长白山阔叶红松林红松种群年龄结构与数量动态特征

doi:10.17521/cjpe.2021.0498

摘要/Abstract

摘要：

该研究以分布于长白山阔叶红松林内的红松(Pinus koraiensis)种群为对象, 通过编制种群静态生命表, 计算数量动态指数, 绘制存活曲线、死亡率曲线和消失率曲线, 应用4个生存函数并引入谱分析和时间序列预测模型, 分析红松种群年龄结构, 揭示其天然更新过程及未来发展趋势, 以期为野生红松种群的保护和恢复提供科学依据。结果显示: 红松种群数量变化具有阶段性, 幼龄(I-III龄级)和成龄(VII-X龄级)个体数量多, 中龄(IV-VI龄级)和老龄(XI-XIV龄级)个体数量少, 形成明显间断的两个优势年龄分布区。种群存活曲线趋近于Deevey-Ⅲ型, 表明幼龄个体死亡率高。忽略外部干扰时的总体数量动态变化指数大于>0, 表明红松种群为增长型; 考虑未来外部干扰时的种群动态变化指数趋近于0, 结合死亡率和消失率曲线呈现出连续先增后降的复杂动态变化趋势, 可知该种群受外界随机干扰时增长不明显。生存函数分析显示, 红松种群具有前期锐减、中期稳定、后期衰退的特点。谱分析表明红松种群天然更新呈周期性波动。未来2、3、4、5、6、7、8、9、10个龄级时间后, 红松幼、中龄个体数逐渐减少, 而成、老龄个体数量将逐渐增加。幼龄个体死亡率高、生存空间和资源条件有限, 老龄个体生理衰老明显是限制红松种群增长的主要原因。建议加强幼龄个体的抚育工作, 提高其存活率和生存质量; 保护和改善生存环境, 从而促进红松种群的自然更新和恢复。

关键词: 红松, 种群年龄结构, 静态生命表, 数量动态特征, 保护策略

Abstract:

Aims To provide a theoretical basis for the protection and restoration of broadleaved Korean pine forest in Changbai Mountain area, the population age structure and quantitative dynamics of Pinus koraiensis were investigated.

Methods We used field data to form the static life table, estimate the population quantitative indexes, and to fit the survival and mortality curves for identifying the population characteristics of P. koraiensis. We also used the survival functions and spectral analysis to explore its population age structure and to reveal natural regeneration processes and as well as future developmental trend.

Important findings Based on the distribution of diameter, the population of P. koraiensis could be sorted into 14 age classes and combined into four subpopulations, i.e., juvenile (I-III), middle-age (IV-VI), adult (VII-X) and old (XI-XIV) stages. By contrast, the number of individuals in population was higher in stages of young and adult and lower in middle and old stages. However, the survival curve of population was close to Deevey-III type, reflecting that the high mortality occurred in juvenile stage. In addition, external random interference was crucial for whole population dynamics. With and without such interference, the population of P. koraiensis become the constant type (V′_pi was close to 0) and increasing type (V_pi > 0), respectively. Moreover, the survival function analysis showed that the number of individuals of this species was dramatically decreased in early stage, and then relatively stable in middle-age stage and declined again in late stage. Furthermore, spectral analysis for this population indicated that the pattern of natural regeneration exhibited periodic fluctuation along successional process. With population development, the number of young and middle-aged individuals would gradually decrease while the number of adult and old individuals would gradually increase. High mortality of young individuals, limitation of resources, and physiological aging of old trees seem likely to be the important factors that drove population dynamics in this species. We thus suggest that improving habitat condition from silvicultural way may enhance seedling or sapling survival rate and promote their growth, which may benefit the recovery of population for P. koraiensis in this area.

Key words: Pinus koraiensis, population age structure, static life table, quantitative dynamics characteristics, conservation strategy

张金峰, 葛树森, 梁金花, 李俊清. 长白山阔叶红松林红松种群年龄结构与数量动态特征. 植物生态学报, 2022, 46(6): 667-677. DOI: 10.17521/cjpe.2021.0498

ZHANG Jin-Feng, GE Shu-Sen, LIANG Jin-Hua, LI Jun-Qing. Population age structure and dynamics of Pinus koraiensis in a broadleaved Korean pine forest in Changbai Mountain, China. Chinese Journal of Plant Ecology, 2022, 46(6): 667-677. DOI: 10.17521/cjpe.2021.0498

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0498

https://www.plant-ecology.com/CN/Y2022/V46/I6/667

图/表 10

图1 长白山阔叶红松林红松种群样地分布图。

Fig. 1 Map of sampling locations for Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain.

表1 长白山阔叶红松林红松种群静态生命表

Table 1 Static life table of Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain

龄级 Age class (x)	径级 DBH class (cm)	A_x	a_x	l_x	d_x	q_x	L_x	T_x	e_x	lnl_x	K_x	S_x
I	0-5	30	51	1 000	196	0.20	902	4 971	4.97	6.91	0.22	0.78
II	5-10	52	41	804	196	0.24	706	4 069	5.06	6.69	0.28	0.76
III	10-15	41	31	608	137	0.23	539	3 422	5.63	6.41	0.26	0.84
IV	15-20	23	24	471	39	0.08	451	2 882	6.13	6.15	0.09	0.91
V	20-25	14	22	431	39	0.09	412	2 431	5.64	6.07	0.10	0.90
VI	25-30	11	20	392	39	0.10	373	2 020	5.15	5.97	0.11	0.89
VII	30-35	15	18	353	39	0.11	333	1 647	4.67	5.87	0.12	0.88
VIII	35-40	19	16	314	39	0.13	294	1 313	4.19	5.75	0.13	0.87
IX	40-45	19	14	275	39	0.14	255	1 020	3.71	5.61	0.15	0.85
X	45-50	23	12	235	39	0.17	216	765	3.25	5.46	0.18	0.86
XI	50-55	10	10	196	19	0.10	186	549	2.80	5.28	0.11	0.89
XII	55-60	9	9	176	19	0.11	167	363	2.06	5.17	0.12	0.82
XIII	60-65	8	8	157	39	0.25	137	196	1.25	5.06	0.29	0.43
XIV	>65	6	6	118	-	-	59	-	-	4.77	-	-

表2 长白山阔叶红松林红松种群动态变化指数

Table 2 Dynamic indices of Pinus koraiensis population (%) in a broadleaved Korean pine forest in Changbai Mountain

V_I	V_II	V_III	V_IV	V_V	V_VI	V_VII	V_VIII	V_IX	V_X	V_XI	V_XII	V_XIII	V_pi	V′_pi
-42.31	21.15	43.90	39.13	21.43	-26.67	-21.05	0	-17.39	56.52	10.00	11.11	25.00	13.11	0.12

图2 长白山阔叶红松林红松种群死亡率(qx)和消失率(Kx)曲线。龄级同表1。

Fig. 2 Curves of mortality rate (qx) and vanish rate (Kx) for Pinus koraiensis population in broadleaved Korean pine forest in Changbai Mountain. Age class see Table 1.

图3 长白山阔叶红松林红松种群存活曲线。龄级同表1。

Fig. 3 Survival curves of Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain. Age class see Table 1.

表3 长白山阔叶红松林红松种群存活曲线的检验模型

Table 3 Statistical test model for two survival curves of Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain

存活曲线类型 Type	拟合模型 Test model	方程 Equation	R²	F	p
Deevey-II	a_x = a₀e^-bx	a_x = 7.742e^-0.038^x	0.623	18.918	0.001
Deevey-III	a_x = a₀x^-b	a_x = 1.496x^-0.853	0.718	30.505	0.000

图4 长白山阔叶红松林红松种群生存率(Si)和累计死亡率(Fi)曲线。龄级同表1。

Fig. 4 Curves of survival rate (Si) and cumulative mortality rate (Fi) for Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain. Age class see Table 1.

图5 长白山阔叶红松林红松种群死亡密度(fti)与危险率(λti)曲线。龄级同表1。

Fig. 5 Mortality density (fti) and hazard rate curves (λti) of Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain. Age class see Table 1.

表4 长白山阔叶红松林红松种群的周期性波动

Table 4 Periodic fluctuation of Pinus koraiensis population in a broadleaved Korean pine forest in Changbai Mountain

谐波 Resonance spectra	A₀	A₁	A₂	A₃	A₄	A₅	A₆	A₇
振幅值 Amplitude value	3.18	1.73	0.41	0.15	0.23	0.36	0.13	0.02

表5 长白山阔叶红松林红松种群数量动态时间序列预测

Table 5 The prediction number of Pinus koraiensis population by using time sequence analysis in a broadleaved Korean pine forest in Changbai Mountain

龄级 Age class	>初始个体数 Initial number of individuals	M_t
龄级 Age class	>初始个体数 Initial number of individuals	M₂		M₃		M₄		M₅		M₆		M₇		M₈		M₉		M₁₀
I	30
II	52	41
III	41	47		41
IV	23	32		39		37
V	14	19		26		33		32
VI	11	13		18		22		28		29
VII	15	13		14		16		21		26		27
VIII	19	17		14		15		16		21		25		26
IX	19	19		17		16		16		17		20		24		25
X	23	21	16		19		17		17		18		21		24		25
XI	10	17	11		18		17		16		16		17		19		23
XII	9	10	9		15		16		16		15		15		16		18
XIII	8	9	9		13		14		15		15		14		14		15
XIV	8	8	8		9		12		13		14		14		14		14

参考文献 44

[1]	Chen KY, Zhang HR, Zhang B, He YJ (2021). Population structure and dynamic characteristics of typical constructive species in natural secondary forest on the northern slope of Changbai Mountain. Acta Ecologica Sinica, 41, 5142-5152.
	[陈科屹, 张会儒, 张博, 何友均 (2021). 长白山北坡天然次生林典型建群种的种群结构及动态特征. 生态学报, 41, 5142-5152.]
[2]	Chen XD (1998). A study on the method of quantitative analysis for plant population and community structural dynamics. Acta Ecologica Sinica, 18, 214-217.
	[陈晓德 (1998). 植物种群与群落结构动态量化分析方法研究. 生态学报, 18, 214-217.]
[3]	Chhetri PK, Bista R, Cairns DM (2016). Population structure and dynamics of Abies spectabilis at treeline ecotone of Barun Valley, Makalu Barun National Park, Nepal. Acta Ecologica Sinica, 36, 269-274. DOI URL
[4]	Deevey ES Jr (1947). Life tables for natural populations of animals. The Quarterly Review of Biology, 22, 283-314. DOI URL
[5]	Ding JM, Zhang XD, Li GL, Wang J, Huang J, Zhang ZX, Gao PX (2018). Genetic considerations in recovery of endangered plants. Plant Science Journal, 36, 452-458.
	[丁剑敏, 张向东, 李国梁, 王隽, 黄江, 张兆祥, 高浦新 (2018). 濒危植物居群恢复的遗传学考量. 植物科学学报, 36, 452-458.]
[6]	Dong X, Du X, Sun ZH, Gu HY, Chen XW (2020). Spatial pattern and intraspecific association of natural Korean pine population under the influence of habitat gradient. Acta Ecologica Sinica, 40, 5239-5246.
	[董雪, 杜昕, 孙志虎, 谷会岩, 陈祥伟 (2020). 生境梯度影响下的天然红松种群空间格局与种内关联. 生态学报, 40, 5239-5246.]
[7]	Fu JQ (2009). Current situation of natural Korean pine forests in northeast China and their management countermeasures. Journal of Northeast Forestry University, 37(2), 75-78.
	[傅俊卿 (2009). 东北天然红松林资源现状与保护经营对策. 东北林业大学学报, 37(2), 75-78.]
[8]	Ge JP, Guo HY, Chen D, (1990). Study on age structure and spatial pattern of old-growth Korean pine forest in Lesser Xingan Mountain. Journal of Northeast Forestry University, 18(6), 26-32.
	[葛剑平, 郭海燕, 陈动 (1990). 小兴安岭天然红松林种群结构的研究. 东北林业大学学报, 18(6), 26-32.]
[9]	Harcombe PA (1987). Tree life tables. BioScience, 37, 557-568. DOI URL
[10]	Hett JM, Loucks OL (1976). Age structure models of balsam fir and eastern hemlock. Journal of Ecology, 64, 1029-1044. DOI URL
[11]	Jiang ZM, He ZS, Su H, Zhao H, Cai J (2018). Population structure and dynamic characteristics of endangered Syringa pinnatifolia Hemsl. Acta Ecologica Sinica, 38, 2471- 2480.
	[姜在民, 和子森, 宿昊, 赵涵, 蔡靖 (2018). 濒危植物羽叶丁香种群结构与动态特征. 生态学报, 38, 2471-2480.]
[12]	Kang D, Guo YX, Ren CJ, Zhao FZ, Feng YZ, Han XH, Yang GH (2014). Population structure and spatial pattern of main tree species in secondary Betula platyphylla forest in Ziwuling Mountains, China. Scientific Reports, 4, 6873-6881. DOI PMID
[13]	Li JQ, Wang YJ (1986). Wave features of population changes of Pinus koraiensis in natural forest. Chinese Journal of Ecology, 5(5), 1-5.
	[李俊清, 王业蘧 (1986). 天然林内红松种群数量变化的波动性. 生态学杂志, 5(5), 1-5.]
[14]	Li JQ, Zhu N (1990). Structure of Korean pine and its dynamics. Chinese Journal of Ecology, 9(1), 6-10.
	[李俊清, 祝宁 (1990). 红松的种群结构与动态过程. 生态学杂志, 9(1), 6-10.]
[15]	Li YL, Yang H, Kang XG, Qiu S (2014). Population structures and dynamics of Abies nephrolepis and Picea koraiensis in the Changbai Mountains of northeastern China. Journal of Beijing Forestry University, 36(3), 18-25.
	[李艳丽, 杨华, 亢新刚, 邱实 (2014). 长白山云冷杉种群结构和动态分析. 北京林业大学学报, 36(3), 18-25.]
[16]	Liu HY, Jin XL, Shen SY, Zhang RQ (2012). Population quantitative characteristics and dynamics of rare and endangered plant Davidia involucrata in Hunan Province. Acta Ecologica Sinica, 32, 7738-7746. DOI URL
	[刘海洋, 金晓玲, 沈守云, 张日清 (2012). 湖南珍稀濒危植物--珙桐种群数量动态. 生态学报, 32, 7738-7746.]
[17]	Liu S, Hou MM, Liao JX, Wang JW, Xiao C, Fan XH (2016). Local biotic neighborhood and habitat heterogeneity drive tree seedling survival dynamics in a temperate forest. Chinese Journal of Applied and Environmental Biology, 22, 639-645.
	[刘帅, 侯嫚嫚, 廖嘉星, 王均伟, 肖翠, 范秀华 (2016). 生物邻体和生境异质性共同驱动乔木幼苗存活动态. 应用与环境生物学报, 22, 639-645.]
[18]	Ma JL, Zhuang LW, Chen D, Li JW (1992). Geographic distribution of Pinus koraiensis in the world. Journal of Northeast Forestry University, 20(5), 40-48.
	[马建路, 庄丽文, 陈动, 李景文 (1992). 红松的地理分布. 东北林业大学学报, 20(5), 40-48.]
[19]	Ma L, Li JQ (2019). Systematic conservation planning optimization for the broad-leaved Korean pine mixed forest conservation network system in the Changbai Mountain Region, China. Acta Ecologica Sinica, 39, 8547-8555.
	[马琳, 李俊清 (2019). 基于系统保护规划的长白山阔叶红松林保护网络优化研究. 生态学报, 39, 8547-8555.]
[20]	Meng LJ, Yao J, Qin JH, Fan CY, Zhang CY, Zhao XH (2018). Drivers of composition and density pattern of tree seedlings in a secondary mixed conifer and broad-leaved forest, Jiaohe, Jilin, China. Chinese Journal of Plant Ecology, 42, 653-662. DOI URL
	[孟令君, 姚杰, 秦江环, 范春雨, 张春雨, 赵秀海 (2018). 吉林蛟河针阔混交林乔木幼苗组成及其密度格局影响因素. 植物生态学报, 42, 653-662.] DOI
[21]	Nunes YRF, Petrere M Jr (2012). Structure and dynamics of a Cariniana estrellensis (Lecythidaceae) population in a fragment of Atlantic Forest in Minas Gerais, Brazil. Rodriguésia, 63, 257-267. DOI URL
[22]	Qin AL, Ma FQ, Xu GX, Shi ZM, Chen QY (2020). Population structure and dynamic characteristics of a rare and endangered tree species Michelia wilsonii Finet et Gagn. Acta Ecologica Sinica, 40, 4445-4454.
	[秦爱丽, 马凡强, 许格希, 史作民, 陈其勇 (2020). 珍稀濒危树种峨眉含笑种群结构与动态特征. 生态学报, 40, 4445-4454.]
[23]	Qiu H, Shu H, Wu ZF, Li ML, Yin Z, Zhang CY, Zhao XH (2020). Influencing factors of composition and abundance pattern of tree seedlings in broad-leaved Korean pine (Pinus koraiensis) mixed forest, Changbai Mountain, China. Acta Ecologica Sinica, 40, 2049-2056.
	[邱华, 舒皓, 吴兆飞, 李明鲁, 殷正, 张春雨, 赵秀海 (2020). 长白山阔叶红松林乔木幼苗组成及多度格局的影响因素. 生态学报, 40, 2049-2056.]
[24]	Tan YB, Zhan CA, Xiao ZX, Yang HD, Peng JH, Wu KS (2010). Population structure and dynamic characteristics of Machilus chinensis in Nan'ao Island, Guangdong Province. Chinese Journal of Ecology, 29, 1901-1906.
	[谭一波, 詹潮安, 肖泽鑫, 杨海东, 彭剑华, 吴凯胜 (2010). 广东南澳岛中华楠种群结构及动态特征. 生态学杂志, 29, 1901-1906.]
[25]	Tasiken H, Cai HY, Jin GZ (2021). Effects of canopy structure on productivity in a typical mixed broadleaved Korean pine forest. Chinese Journal of Plant Ecology, 45, 38-50. DOI URL
	[哈努拉·塔斯肯, 蔡慧颖, 金光泽, (2021). 树冠结构对典型阔叶红松林生产力的影响. 植物生态学报, 45, 38- 50.]
[26]	Wang T, Liang Y, Ren HB, Yu D, Ni J, Ma KP (2004). Age structure of Picea schrenkiana forest along an altitudinal gradient in the central Tianshan Mountains, northwestern China. Forest Ecology and Management, 196, 267-274. DOI URL
[27]	Wu HQ (1989). Dynamics of Korean pine population in birch forests of the Changbai Mountain. Chinese Journal of Ecology, 8(5), 16-18.
	[乌弘奇 (1989). 长白山白桦林中红松种群动态的研究. 生态学杂志, 8(5), 16-18.]
[28]	Wu JX, Zhang XM, Deng CZ, Liu GJ, Yan HL (2010). Characteristics and dynamics of Populus euphratica population at lower reaches of Tarim River. Journal of Arid Land Resources and Environment, 24, 164-169.
	[吴俊侠, 张希明, 邓潮州, 刘国军, 阎海龙 (2010). 塔里木河下游胡杨种群特征与动态分析. 干旱区资源与环境, 24, 164- 169.]
[29]	Wu QC, Zang FQ, Li CC, Ma Y, Gao Y, Zheng YQ, Zang DK (2021). Population structure and dynamics of endangered Populus wulianensis. Acta Ecologica Sinica, 41, 5016- 5025.
	[吴其超, 臧凤岐, 李呈呈, 马燕, 高燕, 郑勇奇, 臧德奎 (2021). 濒危树种五莲杨种群结构与动态特征. 生态学报, 41, 5016-5025.]
[30]	Wu YG, Han JX (1988). A spectral analysis of the population dynamics of Korean pine in the mixed broad-leaved Pinus koraiensis forest. Chinese Journal of Ecology, 7(1), 19-23.
	[伍业钢, 韩进轩 (1988). 阔叶红松林红松种群动态的谱分析. 生态学杂志, 7(1), 19-23.]
[31]	Xiao C, Liu S, Huang Z, Fan Y, Wang JW, Zhao XH, Tang JY (2015). Effects of biotic and abiotic factors on tree seedling survival in a broad leaved Korean pine (Pinus koraiensis) mixed forest on Changbai Mountain. Acta Ecologica Sinica, 35, 6557-6565.
	[肖翠, 刘帅, 黄珍, 樊莹, 王均伟, 赵秀海, 唐景毅 (2015). 长白山阔叶红松林中影响乔木幼苗存活的关键因素. 生态学报, 35, 6557-6565.]
[32]	Xiao YA, He P, Li XH, Deng HP (2004). Study on numeric dynamics of natural populations of the endangered species Disanthus cercidifolius var.longipes. Acta Phytoecologica Sinica, 28, 252-257.
	[肖宜安, 何平, 李晓红, 邓洪平 (2004). 濒危植物长柄双花木自然种群数量动态. 植物生态学报, 28, 252-257.] DOI
[33]	Xu WZ, Liu QJ, Meng SW, Zhou G (2018). Long-term monitoring of tree population dynamics of broad-leaved Korean pine forest in Changbai Mountains, China. Chinese Journal of Applied Ecology, 29, 3159-3166.
	[徐玮泽, 刘琪璟, 孟盛旺, 周光 (2018). 长白山阔叶红松林树木种群动态的长期监测. 应用生态学报, 29, 3159-3166.] DOI
[34]	Yang CP, Wei L, Jiang J, Liu GF, Zhao GY (2005). Analysis of genetic diversity for nineteen populations of Pinus sibirica Du Tour with technique of ISSR. Journal of Northeast Forestry University, 33(1), 1-3.
	[杨传平, 魏利, 姜静, 刘桂丰, 赵光仪 (2005). 应用ISSR PCR对西伯利亚红松19个种源的遗传多样性分析. 东北林业大学学报, 33(1), 1-3.]
[35]	Yang FX, Wang SQ, Xu HG, Li BQ (1991). The theory of survival analysis and its application to life table. Acta Ecologica Sinica, 11, 153-158.
	[杨凤翔, 王顺庆, 徐海根, 李邦庆 (1991). 生存分析理论及其在研究生命表中的应用. 生态学报, 11, 153-158.]
[36]	Yang LR, Zhang ZL, Yun Y, Yan WP, Chen X, Zhang L, Zheng DJ, Chen JL (2018). The population structure and dynamics of Dracaena cambodiana, an endangered tree on Hainan Island. Acta Ecologica Sinica, 38, 2802-2815.
	[杨立荣, 张治礼, 云勇, 严武平, 陈宣, 张浪, 郑道君, 陈加利 (2018). 濒危植物海南龙血树的种群结构与动态. 生态学报, 38, 2802-2815.]
[37]	Yu DP, Zhou WM, Bao Y, Qi L, Zhou L, Dai LM (2015). Forest management of Korean pine and broadleaf mixed forest in Northeast China since the implementation of Natural Forest Protection Project. Acta Ecologica Sinica, 35, 10- 17. DOI URL
	[于大炮, 周旺明, 包也, 齐麟, 周莉, 代力民 (2015). 天保工程实施以来东北阔叶红松林的可持续经营. 生态学报, 35, 10-17.]
[38]	Yu DP, Zhou WM, Zhou L, Dai LM (2019). Exploring the history of the management theory and technology of broad leaved Korean pine (Pinus koraiensis Sieb. et Zucc.) forest in Changbai Mountain Region, Northeast China. Chinese Journal of Applied Ecology, 30, 1426-1434.
	[于大炮, 周旺明, 周莉, 代力民 (2019). 长白山区阔叶红松林经营历史与研究历程. 应用生态学报, 30, 1426-1434.] DOI
[39]	Zhang Q, Fan SH, Shen HL (2003). Research and development on the growth environment of the young tree of Pinus koraiensis in Pinus koraiensis-broadleaved mixed forest. Forest Research, 16, 216-224.
	[张群, 范少辉, 沈海龙 (2003). 红松混交林中红松幼树生长环境的研究进展及展望. 林业科学研究, 16, 216-224.]
[40]	Zhang XP, Yu LZ, Yang XY, Huang JQ, Yin Y (2021). Population structure and dynamics of Pinus koraiensis seedlings regenerated from seeds in a montane region of eastern Liaoning Province, China. Chinese Journal of Applied Ecology, 33, 289-296.
	[张晓鹏, 于立忠, 杨晓燕, 黄佳琪, 殷有 (2021). 辽东山区天然更新红松幼苗种群结构与动态. 应用生态学报, 33, 289-296.]
[41]	Zhang Y, Yi XM, Wang YX, Ji LZ, Wu PL (2015). Impact of tree harvesting on the population structure and dynamics of Pinus koraiensis (Pinaceae). Acta Ecologica Sinica, 35, 38-45.
	[张悦, 易雪梅, 王远遐, 姬兰柱, 吴培莉 (2015). 采伐对红松种群结构与动态的影响. 生态学报, 35, 38-45.]
[42]	Zhang YF, Li DW, Wang M, Liu P (2015). Population structure and dynamics of Juniperus rigida in different regions of Loess Plateau. Scientia Silvae Sinicae, 51(2), 1-10.
	[张亚芳, 李登武, 王梅, 刘盼 (2015). 黄土高原不同地区杜松种群结构与动态. 林业科学, 51(2), 1-10.]
[43]	Zhao Y, Liu JQ, Chen XL, Yang MM, Cao JH, Qi R, Cao XW (2020). Population structure characteristics of Picea purpurea in the upstream of Taohe River. Chinese Journal of Plant Ecology, 44, 266-276. DOI
	[赵阳, 刘锦乾, 陈学龙, 杨萌萌, 曹家豪, 齐瑞, 曹秀文 (2020). 洮河上游紫果云杉种群结构特征. 植物生态学报, 44, 266-276.] DOI
[44]	Zhou SX, Peng YS, Zhan XH, Zhang L, Liang TJ, Liu XP, Pan FX (2020). Population structure and numeric dynamics of Pinus taiwanensis in Lushan Mountain of Jiangxi Province. Guihaia, 40, 247-254.
	[周赛霞, 彭焱松, 詹选怀, 张丽, 梁同军, 刘向平, 潘福兴 (2020). 庐山黄山松种群结构及数量动态研究. 广西植物, 40, 247-254.]