中域效应假说: 模型、证据和局限性

doi:10.3724/SP.J.1003.2009.09140

生物多样性 ›› 2009, Vol. 17 ›› Issue (6): 568-578. DOI: 10.3724/SP.J.1003.2009.09140

所属专题：群落中的物种多样性：格局与机制

中域效应假说: 模型、证据和局限性

王襄平¹^,²^,^*(), 方精云¹, 唐志尧¹

1 北京大学城市与环境学院生态学系, 北京大学地表过程与分析模拟教育部重点实验室, 北京 100871
2 北京林业大学省部共建森林培育与保护教育部重点实验室, 北京 100083

收稿日期:2009-06-03 接受日期:2009-11-12 出版日期:2009-11-20 发布日期:2009-11-20
通讯作者: 王襄平
作者简介:*E-mail: wangxiangping@bjfu.edu.cn
基金资助:
北京林业大学科技创新计划(BLJC200904);国家自然科学基金(40638039)

The mid-domain effect hypothesis: models, evidence and limitations

Xiangping Wang¹^,²^,^*(), Jingyun Fang¹, Zhiyao Tang¹

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
2 The Key Laboratory of Silviculture and Conservation of the Ministry of Education, Beijing Forestry University, Beijing 100083

Received:2009-06-03 Accepted:2009-11-12 Online:2009-11-20 Published:2009-11-20
Contact: Xiangping Wang

摘要/Abstract

摘要：

物种多样性的地理格局是宏观生态学和生物地理学的核心问题之一, 中域效应假说是该领域的一项重要的理论进展。中域效应是指由于边界对物种分布构成限制, 使不同物种分布区在区域中间重叠程度较大, 而在边界附近重叠较少, 从而形成物种丰富度从边界向中心逐渐增加的格局。现有研究表明, 中域效应是影响物种丰富度格局的一种重要机制, 但其作用大小受很多因素的影响。本文介绍该假说的基本假设和模型, 模拟分析不同模型之间的差异, 并就中域效应假说所展开的争论进行综述, 指出该假说的合理性和局限性。我们认为, 中域效应假说的重要性在于揭示了几何(边界)限制和随机过程在物种多样性地理格局中的作用, 但其目前的模型和假设还过于简单, 需要在深入理解物种分布机制的基础上进行完善和发展。

关键词: 物种丰富度格局, 中域效应, 几何限制, 零模型, 纬度、海拔梯度

Abstract

Geographic patterns of species diversity and their underlying mechanisms have long been the focus of macro-ecology and biogeography. Recently, the mid-domain effect (MDE) hypothesis has been proposed to explain geographical diversity patterns. The hypothesis states that if the ranges of the species are randomly distributed within a bounded domain then more ranges will overlap near the middle of the domain than at the edges, and thus decreasing species richness will be observed from the mid-domain to the edges. Many studies have shown that the MDE is an important mechanism affecting geographic richness pattern. However, its relative role in such patterns differs markedly depending on many factors. In this paper, we introduced the assumptions and basic models of the hypothesis and illustrate that the models differ in their predictions as a result of different assumptions. We also review the debate on the MDE hypothesis, and discuss the limitations of present mid-domain models. Although the hypothesis has improved our understanding of the effects of geometric constraints and random process on geographic richness gradients, current MDE models are too simplistic to describe biodiversity patterns in the real world. Improvements to mid-domain models should be based on a better understanding of the mechanisms underlying species distribution.

Key words: species richness pattern, mid-domain effect(MDE), geometric constraints, null model, latitudinal and altitudinal gradients

王襄平, 方精云, 唐志尧 (2009) 中域效应假说: 模型、证据和局限性. 生物多样性, 17, 568-578. DOI: 10.3724/SP.J.1003.2009.09140.

Xiangping Wang, Jingyun Fang, Zhiyao Tang (2009) The mid-domain effect hypothesis: models, evidence and limitations. Biodiversity Science, 17, 568-578. DOI: 10.3724/SP.J.1003.2009.09140.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2009.09140

https://www.biodiversity-science.net/CN/Y2009/V17/I6/568

图/表 5

图1 中域效应假说示意图, 横坐标上0和1.0分别代表区域两端的边界。a: 在“物种分布区不超出边界”的假设限制下, 物种分布宽度和中心点之间的关系(中心点只能分布于三角形内), 图中横线段表示各物种的分布区, 其中点为物种中心点。b: 双随机模型模拟的物种丰富度格局(纵坐标为物种数占区域总物种数的比例), 及最大分布宽度限制对模拟结果的影响, 物种的最大分布宽度分别取区域宽度的0.25、0.5 和1.0 (引自Colwell & Lees, 2000)。

Fig. 1 The mid-domain hypothesis for species richness gradients within a bounded domain, the boundaries are denoted by 0 and 1 on the x axis. a, All midpoint-range coordinate pairs must lie within the isosceles triangle under the assumption that species should be distributed within the boundaries. The ranges for species are shown as horizontal lines centered on their midpoints. b, Patterns of species richness across the domain when maximum range size is set to be 0.25, 0.5 and 1.0 of the domain width in the bivariate model. Note that richness is scaled as the proportion of all species in the simulation (from Colwell & Lees, 2000).

图2 不同中域效应模型产生的物种分布宽度和中心点关系差异。a: 双随机模型; b: 中心点随机模型; c: 分布宽度随机模型。横坐标上0和1.0分别代表区域两端的边界, 因此纵坐标的两端也为0和1.0 (物种分布宽度不能大于区域宽度)。

Fig. 2 Patterns of midpoint-range pairs generated by different mid-domain models. a, the bivariate random model; b, the random midpoint model; c, the random range model. The domain boundaries are denoted by 0 and 1.0 on the x axis, and thus the range of the y axis is also between 0 and 1.0 (the species ranges can not be wider than the domain range because of the geometric constraints).

图3 不同中域效应模型模拟结果的差异。a: 物种中心点的频度分布; b: 物种分布宽度的频度分布; c: 物种丰富度格局; d:平均物种分布宽度(MRS)格局。模拟中使用的参数: 总物种数为50, 区域宽度划分为50段, 图中物种丰富度和MRS为1,000次运算的平均值。

Fig. 3 Comparison of patterns generated by different mid-domain models. a, midpoint frequency distribution; b, range size frequency distribution; c, species richness; d, mean range size (MRS). Each simulation was run with 50 species and 50 bins. Species richness and MRS were the means of 1,000 iterations.

图4 云南丽江地区物种丰富度的垂直格局及不同中域效应模型的模拟结果。物种数据来自吕正伟(1998), 模型的预测值均为500次运算的平均值。

Fig. 4 Altitudinal patterns of species richness in Lijiang region in comparison with patterns simulated by different mid-domain models. The species data were compiled from Lü (1998). The model predictions were the means of 500 iterations.

图5 物种丰富度沿纬度和海拔梯度的变化, 每个梯度各举一个符合和偏离中域效应预测的例子, 图a和c与中域效应预测值一致, 而图b和d则不符。a: 马达加斯加岛10类动物物种丰富度的纬度格局, ■ 所有物种, □ 分布区宽的物种, ○ 分布区窄的物种, 实线为中域效应预测值, 纵坐标为物种丰富度占用于模拟的总物种数的比例(Colwell & Lees, 2000)。b: 非洲Cape植物区的山龙眼科物种丰富度的纬度格局, 实线为物种丰富度, 虚线为中域效应预测值(Laurie & Silander, 2002)。c: 美国Aquarius山区小型哺乳动物的物种丰富度垂直格局(McCain, 2005); d: 婆罗洲Kinabalu山小型哺乳动物的物种丰富度格局(McCain, 2005)。图c、d中给出了中域效应预测值的95%置信区间(无数据点的曲线)。

Fig. 5 Empirical patterns of species richness along latitude or altitude gradients. Two of them were consistent with mid-domain model predictions (a and c), while the other two showed remarkable deviation. a, Latitudinal richness pattern for ten faunal groups in Madagascar. ■ Richness pattern for all species; □ Large ranged species; ○ Small ranged species; solid lines, model predictions (Colwell & Lees, 2000). b, Latitudinal richness pattern for Proteaceae in the Cape Floristic Region of Africa. Solid line, observed pattern; dashed line, model prediction (Laurie & Silander, 2002). c, Altitudinal richness patterns of small mammals in Aquarius Mountains of USA. d, Richness patterns of small mammals in Mt. Kinabalu, Borneo. Lines without data points were the 95% confidence limits of mid-domain model predictions (McCain, 2005).

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中域效应假说: 模型、证据和局限性

The mid-domain effect hypothesis: models, evidence and limitations

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