亚高寒草甸植物群落的中性理论验证

doi:10.3773/j.issn.1005-264x.2008.02.011

摘要/Abstract

摘要：

该文以物种组成较为复杂的青藏高原东部亚高寒草甸为背景,结合最新的群落中性理论,以解释亚高寒草甸草本植物群落的物种分布格局和生物多样性的维持机制。通过对阴坡、阳坡和滩地3个生境进行随机取样调查,用中性模型对所得多样性数据进行拟合,并分别应用置信区间检验、拟合优度检验和多样性指数检验3种方法对拟合效果进行检验。研究结果表明,在拟合优度检验中,3个生境中中性理论预测和实际物种多度分布之间没有显著差异(p>0.05);实际观测值基本全部落入模型预测分布的95%的置信区间之内(仅滩地草本植物群落的63个物种中的1个以及阴坡草本植物群落75个物种中的2个偏离95%的置信区间);对群落多样性的预测也和实际观测没有显著差异,其中丰富度预测拟合得最好(0.49<p<0.56),均匀度拟合得相对较差。比较3种不同的生境,阴坡的3种指数拟合得都非常好,p值均在0.49~0.70之间变化,而滩地拟合相对较差(其中Simpson多样性指数的p值小于0.1)。尽管3种不同的检验方法和生境在对中性理论的检验上存在着差异,但是最终的结论是一致的,即中性模型能够很好地预测3种不同空间生境中的亚高寒草甸群落的多样性分布格局。

关键词: 中性理论, 生物多样性, 亚高寒草甸, 多度

Abstract:

Aims We tested the neutral theory of biodiversity on subalpine meadows of the eastern Tibetan Plateau that exhibited comparatively complicated species composition. Our objective was to explain the species abundance distribution pattern and the underlying mechanism of biodiversity.
Methods We fit the neutral model to a randomly sampled data set obtained in three different habitats (north-facing slope, level field and south-facing slope) and used three methods to test the fitness of the neutral model to the real community: confidence interval, goodness of fit and diversity index.
Important findings We found no significant difference (p>0.05) between the neutral theory predictions and observed species abundance distributions in the three habitats according to the goodness of fit method. The observed data nearly completely fall into 95% confidence intervals of the neutral model predictions (only one out of 63 species in level field communities and 2 out of 75 species in the north-facing slope communities deviate from the 95% confidence interval). There is no significant difference between the neutral theory predictions and observed species abundance patterns, in which the fit of richness predictions is the best (0.49<p<0.56) and the fitness of evenness predictions is relatively poor. However, for the three different habitats, the fitness of these three indices in north-facing slope communities is perfect and the p-values vary between 0.49 and 0.70, but the fitness in level field communities is poorer (p-value of the Simpson diversity index is less than 0.1). Although the test results of the neutral theory by three different test methods and habitats are somewhat different, we conclude that the neutral model can predict species abundance distribution patterns in the three habitats of subalpine meadow.

Key words: neutral theory, biodiversity, subalpine meadow, abundance

杜晓光, 周淑荣. 亚高寒草甸植物群落的中性理论验证. 植物生态学报, 2008, 32(2): 347-354. DOI: 10.3773/j.issn.1005-264x.2008.02.011

DU Xiao-Guang, ZHOU Shu-Rong. TESTING THE NEUTRAL THEORY OF PLANT COMMUNITIES IN SUBALPINE MEADOW. Chinese Journal of Plant Ecology, 2008, 32(2): 347-354. DOI: 10.3773/j.issn.1005-264x.2008.02.011

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链接本文: https://www.plant-ecology.com/CN/10.3773/j.issn.1005-264x.2008.02.011

https://www.plant-ecology.com/CN/Y2008/V32/I2/347

图/表 5

图2 中性模型预测(白色柱图)和实际观测到(黑色柱图)的物种多度分布纵轴是按Preston 方法分组后统计的物种数,即第一个柱形等于Ф1/2,第二个柱形等于Ф1/2+Ф2/2,第三个柱形等于Ф2/2+Ф3+Ф4/2,第四个柱形等于Ф4/2+Ф5+Ф6+Ф7+Ф8/2,以此类推(其中Фi表示多度为i的物种数)

Fig.2 The Preston distribution of expected abundance by the standard neutral model (open bars) and observed abundance (filled bars) Y-axis denote species number by the Preston distribution, i.e.,the first histogram bar represents Ф1/2, the second bar Ф1/2+Ф2/2, the third bar Ф2/2+Ф3+Ф4/2,the fourth bar Ф4/2+Ф5+Ф6+Ф7+Ф8/2,and so on (Фi denote the number of species whose abundance is i)

表1 置信区间检验汇总

Table 1 Summary of confidence envelope analysis

生境类型 Habitat type	置信区间之上的物种数 Over-abundance	置信区间之下的物种数 Under-abundance	个体总数 Total individual	物种数 Species richness	θ	m
阴坡North-facing slope	0	2	14 223	75	31.51	0.003 8
阳坡South-facing slope	0	0	6 696	72	21.36	0.022 5
滩地Plain field	1	0	5 247	63	24.44	0.01

图1 3个生境群落的置信区间检验结果 ——:实际观测多度分布曲线Observed rank-abundance curves....:中性预测分布的95%置信区间的多度分布95%-confidence intervals ---:中性模型预测多度分布Expected rank-abundance curves by the standard neutral model

Fig.1 The tests of the neutral model by confidence interval method in plants communities in three different types of habitats

表2 中性预测多度和实际观测多度的拟合检验

Table 2 The goodness of fit between the neutral theory predictions and observed species abundance patterns

生境类型 Habitat type	df	χ²	p
阴坡 North-facing slope	6	8.661 6	0.193 5
阳坡 South-facing slope	6	3.437 5	0.752 3
滩地 Plain field	4	1.679 4	0.794 5

图3 中性理论预测多样性指数的p值其中中空的方块、圆和三角分别代表滩地、阳坡和阴坡的均匀度指数和多样性指数的p值。黑色的方块、圆和三角分别代表滩地、阳坡和阴坡的均匀度指数和丰富度指数的p值。虚线围成的区域是95%的置信区间的接受域

Fig.3 The p-value of the expected diversity indexes by neutral model The open squares, open circles and open triangles denote p-values of evenness index and richness index in the plain field, south-facing slope and north-facing slope, respectively. The filled squares, filled circles and filled triangles denote p-values of evenness index and diversity index respectively in the plain field, south-facing slope and north-facing slope. The area enclosed by dashed lines indicate the 95%-confidence interval

参考文献 53

[1]	Abrams PA (2001). A world without competition. Nature, 412, 858-859.
[2]	Alonso D, Etienne RS, McKane AJ (2006). The merits of neutral theory. Trends in Ecology & Evolution, 21, 451-457. DOI URL PMID
[3]	Bell G (2000). The distribution of abundance in neutral communities. The American Naturalist, 155, 606-617.
[4]	Bell G (2001). Neutral macroecology. Science, 293, 2413-2418.
[5]	Chapin FS Ⅲ, Zavaleta ES, Eviner VT, Naylor RL, Vitousek PM, Reynolds HL, Hooper DU, Lavorel S, Sala OE, Hobbie SE, Mack C, Díaz S (2000). Consequences of changing biodiversity. Nature, 405, 234-242. DOI URL PMID
[6]	Chase JM (2005). Towards a really unified theory for metacommunities. Functional Ecology, 19, 182-186.
[7]	Chave J (2004). Neutral theory and community ecology. Ecology Letters, 7, 241-253.
[8]	Chesson P (2000). Mechanisms of maintenance of species diversity. Annual Review of Ecology and Systematics, 31, 343-366.
[9]	Costanza R, D'Arge R, De Groot R, Farber S, Grasso M, Hannon B, Limburg K, Naeem S, O'Neill VR, Paruelo J, Raskin RG, Sutton P, Belt VDM (1997). The value of the world's ecosystem services and natural capital. Nature, 387, 253-260.
[10]	Du GZ (杜国祯), Qin GL (覃光莲), Li ZZ (李自珍), Liu ZH (刘正恒), Dong GS (董高生) (2003). Relationship betweenspecies richness and productivity in an Alpine plant community. Acta Phytoecologica Sinica (植物生态学报), 27, 125-132. (in Chinese with English abstract)
[11]	Du GZ (杜国祯), Wang G (王刚) (1995). Succession and changes of grassland quality of the artificial grassland communities in subalpine meadow in Gannan. Acta Botanica Sinica (植物学报), 37, 306-313. (in Chinese with English abstract)
[12]	Efron B, Tibshirani RJ (1993). An Introduction to the Bootstrap. Chapman and Hall, London.
[13]	Etienne RS (2005). A new sampling formula for neutral biodiversity. Ecology Letters, 8, 253-260.
[14]	Etienne RS, Olff H (2004). A novel genealogical approach to neutral biodiversity theory. Ecology Letters, 7, 170-175.
[15]	Etienne RS, Olff H (2005). Confronting different models of community structure to species-abundance data: a Bayesian model comparison. Ecology Letters, 8, 493-504.
[16]	Gravel D, Canham CD, Beaudet M, Messier C (2006). Reconciling niche and neutrality: the continuum hypothesis. Ecology Letters, 9, 399-409.
[17]	Harpole WS, Tilman D (2006). Non-neutral patterns of species abundance in grassland communities. Ecology Letters, 9, 15-23.
[18]	Harte J (2003). Tail of death and resurrection. Nature, 424, 1006-1007.
[19]	He F (2005). Deriving a neutral model of species abundance from fundamental mechanisms of population dynamics. Functional Ecology, 19, 187-193.
[20]	Hubbell SP (1979). Tree dispersion,abundance,and diversity in a tropical dry forest. Science, 203, 1299-1309. URL PMID
[21]	Hubbell SP (2001). The Unified Neutral Theory of Biodiversity and Biogeography. Princeton University Press, Princeton, NJ.
[22]	Hubbell SP (2005a). Neutral theory in community ecology and the hypothesis of functional equivalence. Functional Ecology, 19, 166-172.
[23]	Hubbell SP (2005b). The neutral theory of biodiversity and biogeography and Stephen Jay Gould. Paleobiology, 31, 122-132.
[24]	Hubbell SP (2006). Neutral theory and the evolution of ecological equivalence. Ecology, 87, 1387-1398.
[25]	Leibold MA, McPeek MA (2006). Coexistence of the niche and neutral perspectives in community ecology. Ecology, 87, 1399-1410. DOI URL PMID
[26]	Levins R (1970). Extinction some mathematical questions in biology. In: Gerstenhaber M ed. Lectures on Mathematics in the Life Sciences. Vol. 2, 75-107.
[27]	MacArthur RH, Wilson EO (1963). An equilibrium theory of insular zoogeography. Evolution, 17, 373-387.
[28]	MacArthur RH, Wilson EO (1967). The Theory of Island Biogeography. Princeton University Press, Princeton, NJ.
[29]	Magurran AE, Henderson PA (2003). Explaining the excess of rare species in natural species abundance distributions. Nature, 422, 881-885. URL PMID
[30]	May RM (1997). Ecology and Evolution of Communities. Harvard University Press, Cambridge. 81-120.
[31]	McGill BJ (2003). A test fo the unified neutral theory of biodiversity. Nature, 422, 881-885.
[32]	McGill BJ, Maurer BA, Weiher E (2006). Empirical evaluation of neutral theory. Ecology, 87, 1411-1423.
[33]	McKane AJ, Alonso D, Sole RV (2004). Analytic solution of Hubbell's model of local community dynamics. Theoretical Population Biology, 65, 67-73.
[34]	Nee S (2005). The neutral theory of biodiversity: do the numbers add up? Functional Ecology, 19, 173-176.
[35]	Olszewski TD, Erwin DH (2004). Dynamic response of Permian brachiopod communities to long-term environmental change. Nature, 428, 738-741. DOI URL PMID
[36]	Pacala SW, Tilman D (1993). Limiting similarity in mechanistic and spatial models of plant competition in heterogeneous environments. The American Naturalist, 143, 222-257.
[37]	Poulin R (2004). Parasites and the neutral theory of biodiversity. Ecography, 27, 119-123.
[38]	Purves DW, Pacala SW (2005). Biotic Interactions in the Tropics. Cambridge University Press, Cambridge. 107-138.
[39]	Qiu B (邱波), Ren QJ (任青吉), Luo YJ (罗燕江), Du GZ (杜国祯) (2004). Study on α diversity and β diversity of plant community of different habitats in alpine meadow. Acta Botanica Boreali-Occidentalisa Sinica (西北植物学报), 24, 655-661. (in Chinese with English abstract)
[40]	Silvertown J (2004). Plant coexistence and the niche. Trends in Ecology & Evolution, 19, 605-611.
[41]	Tilman D (2004). Niche tradeoffs, neutrality, and community structure: a stochastic theory of resource competition, invasion, and community assembly. Proceedings of the National Academy of Sciences of the United States of America, 101, 10854-10861.
[42]	Turnbull LA, Manley L, Rees M (2005). Niches, rather than neutrality, structure a grassland pioneer guild. Proceedings of the Royal Society B, 272, 1357-1364.
[43]	Vallade M, Houchmandzadeh B (2003). Analytical solution of a neutral model of biodiversity. Physical Review E, 68, 61902-61905.
[44]	Volkov I, Banavar JR, He F, Hubbell SP, Maritan A (2005). Density dependence explains tree species abundance and diversity in tropical forests. Nature, 438, 658-661.
[45]	Volkov I, Banavar JR, Hubbell SP, Maritan A (2003). Neutral theory and relative species abundance in ecology. Nature, 424, 1035-1037.
[46]	Walker SC, Cyr H (2006). Testing the standard neutral model of biodiversity in lake communities. Oikos, 116, 143-155.
[47]	Whitfield J (2002). Neutrality versus the niche. Nature, 417, 480-481.
[48]	Wootton JT (2005). Field parameterization and experimental test of the neutral theory of biodiversity. Nature, 433, 309-312.
[49]	Yu DW, Terborgh JW, Potts MD (1998). Can high tree species richness be explained by Hubbell's null model? Ecology Letters, 1, 193-199.
[50]	Zhang DY (张大勇) (2000). Researches on Theoretical Ecology (理论生态学研究). Higher Education Press, Beijing. (in Chinese)
[51]	Zhang DY, Lin K (1997). The effects of competitive asymmetry on the rate of competitive displacement: how robust is Hubbell's community drift model? Journal of Theoretical Biology, 188, 361-367.
[52]	Zhou SR, Zhang DY (2006). Allee effects and the neutral theory of biodiversity. Functional Ecology, 20, 509-513.
[53]	Zhou SR (周淑荣), Zhang DY (张大勇) (2006). Neutral theory in community ecology. Journal of Plant Ecology (Chinese Version) (formerly Acta Phytoecologica Sinica) (植物生态学报). 30, 868-877. (in Chinese with English abstract)