四明山常绿阔叶次生林不同海拔群落生态位与物种联结特征

Ecological niches and species association characteristics of different altitude communities in the evergreen broad-leaved secondary forest of Siming Mountain

摘要

【目的】四明山作为浙东大运河山水林田湖草生态保护修复工程的核心区域，在维护局域生物多样性、涵养水源、调节气候与保持土壤等方面意义重大。然而，当前针对当地森林群落构建机制的研究相对不足。本研究旨在揭示四明山高低海拔梯度间森林群落组成、生态位特征及种间关系的分异规律，进而阐明森林群落的构建机制。【方法】在四明山低和高海拔梯度上共设置了20个森林样地，调查群落学特征后，通过物种生态位宽度和重叠度，以及种间联结性分析，阐明海拔驱动的群落构建机制。【结果】森林乔灌层优势种的重要值在高低海拔表现出差异，佐证了“海拔生态位分化假说”。低海拔乔木层以青冈为主、灌木层以连蕊茶为主导。高海拔乔木层以小叶白辛树占优，灌木层以尖连蕊茶和马银花为核心。豹皮樟和短柄枹栎是跨海拔乔木物种，且它们的重要值随海拔上升而增加。优势种的生态位宽度随海拔升高不断扩大。物种间生态位分化在高海拔比在低海拔更为明显。森林群落内物种间整体呈正联结(VR>1)，且在高海拔正关联更强(VR= 1.08 vs 1.49)。乔木层普遍维持高Ochiai指数(OI>0.5)与低负关联率，而灌木层呈现低Ochiai指数(OI<0.5)与高负关联率。乔木层物种通过生态位分化避免竞争，而灌木层呈现明显竞争排斥。【结论】海拔通过调节关键物种的生态位分化和种间互作强度驱动森林群落构建。高海拔以生态位分化与强物种正关联协同维持群落稳定性，而低海拔依赖高重叠物种的竞争互作维系群落结构。

Abstract

[Objective] As the core area of the ecological restoration project of the Grand Canal in eastern Zhejiang, Siming Mountain holds significant importance for maintaining local biodiversity, conserving water resources, regulating climate, and preventing soil erosion. Nevertheless, studies specifically targeting the community assembly mechanisms in this area are still relatively insufficient. This paper aimed to uncover the differentiation patterns of forest community composition, niche and inter-specific relationships at different altitudes of Siming Mountain, thus clarifying the construction mechanism of forest communities. [Method] A total of 20 forest sample plots were set up at different altitudes (low and high) of Siming Mountain. After surveying the community characteristics, the altitude-driven community construction mechanisms were expounded through the analysis of species niche breadth, niche overlap, and inter-specific associations. [Result] The importance values of dominant species across tree and shrub layers showed differences between high and low altitudes, supporting the "altitude niche differentiation hypothesis". In low-altitude zones, Cyclobalanopsis glauca dominated the tree layer, while Camellia fraterna prevailed in the shrub layer. At high altitudes, Pterostyrax corymbosus became the tree-layer dominant, while C. fraterna and Rhododendron ovatum comprised the main body of shrub-layer. Litsea coreana and Quercus glandulifera were arboreal species spanning across altitudes, and their importance values increased with altitude. Niche breadth of dominant species expanded consistently with altitude, and niche differentiation among species was more pronounced at high altitudes than at low elevations. The forest community exhibited overall positive interspecific associations (VR > 1), with association strength strengthening at high altitudes (VR = 1.08 vs. 1.49). The tree layer maintained high Ochiai indices (OI > 0.5) and low negative association rates, suggesting competition avoidance via niche differentiation, whereas the shrub layer showed low OI (OI < 0.5) and high negative association rates, indicating obvious competitive exclusion. [Conclusion] Altitude affects forest community assembly by regulating niche differentiation of key species and the intensity of interspecific interactions. At high altitudes, community stability is maintained by the synergy between niche differentiation and strong positive interspecific associations, whereas low-altitude communities rely on competitive interactions among species with high niche overlap to sustain structural organization.

关键词

群落构建 / 生态位 / 生态位重叠 / 物种联结 / 海拔生态位分化假说

Key words

community assembly / ecological niche / ecological niche overlap / species association / altitude niche differentiation hypothesis

参考文献

[1] 陈磊, 米湘成, 马克平. 生态位分化与森林群落物种多样性维持研究展望[J]. 生命科学, 2014, 26(2): 112-117.

[2] 雷德芳, 王志杰. 黔中喀斯特地区马尾松群落主要物种种间联结、生态位及群落稳定性特征[J]. 生态学报, 2024, 44(10): 4353-4367.

[3] 李德志, 石强, 臧润国, 等. 物种或种群生态位宽度与生态位重叠的计测模型[J]. 林业科学, 2006, 42(7): 95-103.

[4] BOND W J, MIDGLEY J J. Ecology of sprouting in woody plants: the persistence niche[J]. Trends Ecol Evol, 2001, 16(1): 45-51

[5] 郝建锋, 李艳, 齐锦秋, 等. 人为干扰对碧峰峡栲树次生林群落物种多样性及其优势种群生态位的影响[J]. 生态学报, 2016, 36(23): 7678-7688.

[6] PAZOS F, JUAN D, IZARZUGAZA J M G, et al. Prediction of protein interaction based on similarity of phylogenetic trees[J]. Methods Mol Biol, 2008, 484: 523-535.

[7] YODER J B, NUISMER S L. When does coevolution promote diversification?[J]. Am Nat, 2010, 176(6): 802-817.

[8] 文安邦, 汤青, 欧阳朝军, 等. 中国山地保护与山区发展: 回顾与展望[J]. 中国科学院院刊, 2023, 38(3): 376-384.

[9] XIAO Y L, WANG Q M, ZHANG H K. Global natural and planted forests mapping at fine spatial resolution of 30 m[J]. J Remote Sens, 2024, 4: 204.

[10] 冯云, 马克明, 张育新, 等. 北京东灵山辽东栎(Quercus liaotungensis)林沿海拔梯度的物种多度分布[J]. 生态学报, 2007, 27(11): 4743-4750.

[11] 张悦, 郭利平, 易雪梅, 等. 长白山北坡3个森林群落主要树种种间联结性[J]. 生态学报, 2015, 35(1): 106-115.

[12] 郭昊, 赵厚本, 高亿波, 等. 南岭中段森林优势树种及其生态位随海拔梯度变化特征[J]. 植物科学学报, 2024, 42(5): 582-594.

[13] 倪穗, 胡建军, 陈开超, 等. 四明山国家森林公园野生植物资源的开发利用与保护对策[J]. 宁波大学学报(理工版), 2006, 19(1): 85-89.

[14] 苗国丽, 王卫兵, 陈开超, 等. 四明山国家森林公园景观质量评价[J]. 浙江林业科技, 2012, 32(2): 76-80.

[15] 王翰华, 罗淑青, 倪梦茜, 等. 四明山国家森林公园药用植物资源调查与分析[J]. 浙江林业科技, 2018, 38(1): 91-95.

[16] 金则新. 浙江天台山常绿阔叶林次生演替序列群落物种多样性[J]. 浙江林学院学报, 2002, 19(2): 133-137.

[17] 陈水飞, 李垚, 葛晓敏, 等. 浙江仙居典型中亚热带常绿阔叶林样地的群落特征[J]. 生态与农村环境学报, 2017, 33(12): 1109-1116.

[18] 张崇邦, 金则新, 柯世省. 天台山不同林型土壤酶活性与土壤微生物、呼吸速率以及土壤理化特性关系研究[J]. 植物营养与肥料学报, 2004, 10(1): 51-56.

[19] 陈海生. 天台山植物功能群光合生理特性研究[J]. 安徽农学通报, 2012, 18(9): 41-42.

[20] 刘鹏, 康华靖, 陈子林, 等. 浙江六十田自然保护小区常绿阔叶林群落特征及优势乔木种的动态[J]. 浙江师范大学学报(自然科学版), 2007, 30(2): 128-134.

[21] 哀建国, 丁炳扬, 吴谷汉. 浙江植被生态学研究现状与展望[J]. 浙江林业科技, 2004, 24(6): 46-53.

[22] 蒋有绪, 王丽丽, 王兵. 中国森林植物群落信息管理系统的建立[J]. 林业科学研究, 1994, 7(5): 569-573.

[23] 孙小伟, 杨庆松, 刘何铭, 等. 基于浙江天童20 hm²常绿阔叶林动态监测样地的群丛划分[J]. 植物生态学报, 2018, 42(5): 550-561.

[24] 张光富. 浙江天童山区灌丛群落的物种多样性及其与演替的关系[J]. 生物多样性, 2000, 8(3): 271-276.

[25] 王育松, 上官铁梁. 关于重要值计算方法的若干问题[J]. 山西大学学报(自然科学版), 2010, 33(2): 312-316.

[26] 李林, 魏识广, 叶万辉, 等. 南亚热带常绿阔叶林优势种组的空间分布格局及组间关联[J]. 林业科学, 2022, 58(9): 16-24.

[27] LEVINS R. Evolution in Changing Environments[M]. Princeton, N.J.: Princeton University Press, 1968.

[28] PIANKA E R. The structure of lizard communities[J]. Annu Rev Ecol Syst, 1973, 4 ( 1973): 53-74.

[29] 王伯荪, 彭少麟. 鼎湖山森林群落分析: Ⅱ.物种联结性[J]. 中山大学学报(自然科学版), 1983, 22(4): 29-37.

[30] SCHLUTER D. A variance test for detecting species associations, with some example applications[J]. Ecology, 1984, 65(3): 998-1005.

[31] HUBÁLEK Z. Coefficients of association and similarity, based on binary (presence-absence) data: an evaluation[J]. Biol Rev, 1982, 57(4): 669-689.

[32] 高亮, 周建广, 丁晖, 等. 武夷山国家公园黄山松林沿海拔梯度生态位及种间联结性[J]. 生态学报, 2024, 44(21): 9836-9847.

[33] 张忠华, 胡刚, 刘立斌, 等. 黔中北亚热带喀斯特次生林动态监测样地: 物种组成与群落结构[J]. 生态学报, 2022, 42(2): 742-754.

[34] 王宇超, 周亚福, 王得祥. 秦岭南坡中段主要森林群落类型划分及环境梯度解释[J]. 生态环境学报, 2016, 25(6): 965-972.

[35] 唐志尧, 柯金虎. 秦岭牛背梁植物物种多样性垂直分布格局[J]. 生物多样性, 2004, 12(1): 108-114.

[36] 陈海生, 丁坚钢. 海拔高度对浙江省天台山森林生态系统植物功能群组成的影响[J]. 浙江林业科技, 2013, 33(5): 91-93.

[37] 程瑞梅, 王瑞丽, 刘泽彬, 等. 三峡库区栲属群落主要乔木种群的种间联结性[J]. 林业科学, 2013, 49(5): 36-42.

[38] 黄石德, 聂森, 肖祥希, 等. 武夷山米槠群落优势种群生态位与种间联结[J]. 植物科学学报, 2023, 41(3): 291-300.

[39] MAESTRE F T, CALLAWAY R M, VALLADARES F, et al. Refining the stress-gradient hypothesis for competition and facilitation in plant communities[J]. J Ecol, 2009, 97(2): 199-205.

[40] 牛克昌, 刘怿宁, 沈泽昊, 等. 群落构建的中性理论和生态位理论[J]. 生物多样性, 2009, 17(6): 579-593.

[41] 霍兵兵, 孙哲明, 欧文慧, 等. 环境筛选和扩散限制对长江流域湖北段湿地植物群落构建的共同影响[J]. 生态学报, 2023, 43(5): 1804-1811.

[42] 涂洪润, 李娇凤, 刘润红, 等. 桂林岩溶石山檵木种群空间格局及其关联性[J]. 应用生态学报, 2019, 30(8): 2621-2630.

[43] 丁晖, 陈水飞, 徐辉, 等. 2013-2018年武夷山亚热带常绿阔叶林乔木层动态[J]. 生态学报, 2022, 42(8): 3458-3469.

[44] 林波, 刘庆. 四种亚高山针叶林树种的表型可塑性对不同光照强度的响应[J]. 生态学报, 2008, 28(10): 4665-4675.

[45] LIU Y P, WANG S Y, LI M L, et al. The emergence of adaptive diversification from plant’s light competition[J]. Chaos Solitons Fractals, 2021, 152: 111366.

[46] 安栩立, 张振华, 杨建平. 青藏高原高寒灌木金露梅扩张态势分析[J]. 草业科学, 2024, 41(12): 2823-2832.

[47] 江常春, 梁希妮, 田磊, 等. 浙江省常绿阔叶林优势树种种间联结分析[J]. 浙江师范大学学报(自然科学版), 2022, 45(4): 437-445.

[48] 梁国新, 秦娇, 余臻, 等. 天台县濒危植物夏蜡梅野生分布及种群特征调查[J]. 福建林业科技, 2025, 52(2): 54-61.

[49] 孔垂华, 徐涛, 胡飞, 等. 环境胁迫下植物的化感作用及其诱导机制[J]. 生态学报, 2000, 20(5): 849-854.

基金资助

宁波市科技计划项目(2024S123)

国家自然科学基金项目(42371027)