甘肃小陇山暖温带针阔混交林大样地树木木材密度的空间变化

苏巧灵; 李梦琳; 田立新; 李安民; 袁士云; 刘文桢; 米湘成; 黄继红; 许玥; 丁易; 臧润国

doi:10.12356/j.2096-8884.2021-0009

甘肃小陇山暖温带针阔混交林大样地树木木材密度的空间变化

doi: 10.12356/j.2096-8884.2021-0009

苏巧灵^{1, 2,},
李梦琳¹,
田立新¹,
李安民³,
袁士云³,
刘文桢³,
米湘成⁴,
黄继红^{1, 5},
许玥^{1, 5},
丁易^{1, 5, ,},
臧润国^{1, 5}

1.
中国林业科学研究院森林生态环境与自然保护研究所，国家林业和草原局森林生态环境重点实验室，北京 100091
2.
中国科学院地理科学与资源研究所，生态系统网络观测与模拟院重点实验室，北京 100101
3.
甘肃省小陇山林业实验局，甘肃天水 741020
4.
中国科学院植物研究所，植被与环境变化国家重点实验室，北京 100093
5.
南京林业大学南方现代林业协同创新中心，江苏南京 210037

基金项目: 中央级公益性科研院所基本科研业务费专项资金中国林科院重点项目(CAFYBB2019ZA002)；国家自然科学基金面上项目(41671047)

详细信息

作者简介:
苏巧灵：E-mail: mysql94@163.com

通讯作者:
E-mail: dingy@caf.ac.cn

中图分类号: S718.54
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- 被引次数: 0
出版历程
- 收稿日期: 2021-05-28
- 网络出版日期: 2021-10-22
- 刊出日期: 2021-10-30

Spatial Heterogeneity of Wood Density in Coniferous-broadleaved Mixed Forest Dynamics Plot in Xiaolong Mountains, Gansu Province

Qiaoling SU^{1, 2
,},
Menglin LI¹,
Lixin TIAN¹,
Anmin LI³,
Shiyun YUAN³,
Wenzhen LIU³,
Xiangcheng MI⁴,
Jihong HUANG^{1, 5},
Yue XU^{1, 5},
Yi DING^{1, 5
, ,},
Runguo ZANG^{1, 5}

1.
Key Laboratory of Forest Ecology and Environment of National Forestry and Grassland Administration, Institute of Forest Ecology, Environment and Nature Conservation, Chinese Academy of Forestry, Beijing 100091, China
2.
Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China
3.
Xiaolong Moutains Forest Experiment Bureau of Gansu Province, Tianshui 741020, China
4.
State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
5.
Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing Forestry University, Nanjing 210037, China

摘要

摘要: 目的分析暖温带针阔混交林内木材密度的变化程度；探讨多元生境因子对不同群落木材密度空间分布的影响，揭示木材密度的种间和种内差异及其影响因素，为研究暖温带针阔混交林的物种共存与群落构建机制奠定基础。方法在甘肃小陇山地区的暖温带针阔混交林内布设大样地，依据样地内所有胸径≥ 10 cm独立个体的木材密度实测数据，利用方差分解分析种内、种间以及群落水平上木材密度的变化程度；利用多元回归分析探讨多元生境因子对群落水平木材密度空间分布的影响。结果样地内木材密度的差异主要来源于不同群落里树种间的差异，种间差异（71.70%）大于种内差异（28.30%）。在影响群落水平木材密度空间分布的因素中，非生物因子的影响大于生物因子。土壤因子中的pH值和地形因子中的海拔是影响木材密度的主要因子。在土壤pH值较低、海拔较高的群落中，木材密度的加权平均值较高。木材密度方差主要受土壤pH值影响，并随pH值的增大而增大。种内差异对木材密度加权平均值无显著影响，但对木材密度方差影响显著。结论甘肃小陇山针阔混交林木材密度空间异质性较强，非生物因子通过影响功能性状的总体分布来影响植物适应性，木材密度较高、种内差异较低的树种倾向于分布在海拔较高、土壤pH值较低的区域；相反，木材密度较低、种内差异较高的树种倾向于分布在海拔较低、土壤pH值较高的区域。在局域尺度下，忽视种内差异会导致木材密度方差被低估，影响对真实群落构建机制的理解。
- 木材密度 /
- 种内差异 /
- 生境空间异质性 /
- 针阔混交林 /
- 小陇山
Abstract: Objective This project was conducted to determine the variation of wood density within coniferous-broadleaved mixed forest dynamics plot in Xiaolong Mountains. We also investigated the relative significance of multiple habitat factors on the spatial distribution of wood density. Finally, it was revealed that inter- and intraspecific differences in wood density and their driving factors. This research would provide the basis for studying species coexistence and community assembly mechanism in coniferous-broadleaved mixed forests in warm temperate regions. Method The wood density of all living stems DBH ≥ 10 cm in the plot were sampled and measured. The variance decomposition was applied to differentiate the variance of wood density among individuals, species, and quadrat levels. The relative significance of various habitat factors on the wood density spatial distribution was investigated by multiple regression analysis, specifically community weighted mean (CWM) and community weighted variance (CWV). Result The variation of wood density was mainly contributed by the interspecies differences in the Xiaolong dynamics plot, where interspecific variation (71.70%) in wood density was significantly higher than intraspecific variation (28.30%). In addition, abiotic factors showed more significant influences than biotic factors on the wood density spatial distribution. The CWM of wood density was found significantly associated with topographic (e.g. elevation) and edaphic (e.g. pH value) factors. The trees with higher CWM values usually distributed in the areas at higher elevations with lower soil pH values. However, the CWV value of wood density was significantly positively related to soil pH value. Finally, the intraspecific variation had insignificant effects on the CWM of wood density, while significant effects on the CWV of wood density. Conclusion Our results demonstrated the significant spatial heterogeneity of wood density in coniferous-broadleaved mixed forest dynamics plot in Xiaolong Mountains, Gansu province. Plant adaptation was significantly affected by abiotic factors through overall distribution of functional traits. The species with higher wood densities and unobvious intraspecific differences preferred the habitats at higher altitudes with lower soil pH values. On the contrary, those species with lower wood densities and significant intraspecific variation were observed on the habitats at lower altitudes with higher soil pH values. Therefore, the intraspecific variation in wood density at local scales should not be overlooked, which would affect the understandings of natural community assembly mechanisms.
- wood density /
- intraspecific variation /
- habitat spatial heterogeneity /
- coniferous-broadleaved mixed forest /
- Xiaolong Mountains

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4 附录D 甘肃小陇山暖温带针阔混交林6 hm²样地生境因子（生物和非生物因子）分布

4. Appendix D. Spatial distribution of habitat factors (biotic and abiotic factors) in the 6 hm²Xiaolong Mountain forest dynamics plot, Gansu province

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图 1 甘肃小陇山6 hm²样地内40个物种的物种水平（a）及个体水平（b）木材密度频数分布图

Figure 1. Frequency distribution of species-level (a) and individual-level (b) wood densities for 40 common tree species in the 6 hm² plot Xiaolong Mountains forest dynamics plot, Gansu province

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图 2 甘肃小陇山6 hm²样地内40个物种的平均木材密度与种内木材密度变异系数的相关性

注：***: P＜0.001；黑色或红色实心分别代表木材密度低于或高于平均值的主要树种。Black or red solid circles indicate species with wood density below or above average.

Figure 2. The correlation between species-based mean wood density and the coefficient of variation in the 6 hm²Xiaolong Mountains forest dynamics plot, Gansu province

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图 3 甘肃小陇山针阔混交林6 hm²样地木材密度群落加权平均值（群落加权方差）在考虑（不考虑）种内差异时的空间分异

Figure 3. Spatial distribution of community-weighted mean (or community-weighted variance) of wood density (individual-based or species-based) of adults (DBH ≥ 10 cm) in the 6 hm² Xiaolong Mountains forest dynamics plot, Gansu province

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图 4 10个非生物因子-木材密度CWM（不考虑种内差异时）回归趋势

注：***: P ＜0.001; **: P ＜0.01; *: P ＜0.05; ns: 无显著差异 no significant difference.

Figure 4. Relationship between community-weighted mean of wood density（species-based）and 10 abiotic factors in 20 m × 20 m quadrats（DBH ≥ 10 cm）in the 6 hm² plot（n = 150）

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图 5 3个生物因子-木材密度CWV（考虑、不考虑种内差异）回归趋势

注：***: P ＜0.001; **: P ＜0.01; *: P ＜0.05; ns: 无显著差异 no significant difference.

Figure 5. Relationship between community-weighted variance of wood density (individual-based or species-based) and three biotic factors in 20 m × 20 m quadrats (DBH ≥ 10 cm) in the 6 hm² plot (n = 150)

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图 6 木材密度群落加权平均值与方差在考虑与不考虑种内差异时的差异

注：***: P ＜0.001; ns: 无显著差异 no significant difference.

Figure 6. The differences between species-based wood density values and individual-based values in the 6 hm² Xiaolong Mountains forest dynamics plot, Gansu province

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图 7 生境因子对基于物种水平与个体水平的木材密度群落加权平均值与方差的影响的标准化平均效应

注：***: P ＜0.001; **: P ＜0.01; *: P ＜0.05; ns: 无显著差异 no significant difference.

Figure 7. The standardized mean effects of predictor variables for species-based (and individual-based) community-weighted mean wood density and community-weighted variance in the 6 hm² Xiaolong Mountains forest dynamics plot, Gansu province

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2 附录B 非生物因子（10个）-木材密度CWV（不考虑种内差异时）回归趋势

注：***: P ＜0.001; **: P ＜0.01; *: P ＜0.05; ns: 无显著差异 no significant difference.

2. Appendix B. Relationship between community-weighted variance of wood density（species-based）and 10 abiotic factors in 20 m × 20 m quadrats（DBH ≥ 10 cm）in the 6 hm² plot (n = 150)

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3 附录C 非生物因子（10个）-木材密度CWV（考虑种内差异时）回归趋势

注：***: P ＜0.001; **: P ＜0.01; *: P ＜0.05; ns: 无显著差异 no significant difference.

3. Appendix C. Relationship between community-weighted variance of wood density (individual-based) and 10 abiotic factors in 20 m × 20 m quadrats (DBH ≥ 10 cm) in the 6 hm² plot (n = 150)

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5 附录E 甘肃小陇山暖温带针阔混交林6 hm²样地5个物种（DBH≥10 cm）空间分布

5. Appendix E. Spatial distribution of five species (DBH ≥ 10 cm) in a 6 hm² plot Xiaolong Mountain forest dynamics plot, Gansu province

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1 附录A 非生物因子-木材密度CWM（考虑种内差异时）回归趋势

注：***: P ＜0.001; **: P ＜0.01; *: P ＜0.05; ns: 无显著差异 no significant difference.

1. Appendix A. Correlations between community-weighted mean of wood density（individual-based）and abiotic factors in the 6 hm² plot (n = 150)

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1 附录F 甘肃小陇山6 hm²样地40个物种的平均木材密度及种内变异系数

1. Appendix F. The species-based mean wood densities and the coefficient of variation in the 6 hm² Xiaolong Mountain forest dynamics plot, Gansu province

物种Species	木材密度Wood density/(g/cm³)	变异系数CV/%	样本数n
华山松 Pinus armandii	0.407±0.051	12.64	108
冬瓜杨 Populus purdomii	0.420±0.061	14.46	7
粉椴 Tilia oliveri	0.429±0.071	16.43	21
山杨 Populus davidiana	0.454±0.042	9.35	102
光叶泡花树 Meliosma cuneifolia var. glabriuscula	0.459±0.079	17.20	14
红皮柳 Salix sinopurpurea	0.472±0.037	7.89	15
膀胱果 Staphylea holocarpa	0.502±0.129	25.62	8
香椿 Toona sinensis	0.480±0.034	7.14	8
刺楸 Kalopanax septemlobus	0.480±0.034	7.18	15
青榨槭 Acer davidii	0.492±0.043	8.82	41
坚桦 Betula chinensis	0.511±0.061	12.04	18
青皮槭 Acer cappadocicum	0.514±0.037	7.17	11
红麸杨 Rhus punjabensis var. sinica	0.517±0.053	10.16	127
青麸杨 Rhus potaninii	0.521±0.074	14.26	75
红桦 Betula albosinensis	0.523±0.024	4.63	10
建始槭 Acer henryi	0.529±0.050	9.42	15
灯台树 Cornus controversa	0.534±0.056	10.41	54
领春木 Euptelea pleiosperma	0.542±0.032	5.81	38
榆树 Ulmus pumila	0.562±0.060	10.58	39
白桦 Betula platyphylla	0.568±0.057	10.04	567
春榆 Ulmus davidiana var. japonica	0.568±0.043	7.62	183
小叶乌药 Lindera aggregata var. playfairii	0.578±0.047	8.21	190
木姜子 Litsea pungens	0.575±0.047	8.19	7
象蜡树 Fraxinus platypoda	0.589±0.059	10.02	23
湖北花楸 Sorbus hupehensis	0.594±0.040	6.78	14
苦树 Picrasma quassioides	0.596±0.092	15.36	7
甘肃山楂 Crataegus kansuensis	0.599±0.050	8.30	18
多毛樱桃 Cerasus polytricha	0.601±0.039	6.45	67
水榆花楸 Sorbus alnifolia	0.621±0.049	7.92	64
白蜡树 Fraxinus chinensis	0.630±0.065	10.29	6
五角枫 Acer pictum subsp. mono	0.632±0.038	5.95	23
三桠乌药 Lindera obtusiloba	0.641±0.047	7.37	105
微毛樱桃 Cerasus clarofolia	0.639±0.075	11.69	16
铁木 Ostrya japonica	0.646±0.038	5.88	75
湖北海棠 Malus hupehensis	0.647±0.046	7.15	18
红椋子 Cornus hemsleyi	0.661±0.060	9.02	98
锐齿槲栎 Quercus aliena var. acutiserrata	0.663±0.050	7.60	1827
蒙古栎 Quercus mongolica	0.672±0.044	6.58	29
唐棣 Amelanchier sinica	0.702±0.022	3.14	40
刺叶高山栎 Quercus spinosa	0.784±0.031	3.90	19
平均值 average value	0.564±0.092	9.46	—

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