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