adler, 2004; Harpole & Tilman, 2006; Chu et al., 2007; Kraft et al., 2008; Niu et al., 2008
	物种是否可相互替代以及物种间相互作用是否等价性	热带雨林乔木	物种不可替代, 种间作用不等价, 如同属内物种相互作用强于异属物种。	Kelly et al., 2008
	物种入侵和拓殖成功率是否与入侵物种的属性以及群落结构有关	各类入侵物种和被入侵植物群落	大多研究表明物种的成功拓殖与物种本身属性和入侵群落属性有密切关系; 也有研究表明入侵与物种和群落属性无关。	Fargione et al., 2003; Cadotte et al., 2006; MacDougall et al., 2009
零和多项模型	自然群落物种分布模式是否与中性理论预测一致。	热带雨林、温带针叶林、亚高寒草地、鱼类、腕足类生物化石、鸟类。	大多数的群落中, 物种多度分布符合中性理论预测。	Condit et al., 2002; Volkov et al., 2003; Alonso & McKane, 2004; Etienne & Olff, 2004a; Olszewski & Erwin, 2004; He, 2005; Houlahan et al., 2007; Volkov et al., 2007; Muneepeerakul et al., 2008; 杜晓光和周淑荣, 2008
零和多项模型	比较中性模型和其他模型种对物种多度分布的拟合。	重新检验之前经验检验数据	中性模型的拟合符合程度并不比其他模型拟合的好。	McGill, 2003; McGill et al., 2006; Volkov et al., 2007
扩散限制	扩散对现实群落构建有作用	热带雨林、亚热带常绿阔叶林乔木、温带落叶阔叶林	不同的群落类型、不同研究尺度, 生态位构建和扩散限制相对作用贡献不同, 如图4所示.	Condit et al., 2002; Tuomisto, 2003; Gilbert et al., 2004; Karst et al., 2005; Jones et al., 2006; Cielo et al., 2007; Chave, 2008; Jones et al., 2008; Legendre et al., 2009
扩散限制	岛屿中的物种数是否与物种属性有关。	岛屿生物的重新拓殖, 植物和蝴蝶种群等。	岛屿植物拓殖能力与其叶片坚韧程度相关; 蝴蝶多度与其寄主植物生物量相关	Yamamoto et al., 2007; Burns & Neufeld, 2009

验证项	待检验假设	检验区域	主要结论	相关文献
物种等价性	检验群落物种的出生率、死亡率和拓殖能力是否相同	热带雨林乔木	大部分物种的出生率、死亡率和拓殖能力不符合中性假设	Chave, 2004; Hubbell, 2006
	自然群落中, 物种的功能属性与群落结构是否有关	高寒草地、温带草原、热带雨林	群落组分种对环境响应各异, 物种功能性状与群落结构变化有关, 不符合中性假设	Fargione et al., 2003; Adler, 2004; Harpole & Tilman, 2006; Chu et al., 2007; Kraft et al., 2008; Niu et al., 2008
	物种是否可相互替代以及物种间相互作用是否等价性	热带雨林乔木	物种不可替代, 种间作用不等价, 如同属内物种相互作用强于异属物种。	Kelly et al., 2008
	物种入侵和拓殖成功率是否与入侵物种的属性以及群落结构有关	各类入侵物种和被入侵植物群落	大多研究表明物种的成功拓殖与物种本身属性和入侵群落属性有密切关系; 也有研究表明入侵与物种和群落属性无关。	Fargione et al., 2003; Cadotte et al., 2006; MacDougall et al., 2009
零和多项模型	自然群落物种分布模式是否与中性理论预测一致。	热带雨林、温带针叶林、亚高寒草地、鱼类、腕足类生物化石、鸟类。	大多数的群落中, 物种多度分布符合中性理论预测。	Condit et al., 2002; Volkov et al., 2003; Alonso & McKane, 2004; Etienne & Olff, 2004a; Olszewski & Erwin, 2004; He, 2005; Houlahan et al., 2007; Volkov et al., 2007; Muneepeerakul et al., 2008; 杜晓光和周淑荣, 2008
零和多项模型	比较中性模型和其他模型种对物种多度分布的拟合。	重新检验之前经验检验数据	中性模型的拟合符合程度并不比其他模型拟合的好。	McGill, 2003; McGill et al., 2006; Volkov et al., 2007
扩散限制	扩散对现实群落构建有作用	热带雨林、亚热带常绿阔叶林乔木、温带落叶阔叶林	不同的群落类型、不同研究尺度, 生态位构建和扩散限制相对作用贡献不同, 如图4所示.	Condit et al., 2002; Tuomisto, 2003; Gilbert et al., 2004; Karst et al., 2005; Jones et al., 2006; Cielo et al., 2007; Chave, 2008; Jones et al., 2008; Legendre et al., 2009
扩散限制	岛屿中的物种数是否与物种属性有关。	岛屿生物的重新拓殖, 植物和蝴蝶种群等。	岛屿植物拓殖能力与其叶片坚韧程度相关; 蝴蝶多度与其寄主植物生物量相关	Yamamoto et al., 2007; Burns & Neufeld, 2009

群落构建的中性理论和生态位理论

Community assembly: the relative importance of neutral theory and niche theory

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