面积、温度及分布区限制对物种丰富度海拔格局的影响: 以秦岭太白山为例

doi:10.3724/SP.J.1258.2011.00362

摘要/Abstract

摘要：

物种丰富度的分布格局及其形成机制是生态学研究的热点。以往的研究主要描述丰富度的格局, 而对其形成机制研究较少, 且主要集中于探讨单个因子或过程的影响。物种丰富度同时受到多个因子和过程的综合作用, 面积、温度及物种分布区限制被认为是控制山地物种丰富度海拔格局的主要因素, 三者同时沿海拔梯度而变化, 同时作用于丰富度的海拔格局。幂函数种-面积关系(SAR)、生态学代谢理论(MTE)及中域效应假说(MDE)分别基于以上3个因素, 从机制上解释了物种丰富度的海拔格局。探讨这些假说的相对影响对研究物种丰富度的大尺度格局及其形成机制具有重要意义。方差分离方法有利于分解不同因素的影响, 为此, 该文以秦岭太白山的植物物种丰富度为例, 采用方差分离和逐步回归方法, 分析了SAR、MTE及MDE对物种丰富度海拔格局的影响。结果表明, 太白山的植物物种丰富度沿海拔梯度呈单峰分布格局, 但丰富度峰值存在类群差异; 对太白山所有植物物种丰富度的垂直格局而言, SAR、MTE及MDE分别解释了其物种丰富度随海拔变化的66.4%、19.8%和37.9%, 共同解释了84.6%, 在消除其他因素的影响后, SAR和MTE的独立影响较高(分别为25.5%和17.7%), 而MDE的独立影响不显著; 分类群研究则发现, 苔藓植物丰富度的海拔格局主要受MDE的影响, 蕨类植物丰富度的海拔格局同时受到SAR、MTE以及MDE的影响, 而种子植物物种丰富度的海拔格局主要受SAR和MTE影响。

关键词: 海拔格局, 生态学代谢理论, 中域效应, 种-面积关系, 物种丰富度

Abstract:

Aims Recent studies have illustrated that area, climate and geometric constraints were the main determinants of elevational patterns of species richness; however, the relative importance of these factors is unknown because these factors co-vary with elevation. Based on these factors, the power law species-area relationship (SAR), the metabolic theory of ecology (MTE) and the mid-domain effect hypothesis (MDE) have been proposed to explain elevational patterns of species richness. Our objective is to compare the relative performance of SAR, MTE and MDE in shaping elevational patterns of species richness, using vegetation and climatic data from Mt. Taibai, Qinling Mountains as a case study.
Methods We used a database, “Distribution of Plants in Mt. Taibai”, composed of 2 132 plant species, to derive species richness values for different plant groups. A digital elevation model was used to calculate the area of each elevational band. Eighteen data loggers were set along south and north slopes and at different elevations to record air temperature. RangeModel software was applied to predict species richness of each elevational band, based on the MDE hypothesis. We then compared the explanatory strengths of SAR, MTE and MDE models and used variation partition and stepwise regression to assess their relative importance in shaping elevational patterns of species richness.
Important findings Plant species richness followed a unimodal pattern; however, species richness of each plant group peaked at a different elevational band. As single predictors, SAR, MTE, and MDE explained 66.4%, 19.8% and 37.9% of the variation in species richness, respectively. Together they explained 84.6% of the variation. When the confounding effects of all other factors were eliminated, SAR and MTE explained most (25.5% and 17.7%), whereas MDE played a minor role. Furthermore, we found differences among taxonomic groups: the elevational pattern of lichen richness was primarily explained by MDE, that of ferns was jointly controlled by all three mechanisms and that of seed plants was mainly controlled by SAR and MTE.

Key words: elevational pattern, metabolic theory of ecology, mid-domain effect, species-area relationship, species richness

池秀莲, 唐志尧. 面积、温度及分布区限制对物种丰富度海拔格局的影响: 以秦岭太白山为例. 植物生态学报, 2011, 35(4): 362-370. DOI: 10.3724/SP.J.1258.2011.00362

CHI Xiu-Lian, TANG Zhi-Yao. Effects of area, temperature and geometric constraints on elevational patterns of species richness: a case study in the Mountain Taibai, Qinling Mountains, China. Chinese Journal of Plant Ecology, 2011, 35(4): 362-370. DOI: 10.3724/SP.J.1258.2011.00362

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2011.00362

https://www.plant-ecology.com/CN/Y2011/V35/I4/362

图/表 6

图1 秦岭太白山海拔带面积(实线)及气温(虚线)的垂直变化。

Fig. 1 Variation of area (solid line) and air temperature (dash line) along the elevational gradient in the Mountain Taibai, Qinling Mountains.

图2 3种假说对物种丰富度格局的解释量比较, a、b、c分别表示种面积-关系(SAR)、生态学代谢理论(MTE)及中域效应假说(MDE)的独立贡献; d、e、f分别表示它们两两之间的共同作用; g表示3种假说的共同作用。

Fig. 2 Relative contributions of three hypotheses in explaining species richness patterns; a, b and c represent independent effects of species-area relationship (SAR), metabolic theory of ecology (MTE) and mid-domain effect (MDE); d, e and f represent the joint effects of SAR-MTE, SAR-MDE, and MTE-MDE, and g represents joint effect of the three, respectively.

图3 秦岭太白山不同类群植物物种丰富度的垂直分布格局(实心点表示实际的物种丰富度, 实线为中域效应模型预测的物种丰富度, 虚线间表示95%置信区间)。

Fig. 3 Elevational patterns of species richness for different plant groups in the Mountain Taibai, Qinling Mountains. In each subset figure, dot represents observed richness, solid line represents predicted richness by RangeModel based on mid-domain effect, dashed lines represent the interval of predicted richness with 95% confidential.

表1 不同假说对秦岭太白山植物物种丰富度垂直格局的解释量(R2%)

Table 1 Explanatory strengths of different hypotheses for the elevational patterns of plant richness in the Mountain Taibai, Qinling Mountains (R2%)

类群 Group	种-面积关系 Species-area relationship	生态学代谢理论 Metabolic theory of ecology	中域效应假说 Mid-domain effect hypothesis
所有植物 Overall plants	66.4	19.8	37.9
苔藓植物 Lichens	57.3	2.3	69.5
蕨类植物 Ferns	66.3	13.5	67.7
种子植物 Seed plants	58.3	26.9	32.3

图4 不同假说对秦岭太白山不同类群及不同生活型植物丰富度垂直格局的相对影响, 圆的大小代表对应假说的总解释量。a、b、c分别表示种-面积关系(SAR)、生态学代谢理论(MTE)及中域效应假说(MDE)的独立贡献; d、e、f分别表示SAR与MTE、SAR与MDE及MTE与MDE的共同作用; g表示三者的共同作用; U表示未解释部分。

Fig. 4 Relative importance of different hypotheses in explaining elevational patterns of plant species richness in the Mountain Taibai, Qinling Mountains. In the subsets, the diameters of the circles represent total variance explained by the specific hypothesis, a, b and c represent independent effects species-area relationship (SAR), metabolic theory of ecology (MTE) and mid-domain effect (MDE); d, e and f represent the joint effects of SAR-MTE, SAR-MDE, and MTE-MDE; g represents joint effect of the three; and U represents unexplained, respectively.

表2 秦岭太白山不同类群物种丰富度与不同因子的最优线性模型

Table 2 Optimal linear model for the elevational patterns of species richness and different factors in the Mountain Taibai, Qinling Mountains

丰富度 Richness	变量 Variable	AIC	累积R² Cumulative R²
所有植物 Overall plants	logA	48.9	66.4
	1/kT	24.7	84.5
苔藓植物 Lichens	logPR	118.1	69.5
蕨类植物 Ferns	logPR	119.6	67.7
	1/kT	107.0	78.9
	logA	95.3	85.9
种子植物 Seed plants	logA	54.0	58.3
	1/kT	24.9	83.3

参考文献 51

[1]	Allen AP, Brown JH, Gillooly JF (2002). Global biodiversity, biochemical kinetics, and the energetic- equivalence rule. Science, 297, 1545-1548. DOI URL PMID
[2]	Arrhenius O (1921). Species and area. Journal of Ecology, 9, 95-99. DOI URL
[3]	Bachman S, Baker WJ, Brummitt N, Dransfield J, Moat J (2004). Elevational gradients, area and tropical island diversity: an example from the palms of New Guinea. Ecography, 27, 299-310. DOI URL
[4]	Beck J, Chey VK (2008). Explaining the elevational diversity pattern of geometrid moths from Borneo: a test of five hypotheses. Journal of Biogeography, 35, 1452-1464. DOI URL
[5]	Bhattarai KR, Vetaas OR, Grytnes JA (2004). Fern species richness along a central Himalayan elevational gradient, Nepal. Journal of Biogeography, 31, 389-400. DOI URL
[6]	Brown JH, Gillooly JF, Allen AP, Savage VM, West GB (2004). Toward a metabolic theory of ecology. Ecology, 85, 1771-1789. DOI URL
[7]	Cardelús CL, Colwell RK, Watkins JE (2006). Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. Journal of Ecology, 94, 144-156. DOI URL
[8]	Colwell RK (2006). RangeModel: A Monte Carlo Simulation Tool for Assessing Geometric Constraints on Species Richness. Version 5. User’s Guide and application published at: http://viceroy.eeb.uconn.edu/rangemodel.
[9]	Colwell RK (2008). RangeModel: tools for exploring and assessing geometric constraints on species richness (the mid-domain effect)along transects. Ecography, 31, 4-7. DOI URL
[10]	Colwell RK, Lees DC (2000). The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology and Evolution, 15, 70-76. DOI URL PMID
[11]	Colwell RK, Rahbek C, Gotelli NJ (2004). The mid-domain effect and species richness patterns: What have we learned so far? The American Naturalist, 163, E1-E23. DOI URL PMID
[12]	Connor EF, McCoy ED (1979). The statistics and biology of the species-area relationship. The American Naturalist, 113, 791-833. DOI URL
[13]	Currie DJ, Kerr JT (2008). Tests of the mid-domain hypothesis: a review of the evidence. Ecological Monographs, 78, 3-18. DOI URL
[14]	Currie DJ, Mittelbach GG, Cornell HV, Field R, Guégan JF, Hawkins BA, Kaufman DM, Kerr JT, Oberdorff T, O’Brien E, Turner JRG (2004). Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness. Ecology Letters, 7, 1121-1134. DOI URL
[15]	Feng JM (冯建孟), Wang XP (王襄平), Li J (李晶), Fang JY (方精云) (2006). Effects of area and mid-domain effect on altitudinal pattern of seed plants richness in Lijiang, Yunnan, China. Biodiversity Science (生物多样性), 14, 107-113. (in Chinese with English abstract)
[16]	Gaston KJ (2000). Global patterns in biodiversity. Nature, 405, 220-227. DOI URL PMID
[17]	Grytnes JA, Vetaas OR (2002). Species richness and altitude: a comparison between null models and interpolated plant species richness along the Himalayan altitudinal gradient, Nepal. The American Naturalist, 159, 294-304. DOI URL PMID
[18]	Hawkins BA, Diniz-Filho JAF (2002). The mid-domain effect cannot explain the diversity gradient of Nearctic birds. Global Ecology and Biogeography, 11, 419-426. DOI URL
[19]	Heikkinen RK, Luoto M, Kuussaari M, Pöyry J (2005). New insights into butterfly-environment relationships using partitioning methods. Proceedings of the Royal Society of London (B), 272, 2203-2210.
[20]	Jetz W, Rahbek C (2002). Geographic range size and determinants of avian species diversity. Science, 297, 1548-1551. DOI URL PMID
[21]	Kerr JT, Perring M, Currie DJ (2006). The missing Madagascan mid-domain effect. Ecology Letters, 9, 149-159. DOI URL PMID
[22]	Kessler M, Parris BS, Kessler E (2001). A comparison of the tropical montane pteridophyte floras of Mount Kinabalu, Borneo, and Parque Nacional Carrasco, Bolicia. Journal of Biogeography, 28, 611-622. DOI URL
[23]	Kluge J, Kessler M, Dunn RR (2006). What drives elevational patterns of diversity? A test of geometric constraints, climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica. Global Ecology and Biogeography, 15, 358-371.
[24]	Körner Ch (2007). The use of ‘altitude’ in ecological research. Trends in Ecology and Evolution, 22, 569-574. DOI URL PMID
[25]	McCain CM (2007). Area and mammalian elevational diversity. Ecology, 88, 76-86. DOI URL PMID
[26]	McCain CM (2010). Global analysis of reptile elevational diversity. Global Ecology and Biogeography, 19, 541-553.
[27]	Northwest Institute of Botany, Chinese Academy of Science (中国科学院西北植物研究所) ) (1978). Flora of Qinling (秦岭植物志). Science Press, Beijing. (in Chinese)
[28]	Pautasso M (2007). Scale dependence of the correlation between human population presence and vertebrate and plant species richness. Ecology Letters, 10, 16-24. DOI URL PMID
[29]	Preston FW (1962a). The canonical distribution of commonness and rarity. Part I. Ecology, 43, 185-215.
[30]	Preston FW (1962b). The canonical distribution of commonness and rarity. Part II. Ecology, 43, 410-432.
[31]	R Development Core Team (2009). R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. http://www.r-project.org.
[32]	Rahbek C (1995). The elevational gradient of species richness: A uniform pattern? Ecography, 18, 200-205.
[33]	Ren Y (任毅), Liu MS (刘明时), Tian LH (田联会), Tian XH (田先华) (2006). Research and Management of Biodiversity in the Mt. Taibai Nature Reserve (太白山自然保护区生物多样性研究与管理). China Forestry Publishing House, Beijing. (in Chinese)
[34]	Ricklefs RE (2004). A comprehensive framework for global patterns in biodiversity. Ecology Letters, 7, 1-15.
[35]	Rosenzweig ML (1995). Species Diversity in Space and Time. Cambridge University Press, Cambridge.
[36]	Rowe RJ (2009). Environmental and geometric drivers of small mammal diversity along elevational gradients in Utah. Ecography, 32, 411-422.
[37]	Sanders NJ (2002). Elevational gradients in ant species richness: area, geometry, and Rapoport’s rule. Ecography, 25, 25-32.
[38]	Sanders NJ, Lessard JP, Fitzpatrick MC, Dunn RR (2007). Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. Global Ecology and Biogeography, 16, 640-649.
[39]	Smith TW, Lundholm JT (2010). Variation partitioning as a tool to distinguish between niche and neutral processes. Ecography, 33, 648-655.
[40]	Tang ZY, Fang JY (2006). Temperature variation along the northern and southern slopes of Mt. Taibai, China. Agricultural and Forest Meteorology, 139, 200-207.
[41]	Tang ZY, Wang ZH, Zheng CY, Fang JY (2006). Biodiversity in China’s mountains. Frontiers in Ecology and the Environment, 4, 347-352.
[42]	Tang ZY (唐志尧), Fang JY (方精云), Zhang L (张玲) (2004). Patterns of woody plant species diversity along environmental gradients on Mt. Taibai, Qinling Mountains. Biodiversity Science (生物多样性), 12, 115-122. (in Chinese with English abstract)
[43]	Tang ZY (唐志尧), Fang JY (方精云) (2004). A review on the elevational patterns of plant species diversity. Biodiversity Science (生物多样性), 12, 20-28. (in Chinese with English abstract)
[44]	Tang ZY (唐志尧), Qiao XJ (乔秀娟), Fang JY (方精云) (2009). Species-area relationship in biological communities. Biodiversity Science (生物多样性), 17, 549-559. (in Chinese with English abstract)
[45]	Wang XP (王襄平), Fang JY (方精云), Tang ZY (唐志尧) (2009). The mid-domain effect hypothesis: models, evidence and limitations. Biodiversity Science (生物多样性), 17, 568-578. (in Chinese with English abstract)
[46]	Wang ZH (王志恒), Tang ZY (唐志尧), Fang JY (方精云) (2009). Metabolic theory of ecology: an explanation for species richness patterns based on the metabolic processes of organisms. Biodiversity Science (生物多样性), 17, 625-634. (in Chinese with English abstract)
[47]	Wang ZH, Tang ZY, Fang JY (2007). Altitudinal patterns of seed plant richness in the Gaoligong Mountains, south-east Tibet, China. Diversity and Distributions, 13, 845-854.
[48]	Watkins JE, Cardelús C, Colwell RK, Moran RC (2006). Species richness and distribution of ferns along an elevational gradient in Costa Rica. American Journal of Botany, 93, 73-83.
[49]	Woodward FI, Lomas MR, Kelly CK (2004). Global climate and the distribution of plant biomes. Philosophical Transaction of Royal Society London (B), 359, 1465-1476.
[50]	Worthen WB (1996). Community composition and nested- subset analyses: basic descriptors for community ecology. Oikos, 76, 417-426.
[51]	Zhu Y (朱源), Jiang Y (江源), Liu QR (刘全儒), Xiong M (熊敏), Kang MY (康慕谊) (2007). Altitudinal pattern of vascular plant species richness based on equal-area belts in Mt. Helan. Biodiversity Science (生物多样性), 15, 408-418. (in Chinese with English abstract)