Comparison of plant biomass in conifer and broadleaf mixed artificial forests in south subtropical area and analyses of influential factors

doi:10.17521/cjpe.2024.0398

Abstract

Abstract:

Aims Tropical and subtropical areas are hotspots for the distribution of artificial forests in our country. Clarifying the status and influential factors of plant biomass in typically mixed artificial forests in these areas is important. This can help to uncover the limiting factors for stand productivity and guide the management of artificial forests.

Methods In this study, we selected a conifer-broadleaf mixed forest in Yunyong forestry station located in Foshan, Guangdong. This forest was transformed from a Chinese fir plantation. First, we compared the difference in biomass of each plant organ among tree species configuration modes at the early recovery stage (8-16 a). Second, the shifts in soil physicochemical properties and community-level plant functional traits under different conditions of tree species configuration were characterized. Finally, we explored the relative contributions of tree species configuration, soil properties and plant leaf traits to variations in plant biomass, and evaluated their influential pathways. Based on these analyses, we aimed to evaluate the key influential factors for plant biomass in artificial forests in south subtropical areas.

Important findings Our results showed that both plant and organ biomass differed significantly under different tree species configuration modes. Tree species configurations affected plant biomass mainly via regulating leaf functional traits and soil nutrient content. Moreover, leaf functional traits had greater influence on foliage and branch biomass, while the variation in soil nutrient content was the main factor driving the changes in truck, roots and total plant biomass. Specifically, tree species configurations characterized by a higher ratio of leaf nitrogen (N) : phosphorus (P) and a higher functional diversity had greater biomass in plant communities. An increase in soil total P content significantly promoted the increase in plant biomass, while an excess in soil total potassium would suppress the growth in biomass. This study compared the relative effects of leaf functional traits and soil nutrient content on plant biomass in the typically mixed artificial stands in south subtropical forests, and explored the key influential factors for the biomass in total and in each organ. These results provided the theoretical bases for restoration, operation and management of the artificial forests in south subtropical areas.

Key words: stand biomass, conifer-broadleaf mixed forest, tree species configuration, functional trait, soil nutrient, vegetation restoration

ZHANG Jing, CHEN Jie, LI Yan-Peng, PAN Li-Jun, XU Han, LI Yi-De, HE Hai-Sheng. Comparison of plant biomass in conifer and broadleaf mixed artificial forests in south subtropical area and analyses of influential factors[J]. Chin J Plant Ecol, 2026, 50(2): 400-416.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2024.0398

https://www.plant-ecology.com/EN/Y2026/V50/I2/400

Figures/Tables 7

References 67

[1]	Balazs KR, Munson SM, Butterfield BJ (2022). Functional composition of plant communities mediates biomass effects on ecosystem service recovery across an experimental dryland restoration network. Functional Ecology, 36, 2317-2330. DOI URL
[2]	Bongers FJ, Schmid B, Bruelheide H, Bongers F, Li S, von Oheimb G, Li Y, Cheng AP, Ma KP, Liu XJ (2021). Functional diversity effects on productivity increase with age in a forest biodiversity experiment. Nature Ecology & Evolution, 5, 1594-1603.
[3]	Che Z, Shi J, Xu DL (2016). Aboveground biomass distribution of desert grasslands and its responses to environmental factors in middle regions of Hexi Corridors. Bulletin of Soil and Water Conservation, 36(4), 193-196.
	[车卓, 石菁, 徐德亮 (2016). 河西走廊中段荒漠草地地上生物量分布特征及其与环境因子的关系. 水土保持通报, 36(4), 193-196.]
[4]	Chen BZ, Fang JC, Piao SL, Ciais P, Black TA, Wang F, Niu SL, Zeng ZZ, Luo YQ (2024). A meta-analysis highlights globally widespread potassium limitation in terrestrial ecosystems. New Phytologist, 241, 154-165.
[5]	Chen XL, Ju Q, Lin KL (2014). Development status, issues and countermeasures of China’s plantation. World Forestry Research, 27(6), 54-59.
	[陈幸良, 巨茜, 林昆仑 (2014). 中国人工林发展现状、问题与对策. 世界林业研究, 27(6), 54-59.]
[6]	Cheng DL, Lin N (2009). Research on the specific leaf area of common plants in Fuzhou City. Journal of Anhui Agricultural Sciences, 37, 15131-15133.
	[程栋梁, 林娜 (2009). 福州市常见植物比叶面积研究. 安徽农业科学, 37, 15131-15133.]
[7]	Cui EQ, Lu RL, Xu XN, Sun HF, Qiao Y, Ping JY, Qiu SY, Lin YH, Bao JH, Yong YT, Zheng ZM, Yan ER, Xia JY (2022). Soil phosphorus drives plant trait variations in a mature subtropical forest. Global Change Biology, 28, 3310-3320. DOI PMID
[8]	Deng RM (2018). Relationship between the biomass characteristics and soil nutrients among different forest types in Daiyun Mountain. Forestry Construction, (6), 29-32.
	[邓瑞明 (2018). 戴云山不同类型森林生物量差异特征及其与土壤养分的关系. 林业建设, (6), 29-32.]
[9]	Fang MQ, Yu FZ, Zhang ZQ, Shan YB (2020). Evaluation of soil fertility under typical subtropical forest vegetations in Lushan Mountain Region. Hans Journal of Soil Science, 8, 34-41. DOI URL
	[方明晴, 于法展, 张忠启, 单勇兵 (2020). 庐山亚热带典型森林植被类型下土壤肥力评价. 土壤科学, 8, 34-41.]
[10]	Fu Y (2023). The Leaf Functional Traits of Main Plants on the West Slope of Mount Wuyi Mountain and Their Relationship with Environmental Factors. Master degree dissertation, Jiangxi Agricultural University, Nanchang.
	[付裕 (2023). 武夷山西坡主要植物叶片功能性状及其与环境因子之间的关系. 硕士学位论文, 江西农业大学, 南昌.]
[11]	Garnier E, Laurent G, Bellmann A, Debain S, Berthelier P, Ducout B, Roumet C, Navas ML (2001). Consistency of species ranking based on functional leaf traits. New Phytologist, 152, 69-83. DOI PMID
[12]	Gillespie LM, Hättenschwiler S, Milcu A, Wambsganss J, Shihan A, Fromin N (2021). Tree species mixing affects soil microbial functioning indirectly via root and litter traits and soil parameters in European forests. Functional Ecology, 35, 2190-2204. DOI URL
[13]	Guo Q, Zhao HB, Zhou GY, Long WG, Gan GL, Wu SY, Meng MY, Chen JL (2022). Biomass and its distribution pattern of four tree species plantation in south subtropical China. Forest Research, 35(1), 182-189.
	[郭耆, 赵厚本, 周光益, 隆卫革, 甘谷列, 吴尚勇, 蒙敏燕, 陈金兰 (2022). 南亚热带4个树种人工林生物量及其分配格局. 林业科学研究, 35(1), 182-189.]
[14]	Guo Y, Liu Y, Fan GQ, Chen FQ, Liu JX, Tang XL, Li X (2023). Characteristics of biomass allocation and carbon, nitrogen and phosphorus accumulation in the south subtropical Castanopsis hystrix plantation. Journal of Anhui Agricultural University, 50, 199. DOI: 10.13610/j.cnki.1672-352x.20230511.010.
	[郭勇, 刘岳, 范国强, 陈富强, 刘菊秀, 唐旭利, 李旭 (2023). 南亚热带红锥人工林生物量分配与碳氮磷积累特征. 安徽农业大学学报, 50, 199. DOI: 10.13610/j.cnki.1672-352x.20230511.010.]
[15]	Han TT, Tang X, Ren H, Wang J, Liu N, Guo QF (2021). Community/ecosystem functional diversity: measurements and development. Acta Ecologica Sinica, 41, 3286-3295.
	[韩涛涛, 唐玄, 任海, 王俊, 刘楠, 郭勤峰 (2021). 群落/生态系统功能多样性研究方法及展望. 生态学报, 41, 3286-3295.]
[16]	He B, Wei XM, Feng JQ, Yang JQ, Huang CB, Zhang RS (2023). Soil physicochemical properties in different stand ages of Taiwania flousiana in northwestern Guangxi. Journal of Northeast Forestry University, 51(6), 77-82.
	[何斌, 韦秀媚, 冯建强, 杨家强, 黄承标, 张日施 (2023). 桂西北不同林龄秃杉人工林土壤理化性质. 东北林业大学学报, 51(6), 77-82.]
[17]	He PY, Chen HH (2018). Comparison of water content and biomass of different organs of single plant in Pinus massoniana forest. Jiangsu Agricultural Sciences, 46(20), 139-142.
	[何佩云, 谌红辉 (2018). 连栽马尾松林单株不同器官含水率及生物量的比较. 江苏农业科学, 46(20), 139-142.]
[18]	Hu EC, Wang Z, Li ZH, Yang HF, Li ZF, Wang XJ, Gao RH (2024). Dynamics of understory plant diversity and biomass of Pinus sylvestris var. mongolica plantations with different ages in Mu Us Sandy Land. Chinese Journal of Ecology, 43, 3246-3254.
	[胡尔查, 王铮, 李梓豪, 杨海峰, 李卓凡, 王晓江, 高润红 (2024). 毛乌素沙地不同林龄樟子松人工林林下植物多样性和生物量的动态变化. 生态学杂志, 43, 3246-3254.]
[19]	Huang QF (2011). Structure and Biomass of Dominant Tree of Natural Secondary Broad-leaved Forest in the Low Mountain of Southern Anhui. PhD dissertation, Anhui Agricultural University, Hefei.
	[黄庆丰 (2011). 皖南低山天然次生阔叶林林分结构与优势树种地上生物量. 博士学位论文, 安徽农业大学, 合肥.]
[20]	Huang XM, You YM, Lan JC, Liu SR, Jia HY, Zhu HG, Wen YG (2016). The effect of carbon storage and its allocation in Cunninghamia lanceolata plantations with different thinning intensities. Acta Ecologica Sinica, 36, 156-163.
	[黄雪蔓, 尤业明, 蓝嘉川, 刘世荣, 贾宏炎, 朱宏光, 温远光 (2016). 不同间伐强度对杉木人工林碳储量及其分配的影响. 生态学报, 36, 156-163.]
[21]	Huang YH, Zhang WQ, Gan XH, Tang HH, Pan LJ, Xian GB (2016). Evaluation of different tree species composition patterns in stand conversion of Cunninghamia lanceolata in southern subtropical region of China. Ecology and Environmental Sciences, 25, 956-964.
	[黄钰辉, 张卫强, 甘先华, 唐洪辉, 盘李军, 冼杆标 (2016). 南亚热带杉木林分改造中不同树种组合模式评价. 生态环境学报, 25, 956-964.] DOI
[22]	Jiang J, Cao NN, Yu MX, Chang ZB, Chen Y, Zhang S, Yan JH (2019). A review of the key mechanisms of acidic forest soils to buffer acid deposition. Journal of Tropical and Subtropical Botany, 27, 491-499.
	[江军, 曹楠楠, 俞梦笑, 常中兵, 陈洋, 张硕, 闫俊华 (2019). 酸性森林土壤缓冲酸沉降关键机理研究进展. 热带亚热带植物学报, 27, 491-499.]
[23]	Jiang XL, Zhang WG (2010). Functional diversity and its research method. Acta Ecologica Sinica, 30, 2766-2773.
	[江小雷, 张卫国 (2010). 功能多样性及其研究方法. 生态学报, 30, 2766-2773.]
[24]	Jiang YD, Guo WX, Qin YY, Zhao LL, Wang YJ, Guo WY (2023). Study on aboveground biomass and soil physicochemical properties of different shrub types in Western Ordos. Environment and Development, 35(3), 38-47.
	[姜亚东, 郭威星, 秦玉英, 赵利龙, 王玉婕, 郭威艳 (2023). 西鄂尔多斯不同灌丛类型地上生物量与土壤理化性质研究. 环境与发展, 35(3), 38-47.]
[25]	Kang B, Liu SR, Cai DX, Lu LH (2009). Characteristics of biomass, carbon accumulation and its spatial distribution in Cunninghamia lanceolata forest ecosystem in low subtropical area. Scientia Silvae Sinicae, 45(8), 147-153.
	[康冰, 刘世荣, 蔡道雄, 卢立华 (2009). 南亚热带杉木生态系统生物量和碳素积累及其空间分布特征. 林业科学, 45(8), 147-153.]
[26]	Kaur H (2019). Forms of potassium in soil and their relationship with soil properties—A review. International Journal of Current Microbiology and Applied Sciences, 8, 1580-1586.
[27]	Laliberté E, Lambers H, Burgess TI, Wright SJ (2015). Phosphorus limitation, soil-borne pathogens and the coexistence of plant species in hyperdiverse forests and shrublands. New Phytologist, 206, 507-521. DOI PMID
[28]	Lambers H, Raven JA, Shaver GR, Smith SE (2008). Plant nutrient-acquisition strategies change with soil age. Trends in Ecology & Evolution, 23(2), 95-103. DOI URL
[29]	Lewis SL, Edwards DP, Galbraith D (2015). Increasing human dominance of tropical forests. Science, 349, 827-832. DOI PMID
[30]	Li KJ, Hu ZY, Liu P, Xu ZC (2022). Evaluation on forest biomass and carbon storage in Pearl River Delta in Guangdong Province. Forest Resources Management, (4), 54-60.
	[李康杰, 胡中岳, 刘萍, 徐正春 (2022). 广东省珠三角乔木林生物量碳储量评估. 林业资源管理, (4), 54-60.]
[31]	Li XN, Qi JH, Xu ZX, Lu ZY, Fan ZX, Li ZF, Zhang SB (2024). Recovery of tree species diversity and biomass of degraded evergreen broad-leaved forest in Ailao Mountain, Yunnan Province, southwest China. Journal of Tropical and Subtropical Botany, 32, 500-510.
	[李雪楠, 杞金华, 徐志雄, 鲁志云, 范泽鑫, 栗忠飞, 张树斌 (2024). 云南哀牢山退化常绿阔叶林乔木多样性和生物量的恢复. 热带亚热带植物学报, 32, 500-510.]
[32]	Liu S, Luo Y, Huang YH, Zhou GY (2007). Studies on the community biomass and its allocations of five forest types in Dinghushan Nature Reserve. Ecological Science, 26, 387-393.
	[刘申, 罗艳, 黄钰辉, 周国逸 (2007). 鼎湖山五种植被类型群落生物量及其径级分配特征. 生态科学, 26, 387-393.]
[33]	Liu WJ, Chen SY, Hu FZ, Sha N (2012). Distributions pattern of phosphorus, potassium and influencing factors in the upstream of Shule river basin. Acta Ecologica Sinica, 32, 5429-5437. DOI URL
	[刘文杰, 陈生云, 胡凤祖, 莎娜 (2012). 疏勒河上游土壤磷和钾的分布及其影响因素. 生态学报, 32, 5429-5437.]
[34]	Lu WY, Ye JY, Fu J, Chen ZH, Guo F, Xiang ZF, Li Y, Zhou YL, Chen H, Li P, Li XT (2023). Study on soil physical and chemical properties of pure and mixed forest of Pinus massoniana and Castanopsis hystrix Journal of Green Science and Technology, 25(13), 119-124.
	[陆卫勇, 叶家义, 付军, 陈振华, 郭飞, 项栽芳, 李苑, 周艳玲, 陈虎, 李鹏, 李学团 (2023). 马尾松红锥纯林及混交林土壤理化性质研究. 绿色科技, 25(13), 119-124.]
[35]	Ming AG, Zhang ZJ, Chen HH, Zhang XQ, Tao Y, Su Y (2013). Effects of thinning on the biomass and carbon storage in Pinus massoniana plantation. Scientia Silvae Sinicae, 49(10), 1-6.
	[明安刚, 张治军, 谌红辉, 张显强, 陶怡, 苏勇 (2013). 抚育间伐对马尾松人工林生物量与碳贮量的影响. 林业科学, 49(10), 1-6.]
[36]	Ming AG, Zheng L, Ma J, Tao Y, Lao QX, Lu LH (2015). Biomass, carbon stock and allocation characteristics in Mesua ferrea plantation. Journal of Beijing Forestry University, 37(2), 32-39.
	[明安刚, 郑路, 麻静, 陶怡, 劳庆祥, 卢立华 (2015). 铁力木人工林生物量与碳储量及其分配特征. 北京林业大学学报, 37(2), 32-39.]
[37]	Niu YJ, Squires V, Jentsch A (2023). Risks of China’s increased forest area. Science, 379, 447-448.
[38]	Pan LJ, Xu H, Li YP, Ye XP, Tan S, Wu HJ, He ZL (2023). The effects of configuration modes of different trees on the restoration of species diversity and functional diversity in planted forests. Forestry and Environmental Science, 39(1), 55-64.
	[盘李军, 许涵, 李艳朋, 叶小萍, 谭莎, 吴华俊, 何增丽 (2023). 不同树种配置模式对人工林物种和功能多样性恢复的影响. 林业与环境科学, 39(1), 55-64.]
[39]	Paterno GB, Brambach F, Guerrero-Ramírez N, Zemp DC, Cantillo AF, Camarretta N, Moura CCM, Gailing O, Ballauff J, Polle A, Schlund M, Erasmi S, Iddris NA, Khokthong W, Sundawati L, et al. (2024). Diverse and larger tree islands promote native tree diversity in oil palm landscapes. Science, 386, 795-802. DOI PMID
[40]	Poorter H, Niklas KJ, Reich PB, Oleksyn J, Poot P, Mommer L (2012). Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control. New Phytologist, 193, 30-50. DOI PMID
[41]	Puissant J, Jones B, Goodall T, Mang DN, Blaud A, Gweon HS, Malik A, Jones DL, Clark IM, Hirsch PR, Griffiths R (2019). The pH optimum of soil exoenzymes adapt to long term changes in soil pH. Soil Biology & Biochemistry, 138, 107601. DOI: 10.1016/j.soilbio.2019.107601.
[42]	Schmidt MWI, Torn MS, Abiven S, Dittmar T, Guggenberger G, Janssens IA, Kleber M, Kögel-Knabner I, Lehmann J, Manning DAC, Nannipieri P, Rasse DP, Weiner S, Trumbore SE (2011). Persistence of soil organic matter as an ecosystem property. Nature, 478, 49-56. DOI
[43]	Shi ZM, Tang JC, Cheng RM, Luo D, Liu SR (2015). A review of nitrogen allocation in leaves and factors in its effects. Acta Ecologica Sinica, 35, 5909-5919.
	[史作民, 唐敬超, 程瑞梅, 罗达, 刘世荣 (2015). 植物叶片氮分配及其影响因子研究进展. 生态学报, 35, 5909-5919.]
[44]	Song YT, Wang P, Zhou DW (2011). Methods of measuring plant community functional diversity. Chinese Journal of Ecology, 30, 2053-2059.
	[宋彦涛, 王平, 周道玮 (2011). 植物群落功能多样性计算方法. 生态学杂志, 30, 2053-2059.]
[45]	Su MR, Zhang WQ, Xian WG, Tang HH, Yin ZY, Chen WG, Xu XJ, Pan LJ (2014). Physical and chemical characteristics of soil of Cunninghamia lanceolata conifer-broadleaved mixed forest with different age. Guangdong Forestry Science and Technology, 30(5), 43-47.
	[苏木荣, 张卫强, 冼伟光, 唐洪辉, 殷祚云, 陈伟光, 许雄坚, 盘李军 (2014). 南亚热带不同林龄杉木针阔混交林土壤理化性质分析. 广东林业科技, 30(5), 43-47.]
[46]	Sun J, Zhan TY, Liu M, Zhang ZC, Wang Y, Liu SL, Wu GL, Liu GH, Tsunekawa A (2021). Verification of the biomass transfer hypothesis under moderate grazing across the Tibetan Plateau: a meta-analysis. Plant and Soil, 458, 139-150. DOI
[47]	Tan XP, Tang X, Guo HB, He JH, Wang WR, Wang SH, Nie YX, Zhang W, Ye Q, Shen WJ (2022). Status and strategy of sustainable management of planted forests in South China. Terrestrial Ecosystem and Conservation, 2(5), 102-108.
	[谭向平, 汤萱, 郭惠斌, 何金红, 王玮韧, 王森浩, 聂彦霞, 张炜, 叶清, 申卫军 (2022). 华南地区人工林现状与可持续经营对策. 陆地生态系统与保护学报, 2(5), 102-108.]
[48]	Tobner CM, Paquette A, Gravel D, Reich PB, Williams LJ, Messier C (2016). Functional identity is the main driver of diversity effects in young tree communities. Ecology Letters, 19, 638-647. DOI PMID
[49]	Villéger S, Mason NWH, Mouillot D (2008). New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology, 89, 2290-2301. DOI PMID
[50]	Wang H, Chen L, Song M, Song TQ, Zeng FP, Peng WX, Du H, Su L (2017). Spatial heterogeneity of soil phosphorus and potassium in a mixed evergreen and deciduous broad-leaved forest in karst region of southwest China. Acta Ecologica Sinica, 37, 8285-8293.
	[王华, 陈莉, 宋敏, 宋同清, 曾馥平, 彭晚霞, 杜虎, 苏樑 (2017). 喀斯特常绿落叶阔叶混交林土壤磷钾养分空间异质性. 生态学报, 37, 8285-8293.]
[51]	Wang JL, Wang ZG, Gao Y, Xu WS (2022). Effects of plant allocation patterns on soil nutrients and their stoichiometry in red soil slope. Chinese Science of Soil and Water Conservation, 20(2), 89-98.
	[王家乐, 王志刚, 高阳, 许文盛 (2022). 植物配置模式对红壤坡地土壤养分及其化学计量特征的影响. 中国水土保持科学(中英文), 20(2), 89-98.]
[52]	Wang JR, Shi WK, Wu HF, Hu BA, Cheng XQ, Han HR (2023). Characteristics of leaf functional traits of typical coniferous plantations in northern Shanxi and their relationship with soil factors. Acta Botanica Boreali-Occidentalia Sinica, 43, 835-845.
	[王洁茹, 石文凯, 吴会峰, 胡保安, 程小琴, 韩海荣 (2023). 晋北典型针叶人工林叶功能性状特征及其与土壤因子的关系. 西北植物学报, 43, 835-845.]
[53]	Xiong L, Long CL, Liang S, Wu TH, Liu Q, Liao QL, Xue F (2024). Response of leaf functional traits of woody plants to soil characteristics in karst forests. Journal of Tropical and Subtropical Botany, 32, 310-318.
	[熊玲, 龙翠玲, 梁盛, 吴陶红, 刘奇, 廖全兰, 薛飞 (2024). 喀斯特森林木本植物叶片功能性状对土壤特性的响应. 热带亚热带植物学报, 32, 310-318.]
[54]	Xu QH, Lin LP, Xue CQ, Luo Y, Lei YC (2019). Component specific carbon content and storage of Cinnamomum camphora in Guangdong Province. Journal of Zhejiang A&F University, 36(1), 70-79.
	[徐期瑚, 林丽平, 薛春泉, 罗勇, 雷渊才 (2019). 广东樟树各器官含碳率及碳储量. 浙江农林大学学报, 36(1), 70-79.]
[55]	Xue W, Rillig MC, Yu MF, Hu JN, Huang L, Yu FH (2023). Soil nutrient heterogeneity alters productivity and diversity of experimental plant communities under multiple global change factors. Oikos, 2023, e10189. DOI: 10.1111/oik.10189.
[56]	Yang L, Wang MT, Chen XP, Sun J, Zhong QL, Cheng DL (2020). Relationship between leaf area and leaf biomass of different canopies in subtropical evergreen forest. Acta Ecologica Sinica, 40, 7745-7754.
	[杨力, 王满堂, 陈晓萍, 孙俊, 钟全林, 程栋梁 (2020). 亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究. 生态学报, 40, 7745-7754.]
[57]	Yin QL, Tian TT, Han XH, Xu JS, Chai YF, Mo J, Lei ML, Wang L, Yue M (2019). The relationships between biomass allocation and plant functional trait. Ecological Indicators, 102, 302-308. DOI
[58]	Zhang GJ, Chao L, Zhang WD, Wang SL, Yu X, Zheng WH (2018). Heterogeneity and optimal sample size of soil physicochemical properties in subtropical forest. Chinese Journal of Applied Ecology, 29, 2139-2148.
	[张广杰, 巢林, 张伟东, 汪思龙, 余鑫, 郑文辉 (2018). 亚热带森林土壤理化性质的异质性和最佳取样量. 应用生态学报, 29, 2139-2148.] DOI
[59]	Zhang K, Hou JH, He NP (2017). Leaf functional trait distribution and controlling factors of Pinus tabuliformis. Acta Ecologica Sinica, 37, 736-749.
	[张凯, 侯继华, 何念鹏 (2017). 油松叶功能性状分布特征及其控制因素. 生态学报, 37, 736-749.]
[60]	Zhang Q, Li LJ, Chen YR, Lu HF, Liu BX, Zhao WH, Su RN (2024). Species diversity, biomass, and their relationship in mountainous areas of Beijing. Chinese Environmental Science, 44, 5180-5191.
	[张蔷, 李令军, 陈韵如, 鹿海峰, 刘保献, 赵文慧, 苏日娜 (2024). 北京森林群落物种多样性、生物量及关系分析. 中国环境科学, 44, 5180-5191.]
[61]	Zhang YT, Guo SF, Zhang XP, Shi QD, Zhao FS, Chang SL, Peng JB (2015). Biomass and soil physical-chemical properties of poplar plantations in Karamay. Arid Zone Research, 32, 402-409.
	[张毓涛, 郭树芳, 张新平, 师庆东, 赵福生, 常顺利, 彭佳宾 (2015). 克拉玛依人工林生物量与土壤理化性质. 干旱区研究, 32, 402-409.]
[62]	Zhao JP (2023). Biomass and productivity characteristics of natural Larix gmelinii with different structures. South China Agriculture, 17(4), 101-104.
	[赵吉平 (2023). 不同结构落叶松天然林生物量及生产力特征. 南方农业, 17(4), 101-104.]
[63]	Zhao W, Faust F, Schubert S (2020). Potassium is a potential toxicant for Arabidopsis thaliana under saline conditions. Journal of Plant Natural Nutrition and Soil Science, 183, 455-467.
[64]	Zhao YH (2024). The effect of different thinning intensities on the growth of Pinus koraiensis plantations. South China Agriculture, 18(4), 197-199.
	[赵玉红 (2024). 不同抚育间伐强度对红松人工林生长的影响. 南方农业, 18(4), 197-199.]
[65]	Zhe HY, Luo YY, Zhang HL, Sun FX (2011). The relationship between the root biomass of Fraxinus mandschurica Rupr., Ulums Macrocarpa Hance., Quercus mongolica Fisch. ex Turcz. and soil nutrient and pH value in Daqinggou. Journal of Arid Land Resources and Environment, 25(12), 183-189.
	[折红燕, 罗于洋, 张海龙, 孙福星 (2011). 大青沟水曲柳、蒙古栎、大果榆苗木根系生物量与土壤养分及pH值关系的研究. 干旱区资源与环境, 25(12), 183-189.]
[66]	Zhen GY (2023). Restoration technology and model of natural secondary forest in mountainous areas of eastern Liaoning Province. Forest Science and Technology, (10), 78-81.
	[甄广韵 (2023). 辽东山区天然次生林修复技术与模式. 林业科技通讯, (10), 78-81.]
[67]	Zhu JB, Zhang ZC, Li HQ, Wu QH, He HD, Zhang FW, Li YN (2023). Characteristics and interrelationship of vegetation biomass and soil properties in alpine meadow under grazing gradients. Scientia Sinica Vitae, 53, 1166-1175. DOI URL
	[祝景彬, 张志成, 李红琴, 吴启华, 贺慧丹, 张法伟, 李英年 (2023). 牧压梯度下高寒草甸植被生物量与土壤理化性质变化特征及相互关系. 中国科学: 生命科学, 53, 1166-1175.]

模式 Mode	树种配置 Species composition	修复年限 Recovery year (a)	树高 Tree height (m)	物种数 No. of species	个体数 No. of individuals	胸径 Diameter at breast height (cm)	杉木个体数占比 Proportion of No. of Chinese fir individuals (%)
M1	米老排、桂南木莲、樟树、阴香、长芒杜英、枫香、盆架树 Mytilaria laosensis, Manglietia conifera, Cinnamomum camphora, Cinnamomum burmanni, Elaeocarpus rugosus, Liquidambar formosana, Winchia calophylla	13	6.39 ± 0.65	15.00 ± 4.36	187.00 ± 44.31	8.15 ± 0.23	37.22 ± 7.59
M2	观光木、降香黄檀、樟树、桂南木莲、非洲楝、红锥 Michelia odora, Dalbergia odorifera, Cinnamomum camphora, Mangletia glauca, Khaya senegalensis, Castanopsis hystrix	9	6.93 ± 0.32	16.67 ± 0.58	130.00 ± 2.00	7.31 ± 0.24	51.62 ± 8.80
M3	红锥、非洲楝、黧蒴锥、阴香、杨梅、山杜英、海南红豆 Castanopsis hystrix, Khaya senegalensis, Castanopsis fissa, Cinnamomum burmanni, Myrica rubra, Elaeocarpus sylvestris, Ormosia pinnata	14	6.66 ± 0.67	12.67 ± 4.04	76.00 ± 23.00	8.12 ± 1.09	33.96 ± 10.30
M4	红锥、非洲楝、黧蒴锥、阴香、红花荷、山杜英、木荷、壳菜果 Castanopsis hystrix, Khaya senegalensis, Castanopsis fissa, Cinnamomum burmanni, Rhodoleia championii, Elaeocarpus sylvestris, Schima superba, Mytilaria laosensis	13	7.95 ± 0.33	15.33 ± 5.69	101.33 ± 25.58	10.31 ± 0.08	41.79 ± 5.54
M5	人面子、格木、阴香、降香黄檀、醉香含笑、苦梓、任豆、铁刀木 Dracontomelon duperreanum, Erythrophleum fordii, Cinnamomum burmanni, Dalbergia odorifera, Michelia macclurei, Gmelina hainanensis, Zenia insignis, Cassia siamea	13	8.53 ± 0.28	9.00 ± 0.00	88.00 ± 26.06	10.42 ± 0.80	47.32 ± 7.77
M6	枫香、乌桕、喜树、黧蒴锥、楝叶吴萸、山杜英 Liquidambar formosana, Sapium sebiferum, Camptotheca acuminata, Castanopsis fissa, Evodia glabrifolia, Elaeocarpus sylvestris	8	5.73 ± 0.12	14.67 ± 0.58	126.67 ± 34.70	6.63 ± 0.14	54.30 ± 1.96
M7	凤凰木、西南木荷、醉香含笑、木荷、海南蒲桃、大花紫薇、红花羊蹄甲 Delonix regia, Schima wallichii, Michelia macclurei, Schima superba, Syzygium hainanense, Lagerstroemia speciosa, Bauhinia blakeana	16	6.88 ± 0.06	7.33 ± 2.52	136.33 ± 27.61	9.51 ± 0.50	25.83 ± 2.59
M8	枫香、观光木、醉香含笑、壳菜果、山杜英、黧蒴锥、阴香、格木 Liquidambar formosana, Tsoongiodendron odorum, Michelia macclurei, Mytilaria laosensis, Elaeocarpus sylvestris, Castanopsis fissa, Cinnamomum burmanni, Erythrophleum fordii	15	7.88 ± 0.36	14.33 ± 2.08	262.33 ± 9.29	8.65 ± 0.58	19.57 ± 7.28
M9	毛果青冈、樟树、阴香、香椿、木荷、红锥、醉香含笑 Cyclobalanopsis patelliformis, Cinnamomum camphora, Cinnamomum burmanni, Toona sinensis, Schima superba, Castanopsis hystrix, Michelia macclurei	9	5.78 ± 0.13	13.33 ± 1.53	220.67 ± 37.69	6.79 ± 0.35	65.71 ± 3.36
M10	铁刀木、黧蒴锥、樟树、格木、降香黄檀、醉香含笑、非洲楝、红锥、任豆、桂南木莲 Cassia siamea, Castanopsis fissa, Cinnamomum camphora, Erythrophleum fordii, Dalbergia odorifera, Michelia macclurei, Khaya senegalensis, Castanopsis hystrix, Zenia insignis, Mangletia glauca	14	7.81 ± 0.04	19.33 ± 0.67	150.00 ± 1.53	12.23 ± 1.53	56.06 ± 5.05
M11	红花荷、木棉、醉香含笑、盆架树、火焰树、朱槿 Rhodoleia championii, Bombax malabaricum, Michelia macclurei, Winchia calophylla, Spathodea campanulate, Hibiscus rosa-sinensis	15	10.66 ± 1.03	9.67 ± 2.91	97.33 ± 29.87	5.43 ± 0.15	69.05 ± 8.26

模式 Mode	树种配置 Species composition	修复年限 Recovery year (a)	树高 Tree height (m)	物种数 No. of species	个体数 No. of individuals	胸径 Diameter at breast height (cm)	杉木个体数占比 Proportion of No. of Chinese fir individuals (%)
M1	米老排、桂南木莲、樟树、阴香、长芒杜英、枫香、盆架树 Mytilaria laosensis, Manglietia conifera, Cinnamomum camphora, Cinnamomum burmanni, Elaeocarpus rugosus, Liquidambar formosana, Winchia calophylla	13	6.39 ± 0.65	15.00 ± 4.36	187.00 ± 44.31	8.15 ± 0.23	37.22 ± 7.59
M2	观光木、降香黄檀、樟树、桂南木莲、非洲楝、红锥 Michelia odora, Dalbergia odorifera, Cinnamomum camphora, Mangletia glauca, Khaya senegalensis, Castanopsis hystrix	9	6.93 ± 0.32	16.67 ± 0.58	130.00 ± 2.00	7.31 ± 0.24	51.62 ± 8.80
M3	红锥、非洲楝、黧蒴锥、阴香、杨梅、山杜英、海南红豆 Castanopsis hystrix, Khaya senegalensis, Castanopsis fissa, Cinnamomum burmanni, Myrica rubra, Elaeocarpus sylvestris, Ormosia pinnata	14	6.66 ± 0.67	12.67 ± 4.04	76.00 ± 23.00	8.12 ± 1.09	33.96 ± 10.30
M4	红锥、非洲楝、黧蒴锥、阴香、红花荷、山杜英、木荷、壳菜果 Castanopsis hystrix, Khaya senegalensis, Castanopsis fissa, Cinnamomum burmanni, Rhodoleia championii, Elaeocarpus sylvestris, Schima superba, Mytilaria laosensis	13	7.95 ± 0.33	15.33 ± 5.69	101.33 ± 25.58	10.31 ± 0.08	41.79 ± 5.54
M5	人面子、格木、阴香、降香黄檀、醉香含笑、苦梓、任豆、铁刀木 Dracontomelon duperreanum, Erythrophleum fordii, Cinnamomum burmanni, Dalbergia odorifera, Michelia macclurei, Gmelina hainanensis, Zenia insignis, Cassia siamea	13	8.53 ± 0.28	9.00 ± 0.00	88.00 ± 26.06	10.42 ± 0.80	47.32 ± 7.77
M6	枫香、乌桕、喜树、黧蒴锥、楝叶吴萸、山杜英 Liquidambar formosana, Sapium sebiferum, Camptotheca acuminata, Castanopsis fissa, Evodia glabrifolia, Elaeocarpus sylvestris	8	5.73 ± 0.12	14.67 ± 0.58	126.67 ± 34.70	6.63 ± 0.14	54.30 ± 1.96
M7	凤凰木、西南木荷、醉香含笑、木荷、海南蒲桃、大花紫薇、红花羊蹄甲 Delonix regia, Schima wallichii, Michelia macclurei, Schima superba, Syzygium hainanense, Lagerstroemia speciosa, Bauhinia blakeana	16	6.88 ± 0.06	7.33 ± 2.52	136.33 ± 27.61	9.51 ± 0.50	25.83 ± 2.59
M8	枫香、观光木、醉香含笑、壳菜果、山杜英、黧蒴锥、阴香、格木 Liquidambar formosana, Tsoongiodendron odorum, Michelia macclurei, Mytilaria laosensis, Elaeocarpus sylvestris, Castanopsis fissa, Cinnamomum burmanni, Erythrophleum fordii	15	7.88 ± 0.36	14.33 ± 2.08	262.33 ± 9.29	8.65 ± 0.58	19.57 ± 7.28
M9	毛果青冈、樟树、阴香、香椿、木荷、红锥、醉香含笑 Cyclobalanopsis patelliformis, Cinnamomum camphora, Cinnamomum burmanni, Toona sinensis, Schima superba, Castanopsis hystrix, Michelia macclurei	9	5.78 ± 0.13	13.33 ± 1.53	220.67 ± 37.69	6.79 ± 0.35	65.71 ± 3.36
M10	铁刀木、黧蒴锥、樟树、格木、降香黄檀、醉香含笑、非洲楝、红锥、任豆、桂南木莲 Cassia siamea, Castanopsis fissa, Cinnamomum camphora, Erythrophleum fordii, Dalbergia odorifera, Michelia macclurei, Khaya senegalensis, Castanopsis hystrix, Zenia insignis, Mangletia glauca	14	7.81 ± 0.04	19.33 ± 0.67	150.00 ± 1.53	12.23 ± 1.53	56.06 ± 5.05
M11	红花荷、木棉、醉香含笑、盆架树、火焰树、朱槿 Rhodoleia championii, Bombax malabaricum, Michelia macclurei, Winchia calophylla, Spathodea campanulate, Hibiscus rosa-sinensis	15	10.66 ± 1.03	9.67 ± 2.91	97.33 ± 29.87	5.43 ± 0.15	69.05 ± 8.26

树种 Tree species	树干生物量 Stem biomass (kg)	树枝生物量 Branch biomass (kg)	树叶生物量 Leaf biomass (kg)	树根生物量 Root biomass (kg)
杉木 Cunninghamia lanceolata	W = 0.0556D^2.3579	W = 0.0198D^2.1333	W = 0.0070D^1.5090	W = 0.3366D^1.1952
木荷 Schima superba	W = 0.0310D^2.7140	W = 0.2120D^1.6440	W = 0.0181D^1.9945	W = 0.0310D^2.2582
黧蒴锥 Castanopsis fissa	W = 0.153D^2.2007	W = 0.0117D^2.7605	W = 0.0133D^2.3320	W = 0.0255D^2.3650
山杜英 Elaeocarpus sylvestris	W = 0.0367D^2.6863	W = 0.0058D^3.1688	W = 0.0227D^1.8112	W = 0.0175D^2.5561
樟树 Cinnamomum camphora	W = 0.119122D^2.1521	W = 0.015418D^2.7002	W = 0.095227D^1.4724	W = 0.157486D^1.7732
阔叶树 Broadleaf forest	W = 0.1098D^2.2797	W = 0.0130D^2.6923	W = 0.0144D^2.2200	W = 0.0422D^2.2333

树种 Tree species	树干生物量 Stem biomass (kg)	树枝生物量 Branch biomass (kg)	树叶生物量 Leaf biomass (kg)	树根生物量 Root biomass (kg)
杉木 Cunninghamia lanceolata	W = 0.0556D^2.3579	W = 0.0198D^2.1333	W = 0.0070D^1.5090	W = 0.3366D^1.1952
木荷 Schima superba	W = 0.0310D^2.7140	W = 0.2120D^1.6440	W = 0.0181D^1.9945	W = 0.0310D^2.2582
黧蒴锥 Castanopsis fissa	W = 0.153D^2.2007	W = 0.0117D^2.7605	W = 0.0133D^2.3320	W = 0.0255D^2.3650
山杜英 Elaeocarpus sylvestris	W = 0.0367D^2.6863	W = 0.0058D^3.1688	W = 0.0227D^1.8112	W = 0.0175D^2.5561
樟树 Cinnamomum camphora	W = 0.119122D^2.1521	W = 0.015418D^2.7002	W = 0.095227D^1.4724	W = 0.157486D^1.7732
阔叶树 Broadleaf forest	W = 0.1098D^2.2797	W = 0.0130D^2.6923	W = 0.0144D^2.2200	W = 0.0422D^2.2333