浙江岩礁潮间带大型底栖动物主要优势种的生态位及种间联结性

徐坪; 张翔玉; 周晓东; 赵寒冰; 韩庆喜; 王一农; 尤仲杰

doi:10.12284/hyxb2025142

浙江岩礁潮间带大型底栖动物主要优势种的生态位及种间联结性

doi: 10.12284/hyxb2025142

宁波大学海洋学院浙江宁波 315832

基金项目: 国家自然科学基金（42076156）。

详细信息

作者简介:
徐坪（2001—），男，XX省XX市人，研究方向为大型底栖动物生态学。E-mail：1743847895@qq.com

通讯作者:
王一农，教授，主要从事贝类学、海洋生态学研究。E-mail：wangyinong@nbu.edu.cn

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出版历程
- 收稿日期: 2025-07-17
- 修回日期: 2025-09-30
- 网络出版日期: 2025-11-04

Niche and interspecies association of major dominant species in the rocky intertidal zone of Zhejiang province, China

School of Marine Sciences, Ningbo University, Ningbo 315832, China

摘要

摘要: 为了解浙江岩礁潮间带大型底栖动物的种间资源利用情况及生态关系，在2024年3月至2025年4月于浙江设置了10个岩礁潮间带开展采样调查。采用相对重要指数（IRI）、生态位测定、方差比率（VR）、卡方检验、联结系数（AC）、共同出现百分率（PC）、Spearman秩相关系数以及冗余分析（RDA）对主要优势动物的生态位、种间联结性和环境因子相关性进行了研究。结果表明：（1）浙江岩礁潮间带的物种资源丰富，主要优势种（IRI ＞ 100）共17种；（2）疣荔枝螺（Thais clavigera）和小结节滨螺（Nodilittorina exigua）的生态位宽度较大，为广生态位种；东方小藤壶（Chthamalus challengeri）和葡萄牙牡蛎（Crassostrea angulata）等生物的分布局限性较大，为窄生态位种；（3）疣荔枝螺、短滨螺（Littorina brevicula）、黄口荔枝螺（Thais luteostoma）和史氏背尖贝（Notoacmea schrenckii）等与其他主要优势种的生态位重叠显著；（4）盐度和叶绿素a是影响岩礁潮间带生物丰度的主要环境因子。同时，盐度与叶绿素a之间存在显著负相关关系；（5）浙江岩礁潮间带大型底栖动物的群落结构松散，物种组成不稳定，可能处于演替恢复阶段。
- 岩礁潮间带 /
- 主要优势种 /
- 生态位宽度 /
- 生态位重叠 /
- 种间联结性 /
- 环境因子
Abstract: To investigate the resource utilization and interspecific relationships of macrobenthic fauna in the rocky intertidal zones of Zhejiang Province, surveys were conducted at 10 rocky intertidal sites from March 2024 to April 2025. The index of relative importance (IRI), niche breadth (B_i), niche overlap (O_ik), variance ratio (VR), chi-square test, association coefficient (AC), percentage of co-occurrence (PC), Spearman rank correlation, and redundancy analysis (RDA) were applied to analyze the ecological niches, interspecific associations, and relationships between dominant species and environmental factors. The results showed: (1) the rocky intertidal zones of Zhejiang host a rich diversity of species, with 17 dominant species identified (IRI > 100); (2) Thais clavigera and Nodilittorina exigua had wide niche breadths and were classified as wide niche breadth species, while Chthamalus challengeri and Crassostrea angulata were narrow niche breadth species. (3) significant niche overlap was observed among T. clavigera, Littorina brevicula, Thais luteostoma, and Notoacmea schrenckii, indicating intense resource competition; (4) salinity and chl a were key factors influencing species abundance, and a significant negative correlation was found between salinity and chl a. (5) the community structure of macrobenthos in the rocky intertidal zones appeared to be loosely organized and compositionally unstable, suggesting that it may currently be in a stage of successional recovery.
- rocky intertidal zone /
- dominant species /
- niche width /
- niche overlap /
- interspecific association /
- environmental factors

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图 1 采样站位点

S1：舟山岱山岛燕窝山、S2：舟山朱家尖小乌石塘、S3：宁波象山松兰山、S4：宁波象山花岙岛、S5：台州椒江大陈岛、S6：台州温岭三蒜岛、S7：温州洞头鸽尾礁、S8温州瑞安北麂岛、S9：温州平阳南麂列岛、S10：温州苍南渔寮

Fig. 1 The sampling station

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图 2 主要优势种的生态位宽度（物种编号同表1）

Fig. 2 Niche width of the major dominant species

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图 3 主要优势种的丰度数据聚类分析

Fig. 3 The Cluster analysis of the abundance data of major dominant species

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图 4 主要优势种的联结性检验半矩阵图（物种编号同表1）

Fig. 4 Semi-matrix diagram of association tests for major dominant species

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图 5 主要优势种的Spearman秩相关性检验（物种编号同表1）

Fig. 5 Speraman correlation hot map for major dominant species

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图 6 主要优势种与环境因子的RDA排序

Fig. 6 RDA ordination of major dominant species and environmental factors

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表 1 浙江岩礁潮间带主要优势种的相对重要性指数及生态习性

Tab. 1 The index of relative importance and ecological habits of major dominant species in the rocky intertidal zone of Zhejiang

潮间带主要动物		相对重要性指数IRI	运动方式	适温性	食物类型
编号	种名	相对重要性指数IRI	运动方式	适温性	食物类型
1	日本笠藤壶 Tetraclita japonica	3657.2	固着	广温广布种	浮游生物、微藻、有机碎屑
2	条纹隔贻贝 Septifer virgatus	1847.1	附着	暖水性种	浮游生物、微藻、有机碎屑
3	疣荔枝螺 Thais clavigera	1375.2	爬行	广温广布种	食藤壶或其他贝类
4	小结节滨螺 Nodilittorina exigua	857.6	爬行	广温广布种	藻类为主、偶尔食腐或小型无脊椎动物卵
5	短滨螺 Littorina brevicula	731.9	附着	广温广布种	藻类为主、偶尔食腐或小型无脊椎动物卵
6	青蚶 Barbatia obliquata	698.7	固着	暖水性种	浮游生物、微藻、有机碎屑
7	葡萄牙牡蛎 Crassostrea angulata	586.3	固着	暖水性种	浮游生物、微藻、有机碎屑
8	棘刺牡蛎 Saccostrea echinata	555.7	固着	暖水性种	浮游生物、微藻、有机碎屑
9	鳞笠藤壶 Tetraclita squamosa	471.4	附着	广温广布种	浮游生物、有机碎屑
10	厚壳贻贝 Mytilus coruscus	340.2	固着	暖水性种	浮游生物、微藻、有机碎屑
11	日本花棘石鳖 Acanthopleura japonica	291.4	爬行	暖水性种	刮食藻类、微生物
12	黄口荔枝螺 Thais luteostoma	284.7	爬行	广温广布种	捕食藤壶、双壳类
13	史氏背尖贝 Notoacmea schrenckii	271.5	爬行	暖水性种	刮食藻类、微生物
14	日本菊花螺 Siphonaria japonica	246.9	爬行	广温广布种	藻类为主、偶尔食腐或小型无脊椎动物卵
15	紫贻贝 Mytilus galloprovincialis	226.5	附着	暖水性种	浮游生物、微藻、有机碎屑
16	齿纹蜒螺 Nerita yoldii	212.8	爬行	暖水性种	浮游生物、微藻、有机碎屑
17	东方小藤壶 Chthamalus challengeri	150.4	固着	暖水性种	浮游生物、微藻、有机碎屑
注：优势种的运动方式、适温性和食物类型参考^{[17–18, 32–33]}

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表 2 主要优势种的种对间生态位重叠值

Tab. 2 Niche overlap between major dominant species pairs

编号	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16
2	0.97
3	0.87	0.83
4	0.48	0.33	0.63
5	0.67	0.18	0.73	0.76
6	0.51	0.22	0.56	0.52	0.45
7	0.16	0.01	0.61	0.40	0.12	0.20
8	0.29	0.18	0.69	0.29	0.13	0.26	0.92
9	0.88	0.59	0.82	0.35	0.75	0.31	0.02	0.11
10	0.53	0.48	0.61	0.46	0.26	0.12	0.43	0.56	0.39
11	0.62	0.68	0.76	0.69	0.42	0.23	0.01	0.07	0.47	0.87
12	0.85	0.73	0.91	0.51	0.58	0.64	0.64	0.62	0.84	0.68	0.51
13	0.64	0.35	0.58	0.79	0.66	0.38	0.44	0.45	0.52	0.65	0.69	0.58
14	0.54	0.33	0.73	0.59	0.63	0.28	0.09	0.14	0.44	0.24	0.27	0.39	0.68
15	0.68	0.42	0.77	0.46	0.61	0.20	0.01	0.01	0.66	0.92	0.80	0.86	0.72	0.27
16	0.17	0.02	0.13	0.43	0.15	0.41	0.94	0.96	0.01	0.01	0.02	0.17	0.44	0.08	0.01
17	0.66	0.17	0.64	0.12	0.69	0.19	0	0	0.82	0.45	0.28	0.80	0.50	0.28	0.75	0
注：物种编号同表1。加粗数字为生态位重叠指数大于0.60的种对，表示这两个物种间生态位重叠显著。

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表 3 主要优势种种间的总体关联性

Tab. 3 General interspecific association test of major dominant species

总种数方差	总站位方差	方差比率	统计量	卡方检验置信区间	检验结果
S_T²	σ_T²	VR	W	χ² [χ²_0.05 (N), χ²_0.95 (N)]	检验结果
3.96	2.30	1.72	17.22	3.94, 18.3	不显著正关联

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表 4 主要优势种χ²检验表

Tab. 4 chi-square test table of major dominant species

编号	1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16
2	0.63
3	0.23	0.02
4	0.23	0.02	1.14
5	1.98	0.63	0.23	0.23
6	0.23	0.02	1.14	1.14	0.23
7	0.21	0.03	0.02	0.02	0.21	0.02
8	0.56	0.63	0.35	0.35	0.56	0.35	1.9
9	0.56	0.63	0.35	0.35	0.56	0.35	0.24	0.2
10	0.56	0.63	0.35	0.35	0.56	0.35	0.24	0.2	0.2
11	0.21	2.41	0.02	0.02	0.21	0.02	0.82	0.24	1.9	0.24
12	1.98	0.63	0.23	0.23	1.98	0.23	0.21	0.56	0.56	0.56	0.21
13	0.23	0.02	1.14	1.14	0.23	1.14	0.02	0.35	0.35	0.35	0.02	0.23
14	0.63	0.04	0.02	0.02	0.63	0.02	0.03	0.63	0.63	0.63	2.41	0.63	0.38
15	0.05	1.28	0.13	0.13	0.05	0.13	0.18	0.42	0.42	3.85*	0.18	0.05	0.03	0.23
16	0.21	0.03	0.02	0.02	0.21	0.02	0.36	1.9	1.9	0.24	0.36	0.21	0.14	0.03	0.97
17	1.98	0.63	0.23	0.23	1.98	0.23	0.21	0.56	0.56	0.56	0.21	1.98	1.6	0.63	0.05	0.21
注：物种编号同表1，表中标黑数字表示为正联结，下划线数字表示为无联结性，星号数字表示联结达显著水平。

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浙江岩礁潮间带大型底栖动物名录
物种名	拉丁名	站位点
物种名	拉丁名	S1	S2	S3	S4	S5	S6	S7	S8	S9	S10
腔肠动物	Coelenterata
太平洋侧花海葵	Anthopleura nigrescens		+						+	+
朴素侧花海葵	Anthopleura inornata	+	+	+	+	+	+		+	+	+
绿侧花海葵	Anthopleura midori	+		+	+	+	+	+	+	+	+
纵条肌海葵	Haliplanella lucide		+							+
环节动物	Annelida
短毛海鳞虫	Halosydna brevisetosa									+
巧言虫	Eulalia viridis									+
多齿围沙蚕	Perinereis nuntia	+	+		+		+		+	+	+
花索沙蚕	Arabella iricolor	+							+	+
软体动物	Mollusca
红条毛肤石鳖	Acanthochitona rubrolineatus		+	+		+	+	+	+	+	+
平濑锦石鳖	Onithochiton hirasei									+
花斑锉石鳖	Ischnochiton comptus									+
网纹鬃毛石鳖	Mopalia retifera									+
日本花棘石鳖	Acanthopleura japonica		+			+	+	+	+	+	+
嫁䗩	Cellana toreuma		+		+		+	+	+	+
斗嫁䗩	Cellana grata									+
史氏背尖贝	Notoacmea schrenckii	+	+	+	+	+	+	+			+
北戴河小笠贝	Lottia dorsuosa								+		+
矮拟帽贝	Patelloida pygmaea					+		+	+	+	+
花边拟帽贝	Patelloida heroldi		+
单齿螺	Monodonta labio	+	+	+	+	+	+	+		+	+
锈凹螺	Chlorostoma rustica							+	+	+	+
黑凹螺	Chlorostoma nigerrima						+
单一丽口螺	Calliostoma unicum			+						+
托氏娼螺	Umbonium thomasi				+
角蝾螺	Turbo cornutus									+
朝鲜花冠小月螺	Lunella coronata			+			+
粒花冠小月螺	Lunella coronata granulate				+		+
齿纹蜒螺	Nerita yoldii	+		+	+	+	+	+		+
日本蜒螺	Nerita japonica	+			+	+
渔舟蜒螺	Nerita albicilla							+		+
小结节滨螺	Nodilittorina exigua	+	+	+	+	+	+	+	+	+	+
短滨螺	Littorina brevicula	+	+	+	+	+	+	+	+	+	+
塔结节滨螺	Nodilittorina pyramidalis	+	+	+		+		+	+	+	+
中间拟滨螺	Littoraria intermedia	+			+	+	+	+
覆瓦小蛇螺	Thylacodes adamsii									+	+
扁玉螺	Glossaulax didyma						+
粒蝌蚪螺	Gyrineum natator					+	+	+	+	+	+
玫瑰覆螺	Sandalia triticea	+
刺履螺	Bostrycapulus gravispinosus									+
疣荔枝螺	Thais clavigera	+	+	+	+	+	+	+	+	+	+
黄口荔枝螺	Thais luteostoma	+		+	+	+	+	+	+	+	+
爪哇荔枝螺	Thais javanica			+			+
蛎敌荔枝螺	Thais gradata							+	+		+
瘤荔枝螺	Thais bronni		+							+	+
丽小笔螺	Mitrella albuginosa	+		+		+	+			+
甲虫螺	Cantharus cecillei	+		+				+		+
环唇齿螺	Engina lanceolata							+		+
秀丽织纹螺	Nassarius festivus				+
白带三角口螺	Scalptia scalariformis						+
迷乱环肋螺	Gyroscala commutata		+
石磺海牛	Homoiodorididae sp.								+		+
日本菊花螺	Siphonaria japonica	+	+			+	+	+	+	+
蛛形菊花螺	Siphonaria sirius					+		+	+	+
石磺sp.	Onchidiidae. sp.					+				+
角杯阿地螺	Atys angusta									+
东方壳蛞蝓	Philine orientalis				+
青蚶	Barbatia obliquata	+	+	+	+	+	+	+	+	+	+
棕节毛蚶	Barbatia amygdalumtostum			+
双纹须蚶	Barbatia bistrigata
布氏蚶	Arca boucardia									+
条纹隔贻贝	Septifer virgatus		+	+		+	+	+	+	+	+
厚壳贻贝	Mytilus coruscus			+		+		+	+		+
紫贻贝	Mytilus galloprovincialis		+	+		+		+	+		+
短石蛏	Lithophaga curta									+	+
带偏顶蛤	Modiolus comptus									+
毛贻贝	Trichomya hirsuta									+
黑荞麦蛤	Xenostrobus atratus									+
云石肌蛤	Musculus cupreus									+
翡翠股贻贝	Perna viridis									+
中国不等蛤	Anomia chinensis							+		+	+
棘刺牡蛎	Saccostrea echinata	+		+			+	+		+
团聚牡蛎	Saccostrea glomerata									+
长牡蛎	Magallana gigas			+
葡萄牙牡蛎	Crassostrea angulata	+		+				+
熊本牡蛎	Crassostrea sikamea	+		+
中华牡蛎	Hyotissa sinensis									+
豆荚钳蛤	Isognomon legumen									+
栗色拉沙蛤	Lasaea nipponica									+
敦氏猿头蛤	Chama dunkeri									+
纹斑棱蛤	Neotrapezium liratum							+
异纹心蛤	Cardita variegata							+
东方缝栖蛤	Hiatella orientalis									+
甲壳动物	Crustacea
龟足	Capitulum mitella						+	+	+	+	+
日本笠藤壶	Tetraclita japonica	+	+	+	+	+	+	+	+	+	+
鳞笠藤壶	Tetraclita squamosa		+					+	+	+	+
东方小藤壶	Chthamalus challengeri										+
刺巨藤壶	Megabalanus volcano									+
纹藤壶	Amphibalanus amphitrite			+		+
强壮藻钩虾	Ampithoe valida									+
海蟑螂	Ligia exotica									+
刺额光滑瓷蟹	Lissoporcellana spinuligera									+
光辉圆扇蟹	Sphaerozius nitidus									+
斑点拟相手蟹	Parasesarma pictum					+				+
无齿相手蟹	Chiromantes dehaani		+		+
长足长方蟹	Metaplax longipes		+
肉球近方蟹	Hemigrapsus sanguineus				+
白纹方蟹	Grapsus albolineatus					+
中华蚶豆蟹	Arcotheres sinensis									+
棘皮动物	Echinodermata
近辐蛇尾	Ophiactis affinis									+
紫海胆	Anthocidaris crassispina								+	+
马粪海胆	Hemicentrotus pulcherrimus									+
细雕刻肋海胆	Temnopleurus toreumatica									+
注：S1：舟山岱山岛燕窝山、S2：舟山朱家尖小乌石塘、S3：宁波象山松兰山、S4：宁波象山花岙岛、S5：台州椒江大陈岛、S6：台州温岭三蒜岛、S7：温州洞头鸽尾礁、S8温州瑞安北麂岛、S9：温州平阳南麂列岛、S10：温州苍南渔寮。

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