浙江象山沿海岛屿苔藓植物种-面积关系及物种丰富度影响因素

doi:10.17521/cjpe.2024.0122

摘要/Abstract

摘要：

种-面积关系描述了当取样面积增加时, 物种丰富度也随之变化的规律, 是群落生态学研究的核心问题之一。小岛屿效应(SIE)是种-面积关系的一种特殊现象, 是岛屿生物地理学和生物多样性研究理论框架的重要元素。沿海岛屿是一种实际存在的岛屿系统, 然而以苔藓植物为研究对象的SIE研究还比较少。该研究以浙江省象山地区沿海岛屿为研究区域, 调查了43个沿海岛屿苔藓植物的物种组成。采用了4种连续分段回归模型和2种对照模型, 分别检测了不同类群的苔藓植物是否存在SIE, 并对其产生的阈值大小进行比较。使用广义线性混合效应模型和Mantel检验分析探讨了包括景观特征、人为干扰等环境因子对海岛苔藓物种丰富度的影响, 以及不同类群(苔类、藓类、顶蒴藓类、侧蒴藓类)对海岛环境因子的响应。结果显示, 象山地区沿海43个岛屿共记录苔藓植物209种, 隶属于46科85属。苔藓植物所有类群均存在SIE, 双拐点模型被认为是最优模型, 其面积阈值分布在0.098-4.419 km²之间, 其中, 苔类高于藓类, 顶蒴藓类高于侧蒴藓类。广义线性混合效应模型结果表明, 苔藓植物物种丰富度随着岛屿面积、生境丰富度、形状指数、海岸线长度、人口数量、交通频率增加而增多, 随着隔离程度增大而减少, 生境丰富度是影响苔藓植物物种丰富度的最主要因素。一般而言, 资源需求较低, 传播能力更强的类群(藓类), 其小岛屿面积的阈值较低。远离大陆的几个大岛屿由于植被覆盖率低, 生境丰富度低, 不适合侧蒴藓类生长, 导致其小岛屿面积阈值低于顶蒴藓类。因此, 生境类型的多样性保护是维持物种多样性的重要保障。

关键词: 岛屿生物地理学, 种-面积关系, 小岛屿效应, 生境异质性, 面积阈值, 广义线性混合效应模型

Abstract:

Aims The species-area relationship describes a fundamental pattern in community ecology, illustrating how species richness changes as the sampled area increases. It is a core issue in community ecology research. The island biogeography theory highlights a specific aspect of the species-area relationship known as the small island effect (SIE), which is a key component of the theoretical framework of island biogeography and biodiversity studies. Coastal islands represent a tangible and existing island system, yet there is relatively limited research on the SIE, especially concerning bryophytes.

Methods This study investigated the species composition of bryophytes on 43 coastal islands in the Xiangshan region of Zhejiang. We utilized four segmented regression models and two control models to examine the SIE across different bryophyte taxa. We compared the thresholds at which it occurs. Using generalized linear mixed-effect models and Mantel tests, we explored the impact of environmental factors, such as landscape characteristics and human disturbances, on bryophyte species richness on these maritime islands. Furthermore, we analyzed the response of different bryophyte taxa (liverworts, mosses, acrocarpous mosses, and pleurocarpous mosses) to environmental factors in these island environments.

Important findings Our study documented 209 bryophyte species belonging to 46 families and 85 genera across the 43 coastal islands in the Xiangshan region. All bryophytes taxa exhibited the SIE, with the double-breakpoint model identified as the optimal model. The area thresholds ranged from 0.098 to 4.419 km², with mosses showing higher thresholds than liverworts, and acrocarpous mosses higher than pleurocarpous mosses. Moreover, the richness of bryophytes increased with the island area, habitat richness, shape index, coastline length, population size, and traffic frequency, while it decreased with increasing isolation. Habitat richness was identified as the primary factor influencing bryophytes richness. Generally, taxa with lower resource requirements and higher dispersal abilities (such as mosses) exhibited lower SIE area thresholds. Larger islands with greater distances from the mainland, characterized by lower vegetation cover and habitat richness, were less suitable for the growth of pleurocarpous mosses, resulting in lower SIE area thresholds than acrocarpous mosses. Therefore, the conservation of habitat diversity is crucial for maintaining species diversity.

Key words: island biogeography, species-area relationship, small island effect, habitat heterogeneity, area threshold, generalized linear mixed models

杨子龙, 罗光宇, 雷富民, 余静柔, 蔡圣轩, 于晶. 浙江象山沿海岛屿苔藓植物种-面积关系及物种丰富度影响因素. 植物生态学报, 2024, 48(11): 1486-1500. DOI: 10.17521/cjpe.2024.0122

YANG Zi-Long, LUO Guang-Yu, LEI Fu-Min, YU Jing-Rou, CAI Sheng-Xuan, YU Jing. Species-area relationship of bryophytes and its influencing factors on the coastal islands in Xiangshan, Zhejiang, China. Chinese Journal of Plant Ecology, 2024, 48(11): 1486-1500. DOI: 10.17521/cjpe.2024.0122

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https://www.plant-ecology.com/CN/Y2024/V48/I11/1486

图/表 9

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模型 Model	方程式 Equation
1	S = C + Z₁lnA (lnA ≤ T) + \|Z₁T + Z₂ (lnA - T)\|(lnA > T)
2	S = C + Z₁lnA (lnA ≤ T₁) + \|Z₁T₁ + Z₂ (lnA - T₁)\|(T₂ ≥ lnA > T₁) + \|Z₂ (T₂ - T₁) + Z₃ (lnA - T₂)\|(lnA > T₂)
3	S = C + \|Z₂ (lnA - T)\|(lnA > T)
4	S = C + Z₂ (lnA - T₁)\|(T₂ ≥ lnA > T₁) + \|Z₂ (T₂ - T₁) + Z₃ (lnA - T₂)\|(A > T₂)
5	S = C + Z₁ logA
6	S = C

模型 Model	方程式 Equation
1	S = C + Z₁lnA (lnA ≤ T) + \|Z₁T + Z₂ (lnA - T)\|(lnA > T)
2	S = C + Z₁lnA (lnA ≤ T₁) + \|Z₁T₁ + Z₂ (lnA - T₁)\|(T₂ ≥ lnA > T₁) + \|Z₂ (T₂ - T₁) + Z₃ (lnA - T₂)\|(lnA > T₂)
3	S = C + \|Z₂ (lnA - T)\|(lnA > T)
4	S = C + Z₂ (lnA - T₁)\|(T₂ ≥ lnA > T₁) + \|Z₂ (T₂ - T₁) + Z₃ (lnA - T₂)\|(A > T₂)
5	S = C + Z₁ logA
6	S = C

岛屿编号 Island No.	面积 Area (km²)	生境丰富度 Habitat richness	面积×生境丰富度 Area × habitat richness	海拔 Altitude (m)	形状指数 Shape index	岛屿被海水的隔离程度 Seawater isolation	人口 Population	岛屿的海岸线长度 Coastline length (m)	交通频率 Frequency of traffic
1	0.11	2	0.22	24	2.60	0.96	0	3 061.03	1
2	0.06	3	0.18	9	2.67	0.97	0	2 314.93	1
3	0.003	1	0.003	15	1.49	0.97	0	287.37	1
4	0.10	3	0.30	29	1.59	0.98	0	1 787.68	1
5	0.01	2	0.02	6	1.73	0.98	0	615.10	1
6	0.06	3	0.18	16	2.04	0.97	0	1 769.54	1
7	0.02	3	0.06	10	1.35	0.92	0	675.56	1
8	0.01	4	0.04	7	0.90	0.91	0	317.71	1
9	0.30	5	1.50	16	1.80	0.60	0	3 502.88	1
10	0.84	5	4.20	7	1.53	0.88	0	4 982.03	1
11	0.004	7	0.03	4	1.28	0.83	0	277.98	1
12	0.83	7	5.81	11	2.33	0.77	0	7 551.65	2
13	0.14	13	1.82	14	1.34	0.42	310	1 779.01	4
14	0.03	6	0.18	4	1.06	0.43	10	652.24	3
15	0.06	5	0.30	10	1.62	0.94	0	1 409.28	1
16	0.11	1	0.11	9	1.75	0.89	0	2 056.60	1
17	2.02	18	36.36	103	1.66	0.49	380 6	8 369.43	5
18	0.08	1	0.08	17	1.30	0.88	0	1 303.04	1
19	0.02	2	0.04	8	1.24	0.77	0	620.52	1
20	0.02	3	0.06	33	1.24	0.76	0	619.50	2
21	0.01	2	0.02	5	1.11	0.76	0	393.85	1
22	0.03	1	0.03	3	1.86	0.77	0	1 145.30	1
23	0.003	1	0.003	4	1.24	0.81	0	235.16	1
24	0.02	3	0.06	4	1.28	0.44	0	643.89	1
25	0.10	5	0.50	34	1.60	0.45	5	1 789.21	3
26	0.03	3	0.09	10	1.50	0.78	0	924.41	1
27	0.28	10	2.80	32	1.14	0.24	298	2 138.81	4
28	0.40	13	5.20	35	1.11	0.25	707	2 489.83	4
29	4.42	14	61.88	33	1.82	0.58	120	13 535.31	4
30	11.30	21	237.30	100	4.14	0.85	100	49 331.42	4
31	0.79	15	11.85	41	1.19	0.27	707	3 756.32	4
32	0.01	4	0.04	9	1.63	0.26	0	577.22	1
33	0.82	8	6.56	96	1.47	0.28	20	4 705.48	3
34	1.48	8	11.84	26	1.22	0.27	350	5 248.01	4
35	0.04	11	0.44	57	1.11	0.51	0	788.13	3
36	0.04	11	0.44	61	1.24	0.53	0	876.24	3
37	0.08	14	1.12	48	1.51	0.26	30	1 510.69	3
38	0.02	4	0.08	21	1.61	0.25	0	806.15	2
39	0.02	6	0.12	98	1.40	0.55	0	700.93	2
40	0.21	11	2.31	31	1.46	0.27	6	2 366.80	3
41	47.52	25	118 8.00	129	1.39	0.29	20 000	33 898.23	5
42	89.70	25	224 2.50	117	1.73	0.10	35 000	58 082.13	5
43	16.92	18	304.56	54	1.82	0.58	1 026	26 559.84	4

岛屿编号 Island No.	面积 Area (km²)	生境丰富度 Habitat richness	面积×生境丰富度 Area × habitat richness	海拔 Altitude (m)	形状指数 Shape index	岛屿被海水的隔离程度 Seawater isolation	人口 Population	岛屿的海岸线长度 Coastline length (m)	交通频率 Frequency of traffic
1	0.11	2	0.22	24	2.60	0.96	0	3 061.03	1
2	0.06	3	0.18	9	2.67	0.97	0	2 314.93	1
3	0.003	1	0.003	15	1.49	0.97	0	287.37	1
4	0.10	3	0.30	29	1.59	0.98	0	1 787.68	1
5	0.01	2	0.02	6	1.73	0.98	0	615.10	1
6	0.06	3	0.18	16	2.04	0.97	0	1 769.54	1
7	0.02	3	0.06	10	1.35	0.92	0	675.56	1
8	0.01	4	0.04	7	0.90	0.91	0	317.71	1
9	0.30	5	1.50	16	1.80	0.60	0	3 502.88	1
10	0.84	5	4.20	7	1.53	0.88	0	4 982.03	1
11	0.004	7	0.03	4	1.28	0.83	0	277.98	1
12	0.83	7	5.81	11	2.33	0.77	0	7 551.65	2
13	0.14	13	1.82	14	1.34	0.42	310	1 779.01	4
14	0.03	6	0.18	4	1.06	0.43	10	652.24	3
15	0.06	5	0.30	10	1.62	0.94	0	1 409.28	1
16	0.11	1	0.11	9	1.75	0.89	0	2 056.60	1
17	2.02	18	36.36	103	1.66	0.49	380 6	8 369.43	5
18	0.08	1	0.08	17	1.30	0.88	0	1 303.04	1
19	0.02	2	0.04	8	1.24	0.77	0	620.52	1
20	0.02	3	0.06	33	1.24	0.76	0	619.50	2
21	0.01	2	0.02	5	1.11	0.76	0	393.85	1
22	0.03	1	0.03	3	1.86	0.77	0	1 145.30	1
23	0.003	1	0.003	4	1.24	0.81	0	235.16	1
24	0.02	3	0.06	4	1.28	0.44	0	643.89	1
25	0.10	5	0.50	34	1.60	0.45	5	1 789.21	3
26	0.03	3	0.09	10	1.50	0.78	0	924.41	1
27	0.28	10	2.80	32	1.14	0.24	298	2 138.81	4
28	0.40	13	5.20	35	1.11	0.25	707	2 489.83	4
29	4.42	14	61.88	33	1.82	0.58	120	13 535.31	4
30	11.30	21	237.30	100	4.14	0.85	100	49 331.42	4
31	0.79	15	11.85	41	1.19	0.27	707	3 756.32	4
32	0.01	4	0.04	9	1.63	0.26	0	577.22	1
33	0.82	8	6.56	96	1.47	0.28	20	4 705.48	3
34	1.48	8	11.84	26	1.22	0.27	350	5 248.01	4
35	0.04	11	0.44	57	1.11	0.51	0	788.13	3
36	0.04	11	0.44	61	1.24	0.53	0	876.24	3
37	0.08	14	1.12	48	1.51	0.26	30	1 510.69	3
38	0.02	4	0.08	21	1.61	0.25	0	806.15	2
39	0.02	6	0.12	98	1.40	0.55	0	700.93	2
40	0.21	11	2.31	31	1.46	0.27	6	2 366.80	3
41	47.52	25	118 8.00	129	1.39	0.29	20 000	33 898.23	5
42	89.70	25	224 2.50	117	1.73	0.10	35 000	58 082.13	5
43	16.92	18	304.56	54	1.82	0.58	1 026	26 559.84	4

类群 Group	模型 Model	参数 Parameter						模型比较 Model comparison
类群 Group	模型 Model	C	Z₁	Z₂	Z₃	T₁	T₂	R²	AIC_c	ΔAIC_c
苔藓植物 Bryophytes	1	43.97	8.08	81.57	-	25.43	-	0.87	344.04	1.67
	2	17.37	6.04	24.28	81.69	1.01	34.67	0.89	342.37	0.00
	3	17.36	0.00	11.58	-	0.51	-	0.81	359.73	17.36
	4	37.10	0.00	16.44	81.66	0.07	30.14	0.88	343.40	1.03
	5	54.26	11.43	-	-	-	-	0.72	374.28	31.91
	6	39.96	-	-	-	-	-	0.00	427.63	85.26
藓类植物 Mosses	1	41.04	7.58	72.45	-	27.01	-	0.86	337.61	1.59
	2	34.58	5.67	14.95	72.18	0.92	37.19	0.88	336.02	0.00
	3	13.46	0.00	17.00	-	0.18	-	0.81	350.05	14.03
	4	12.09	0.00	10.70	72.09	0.06	32.01	0.87	337.13	1.11
	5	49.53	10.36	-	-	-	-	0.73	363.00	26.98
	6	36.67	-	-	-	-	-	0.00	418.38	82.36
苔类植物 Liverworts	1	2.89	0.48	8.95	-	15.58	-	0.85	164.48	2.24
	2	2.56	0.30	2.30	9.46	4.42	22.33	0.86	162.24	0.00
	3	1.54	0.00	6.89	-	7.07	-	0.82	168.45	6.21
	4	1.09	0.00	0.77	9.46	0.08	18.84	0.86	163.50	1.26
	5	4.72	1.08	-	-	-	-	0.51	209.95	47.71
	6	3.29	-	-	-	-	-	0.00	238.99	76.75
顶蒴藓类植物 Acrocarpous mosses	1	24.06	3.61	13.65	-	1.49	-	0.88	287.54	0.45
	2	24.56	3.75	10.64	39.42	1.23	43.21	0.90	287.09	0.00
	3	9.97	0.00	9.23	-	0.10	-	0.83	302.17	15.08
	4	4.66	0.00	3.77	13.53	0.01	1.49	0.88	290.06	2.97
	5	33.07	6.41	-	-	-	-	0.77	312.51	25.42
	6	25.10	-	-	-	-	-	0.00	374.89	87.80
侧蒴藓类植物 Pleurocarpous mosses	1	12.45	2.64	32.93	-	26.74	-	0.79	280.45	1.11
	2	5.95	0.91	3.86	32.94	0.10	31.06	0.81	279.34	0.00
	3	2.89	0.00	7.06	-	0.24	-	0.72	290.86	11.52
	4	2.38	0.00	3.90	33.13	0.07	31.37	0.81	281.56	2.22
	5	16.46	3.95	-	-	-	-	0.62	302.19	22.85
	6	11.51	-	-	-	-	-	0.00	342.39	63.05