云南松混交林火烧迹地更新早期草本层物种多样性的空间格局

doi:10.17521/cjpe.2015.0161

摘要/Abstract

摘要：

云南松(Pinus yunnanensis)林是我国西南地区最为广布的针叶林。火烧是云南松林常见的一种自然(或人为)干扰, 但对云南松林火后更新群落的物种多样性特征很少研究。该研究以云南禄丰县勤丰镇2013年“4.13”火灾后云南松混交林火烧迹地更新早期群落草本层为研究对象, 设置了47条1000 m²样带进行群落和环境因子调查, 探讨火烧后更新早期群落草本层的物种构成与丰富度的空间格局及其影响因素。结果表明: 火后更新草本层按物种构成可分为4个群落类型; 草本层物种丰富度受地形和火前群落冠层的影响显著, 与立木生物量和地形坡度正相关, 而与坡位和海拔负相关, 但与火烧强度没有显著的相关性。样带内草本群落β多样性与火前立木胸面积及火烧强度显著正相关, 而与海拔和坡位负相关。总体上, 生境地形特征、火前冠层和火烧强度构成了火烧后早期群落草本层物种构成与丰富度变化的主要控制因子。此外, 一年生和多年生草本物种的更新格局及其对环境的响应存在显著差异。

关键词: 火后更新, 草本多样性, 物种构成, 火前植被, 地形, 火烧强度

Abstract: Aims

Herb layer plays an important role in maintaining ecosystem functioning of forests. The aims of this study were to determine the pattern of species diversity in early post-fire regeneration of the herb layer in mixed Yunnan pine forests and to identify the effects of topography, fire severity, pre-fire vegetation and herb species life-history feature on post-fire regeneration.

Methods

We conducted field investigations of post-fire regeneration by randomly setting 47 transects of 10 m × 100 m in a burned forest stand in Qinfeng Township, Lufeng County, Yunnan Province, from September to October, 2013. The Two-Way Indicator Species Analysis (TWINSPAN) was used to classify the herb community types from the 47 transects, and species richness and two indexes of β-diversity were used to analyze herb species diversity and compositional change within sampling transects. Generalized linear model and hierarchical variation partitioning were applied to estimate factors affecting the pattern of herb species diversity of the post-fire communities.

Important findings

Four herb community types were identified for the 47 transects, which are differentiated by topographic features and fire severities. The species richness in the herb layer of early post-fire regeneration was primarily affected by elevation, topography, fire severity and the pre-fire forest canopy. Specifically, the species richness decreased with increasing elevation, and was positively correlated with biomass of the standing trees and steepness of site slope and negatively correlated with slope position. However, the species richness of herbs was not correlated with fire severity significantly. The within community β-diversity of the post-fire herb layer had a positive correlation with both basal area of pre-fire canopy trees and fire severity. Moreover, the β-diversity was lower at higher elevations. The results also highlighted the differences between annual and perennial species in their distribution patterns.

Key words: post-fire regeneration, herb diversity, species composition, pre-fire vegetation, topography, fire severity

韩杰, 应凌霄, 李贵祥, 沈泽昊. 云南松混交林火烧迹地更新早期草本层物种多样性的空间格局. 植物生态学报, 2016, 40(3): 200-211. DOI: 10.17521/cjpe.2015.0161

Jie HAN, Ling-Xiao YING, Gui-Xiang LI, Ze-Hao SHEN. Spatial patterns of species diversity in the herb layer of early post-fire regeneration in mixed Pinus yunnanensis forests. Chinese Journal of Plant Ecology, 2016, 40(3): 200-211. DOI: 10.17521/cjpe.2015.0161

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0161

https://www.plant-ecology.com/CN/Y2016/V40/I3/200

图/表 7

图1 研究区域的地理区位、气候、地貌特征和火烧面积(2013年)及其调查样点分布。

Fig. 1 The geographical location, climate, geomorphic feature of the burnt area (in 2013) and the distribution of sampling sites.

表1 火后植被与生境调查的变量

Table 1 Variables of post-fire vegetation and habitat investigations

类型 Type	变量 Variable	定义及范围 Definition & value range
火烧烈度 Fire severity	火烧比例 Burn ratio	0-1
火前植被 Pre-fire vegetation	ln(胸面积和) ln(Sum of basal area)	样方中死树胸面积和的自然对数值 Natural logarithm of sum of basal area of dead trees in sample plot
地形 Topography	海拔 Elevation (m)	1 928-2 331 m
	坡度 Slope gradient (°)	5°-52°
	坡向 Slope aspect (°)	0°(N) -180°(S)
	坡位 Slope position	2-下坡位; 3-中坡位; 4-上坡位; 5-山顶 2-lower slope; 3-midslope; 4-upper slope; 5-top
	坡形 Slope shape	1-凹; 2-平; 3-凸 1-concave; 2-flat; 3-convex

图2 火后更新草本层的TWINSPAN群落分类。N代表样方个数。A、B、C、D代表4种火后草本更新群落类型。A代表蕨+紫茎泽兰+头花龙胆群落; B代表西南野古草+紫马唐+白羊草群落; C代表鸡眼草+丝叶球柱草+白羊草群落; D代表荩草+鸡眼草+白羊草群落。(+)表示与分类聚组正相关; (-)表示与分类聚组负相关。

Fig. 2 The classification diagram of TWINSPAN for the herb layer of post-fire regenerated communities. N represents the number of transects. A, B, C, and D designate four types of post-fire regenerated herb communities. A, Pteridium aquilinum var. latiusculum + Ageratina adenophora + Gentiana cephalantha community; B, Arundinella hookeri + Digitaria violascens + Bothriochloa ischaemum community; C, Kummerowia striata + Bulbostylis densa + Bothriochloa ischaemum community; D, Arthraxon hispidus + Kummerowia striata + Bothriochloa ischaemum community. (+) means it is positive with classification; (-) means it is negetive with classification.

图3 火后更新植物群落中一年生(∆)和多年生(○)草本物种的典范对应分析排序格局。

Fig. 3 The canonical correspondence analysis ordination pattern for the annual (∆) and perennial (○) plant species in the early post-fire regenerated communities.

图4 火后植物群落样带(1 000 m2)草本层物种丰富度与生境因子的关系。所有小图中标注的a、b代表类型间t检验显著的不同水平。A、B、C、D代表4种火后草本更新群落类型。A代表蕨+紫茎泽兰+头花龙胆群落; B代表西南野古草+紫马唐+白羊草群落; C代表鸡眼草+丝叶球柱草+白羊草群落; D代表荩草+鸡眼草+白羊草群落。

Fig. 4 The relationships between habitat factors and the species richness in herb layer of the transects of post-fire plant communities. The letters a and b above all box-plots refer to significantly different levels in t-test. A, B, C, and D designate four types of post-fire regenerated herb communities. A, Pteridium aquilinum var. latiusculum + Ageratina adenophora + Gentiana cephalantha community; B, Arundinella hookeri + Digitaria violascens + Bothriochloa ischaemum community; C, Kummerowia striata + Bulbostylis densa + Bothriochloa ischaemum community; D, Arthraxon hispidus + Kummerowia striata + Bothriochloa ischaemum community.

图5 火后植物群落草本层β多样性格局。所有小图中标注的a、b表示类型间t检验显著的不同水平。A、B、C、D 代表4种火后草本更新群落类型。第一幅小图中A代表蕨+紫茎泽兰+头花龙胆群落; B代表西南野古草+紫马唐+白羊草群落; C代表鸡眼草+丝叶球柱草+白羊草群落; D代表荩草+鸡眼草+白羊草群落。

Fig. 5 The patterns of β-diversity in herb layer of post-fire plant communities. The letters a and b above all box-plots refer to significantly different levels in t-test. A, B, C, and D in the first graph panel designate four types of post-fire regenerated herb communities. A, Pteridium aquilinum var. latiusculum + Ageratina adenophora + Gentiana cephalantha community; B, Arundinella hookeri + Digitaria violascens + Bothriochloa ischaemum community; C, Kummerowia striata + Bulbostylis densa + Bothriochloa ischaemum community; D, Arthraxon hispidus + Kummerowia striata + Bothriochloa ischaemum community.

表2 生境因子对火后植物群落草本层物种多样性的影响

Table 2 Effects of habitat factors on the species diversity in herb layer of post-fire plant communities

生境因子 Habitat factor	α多样性 α diversity		β多样性 β diversity
生境因子 Habitat factor	样方尺度 Plot scale	样带尺度 Transect scale	Jaccard系数 Jaccard index	Morisita-Horn系数 Morisita-Horn index
海拔 Elevation	3.8	12.0	16.8^*	1.9
坡向 Aspect	1.6	1.9	1.6	9.2
坡形 Slope shape	17.9^*	6.4	1.5	5.3
坡位 Slope position	54.7^*	30.6^*	20.7^*	13.4
乔木胸面积对数 ln(sum of tree basal area)	3.4	26.6^*	38.6^*	41.6^*
坡度 Slope	8.7^*	20.9^*	11.9^*	19.5^*
火烧强度 Fire severity	9.9^*	1.5	8.9^*	9.0
调整的决定系数 Adjusted coefficient of determination	0.172	0.645	0.678	0.638

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