Characteristics of plant diversity and community structure in Shuanghe Nature Reserve in Daxing’anling area of northeastern China
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摘要:目的 植物多样性保护与良好林分结构是天然林保护的重要目标，明确林分个体大小、群落特征及乔灌草垂直结构对其变化的作用，将有助于基于林分结构调整来提升植物多样性。方法 本文于2016年8—9月对我国最北端寒温带针叶林双河国家级自然保护区的乔灌草更新层个体大小特征、群落特征和植物多样性特征进行详细调查，冗余分析和方差分解分析相结合解析植物多样性与群落结构特征的耦合关系，排序影响多样性变化的最重要个体与群落特征指标。结果 （1）研究区域共记录9个树种，其中白桦和兴安落叶松的株数占全部树种的80%以上；灌木树种19种，其中越桔数量占灌木总量的93%；草本植物记录了74种，最优势植物小叶樟占全部草本数量15%；更新层以先锋树种白桦数量最多，占比60%。（2）双河自然保护区树木胸径和树高仅相当于20世纪70年代的一半；双河自然保护区的物种丰富度、多样性和均匀度都表现出草本层 > 乔木层 > 灌木层，均匀度指数在0.53 ~ 0.73之间，更新层物种丰富度较低，乔木和草本多样性较大兴安岭核心地区（呼中）高出50%左右。（3）借助冗余和方差分解分析得出，群落特征、草本层对多样性的差异解释功能最大，而且更新层株高对多样性差异具有最大解释功能（P < 0.05）。结论 双河自然保护区具有本区域较高的乔木和草本植物丰富度，但尚需要更长的保护时间来恢复森林质量，而通过调整林分结构是实现植物物种演替更新以达到植物多样性保护的有效途径。Abstract:Objective Plant diversity protection and good forest structure are important targets for natural forest protection. It is helpful to improve plant diversity based on stand structure adjustment to clarify the effects of stand size, community characteristics and vertical structure of arbor, shrub and herb on its change.Method In this paper, from August to September of 2016, the individual size characteristics, community characteristics and plant diversity characteristics of arbor, shrub, herb and regeneration layer were investigated in China northernmost Shuanghe National Nature Reserve in the cold temperate zone of China, and redundant analysis and variance partitioning analysis were conducted to decouple the complex association and possible factors responsible for diversity variations.Result (1) A total of 9 tree species were found in the study area, with over 80% of white birch (Betula platyphylla) and larch (Larix gmelinii) in total number;19 shrub species were found and bilberry (Vaccinium vitis-idaea) accounted for 93% of the total shrubs. There were 74 herbaceous plants, and the most dominant was Deyeuxia langsdorffii (15%); pioneer tree species of Betula platyphylla was the most abundant in the regeneration layer (60%). (2) The diameter and height of trees were only half of those in 1970s. The species richness, diversity and evenness of Shuanghe National Nature Reserve all showed as herb layer > tree layer > shrub layer, and evenness index was 0.53−0.73, with very lower richness in the regeneration layer. The diversity of trees and herbs was about 50% higher than that in the central Daxing’anling at Huzhong Nature Reserve. (3) Redundancy ordination and variance partitioning found that community characteristics and herb characteristics had the greatest interpretation power for diversity variation, and the regenerated tree height showed the most statistical significant indicating function (P < 0.05).Conclusion Shuanghe National Nature Reserve has a rather high richness of arbor and herb plants in this area, but it still needs a long time to recover the forest quality related to tree size. Adjusting the stand structure should be an effective way to approach species diversity conservation via the plant species succession.
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图 1 乔木层、灌木层、草本层和更新层主要种相对多度
Figure 1. Relative abundance of main species in arbor layer, shrub layer, herb layer and regeneration layer
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图 2 区分个体大小(A)与群落特征和乔灌草的方差分解分析(B)
Figure 2. Variation partition between individual sizes and community characteristics (A) and among arbor layer, shrub layer and herb layer (B)
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图 3 群落结构特征与植物多样性的RDA排序(A)、不同林型样地分布图(B)和不同生境分布图(C)
Figure 3. RDA ordinations between species diversity and various factors as well as their statistics (A), non-differentiation of sampling sites of 4 different forest types (B), and distribution of sample plots in different environments (C)
表 1 个体大小、群落特征和植物多样性的统计表
Table 1 Statistics of individual size, community characteristics and species diversity indices
项目
Item均值
Mean中位数
Median标准误差
Standard
error25%
中位数
25%
median75%
中位数
75%
median最小50%区间
Min. 50%
range最大分布区间
Max.
distribution
interval分布区间
Distribution
interval类型
Type乔木丰富度
Arbor richness (T-R)3.61 3.00 0.23 3.00 5.00 2.00 ~ 3.00 3.00 ~ 4.00 2.00 ~ 6.00 单峰 Single peak 乔木 Simpson 指数
Arbor Simpson index (T-D)0.43 0.49 0.03 0.28 0.57 0.46 ~ 0.65 0.50 ~ 0.60 0.07 ~ 0.74 单峰 Single peak 乔木 Alatalo 指数
Arbor Alatalo index (T-Ea)0.71 0.68 0.03 0.59 0.85 0.67 ~ 0.88 0.60 ~ 0.70 0.38 ~ 0.99 多峰 Multi peak 灌木丰富度
Shrub richness (S-R)3.09 3.00 0.19 2.00 4.00 2.00 ~ 3.00 3.00 ~ 4.00 1.00 ~ 6.00 多峰 Multi peak 灌木 Simpson 指数
Shrub Simpson index (S-D)0.21 0.11 0.04 0.02 0.43 0 ~ 0.10 0 ~ 0.11 0 ~ 0.71 多峰 Multi peak 灌木 Alatalo 指数
Shrub Alatalo index (S-Ea)0.53 0.44 0.04 0.36 0.67 0.27 ~ 0.46 0.30 ~ 0.40 0.26 ~ 1.00 单峰 Single peak 草木丰富度
Herb richness (H-R)9.39 10.00 0.64 7.00 12.50 6.00 ~ 11.00 10.00 ~ 12.50 2.00 ~ 18.00 多峰 Multi peak 草木 Simpson 指数
Herb Simpson index (H-D)0.75 0.81 0.04 0.67 0.85 0.83 ~ 0.88 0.80 ~ 0.90 0.28 ~ 0.91 单峰 Single peak 草木 Alatalo 指数
Herb Alatalo index (H-Ea)0.73 0.74 0.02 0.63 0.84 0.75 ~ 0.85 0.70 ~ 0.80 0.42 ~ 1.00 单峰 Single peak 更新丰富度
Regenerated richness (R-R)2.33 2.00 0.15 2.00 3.00 2.00 ~ 3.00 1.00 ~ 5.00 单峰 Single peak 更新 Simpson 指数
Regenerated Simpson index (R-D)0.34 0.38 0.04 0.15 0.53 0.25 ~ 0.55 0 ~ 0.10 0 ~ 0.75 多峰 Multi peak 更新 Alatalo 指数
Regenerated Alatalo index (R-Ea)0.78 0.82 0.03 0.62 0.98 0.82 ~ 1.00 0.90 ~ 1.00 0.46 ~ 1.00 多峰 Multi peak 乔木株高
Arbor height (TH)/m12.82 12.00 0.36 10.99 13.36 10.30 ~ 12.20 10.00 ~ 12.00 7.74 ~ 19.14 单峰 Single peak 乔木胸径
Arbor DBH (TDBH)/cm13.99 13.53 0.75 11.10 16.59 11.10 ~ 14.80 12.50 ~ 15.00 7.21 ~ 13.53 单峰 Single peak 乔木枝下高
Arbor underbranch height (TCBH)/m6.96 6.95 0.26 5.91 8.00 6.30 ~ 8.20 6.00 ~ 8.00 3.57 ~ 9.69 单峰 Single peak 灌木高度
Shrub height (SH)/m0.36 0.19 0.07 0.11 0.50 0.10 ~ 0.21 0 ~ 0.25 0.10 ~ 1.93 单峰 Single peak 灌木地径
Shrub ground diameter (SGD)/cm1.03 0.95 0.07 0.80 1.10 0.80 ~ 1.11 0.75 ~ 1.00 0.48 ~ 2.18 单峰 Single peak 灌木冠幅
Shrub canopy width (SCD)/cm65.82 62.75 4.36 48.60 83.65 35.50 ~ 66.30 60.00 ~ 70.00 27.50 ~ 113.60 单峰 Single peak 草本高度
Herb height (HH)/cm29.04 28.83 1.03 25.11 32.77 27.10 ~ 34.00 25.00 ~ 30.00 16.20 ~ 41.00 单峰 Single peak 更新株高
Regenerated tree height (RH)/m1.66 1.60 0.03 1.40 2.11 1.60 ~ 2.00 1.50 ~ 1.75 0.50 ~ 3.10 多峰 Multi peak 更新地径
Regenerated ground diameter (RGD)/cm1.08 1.00 0.03 0.52 1.50 0.40 ~ 1.00 1.00 ~ 1.25 0.31 ~ 2.40 多峰 Multi peak 乔木郁闭度
Arbor canopy density (TC)0.73 0.80 0.02 0.65 0.80 0.71 ~ 0.80 0.80 ~ 0.85 0.50 ~ 0.90 单峰 Single peak 灌木盖度
Shrub coverage (SC)/%29.30 19.33 4.41 8.00 17.10 10.20 ~ 27.30 15.00 ~ 17.50 3.20 ~ 100.00 多峰 Multi peak 草本盖度
Herb coverage (HC)/%38.40 38.33 2.78 12.30 48.30 38.40 ~ 54.30 40.00 ~ 50.00 0.03 ~ 0.98 单峰 Single peak 乔木密度/(株·m−2)
Arbor density (TD)/(plant·m−2)0.13 0.13 0.01 0.08 0.16 0.11 ~ 0.17 0.15 ~ 0.175 0.04 ~ 0.28 多峰 Multi peak 灌木密度/(株·m−2)
Shrub density (SD)/(plant·m−2)22.70 14.08 4.00 2.33 36.17 0.25 ~ 14.08 0 ~ 10.00 0.25 ~ 84.08 单峰 Single peak 更新密度/(株·m−2)
Regenerated density (RD)/(plant·m−2)0.19 0.14 0.02 0.07 0.32 0.05 ~ 0.19 0.05 ~ 0.10 0.05 ~ 0.46 多峰 Multi peak 
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表 2 简单效应与条件效应统计表
Table 2 Statistics of simple-term effects and conditional term effects
简单效应 Simple term effect 条件效应 Conditional term effect 项目 Item 解释量 Explanatory quantity/% F P 项目 Item 解释量 Explanatory quantity/% F P RH 12.8 3.9 0.002 RH 12.8 3.9 0.002 SH 8.1 2.4 0.010 HC 6.2 2.0 0.048 HC 6.8 2.0 0.050 TDBH 5.0 1.6 0.106 HH 6.4 1.8 0.082 TD 5.0 1.7 0.108 SCD 6.2 1.8 0.068 SD 4.8 1.7 0.096 TDBH 5.0 1.4 0.174 RGD 5.0 1.8 0.088 SD 4.5 1.3 0.264 HH 4.1 1.5 0.154 TC 4.5 1.3 0.250 SH 3.1 1.2 0.344 RD 4.5 1.3 0.246 RD 2.4 0.9 0.530 TCBH 4.5 1.3 0.246 TC 2.3 0.9 0.584 TH 4.1 1.1 0.348 TCBH 2.2 0.8 0.608 TD 3.7 1.0 0.426 SCD 1.9 0.7 0.678 SC 3.3 0.9 0.512 SGD 2.6 0.9 0.476 RGD 3.2 0.9 0.542 SC 1.7 0.6 0.768 SGD 1.6 0.4 0.920 TH 1.2 0.4 0.882
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