基于地形因子的天童地区常绿树种和落叶树种共存机制研究

doi:10.3724/SP.J.1003.2012.09242

摘要/Abstract

摘要：

位于亚热带的浙江天童和古田山常绿阔叶林大样地分布有较高比例的落叶树种, 那么它们与常绿树种的共存机制是什么? 常绿树种和落叶树种生态习性差异较大, 二者对生境的选择应有所不同, 我们推测生境分化可能是两类植物实现共存的主要机制。为检验该假设, 我们以天童20 ha动态样地调查数据为依托, 选择个体数≥20的55个常绿树种和42个落叶树种作为分析对象, 用典范对应分析(CCA)研究了地形因子对二者分布的影响差异, 用torus转换检验来分析常绿树种和落叶树种与各类地形生境的关联。结果如下: (1)CCA分析表明地形因子对常绿树种分布的解释量为19.2%, 对落叶树种分布的解释量为7.0%。(2)torus转换检验结果表明, 与沟谷成正关联的常绿树种和落叶树种的比例分别为16.4%和28.6%, 成负关联的比例分别为40%和7%; 与山脊成正关联的常绿树种和落叶树种的比例分别为41.8%和4.8%, 成负关联的比例分别为10.9%和47.6%; 与受干扰生境成正关联的常绿树种和落叶树种的比例分别为16.4%和42.9%。上述结果说明地形对常绿树种分布的影响大于落叶树种; 两个植物类群对生境的选择多呈现相反格局, 尤其是在沟谷生境和山脊生境, 这进一步表明生境分化是常绿树种和落叶树种共存的重要机制之一, 生态位理论在一定程度上能较好地解释亚热带常绿阔叶林物种多样性的维持。

关键词: 地形, 常绿树种, 落叶树种, 常绿阔叶林, 生态位理论, 生境关联, 共存机制

Abstract

The zonal vegetation of subtropical regions in China is recognized as the evergreen broad-leaved forest (EBLF), a forest that usually contains many deciduous tree species. Preliminary investigation of a 20-ha plot at Tiantong (East China) showed that the proportion of deciduous trees reached 52.6%. How do these evergreen and deciduous trees coexist? Up till now, this question lacks a sufficient answer. Evergreen and deciduous tree species often differ in ecological habits, and this niche differentiation may be manifested in distinct habitat preferences. These habitat differences may be an important point in explaining the mechanisms by which these two groups coexist. To test this hypothesis, on the basis of the aforementioned investigation at Tiantong 20 ha plot, 55 evergreen and 42 deciduous tree species with abundance more than 20 individuals were selected for analysis. Canonical Correspondence Analysis (CCA) was used to explore the influence of topographic factors on spatial distribution, and torus-translation tests were employed to test the significance of the species-topography (elevation, convexity, and slope) association. Results were as follows: (1) CCA analysis showed that the three topographic factors (elevation, convexity and slope) explained 19.2% of the distribution of evergreen trees and 7.0% of that for deciduous trees. (2) 16.4% of the 55 evergreen trees and 28.6% of the 42 deciduous trees showed a significantly positive association with the valley habitat, and the proportion of negative association with the valley habitat was 40% and 7% respectively; 41.8% evergreen trees and 4.8% deciduous trees showed a significantly positive association with the ridge habitat, and the proportion of negative association with the ridge habitat was 10.9% and 47.6% respectively; 16.4% evergreen trees and 42.9% deciduous trees were positively associated with the disturbed habitat. Our results showed that topographic factors significantly influence the distribution of evergreen and deciduous trees in the Tiantong plot, and this influence seems greater for evergreen trees. Evergreen and deciduous trees generally showed opposite trends in habitat preferences, especially in valley and ridge habitats. This indicates that habitat partitioning may be a major mechanism allowing the coexistence of evergreen and deciduous trees, and that the niche theory, to some extent, may be sufficient in explaining species diversity maintenance in subtropical evergreen forests.

Key words: topography, evergreen trees, deciduous trees, evergreen broad-leaved forest, niche theory, habitat association, coexistence mechanisms

谢玉彬, 马遵平, 杨庆松, 方晓峰, 张志国, 阎恩荣, 王希华 (2012) 基于地形因子的天童地区常绿树种和落叶树种共存机制研究. 生物多样性, 20, 159-167. DOI: 10.3724/SP.J.1003.2012.09242.

Yubin Xie, Zunping Ma, Qingsong Yang, Xiaofeng Fang, Zhiguo Zhang, Enrong Yan, Xihua Wang (2012) Coexistence mechanisms of evergreen and deciduous trees based on topographic factors in Tiantong region, Zhejiang Province, eastern China. Biodiversity Science, 20, 159-167. DOI: 10.3724/SP.J.1003.2012.09242.

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2012.09242

https://www.biodiversity-science.net/CN/Y2012/V20/I2/159

图/表 10

参考文献 51

[1]	Baker TR, Affum-Baffoe K, Burslem DFRP, Swaine MD (2002) Phenological differences in tree water use and the timing of tropical forest inventories: conclusions from patterns of dry season diameter change. Forest Ecology and Management, 171, 261-274. DOI URL
[2]	Burslem DFRP, Grubb PJ, Turner IM (1995) Responses to nutrient addition among shade-tolerant tree seedlings of lowland tropical rain forest in Singapore. Journal of Ecology, 83, 113-122.
[3]	Chave J (2004) Neutral theory and community ecology. Ecology Letters, 7, 241-253.
[4]	Chiho K, Shimpei O, Takuya K, Kouki H (2010) Light interception in species with different functional groups coexisting in moorland plant communities. Oecologia, 164, 591-599. URL PMID
[5]	Choat B, Ball M, Luly J, Holtum J (2003) Pit membrane porosity and water stress-induced cavitation in four co-existing dry rainforest tree species. Plant Physiology, 131, 41-48. URL PMID
[6]	Clark JS, Silman M, Kern R, Macklin E, HilleRis-Lambers J (1999) Seed dispersal near and far: patterns across temperate and tropical forests. Ecology, 80, 1475-1494.
[7]	Condit R (1998) Tropical Forest Census Plots: Methods and Results from Barro Colorado Island, Panama and a Comparison with Other Plots. Springer Verlag, Berlin.
[8]	Connell JH (1978) Diversity in tropical rain forest and coral reefs. Science, 199, 1302-1309. URL PMID
[9]	Dalling JW, Muller LHC, Wright SJ, Hubbell SP (2002) Role of dispersal in the recruitment limitation of neotropical pioneer species. Journal of Ecology, 90, 714-727.
[10]	De'ath G (2002) Multivariate regression trees: a new technique for modeling species-environment relationships. Ecology, 83, 1105-1117.
[11]	Diaz M, Granadillo E (2005) The significance of episodic rains for reproductive phenology and productivity of trees in semiarid regions of northwestern Venezuela. Trees―Stru- cture and Function, 19, 336-348.
[12]	Enoki T, Kawaguchi H, Iwatsubo G (1997) Nutrient-uptake and nutrient-use efficiency of Pinus thunbergii Parl, along a topographical gradient of soil nutrient availability. Ecological Research, 12, 191-199.
[13]	Fang JY (方精云) (2004) Exploring altitudinal pattern of plant diversity of China’s mountains. Biodiversity Science (生物多样性), 12, 1-4. (in Chinese with English abstract)
[14]	Foster JR (1988) Disturbance history, community organization and vegetation dynamics of the old-growth Pisgah forest, southwestern New Hampshire, U.S.A. Journal of Ecology, 76, 105-134. DOI URL
[15]	Harms KE, Condit R, Hubbell SP, Foster RB (2001) Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology, 89, 947-959. DOI URL
[16]	Hirobe M, Tokuchi N, Iwatsubo G (1998) Spatial variability of soil nitrogen transformation patterns along a forest slope in a Cryptomeria japonica D. Don plantation. European Journal of Soil Biology, 34, 123-131. DOI URL
[17]	Hou JH (侯继华), Ma KP (马克平) (2002) On mechanisms of species coexistence in plant communities. Acta Phytoecologica Sinica (植物生态学报), 26, 1-8. (in Chinese with English abstract)
[18]	Hubbell SP, Foster RB, O’Brien ST, Harms KE, Condit R, Wechsler R, Wright SJ, de Lao SL (1999) Light-gap disturbances, recruitment limitation, and tree diversity in a neotropical forest. Science, 283, 554-557. URL PMID
[19]	Hubbell SP, Foster RB (1983) Diversity of canopy trees in a neotropical forest and implications for conservation. In: Tropical Rain Forest: Ecology and Management (eds Sutton SJ, Whitmore TC, Chadwick AC), pp. 25-41. Blackwell Science, Oxford.
[20]	Hubbell SP, Foster RB (1986a) Biology, chance, and history and the structure of tropical rain forest tree communities. In: Community Ecology (eds Diamond J, Case TJ), pp. 314-329. Harper Row, New York.
[21]	Hunter JC, Parker VT (1993) The disturbance regime of an old-growth forest in coastal California. Journal of Vegetation Science, 4, 19-24.
[22]	Kikuchi T, Miura O (1993) Vegetation patterns in relation to micro-scale landforms in hilly land regions. Vegetatio, 106, 147-154.
[23]	Kilgore BM, Taylor D (1979) Fire history of a Sequoia-mixed conifer forest. Ecology, 60, 129-142.
[24]	Kubota Y (2006) Spatial pattern and regeneration dynamics in a temperate Abies-Tsuga forest in southwestern Japan. Journal of Forest Research, 11, 191-201 DOI URL
[25]	Leigh EG Jr (1999) Tropical Forest Ecology: A View from Barro Colorado Island. Oxford University Press, Oxford.
[26]	Losos EC (1995) Habitat specificity of two palm species: experimental transplantation in Amazonian successional forests. Ecology, 76, 2595-2606.
[27]	Marod D, Kutintara U, Tanaka H, Nakashizuka T (2002) The effects of drought and fire on seed and seedling dynamics in a tropical seasonal forest in Thailand. Plant Ecology, 161, 41-57.
[28]	Nakashizuka T (2001) Species coexistence in temperate, mixed deciduous forests. Trends in Ecology and Evolution, 16, 205-210. DOI URL PMID
[29]	Nee S (2005) The neutral theory of biodiversity: do the numbers add up? Functional Ecology, 19, 173-176. DOI URL
[30]	Ohsawa M (1987) Habitat differentiation and the ecological niche in vegetation. Physiological Ecology (Japan), 24, s15-s27.
[31]	Pollmann W (2002) Effects of natural disturbance and selective logging on Nothofagus forests in south-central Chile. Journal of Biogeography, 29, 955-970.
[32]	Pollmann W (2005) A long-term record of Nothofagus dominance in the southern Andes, Chile. Austral Ecology, 30, 91-102.
[33]	Poorter L (2009) Leaf traits show different relationships with shade tolerance in moist versus dry tropical forests. New Phytologist, 181, 890-900. DOI URL PMID
[34]	Primack RB, Miao SL (1992) Dispersal can limit local plant distribution. Conservation Biology, 6, 513-519.
[35]	Sakai A, Ohsawa M (1993) Vegetation pattern and microtopography on a landslide scar of Mt. Kiyosumi, central Japan. Ecological Research, 8, 47-56. DOI URL
[36]	Sakai A, Ohsawa M (1994) Topographical pattern of the forest vegetation on a river basin in a warm-temperate hilly region, central Japan. Ecological Research, 9, 269-280.
[37]	Sato T, Ito S, Mitsuda Y, Soen N (2010) Impacts of land-use history on the diversity of a riparian forest landscape in warm-temperate Kyushu, southern Japan. Landscape and Ecological Engineering, 6, 89-98.
[38]	Shen ZH, Tang YY, Lu N, Zhao J, Li DX, Wang GF (2007) Community dynamics of seed rain in mixed evergreen broad-leaved and deciduous forests in a subtropical mountain of central China. Journal of Integrative Plant Biology, 49, 1294-1303.
[39]	Song YC (宋永昌) (2001) Vegetation Ecology (植被生态学). East China Normal University Press, Shanghai. (in Chinese)
[40]	Song YC (宋永昌), Wang XR (王祥荣) (1995) Vegetation and Flora of Tiantong National Forest Park, Vegetation and Flora of Tiantong National Forest Park, Zhejiang Province (浙江天童国家森林公园的植被和区系). Shanghai Science and Technology Literature Press, Shanghai. (in Chinese with English summary)
[41]	Tilman D, Pacala S (1993) The maintenance of species richness in plant communities. Species Diversity in Ecological Communities (eds Ricklefs RE, Schluter D), pp. 13-25. University of Chicago Press, Chicago, Illinois.
[42]	Valencia R, Foster R, Villa G, Condit R, Svenning JC, Hernandez C, Romoleroux K, Losos E, Magard E, Balslev H (2004) Tree species distributions and local habitat variation in the Amazon: a large plot in eastern Ecuador. Journal of Ecology, 92, 214-229.
[43]	Wan Y, Cohen J, Guerra R (1997) A permutation test for the robust sib-pair linkage method. Annals of Human Genetics, 61, 79-87. DOI URL PMID
[44]	Wang BC, Smith TB (2002) Closing the seed dispersal loop. Trends in Ecology and Evolution, 17, 379-385.
[45]	Webb CO, Peart DR (2000) Habitat associations of trees and seedlings in a Bornean rain forest. Journal of Ecology, 88, 464-478. DOI URL
[46]	Wright SJ (2002) Plant diversity in tropical forests: a review of mechanisms of species coexistence. Oecologia, 130, 1-14. DOI URL PMID
[47]	Wu ZY (吴征镒) (1980) Vegetation of China (中国植被). Science Press, Beijing. (in Chinese)
[48]	Yamakura T, Kanzake M, Itoh A, Ohkubo T, Ogino K, Chai EOK, Lee HS, Ashton PS (1995) Topography of a large-scale research plot established within a tropical rain forest at Lambir, Sarawak. Tropics, 5, 41-56.
[49]	Yang QS (杨庆松), Ma ZP (马遵平), Xie YB (谢玉彬), Zhang ZG (张志国), Wang ZH (王樟华), Liu HM (刘何铭), Li P (李萍), Zhang N (张娜), Wang DL (王达力), Yang HB (杨海波), Fang XF (方晓峰), Yan ER (阎恩荣), Wang XH (王希华) (2011) Community structure and species composition of an evergreen broad-leaved forest in Tiantong’s 20 ha dynamic plot, Zhejiang Province, eastern China. Biodiversity Science (生物多样性), 19, 215-223. (in Chinese with English abstract)
[50]	Zak DR, Hairston A, Grigal DF (1991) Topographic influences on nitrogen cycling within an upland pin oak ecosystem. Forest Science, 37, 45-53.
[51]	Zhu Y (祝燕), Zhao GF (赵谷风), Zhang LW (张俪文), Shen GC (沈国春), Mi XC (米湘成), Ren HB (任海保), Yu MJ (于明坚), Chen JH (陈建华), Chen SW (陈声文), Fang T (方腾), Ma KP (马克平) (2008) Community composition and structure of Gutianshan forest dynamic plot in a mid-subtropical evergreen broad-leaved forest, east China. Journal of Plant Ecology (植物生态学报), 32, 262-273. (in Chinese with English abstract)

生境类型 Habitat type	海拔 Elevation (m)	凸度 Convexity (m)	总面积 Total area (ha)	总多度 (密度) Total no. of stem (density, no./ha)
受干扰生境 Disturbed habitat	所有 All	所有 All	1.6	4,099 (2,499.4)
低海拔沟谷 Low valley	< 450	[-5.8, -2.0)	3.0	11,634 (3,930.4)
高海拔沟谷 High valley	≥ 450	[-5.8, -2.0)	2.0	8,954 (4,477.0)
低海拔山脊 Low ridge	< 450	(2.0, 6.9]	1.7	8,365 (4,979.2)
高海拔山脊 High ridge	≥ 450	(2.0, 6.9]	1.8	9,862 (5,478.9)
低海拔坡面 Low slope	< 450	[-2.0, 2.0)	6.2	32,555 (5,217.2)
高海拔坡面 High slope	≥ 450	[-2.0, 2.0)	3.7	19,134 (5,199.5)

生境类型 Habitat type	海拔 Elevation (m)	凸度 Convexity (m)	总面积 Total area (ha)	总多度 (密度) Total no. of stem (density, no./ha)
受干扰生境 Disturbed habitat	所有 All	所有 All	1.6	4,099 (2,499.4)
低海拔沟谷 Low valley	< 450	[-5.8, -2.0)	3.0	11,634 (3,930.4)
高海拔沟谷 High valley	≥ 450	[-5.8, -2.0)	2.0	8,954 (4,477.0)
低海拔山脊 Low ridge	< 450	(2.0, 6.9]	1.7	8,365 (4,979.2)
高海拔山脊 High ridge	≥ 450	(2.0, 6.9]	1.8	9,862 (5,478.9)
低海拔坡面 Low slope	< 450	[-2.0, 2.0)	6.2	32,555 (5,217.2)
高海拔坡面 High slope	≥ 450	[-2.0, 2.0)	3.7	19,134 (5,199.5)

植物类群 Plant group	地形因子 Topographic factor	CCA1	CCA2	R²	P_r(>r)	显著性水平 Significance level
常绿树种 Evergreen trees	海拔 Elevation	0.97	0.25	0.72	0.001	***
	坡度 Slope	0.47	-0.88	0.17	0.001	***
	凸度 Convexity	0.94	-0.34	0.40	0.001	***
落叶树种 Deciduous trees	海拔 Elevation	0.91	-0.41	0.46	0.001	***
	坡度 Slope	0.35	0.94	0.22	0.001	***
	凸度 Convexity	0.99	0.13	0.39	0.001	***

植物类群 Plant group	地形因子 Topographic factor	CCA1	CCA2	R²	P_r(>r)	显著性水平 Significance level
常绿树种 Evergreen trees	海拔 Elevation	0.97	0.25	0.72	0.001	***
	坡度 Slope	0.47	-0.88	0.17	0.001	***
	凸度 Convexity	0.94	-0.34	0.40	0.001	***
落叶树种 Deciduous trees	海拔 Elevation	0.91	-0.41	0.46	0.001	***
	坡度 Slope	0.35	0.94	0.22	0.001	***
	凸度 Convexity	0.99	0.13	0.39	0.001	***

植物类群 Plant group	限制性轴特征值 Eigenvalues for constrained axes			非限制性轴特征值 Eigenvalues for unconstrained axes
植物类群 Plant group	CCA1	CCA2	CCA3	CA1	CA2	CA3	CA4	CA5	CA6	CA7	CA8
常绿树种 Evergreen trees	0.27	0.09	0.02	0.18	0.16	0.12	0.10	0.09	0.09	0.08	0.07
落叶树种 Deciduous trees	0.24	0.13	0.04	0.42	0.34	0.33	0.26	0.23	0.19	0.19	0.17
	能被生境因子解释的总变异 Variation explained by topographic factors			未能被生境因子解释的总变异 Variation unexplained by topographic factors
常绿树种 Evergreen trees	0.39 (19.2%)			1.64 (80.8%)
落叶树种 Deciduous trees	0.41 (7.0%)			5.22 (93.0%)