森林与灌丛的灌木性状揭示不同的生活策略

doi:10.17521/cjpe.2020.0024

植物生态学报 ›› 2020, Vol. 44 ›› Issue (7): 715-729.DOI: 10.17521/cjpe.2020.0024

森林与灌丛的灌木性状揭示不同的生活策略

曹嘉瑜¹, 刘建峰¹, 袁泉¹, 徐德宇¹, 樊海东¹, 陈海燕¹, 谭斌¹, 刘立斌¹^,², 叶铎¹^,², 倪健¹^,²^,^*()

¹浙江师范大学化学与生命科学学院, 浙江金华 321004
²浙江金华山亚热带森林生态系统野外科学观测研究站, 浙江金华 321004

收稿日期:2020-01-18 接受日期:2020-05-21 出版日期:2020-07-20 发布日期:2020-06-12
通讯作者: *倪健: ORCID:0000-0001-5411-7050, nijian@zjnu.edu.cn
基金资助:
国家自然科学基金(31870462);浙江省“万人计划”(2019R52014)

Traits of shrubs in forests and bushes reveal different life strategies

CAO Jia-Yu¹, LIU Jian-Feng¹, YUAN Quan¹, XU De-Yu¹, FAN Hai-Dong¹, CHEN Hai-Yan¹, TAN Bin¹, LIU Li-Bin¹^,², YE Duo¹^,², NI Jian¹^,²^,^*()

¹College of Chemistry and Life Sciences, Zhejiang Normal University, Jinhua, Zhejiang 321004, China
²Jinhua Mountain Observation and Research Station for Subtropical Forest Ecosystems, Jinhua, Zhejiang 321004, China

Received:2020-01-18 Accepted:2020-05-21 Online:2020-07-20 Published:2020-06-12
Contact: NI Jian: ORCID:0000-0001-5411-7050, nijian@zjnu.edu.cn
Supported by:
National Natural Science Foundation of China(31870462);Ten Thousand Talents Program of Zhejiang Province (2019R52014).(2019R52014)

摘要/Abstract

摘要：

灌木是森林和灌丛生态系统的重要组成部分, 探究森林与灌丛灌木功能性状的差异, 可揭示灌木在不同生境的适应策略。该研究以金华北山森林群落林下灌木层、低山灌丛和山顶灌丛共24个样地中的优势灌木为研究对象, 分析叶片和小枝9个功能性状在3种生境下的总体差异, 以及种间、种内变异和不同生活型的差异。结果表明: 1) 9个性状在3种不同生境下存在差异。林下灌木具有较大的叶面积和比叶面积, 较小的叶干物质含量、叶和小枝的组织密度, 低山灌丛相较于山顶灌丛具有较大的叶厚度、叶组织密度和较小的比叶面积、小枝干物质含量。2)林下灌木的比叶面积、小枝直径、小枝组织密度和小枝干物质含量的种内种间变异系数最大, 低山灌丛的比叶面积、叶和小枝的干物质含量、叶和小枝的组织密度的种内种间变异系数最小。3)不同生活型间, 林下常绿灌木的叶厚度、叶组织密度、叶干物质含量显著高于落叶灌木, 落叶灌木的比叶面积显著高于常绿灌木, 而山顶灌丛叶厚度和比叶面积的差异规律与林下灌木相同, 叶组织密度和叶干物质含量的差异与其相反。4)影响灌木性状的主要因素是物种以及物种和生境的交互作用。总之, 森林群落林下灌木形成较大叶面积和比叶面积, 较小叶和小枝组织密度、叶干物质含量的性状组合, 以快速生长而适应光照较弱、竞争作用强的林下环境, 是资源获取型策略; 低山灌丛和山顶灌丛具有较大叶厚度、组织密度、干物质含量和小枝组织密度和较小叶面积、比叶面积等一系列储存养分、慢速生长的性状组合, 属于资源保守型策略。灌木植物性状的不同组合及其所反映的不同生活策略, 对亚热带地区退化植被的生态恢复具有指导意义。

关键词: 灌木, 植物功能性状, 亚热带森林, 灌丛, 生境, 生活史对策

Abstract:

Aims Shrubs play important roles in both forest and bushland ecosystems. This study aims to identify the adaptative strategies of shrubs in different habitats by analyzing the differences in functional traits of shrubs grown in understory of forest communities and in two bushlands.
Methods Nine functional traits for leaves and twigs were measured on samples collected from the dominant shrub species in 24 plots distributed in three contrasting habitats: forest understory, low mountain bushes, and bushes on the mountaintop, in Beishan Mountain of Jinhua, Zhejiang Province. The overall differences among habitats, inter- and intra-specific variations, and differences between life forms in the functional traits of shrubs were tested by statistical analysis.
Important findings The nine plant traits differed for shrubs grown in the three habitats. The shrubs in forest understory had higher leaf area (LA) and specific leaf area (SLA), lower leaf dry-matter content (LDMC), leaf tissue density (LTD) and twig tissue density (TTD), while those in low mountain bushes had greater leaf thickness (LT) and LTD, smaller SLA and twig dry-matter content (TDMC), compared with shrubs from bushes on the mountaintop. The inter- and intra-specific variation coefficients of SLA, twig diameter (TD), TTD, and TDMC were greatest in shrubs of the forest understory, whereas the inter- and intra-specific variation coefficients of SLA, LDMC, TDMC, and TTD were smallest in shrubs of low mountain bushes. Among different life forms, the understory evergreen shrubs had significantly higher LT, LTD, and LDMC, and lower SLA, than that of deciduous shrubs. The differences in LT and SLA between evergreen and deciduous shrubs of the mountaintop bushes were the same as the understory shrubs, but the differences in LTD and LDMC were reversed. Species and its interaction with habitat are the major factors affecting the shrub traits. In short, compared to the shrubs from bushes, the understory shrubs in forest communities form a series of trait combinations with greater LA and SLA, and smaller LTD, TTD and LDMC for faster growth in order to adapt to the understory environment with less light and stronger competition; this is a quick investment-return (resource acquisitive) strategy. Shrubs from low mountain bushes and the mountaintop bushes are associated with a series trait combinations with greater LT, LTD, LDMC and TTD, and smaller LA, SLA for storing more nutrients and growing slower; this is a slow investment-return (resource conservative) strategy. Different combinations of shrub functional traits and their various life strategies can provide guidance to the ecological restoration of degraded vegetation in the subtropical region of China.

Key words: shrub, plant functional traits, subtropical forests, shrubland, habitat, life history strategy

曹嘉瑜, 刘建峰, 袁泉, 徐德宇, 樊海东, 陈海燕, 谭斌, 刘立斌, 叶铎, 倪健. 森林与灌丛的灌木性状揭示不同的生活策略. 植物生态学报, 2020, 44(7): 715-729. DOI: 10.17521/cjpe.2020.0024

CAO Jia-Yu, LIU Jian-Feng, YUAN Quan, XU De-Yu, FAN Hai-Dong, CHEN Hai-Yan, TAN Bin, LIU Li-Bin, YE Duo, NI Jian. Traits of shrubs in forests and bushes reveal different life strategies. Chinese Journal of Plant Ecology, 2020, 44(7): 715-729. DOI: 10.17521/cjpe.2020.0024

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0024

https://www.plant-ecology.com/CN/Y2020/V44/I7/715

图/表 6

表1 金华北山24个样地基本特征及优势灌木

Table 1 Site characteristics and dominant shrubs for the 24 plots in Beishan Mountain of Jinhua, Zhejiang Province

表2 灌木主要功能性状及其反映的植物功能及生活策略

Table 2 Main functional traits of shrubs and their implications to plant functions and life strategies

图1 金华北山3种不同生境灌木的功能性状(平均值±标准误差)。 lmb, 低山灌丛; mtb, 山顶灌丛; uss, 林下灌木层。LA, 叶面积; LDMC, 叶干物质含量; LT, 叶厚度; LTD, 叶组织密度; SLA, 比叶面积; TBT, 小枝树皮厚度; TD, 小枝直径; TDMC, 小枝干物质含量; TTD, 小枝组织密度。不同小写字母代表差异显著(p < 0.05)。

Fig. 1 Shrub functional traits (mean ± SE) for three different habitats in Beishan Mountain of Jinhua, Zhejiang Province. lmb, low mountain bushes; mtb, mountaintop bushes; uss, understory shrub layer. LA, leaf area; LDMC, leaf dry-matter content; LT, leaf thickness; LTD, leaf tissue density; SLA, specific leaf area; TBT, twig bark thickness; TD, twig diameter; TDMC, twig dry-matter content; TTD, twig tissue density. Different lowercase letters indicate significant differences (p < 0.05).

图2 金华北山3种不同生境下灌木功能性状的种间和种内变异系数。 lmb, 低山灌丛; mtb, 山顶灌丛; uss, 林下灌木层。LA, 叶面积; LDMC, 叶干物质含量; LT, 叶厚度; LTD, 叶组织密度; SLA, 比叶面积; TBT, 小枝树皮厚度; TD, 小枝直径; TDMC, 小枝干物质含量; TTD, 小枝组织密度。

Fig. 2 Inter- and intra-specific coefficients of variation (CV) in shrub functional traits from three different habitats in Beishan Mountain of Jinhua, Zhejiang Province. lmb, low mountain bushes; mtb, mountaintop bushes; uss, understory shrub layer. LA, leaf area; LDMC, leaf dry-matter content; LT, leaf thickness; LTD, leaf tissue density; SLA, specific leaf area; TBT, twig bark thickness; TD, twig diameter; TDMC, twig dry-matter content; TTD, twig tissue density.

表3 物种、生活型和生境对灌木性状的影响

Table 3 Effects of species, life forms and habitats on shrub functional traits

性状 Trait	物种 Species			生活型 Life form			生境 Habitat			生活型×生境 Life form × Habitat			物种×生境 Species × Habitat
性状 Trait	F	Sig	R²	F	Sig	R²	F	Sig	R²	F	Sig	R²	F	Sig	R²
叶面积 LA	145.24	***	0.79	20.51	***	0.02	16.09	***	0.03	20.13	***	0.02	0.47	***	0.78
叶厚度 LT	82.01	***	0.68	187.76	***	0.16	1.14	ns	0.00	9.19	**	0.01	3.16	**	0.69
叶组织密度 LTD	31.03	***	0.44	0.14	ns	0.00	79.49	***	0.17	15.35	***	0.02	3.83	***	0.53
比叶面积 SLA	140.51	***	0.78	290.78	***	0.17	38.24	***	0.04	1.65	ns	0.00	2.05	**	0.80
叶干物质含量 LDMC	99.58	***	0.72	0.84	ns	0.00	33.02	***	0.08	12.28	***	0.01	6.56	***	0.76
小枝直径 TD	21.82	***	0.36	11.48	**	0.01	7.16	**	0.02	12.68	***	0.01	0.76	**	0.35
小枝树皮厚度 TBT	60.72	***	0.61	0.08	ns	0.00	15.89	***	0.04	0.50	ns	0.00	6.04	***	0.65
小枝组织密度 TTD	43.07	***	0.53	3.28	ns	0.00	11.31	***	0.03	3.24	ns	0.00	3.28	***	0.53
小枝干物质含量 TDMC	39.70	***	0.51	3.23	ns	0.00	21.48	***	0.05	1.70	ns	0.00	2.46	*	0.50

图3 金华北山3种不同生境下不同生活型灌木的功能性状(平均值±标准误差)。 lmb, 低山灌丛; mtb, 山顶灌丛; uss, 林下灌木层。LA, 叶面积; LDMC, 叶干物质含量; LT, 叶厚度; LTD, 叶组织密度; SLA, 比叶面积; TBT, 小枝树皮厚度; TD, 小枝直径; TDMC, 小枝干物质含量; TTD, 小枝组织密度。不同小写字母代表差异显著(p < 0.05)。

Fig. 3 Shrub functional traits (mean ± SE) for different life forms from three different habitats in Beishan Mountain of Jinhua, Zhejiang Province. lmb, low mountain bushes; mtb, mountaintop bushes; uss, understory shrub layer. LA, leaf area; LDMC, leaf dry-matter content; LT, leaf thickness; LTD, leaf tissue density; SLA, specific leaf area; TBT, twig bark thickness; TD, twig diameter; TDMC, twig dry-matter content; TTD, twig tissue density. Different lowercase letters indicate significant differences (p < 0.05).

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森林与灌丛的灌木性状揭示不同的生活策略

Traits of shrubs in forests and bushes reveal different life strategies

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