浑善达克沙地中部斑块格局影响植物多样性及功能性状

doi:10.17521/cjpe.2020.0239

摘要/Abstract

摘要：

景观格局与植物多样性之间的关系已被广泛研究, 然而, 斑块格局如何影响植物多样性, 以及这些影响的边缘类型差异尚不清楚。为从植物功能性状角度揭示斑块格局影响植物多样性的机制, 该研究采用不同边缘类型的斑块, 利用两年调查的705个样方数据, 结合遥感卫星图像解译和空间信息分析, 根据植物群落的空间位置和斑块格局, 按照向内、向外、向内成核和向外成核4种边缘类型将浑善达克沙地中部的斑块进行分类, 采用Duncan新复极差法比较了4种边缘类型斑块格局与植物功能性状多样性关系的差异, 应用Pearson相关系数和冗余分析(RDA)法量化了斑块格局对植物多样性的影响。结果发现, 植物丰富度和多样性指数与斑块格局密切相关, 这些关系存在着边缘类型的显著差异, 表现出不同的功能性状。对于向内边缘, 边缘指数与丰富度负相关, 和C₄植物比例正相关; 对于向外边缘, 边缘指数与C₄植物比例正相关; 对于向内成核边缘, 边缘密度与多年生植物和虫媒植物比例正相关; 对于向外成核边缘, 边缘角度和指数与物种Shannon-Wiener指数和Simpson指数正相关, 与Pielou指数、丰富度和动物传播植物比例负相关。这些发现表明, 斑块格局通过边缘效应影响植物多样性, 这种影响可以通过植物功能性状进行一定的解释。在景观管理、植被恢复和生物多样性保护方面需要考虑这些因素。

关键词: 斑块格局, 植物多样性, 边缘类型, 功能性状, 边缘效应

Abstract:

Aims We address the issue of adapting patch pattern for improved plant biodiversity conservation in sandy grasslands by assessing the role of edge type in the influences of patch pattern on plant diversity and functional traits. We test the hypotheses that plant richness, alpha and beta diversity would vary according to patch pattern but that these effects would vary according to edge type, through adaptation of plant functional traits. Methods Jointing satellite imagery interpretation, 705 plots from two-year surveys and spatial analysis, patch edges in center Hunshandak Sandland were categorized into four types. Duncan’s differences were tested across four edge types. Pearson’s correlation and redundancy analysis (RDA) were used to discern the patch contributions. Important findings Plant richness and diversity indices are closely related with patch patterns, these relations vary significantly across edge types, demonstrating different functional traits. For inward edge, edge metrics attributes to richness reduction and C₄ plant proportion, mean perimeter-area ratio (MPAR) positively related to nitrogen-fixed species proportion. For outward edge, mean patch fractal dimension (MPFD) negatively related to richness, MPAR positively related to the proportion of nitrogen-fixing species, edge metrics positively related to C₄ proportion. For inward nucleating edge, MPAR, total edge (TE) and landscape area (TLA) positively related to perennial plants, TE and TLA positively linked to insect-pollination plants. For outward nucleating edge, angle and pattern metrics positively related to Shannon-Wiener and Simpson index, negatively related to Pielou index, richness and animal-seed dispersing plants. Patch pattern distressing plant diversity through edge effects can be moderately interpreted by plant practical qualities and edge metrics. It must be well thought-out in landscape management, vegetation restoration and biodiversity conservation.

Key words: patch pattern, plant diversity, edge type, functional traits, edge effect

范敏, 卢奕曈, 王照华, 黄颖琪, 彭羽, 尚佳欣, 张杨. 浑善达克沙地中部斑块格局影响植物多样性及功能性状. 植物生态学报, 2022, 46(1): 51-61. DOI: 10.17521/cjpe.2020.0239

Min FAN, Yi-Tong LU, Zhao-Hua WANG, Ying-Qi HUANG, Yu PENG, Jia-Xin SHANG, Yang ZHANG. Effects of patch pattern on plant diversity and functional traits in center Hunshandak Sandland. Chinese Journal of Plant Ecology, 2022, 46(1): 51-61. DOI: 10.17521/cjpe.2020.0239

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0239

https://www.plant-ecology.com/CN/Y2022/V46/I1/51

图/表 5

图1 浑善达克沙地中部斑块4种边缘类型划分示意图(基于Porensky和Young (2013)标准)。

Fig. 1 Diagrams point out four types of patches with dissimilar probable edge effects: inward, outward, inward nucleating and outward nucleating in center Hunshandak Sandland (based on the criterion of Porensky & Young (2013)).

表1 4种边缘类型斑块内的优势植物物种(平均值±标准差)

Table 1 Dominant species in the patches of four edge types (mean ± SD)

边缘类型 Edge type	优势物种 Dominant plant	重要值 IV
向内 Inward	百里香 Thymus mongolicus	0.96 ± 0.09
	鹤虱 Lappula myosotis	0.66 ± 0.13
	柠条锦鸡儿 Caragana korshinskii	0.61 ± 0.15
向外 Outward	披碱草 Elymus dahuricus	0.67 ± 0.13
	寸草 Carex duriuscula	0.59 ± 0.09
	沙芦草 Agropyron mongolicum	0.53 ± 0.11
向内成核 Inward nucleating	猪毛菜 Salsola collina	0.79 ± 0.25
	黑沙蒿 Artemisia ordosica	0.72 ± 0.18
	沙芦草 Agropyron mongolicum	0.55 ± 0.09
向外成核 Outward nucleating	硬阿魏 Ferula bungeana	0.83 ± 0.20
	黑沙蒿 Artemisia ordosica	0.72 ± 0.21
	榆树(幼苗) Ulmus pumila (seedling)	0.61 ± 0.15

表2 浑善达克沙地中部4种边缘类型植物多样性指数、盖度和功能性状的差异(平均值±标准差)

Table 2 Differences of plant diversity indices, coverage and functional traits across four edge types in center Hunshandak Sandland (mean ± SD)

边缘类型 Edge type	向内 Inward	向外 Outward	向内成核 Inward nucleating	向外成核 Outward nucleating
Shannon-Wiener指数 Shannon-Wiener index (H′)	1.69 ± 0.36^Ba	1.89 ± 0.49^ABa	2.07 ± 0.53^Aa	2.10 ± 0.45^Aa
Simpson指数 Simpson index (D)	0.71 ± 0.10^Aa	0.64 ± 0.07^Aa	0.66 ± 0.08^Aa	0.68 ± 0.07^Aa
Pielou指数 Pielou index (J′)	0.68 ± 0.10^Ba	0.70 ± 0.11^Ba	0.77 ± 0.07^Aa	0.78 ± 0.06^Aa
β多样性指数 β diversity index (βw)	2.39 ± 1.01^Ba	3.81 ± 1.20^Aa	3.81 ± 1.39^Aa	4.15 ± 2.23^Aa
物种丰富度 Species richness	12.25 ± 5.40^Ba	18.30 ± 7.53^Aa	20.00 ± 8.03^Aa	18.08 ± 6.20^Aa
盖度 Coverage (%)	18.68 ± 9.76^Aa	23.09 ± 10.76^Aa	21.42 ± 10.34^Aa	24.20 ± 9.88^Aa
一年生植物比例 Annual proportion (%)	28.57 ± 2.21^Aa	8.78 ± 1.34^Ba	20.21 ± 2.22^ABa	17.47 ± 1.47^ABa
两年生植物比例 Biennial proportion (%).	5.24 ± 9.52^Ba	1.78 ± 2.77^ABa	3.14 ± 5.47^Aa	9.81 ± 1.46^ABa
多年生植物比例 Perennial proportion (%)	5.67 ± 1.98^Aa	6.52 ± 2.17^Aa	6.17 ± 2.65^Aa	5.78 ± 2.24^Aa
灌木植物比例 Shrub proportion (%)	1.33 ± 1.02^Aa	2.17 ± 1.94^Aa	1.57 ± 1.62^Aa	1.88 ± 1.29^Aa
乔木植物比例 Tree proportion (%)	0.06 ± 0.14^Aa	0.07 ± 0.18^Aa	0.13 ± 0.37^Aa	0.22 ± 0.41^Aa
CAM植物比例 CAM plant proportion (%)	3.12 ± 4.33^Aa	1.24 ± 2.76^ABa	0.85 ± 1.61^ABa	0.15 ± 0.42^Ba
C₃植物比例 C₃ plant proportion (%)	3.67 ± 2.03^Ba	4.57 ± 2.73^ABa	5.60 ± 2.17^Aa	5.14 ± 1.37^ABa
C₄植物比例 C₄ plant proportion (%)	54.28 ± 27.22^Aa	51.35 ± 28.47^Aa	43.19 ± 21.88^Aa	48.05 ± 13.46^Aa
风传种植物比例 Wind-dispersed proportion (%)	1.33 ± 9.61^Ba	2.48 ± 1.87^ABa	2.44 ± 1.65^ABa	2.84 ± 1.78^Aa
自动传种植物比例 Auto-dispersed proportion (%)	3.20 ± 2.01^Aa	1.60 ± 1.58^Bab	1.38 ± 8.63^Bb	1.35 ± 1.23^Bb
重力传种植物比例 Gravity-dispersed proportion (%)	4.34 ± 1.90^Aa	3.21 ± 2.09^Aa	4.40 ± 1.85^Aa	4.29 ± 1.85^Aa
动物传种植物比例 Animal dispersion proportion (%)	12.78 ± 17.58^Aa	21.63 ± 17.69^Aa	12.61 ± 13.43^Aa	14.02 ± 14.77^Aa
风媒花植物比例 Wind-pollinated proportion (%)	6.71 ± 2.55^Aa	3.72 ± 2.20^Bb	5.34 ± 1.88^ABab	5.41 ± 1.81^ABab
虫媒花植物比例 Insect-pollinated proportion (%)	3.21 ± 2.55^Ba	6.29 ± 2.33^Ab	4.39 ± 1.74^ABa	4.44 ± 1.92^ABa
非固氮植物比例 Non nitrogen-fixing proportion (%)	72.81 ± 25.62^ABa	60.06 ± 19.98^Ba	76.50 ± 16.05^ABa	80.08 ± 16.06^Aa
固氮植物比例 Nitrogen-fixing proportion (%)	21.64 ± 22.84^Ba	39.88 ± 20.02^Aa	26.27 ± 19.02^ABa	20.02 ± 16.00^Ba

表2 浑善达克沙地中部4种边缘类型植物多样性指数、盖度和功能性状的差异(平均值±标准差)

Table 2 Differences of plant diversity indices, coverage and functional traits across four edge types in center Hunshandak Sandland (mean ± SD)

边缘类型 Edge type	向内 Inward	向外 Outward	向内成核 Inward nucleating	向外成核 Outward nucleating
Shannon-Wiener指数 Shannon-Wiener index (H′)	1.69 ± 0.36^Ba	1.89 ± 0.49^ABa	2.07 ± 0.53^Aa	2.10 ± 0.45^Aa
Simpson指数 Simpson index (D)	0.71 ± 0.10^Aa	0.64 ± 0.07^Aa	0.66 ± 0.08^Aa	0.68 ± 0.07^Aa
Pielou指数 Pielou index (J′)	0.68 ± 0.10^Ba	0.70 ± 0.11^Ba	0.77 ± 0.07^Aa	0.78 ± 0.06^Aa
β多样性指数 β diversity index (βw)	2.39 ± 1.01^Ba	3.81 ± 1.20^Aa	3.81 ± 1.39^Aa	4.15 ± 2.23^Aa
物种丰富度 Species richness	12.25 ± 5.40^Ba	18.30 ± 7.53^Aa	20.00 ± 8.03^Aa	18.08 ± 6.20^Aa
盖度 Coverage (%)	18.68 ± 9.76^Aa	23.09 ± 10.76^Aa	21.42 ± 10.34^Aa	24.20 ± 9.88^Aa
一年生植物比例 Annual proportion (%)	28.57 ± 2.21^Aa	8.78 ± 1.34^Ba	20.21 ± 2.22^ABa	17.47 ± 1.47^ABa
两年生植物比例 Biennial proportion (%).	5.24 ± 9.52^Ba	1.78 ± 2.77^ABa	3.14 ± 5.47^Aa	9.81 ± 1.46^ABa
多年生植物比例 Perennial proportion (%)	5.67 ± 1.98^Aa	6.52 ± 2.17^Aa	6.17 ± 2.65^Aa	5.78 ± 2.24^Aa
灌木植物比例 Shrub proportion (%)	1.33 ± 1.02^Aa	2.17 ± 1.94^Aa	1.57 ± 1.62^Aa	1.88 ± 1.29^Aa
乔木植物比例 Tree proportion (%)	0.06 ± 0.14^Aa	0.07 ± 0.18^Aa	0.13 ± 0.37^Aa	0.22 ± 0.41^Aa
CAM植物比例 CAM plant proportion (%)	3.12 ± 4.33^Aa	1.24 ± 2.76^ABa	0.85 ± 1.61^ABa	0.15 ± 0.42^Ba
C₃植物比例 C₃ plant proportion (%)	3.67 ± 2.03^Ba	4.57 ± 2.73^ABa	5.60 ± 2.17^Aa	5.14 ± 1.37^ABa
C₄植物比例 C₄ plant proportion (%)	54.28 ± 27.22^Aa	51.35 ± 28.47^Aa	43.19 ± 21.88^Aa	48.05 ± 13.46^Aa
风传种植物比例 Wind-dispersed proportion (%)	1.33 ± 9.61^Ba	2.48 ± 1.87^ABa	2.44 ± 1.65^ABa	2.84 ± 1.78^Aa
自动传种植物比例 Auto-dispersed proportion (%)	3.20 ± 2.01^Aa	1.60 ± 1.58^Bab	1.38 ± 8.63^Bb	1.35 ± 1.23^Bb
重力传种植物比例 Gravity-dispersed proportion (%)	4.34 ± 1.90^Aa	3.21 ± 2.09^Aa	4.40 ± 1.85^Aa	4.29 ± 1.85^Aa
动物传种植物比例 Animal dispersion proportion (%)	12.78 ± 17.58^Aa	21.63 ± 17.69^Aa	12.61 ± 13.43^Aa	14.02 ± 14.77^Aa
风媒花植物比例 Wind-pollinated proportion (%)	6.71 ± 2.55^Aa	3.72 ± 2.20^Bb	5.34 ± 1.88^ABab	5.41 ± 1.81^ABab
虫媒花植物比例 Insect-pollinated proportion (%)	3.21 ± 2.55^Ba	6.29 ± 2.33^Ab	4.39 ± 1.74^ABa	4.44 ± 1.92^ABa
非固氮植物比例 Non nitrogen-fixing proportion (%)	72.81 ± 25.62^ABa	60.06 ± 19.98^Ba	76.50 ± 16.05^ABa	80.08 ± 16.06^Aa
固氮植物比例 Nitrogen-fixing proportion (%)	21.64 ± 22.84^Ba	39.88 ± 20.02^Aa	26.27 ± 19.02^ABa	20.02 ± 16.00^Ba

图2 浑善达克沙地中部向内边缘(A、B)和向外边缘(C、D)斑块格局指数与植物多样性指数(A、C)和功能性状(B、D)的冗余分析(RDA)。angle, 样方所处角度; depth, 样方距离边缘深度; ED, 边缘密度; MPAR,平均周长-面积比; MPFD, 平均斑块分形维数; MSI, 平均形状指数; TE, 总边缘长度; TLA, 景观面积。βw, β多样性指数; D, Simpson多样性指数; H′, Shannon-Wiener多样性指数; J′, Pielou多样性指数。abundance, 多度; anthesis, 开花时间; coverage, 盖度; ecotype, 生态型; IV, 重要值; life form, 生活型; nitrogen fixation, 固氮类型; pollination, 传粉途径; photosynthesis, 光合途径; richness, 丰富度; seed dispersal, 种子传播方式。

Fig. 2 Redundancy analysis (RDA) indicates the relationships between patch metrics and plant diversity indices (A, C), and functional traits (B, D) for inward edge (A, B) and outward edge (C, D) in center Hunshandak Sandland. angle, angle of sample location; depth, distance to edge; ED, edge density; MPAR, mean perimeter-area ratio; MPFD, mean patch fractal dimension; MSI, mean shape index; TE, total edge; TLA, landscape area. βw, β diversity index; D, Simpson index; H′, Shannon-Wiener index; J′, Pielou index. IV, important value.

图3 浑善达克沙地中部向内成核边缘(A、B)和向外成核边缘(C、D)斑块格局指数与植物多样性指数(A、C)和功能性状(B、D)的冗余分析(RDA)。angle, 样方所处角度; depth, 样方距离边缘深度; ED, 边缘密度; MPAR,平均周长-面积比; MPFD, 平均斑块分形维数; MSI, 平均形状指数; TE, 总边缘长度; TLA, 景观面积。βw, β多样性指数; D, Simpson多样性指数; H′, Shannon- Wiener多样性指数; J′, Pielou多样性指数。abundance, 多度; anthesis, 开花时间; coverage, 盖度; ecotype, 生态型; IV, 重要值; life form, 生活型; nitrogen fixation, 固氮类型; pollination, 传粉途径; photosynthesis, 光合途径; richness, 丰富度; seed dispersal, 种子传播方式。

Fig. 3 Redundancy analysis (RDA) indicate the relationships between patch metrics and plant diversity indices (A, C), and functional traits (B, D) for Inward nucleating edge (A, B) and outward nucleating edge (C, D) in enter Hunshandak Sandland. angle, angle of sample location; depth, distance to edge; ED, edge density; MPAR, mean perimeter-area ratio; MPFD, mean patch fractal dimension; MSI, mean shape index; TE, total edge; TLA, landscape area. βw, β diversity index; D, Simpson index; H′, Shannon-Wiener index; J′, Pielou index; IV, important value.

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