珍稀濒危植物天山梣林龄结构及种群动态

doi:10.17521/cjpe.2024.0438

摘要/Abstract

摘要：

在新疆伊犁小叶白蜡国家级自然保护区对国家二级濒危植物天山梣(也称新疆小叶白蜡, Fraxinus sogdiana)进行大样本调查, 以了解该种群现存状态, 预测种群发展趋势。以径级结构代替年龄结构, 编制种群静态生命表, 分析种群结构, 绘制种群存活曲线、死亡率曲线和生存分析函数曲线, 计算种群动态指数, 采用谱分析探讨种群波动周期。结果表明: 1)喀什河东、喀什河西天山梣种群和种群总体均为增长型年龄结构, 且存活曲线均趋于Deevey-II型; 2)喀什河东种群和种群总体死亡高峰在IV、VII、X龄级, 喀什河西种群死亡高峰仅在V龄级; 3)喀什河东、喀什河西种群和种群总体在忽略外部干扰时的动态变化指数(V_pi)和受外部干扰时的动态变化指数$({{{V}'}_{pi}})$均大于0, 显示均为增长型种群, 喀什河东种群${{{V}'}_{pi}}$更趋近0, 即更能抵御外部随机环境干扰; 4)生存分析显示, 不同生境种群的生存率和累计死亡率均在II-III龄级达到平衡, 喀什河东种群危险率曲线波动幅度较小, 喀什河东相对更适合种群生存; 5)谱分析表明, 天山梣种群的波动主要受其自身生物学特征控制, 喀什河东种群和种群总体还存在大周期内多谐波迭加的小周期波动特征。建议保护和改善天山梣生存环境, 精准抚育第I、II龄级个体, 促进种群的天然更新和发展。

关键词: 天山梣, 濒危植物, 种群结构, 静态生命表, 生存分析, 谱分析

Abstract:

Aims To understand the current state of the population of Fraxinus sogdiana, a second-grade endangered species in China, and predict its development trend, a large sample survey was conducted in the Xinjiang Ili Fraxinus sogdiana National Nature Reserve.

Methods By replacing the age structure with the diameter at breast height structure, a population static life table was compiled and related population structure was analyzed. The survival curve, mortality curve and survival analysis function curve were drawn, and population dynamic index was calculated, and the population fluctuation cycle was explored via using spectral analysis.

Important findings 1) The populations of F. sogdiana on the east and west of the Kax River and total population exhibited growing age structure, with the survival curves towards Deevey-II type. 2) The mortality peaks for the population on the east of the Kax River and total population were in the age classes of IV, VII, and X, while those on the west of Kax River population class had a mortality peak in age class V. 3) The dynamic change index ignoring the external interference (V_pi) and dynamic change index under external interference ${{{V}'}_{pi}}$ of the populations on the east and west of the Kax River and total population were greater than 0, indicating that they were growing-type populations. But the ${{{V}'}_{pi}}$ of population on the east of the Kax River was close to 0, suggesting that it was more resistant to external random environmental interference. 4) Survival analysis revealed that the survival rate and cumulative mortality rate of populations in different habitats reached equilibrium in age classes II-III. The hazard rate curve of the population on the east of the Kax River exhibited relatively smaller fluctuations, indicating that this area is more suitable for the growth of F. sogdiana. 5) Spectral analysis showed that the fluctuations of F. sogdianapopulations are primarily controlled by their own biological characteristics. The population on the east of the Kax River and total population exhibited small cycle fluctuation characteristics with multiple harmonics superimposed during the large cycle. It is recommended protecting and improving habitat of F. sogdiana, accurately nurturing individuals in age classes I and II, and promoting the natural regeneration and development of the population.

Key words: Fraxinus sogdiana, endangered species, population structure, static life table, survival analysis, spectral analysis

贾妍妍, 柳华清, 解欣然, 王博, 张维, 杨允菲. 珍稀濒危植物天山梣林龄结构及种群动态. 植物生态学报, 2025, 49(5): 760-772. DOI: 10.17521/cjpe.2024.0438

JIA Yan-Yan, LIU Hua-Qing, XIE Xin-Ran, WANG Bo, ZHANG Wei, YANG Yun-Fei. Age structure and population dynamics of rare and endangered Fraxinus sogdiana, China. Chinese Journal of Plant Ecology, 2025, 49(5): 760-772. DOI: 10.17521/cjpe.2024.0438

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0438

https://www.plant-ecology.com/CN/Y2025/V49/I5/760

图/表 10

表1 新疆伊犁小叶白蜡国家级自然保护区天山梣种群的生境特征(平均值±标准差)

Table 1 Habitat characteristics of the Fraxinus sogdiana population in the Xinjiang Ili Fraxinus sogdiana National Nature Reserve (mean ± SD)

生境特征 Habitat characteristic	喀什河东 East of the Kax River	喀什河西 West of the Kax River
土壤类型 Soil type	栗钙土 Chestnut soil	灰钙土 Calcareous soil
土壤含水量 Soil water content (%)	24.9 ± 1.4	11.6 ± 1.7
土壤有机质含量 Soil organic matter content (g·kg^-1)	34.9 ± 1.6	23.5 ± 1.3
土层厚度 Soil thickness (cm)	46.8 ± 13.6	21.7 ± 12.2
植被盖度 Vegetation coverage (%)	87.3 ± 6.8	77.3 ± 13.5
郁闭度 Crown density (%)	91.0 ± 7.4	79.0 ± 18.5
种群密度 Population density (m^-2)	0.2 ± 0.0	0.2 ± 0.1
人为干扰强度 Anthropogenic interference intensity	轻度干扰 Slight disturbance	轻度干扰 Slight disturbance
种群胸径 Population DBH (cm)	24.0 ± 19.4	11.0 ± 8.9
种群高度 Population height (m)	11.6 ± 4.9	7.6 ± 5.1
种群冠幅 Population crown width (m)	4.7 ± 2.4	2.5 ± 1.6
主要物种 Dominant species	天山梣、榆树、矮小忍冬、异果小檗、草地早熟禾、短柄草 Fraxinus sogdiana, Ulmus pumila, Lonicera humilis, Berberis heteropoda, Poa pratensis, Brachypodium sylvaticum	天山梣、密叶杨、覆盆子、膜果麻黄、委陵菜、紫菀 Fraxinus sogdiana, Populus talassica, Rubus idaeus, Ephedra przewalskii, Potentilla chinensis, Aster tataricus

表2 天山梣种群样方信息(平均值±标准差)

Table 2 Information on the study sample plots of Fraxinus sogdiana population (mean ± SD)

生境 Habitat	样方 Sample plot	株数 Number	龄级分布 Age class distribution	胸径 DBH (cm)	树高 Tree height (m)	冠幅 Crown width (m)
喀什河东 East of the Kax River	1	82	I-XIII	36.3 ± 26.4	13.5 ± 4.3	6.3 ± 2.4
	2	65	I-VI	17.6 ± 14.0	9.9 ± 5.5	3.4 ± 2.0
	3	70	I-XII	28.9 ± 21.4	12.6 ± 5.0	5.5 ± 3.1
	4	85	I-X	24.7 ± 19.5	14.8 ± 5.3	5.1 ± 2.2
	5	95	I-XIII	25.5 ± 18.6	11.7 ± 4.7	5.4 ± 2.3
	6	102	I-XIII	28.9 ± 23.8	11.6 ± 4.2	5.1 ± 2.2
	7	84	I-VII	21.7 ± 15.4	11.0 ± 4.8	4.2 ± 2.2
	8	100	I-VI	18.0 ± 11.2	10.9 ± 4.7	4.6 ± 2.1
	9	103	I-X	21.9 ± 18.4	10.8 ± 4.7	4.3 ± 2.4
	10	87	I-VIII	17.1 ± 10.6	9.9 ± 4.4	3.3 ± 1.4
喀什河西 West of the Kax River	1	203	I-V	7.1 ± 7.3	5.4 ± 4.0	1.6 ± 0.9
	2	96	I-IV	6.7 ± 5.8	4.7 ± 2.6	1.4 ± 0.7
	3	127	I-IV	11.1 ± 6.0	7.6 ± 4.3	2.4 ± 1.2
	4	79	I-V	16.6 ± 8.6	11.7 ± 4.9	4.0 ± 1.7
	5	73	I-V	19.3 ± 9.7	11.8 ± 4.9	3.1 ± 1.6
	6	57	I-V	15.3 ± 12.3	9.7 ± 6.0	3.1 ± 1.9
	7	65	I-IV	10.3 ± 9.3	7.5 ± 5.7	2.5 ± 1.5
	8	76	I-IV	10.0 ± 7.4	8.2 ± 5.5	2.7 ± 1.7
	9	88	I-IV	10.4 ± 7.3	7.1 ± 4.5	2.7 ± 1.8
	10	65	I-VI	12.5 ± 10.2	7.8 ± 4.7	3.0 ± 2.3

表3 天山梣种群静态生命表

Table 3 Static life table of Fraxinus sogdiana population

生境 Habitat	龄级 Age class (x)	径级 DBH class (cm)	A_x	a_x	l_x	㏑ l_x	d_x	q_x	L_x	T_x	e_x	K_x	S_x
喀什河东 East of the Kax River	I	≤2	60	399	1 000	6.908	170	0.170	915	2 906	2.906	0.187	0.830
	II	2-12	183	331	830	6.721	170	0.205	744	1 991	2.400	0.230	0.795
	III	12-22	263	263	659	6.491	246	0.373	536	1 247	1.892	0.466	0.627
	IV	22-32	165	165	414	6.025	236	0.570	296	711	1.718	0.843	0.430
	V	32-42	71	71	178	5.181	45	0.254	155	415	2.331	0.292	0.746
	VI	42-52	53	53	133	4.889	43	0.321	112	259	1.953	0.387	0.679
	VII	52-62	36	36	90	4.502	48	0.528	66	148	1.639	0.750	0.472
	VIII	62-72	17	17	43	3.752	20	0.471	33	81	1.912	0.636	0.529
	IX	72-82	9	9	23	3.116	0	0.000	23	49	2.167	0.000	1.000
	X	82-92	9	9	23	3.116	15	0.667	15	26	1.167	1.099	0.333
	XI	92-102	3	3	8	2.017	3	0.333	6	11	1.500	0.405	0.667
	XII	102-112	2	2	5	1.612	0	0.000	5	5	1.000	0.000	1.000
	XIII	>112	2	2	5	1.612	-	-	-	-	-	-	-
喀什河西 West of the Kax River	I	≤2	206	321	1 000	6.908	72	0.072	964	2 632	2.632	0.074	0.928
	II	2-12	336	298	928	6.833	72	0.077	893	1 668	1.797	0.080	0.923
	III	12-22	275	275	857	6.753	576	0.673	569	776	0.905	1.117	0.327
	IV	22-32	90	90	280	5.636	215	0.767	173	207	0.739	1.455	0.233
	V	32-42	21	21	65	4.181	62	0.952	34	34	0.524	3.045	0.048
	VI	>42	1	1	3	1.136	-	-	-	-	-	-	-
总体 Total	I	≤2	266	720	1 000	6.908	126	0.126	937	2 785	2.785	0.135	0.874
	II	2-12	519	629	874	6.773	126	0.145	810	1 848	2.115	0.156	0.855
	III	12-22	538	538	747	6.616	393	0.526	551	1 038	1.388	0.747	0.474
	IV	22-32	255	255	354	5.870	226	0.639	241	487	1.375	1.019	0.361
	V	32-42	92	92	128	4.850	53	0.413	101	246	1.924	0.533	0.587
	VI	42-52	54	54	75	4.317	25	0.333	63	144	1.926	0.405	0.667
	VII	52-62	36	36	50	3.912	26	0.528	37	82	1.639	0.750	0.472
	VIII	62-72	17	17	24	3.162	11	0.471	18	45	1.912	0.636	0.529
	IX	72-82	9	9	13	2.526	0	0.000	13	27	2.167	0.000	1.000
	X	82-92	9	9	13	2.526	8	0.667	8	15	1.167	1.099	0.333
	XI	92-102	3	3	4	1.427	1	0.333	3	6	1.500	0.405	0.667
	XII	102-112	2	2	3	1.022	0	0.000	3	3	1.000	0.000	1.000
	XⅢ	>112	2	2	3	1.022	-	-	-	-	-	-	-

表3 天山梣种群静态生命表

Table 3 Static life table of Fraxinus sogdiana population

生境 Habitat	龄级 Age class (x)	径级 DBH class (cm)	A_x	a_x	l_x	㏑ l_x	d_x	q_x	L_x	T_x	e_x	K_x	S_x
喀什河东 East of the Kax River	I	≤2	60	399	1 000	6.908	170	0.170	915	2 906	2.906	0.187	0.830
	II	2-12	183	331	830	6.721	170	0.205	744	1 991	2.400	0.230	0.795
	III	12-22	263	263	659	6.491	246	0.373	536	1 247	1.892	0.466	0.627
	IV	22-32	165	165	414	6.025	236	0.570	296	711	1.718	0.843	0.430
	V	32-42	71	71	178	5.181	45	0.254	155	415	2.331	0.292	0.746
	VI	42-52	53	53	133	4.889	43	0.321	112	259	1.953	0.387	0.679
	VII	52-62	36	36	90	4.502	48	0.528	66	148	1.639	0.750	0.472
	VIII	62-72	17	17	43	3.752	20	0.471	33	81	1.912	0.636	0.529
	IX	72-82	9	9	23	3.116	0	0.000	23	49	2.167	0.000	1.000
	X	82-92	9	9	23	3.116	15	0.667	15	26	1.167	1.099	0.333
	XI	92-102	3	3	8	2.017	3	0.333	6	11	1.500	0.405	0.667
	XII	102-112	2	2	5	1.612	0	0.000	5	5	1.000	0.000	1.000
	XIII	>112	2	2	5	1.612	-	-	-	-	-	-	-
喀什河西 West of the Kax River	I	≤2	206	321	1 000	6.908	72	0.072	964	2 632	2.632	0.074	0.928
	II	2-12	336	298	928	6.833	72	0.077	893	1 668	1.797	0.080	0.923
	III	12-22	275	275	857	6.753	576	0.673	569	776	0.905	1.117	0.327
	IV	22-32	90	90	280	5.636	215	0.767	173	207	0.739	1.455	0.233
	V	32-42	21	21	65	4.181	62	0.952	34	34	0.524	3.045	0.048
	VI	>42	1	1	3	1.136	-	-	-	-	-	-	-
总体 Total	I	≤2	266	720	1 000	6.908	126	0.126	937	2 785	2.785	0.135	0.874
	II	2-12	519	629	874	6.773	126	0.145	810	1 848	2.115	0.156	0.855
	III	12-22	538	538	747	6.616	393	0.526	551	1 038	1.388	0.747	0.474
	IV	22-32	255	255	354	5.870	226	0.639	241	487	1.375	1.019	0.361
	V	32-42	92	92	128	4.850	53	0.413	101	246	1.924	0.533	0.587
	VI	42-52	54	54	75	4.317	25	0.333	63	144	1.926	0.405	0.667
	VII	52-62	36	36	50	3.912	26	0.528	37	82	1.639	0.750	0.472
	VIII	62-72	17	17	24	3.162	11	0.471	18	45	1.912	0.636	0.529
	IX	72-82	9	9	13	2.526	0	0.000	13	27	2.167	0.000	1.000
	X	82-92	9	9	13	2.526	8	0.667	8	15	1.167	1.099	0.333
	XI	92-102	3	3	4	1.427	1	0.333	3	6	1.500	0.405	0.667
	XII	102-112	2	2	3	1.022	0	0.000	3	3	1.000	0.000	1.000
	XⅢ	>112	2	2	3	1.022	-	-	-	-	-	-	-

图1 天山梣种群年龄结构。龄级见表2。

Fig. 1 Age structure of Fraxinus sogdiana population. Age classes see Table 2.

图2 天山梣种群存活和死亡率曲线。A, 喀什河东; B, 喀什河西; C, 总体。龄级见表2。

Fig. 2 Survival and mortality curve of Fraxinus sogdiana population. A, east of the Kax River; B, west of the Kax River; C, total. Age classes see Table 2.

表4 天山梣种群存活曲线的检验模型

Table 4 Test models of survival curve of Fraxinus sogdiana population

生境 Habitat	方程 Equation	R²	F	p	存活曲线类型 Survival curve type
喀什河东 East of the Kax River	N_x = 9.594e^-0.131^x	0.938	166.597	0.000	Deevey-II
喀什河东 East of the Kax River	N_x = 10.574x^-0.584	0.722	28.569	0.000	Deevey-II
喀什河西 West of the Kax River	N_x = 13.153e^-0.305^x	0.659	7.721	0.050	Deevey-II
喀什河西 West of the Kax River	N_x = 9.888x^-0.714	0.452	3.302	0.143	Deevey-II
总体 Total	N_x = 10.785e^-0.171^x	0.935	158.917	0.000	Deevey-II
总体 Total	N_x = 12.217x^-0.764	0.717	27.848	0.000	Deevey-II

表5 天山梣种群动态变化指数

Table 5 Dynamic change index of Fraxinus sogdiana population

动态指数级 Dynamic index class	动态指数值 Dynamic index value (%)
动态指数级 Dynamic index class	喀什河东 East of the Kax River	喀什河西 West of the Kax River	总体 Total
V₁	-67.21	-39.00	-48.75
V₂	-30.42	18.00	-3.53
V₃	37.26	67.00	52.60
V₄	56.97	77.00	63.92
V₅	25.35	95.00	41.30
V₆	32.08	-	33.33
V₇	52.78	-	52.78
V₈	47.06	-	47.06
V₉	0.00	-	0.00
V₁₀	66.67	-	66.67
V₁₁	33.33	-	33.33
V₁₂	0.00	-	0.00
V_pi	18.94	27.51	21.56
$V ′ p i$	0.73	4.59	0.83
P_max	3.85	16.67	3.85

表5 天山梣种群动态变化指数

Table 5 Dynamic change index of Fraxinus sogdiana population

动态指数级 Dynamic index class	动态指数值 Dynamic index value (%)
动态指数级 Dynamic index class	喀什河东 East of the Kax River	喀什河西 West of the Kax River	总体 Total
V₁	-67.21	-39.00	-48.75
V₂	-30.42	18.00	-3.53
V₃	37.26	67.00	52.60
V₄	56.97	77.00	63.92
V₅	25.35	95.00	41.30
V₆	32.08	-	33.33
V₇	52.78	-	52.78
V₈	47.06	-	47.06
V₉	0.00	-	0.00
V₁₀	66.67	-	66.67
V₁₁	33.33	-	33.33
V₁₂	0.00	-	0.00
V_pi	18.94	27.51	21.56
$V ′ p i$	0.73	4.59	0.83
P_max	3.85	16.67	3.85

图3 天山梣种群生存率函数和累积死亡率函数曲线。龄级见表2。

Fig. 3 Survival rate and cumulative mortality rate curve of Fraxinus sogdiana population. Age classes see Table 2.

图4 天山梣种群死亡密度和危险率函数曲线。龄级见表2。

Fig. 4 Mortality density and hazard rate curve of Fraxinus sogdiana population. Age class see Table 2.

表6 天山梣种群的周期性波动

Table 6 Periodic fluctuation of Fraxinus sogdiana population

生境 Habitat	A₀	A₁	A₂	A₃	A₄	A₅	A₆	A₇
喀什河东 East of the Kax River	3.305	1.920	0.798	0.415	0.154	0.168	0.225	0.222
喀什河西 West of the Kax River	4.063	2.216	1.058	1.007	-	-	-	-
总体 Total	3.608	2.256	1.052	0.568	0.319	0.289	0.357	0.355

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