夏、秋季南黄海小型底栖动物空间分布格局及其环境影响因素

宋远柳; 刘晓收

doi:10.12284/hyxb2023020

夏、秋季南黄海小型底栖动物空间分布格局及其环境影响因素

doi: 10.12284/hyxb2023020

宋远柳^{1, 2,},
刘晓收^{1, 2, ,}

1.
中国海洋大学海洋生命学院，山东青岛 266003
2.
中国海洋大学海洋生物多样性与进化研究所，山东青岛 266003

基金项目: 国家自然科学基金（41976131）。

详细信息

作者简介:
宋远柳（1996-），女，山东省日照市人，主要从事小型底栖动物生态学研究。E-mail: songyuanliu16@163.com

通讯作者:
刘晓收，教授，主要从事生物海洋学与海洋底栖生物生态学研究。E-mail: liuxs@ouc.edu.cn

中图分类号: P722.5；Q178.535
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出版历程
- 收稿日期: 2022-07-22
- 修回日期: 2022-08-29
- 网络出版日期: 2022-09-09
- 刊出日期: 2023-01-09

Spatial distribution patterns of meiofauna and the influencing environmental factors in the southern Yellow Sea in summer and autumn

Song Yuanliu^{1, 2
,},
Liu Xiaoshou^{1, 2
, ,}

1.
College of Marine Life Sciences, Ocean University of China, Qingdao 266003, China
2.
Institute of Evolution and Marine Biodiversity, Ocean University of China, Qingdao 266003, China

摘要

摘要: 为研究南黄海小型底栖动物的空间分布格局及其环境影响因素，于2020年8月（夏季）和11月（秋季）对南黄海进行了两个航次的野外观测和采样，对小型底栖动物的类群组成、丰度、生物量、垂直分布、群落结构及其与环境因子的关系进行了研究。结果显示，共鉴定出小型底栖动物类群15个，其中自由生活海洋线虫为最优势类群，在两个航次中分别占小型底栖动物总丰度的75.6%和84.6%。其他较重要的类群还包括底栖桡足类、轮虫类和枝角类等。夏季和秋季小型底栖动物的平均丰度分别为（514.9±32.1）ind./(10 cm²) 和（350.8±30.7）ind./(10 cm²)，平均生物量（干质量）分别为（651.7±98.0）μg/(10 cm²)和（589.2±37.1）μg/(10 cm²)。小型底栖动物在时空分布上存在差异。在季节分布上，小型底栖动物丰度和类群组成存在极显著差异。结合环境因子分析结果可知，沉积物中值粒径是引起差异的主要环境因子。在空间分布上，夏季小型底栖动物丰度和类群组成在不同水深间存在极显著差异，秋季小型底栖动物丰度和类群组成在不同水深间差异不显著。推测黄海冷水团是影响夏季小型底栖动物空间分布差异的主要因素。本研究中小型底栖动物的数量和类群多样性相较于国内其他对南黄海小型底栖动物的研究较低，其中沉积物叶绿素a含量及有机质含量是引起南黄海小型底栖动物丰度变化的重要因素。海洋线虫与桡足类的丰度比值（N/C比值）评估显示秋季该区域存在有机污染，这一结果与应用大型底栖动物对同一区域进行环境评价的结果不一致，对于应用N/C比值评价环境质量还需要进一步的研究。
- 小型底栖动物 /
- 自由生活海洋线虫 /
- 丰度 /
- 生物量 /
- 南黄海
Abstract: To study the spatial distribution patterns of meiofauna in the southern Yellow Sea and the influencing environmental factors, two cruises were conducted in the southern Yellow Sea in August (summer) and November (autumn) 2020 to analyze the taxa composition, abundance, biomass, vertical distribution, community structure of meiofauna and their relationship with environmental factors. The results showed that a total of 15 taxa of meiofauna were identified, among which free-living marine nematode was the most dominant group, accounting for 75.6% and 84.6% of the total abundance of meiofauna in the two cruises, respectively. Other important groups were benthic copepods, rotifers and cladocerans. The average abundance of meiofauna in summer and autumn was (514.9±32.1) ind./(10 cm²) and (350.8±30.7) ind./(10 cm²), and the average biomass (dry weight) was (651.7±98.0) μg/(10 cm²) and (589.2±37.1) μg/(10 cm²), respectively. There were differences both in the spatial and temporal distribution of meiofauna. In terms of seasonal distribution, highly significant differences were found in the abundance and taxa composition of meiofauna. Combined with the analysis results of environmental factors, the sediment median diameter was the main environmental factor causing the differences. In terms of spatial distribution, the abundance of meiofauna and community structure differed significantly along the water depth gradient in summer, while no significant differences were found along the water depth gradient in autumn. The Yellow Sea Cold Water Mass was supposed to be the main factor affecting the spatial distribution of meiofauna in summer. The abundance and taxa diversity of meiofauna in this study were lower compared with other studies on meiofauna in the southern Yellow Sea. The sediment chlorophyll a and organic matter contents were the important factors causing the change in abundance of meiofauna in the southern Yellow Sea. The assessment of the abundance ratio of marine nematodes to copepods (N/C ratio) showed the presence of organic pollution in the area in autumn, while this result was not consistent with that of the macrofaunal assemblages in the environmental quality evaluation of the same area, and further studies are needed for the application of N/C ratio to evaluate the environmental quality.
- meiofauna /
- free-living marine nematodes /
- abundance /
- biomass /
- southern Yellow Sea

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图 1 南黄海夏、秋季小型底栖动物调查站位

Fig. 1 The sampling sites of meiofauna in the southern Yellow Sea in summer and autumn

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图 2 南黄海调查站位夏、秋季环境因子主成分分析

Fig. 2 Principal component analysis of environmental factors at the sampling sites in the southern Yellow Sea in summer and autumn

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图 3 南黄海夏季（a）、秋季（b）小型底栖动物丰度分布

Fig. 3 Distribution of meiofaunal abundance in the southern Yellow Sea in summer (a) and autumn (b)

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图 4 南黄海海域夏、秋季小型底栖动物群落结构非度量多维标度

Fig. 4 Non-metric multidimensional scaling plot for meiofauna community structure in the southern Yellow Sea in summer and autumn

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图 5 南黄海海域夏季不同水深的小型底栖动物群落结构的非度量多维标度

Fig. 5 Non-metric multidimensional scaling plot for different depth meiofauna community structure in the southern Yellow Sea in summer

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图 6 南黄海海域秋季不同水深的小型底栖动物群落结构的非度量多维标度

Fig. 6 Non-metric multidimensional scaling plot for different depth meiofauna community structure in the southern Yellow Sea in autumn

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图 7 南黄海夏（a）、秋季（b）调查站位小型底栖动物丰度在沉积物中的垂直分布

Fig. 7 Vertical distribution of meiofaunal abundance in sediment in the southern Yellow Sea in summer (a) and autumn (b)

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表 1 小型底栖动物各类群的个体平均干质量

Tab. 1 Individual average dry weight of different meiofauna groups

类群	平均干质量/μg	类群	平均干质量/μg
海洋线虫 Nematoda	0.4	动吻类 Kinorhyncha	2
桡足类 Copepoda	1.86	介形类 Ostracoda	26
枝角类 Cladocera	26	端足类 Amphipoda	15
轮虫类 Rotifera	3.5	等足类 Isopoda	15
海螨类 Halacaroidea	1.5	涟虫 Cumacea	3.5
涡虫类 Turbellaria	3.5	原足类 Tanaidacea	15
双壳类 Bivalvia	4.2	其他 Others	3.5
多毛类 Polychaeta	14

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表 2 南黄海海域夏、秋季调查站位环境因子

Tab. 2 Environmental factors at the sampling sites in the southern Yellow Sea in summer and autumn

站位	WD/m	SWT/℃	BWT/℃	BWS	Md/mm	YT/%	W/%	Chl a/（μg·g⁻¹）	Pha/（μg·g⁻¹）	OM/%	底质类型
夏15	41	22.70	10.55	32.22	0.08	42.94	29.81	0.26	1.05	0.53	粉砂质砂
夏17	53	22.24	11.09	32.77	0.01	66.60	45.50	0.23	1.16	0.80	砂−粉砂−黏土
夏19	62	22.37	10.83	32.99	0.02	63.96	45.06	0.10	0.55	0.70	砂−粉砂−黏土
夏22	70	24.05	11.15	32.10	0.02	59.40	49.53	0.32	1.56	1.91	砂−粉砂−黏土
夏24	63	21.76	11.03	33.11	0.01	59.46	52.36	0.51	1.80	3.18	砂−粉砂−黏土
夏27	51	23.23	10.58	32.63	0.02	55.70	51.47	0.50	2.19	2.67	砂−粉砂−黏土
夏28	52	23.63	11.16	32.43	0.01	76.00	49.36	0.28	1.26	2.60	砂−粉砂−黏土
夏31	34	23.51	14.15	32.09	0.01	80.18	39.26	0.29	1.31	1.39	黏土质粉砂
夏33	29	27.16	15.76	32.00	0.03	60.90	35.09	0.47	2.27	1.12	砂质粉砂
夏43	16	26.03	22.84	31.37	0.01	91.77	51.43	0.72	2.22	1.38	黏土质粉砂
秋15	47	17.61	17.85	31.86	0.12	30.15	34.95	0.16	0.69	1.08	粉砂质砂
秋17	53	14.97	14.78	32.00	0.14	32.05	48.90	0.09	0.67	3.02	粉砂质砂
秋18	60	16.11	13.99	32.08	0.11	39.92	49.91	0.03	0.16	2.08	粉砂质砂
秋19	63	19.01	12.72	32.34	0.12	36.74	49.37	0.38	2.26	0.63	粉砂质砂
秋20	67	16.90	11.00	32.97	0.08	46.57	51.15	1.15	7.39	2.36	粉砂质砂
秋22	68	16.19	11.00	32.97	0.19	25.57	51.05	0.17	1.06	0.67	粉砂质砂
秋24	67	17.95	11.02	32.89	0.09	42.25	55.66	0.35	1.49	2.74	粉砂质砂
秋26	57	17.11	12.02	32.45	0.19	24.81	51.16	0.14	0.77	1.96	砂
秋27	53	17.00	12.42	32.23	0.19	23.00	58.93	0.19	0.83	4.65	砂
秋28	51	16.21	17.82	31.66	0.15	30.31	54.65	0.12	0.60	1.63	粉砂质砂
秋30	39	12.34	18.14	31.60	0.10	39.63	43.48	0.14	0.56	0.90	粉砂质砂
秋33	30	18.26	17.85	31.74	0.13	33.27	40.40	0.07	0.40	1.11	粉砂质砂
秋39	20	18.01	17.42	31.60	0.01	92.75	47.30	0.37	1.25	1.30	黏土质粉砂
注：WD. 水深（water depth）；SWT. 表层水温度（surface water temperature）；BWT. 底层水温度（bottom water temperature）；BWS. 底层水盐度（bottom water salinity）；Md. 中值粒径（sediment median diameter）；YT. 粉砂黏土含量（silt-clay percentage）；W. 含水量（water content）；Chl a. 叶绿素a含量（chlorophyll a content）；Pha. 脱镁叶绿酸含量（phaeophorbide content）；OM. 有机质含量（organic matter content）。

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表 3 南黄海海域夏、秋季小型底栖动物各类群丰度和生物量

Tab. 3 Abundance and biomass of each meiofaunal group in the southern Yellow Sea in summer and autumn

类群	航次	丰度/ （ind.·（10 cm²）⁻¹）	丰度占比/%	生物量/ （μg·（10 cm²）⁻¹）	生物量占比/%
海洋线虫 Nematoda	夏	389.2±18.4	75.6	155.7±7.3	23.9
海洋线虫 Nematoda	秋	296.8±28.1	84.6	118.7±11.3	20.1
桡足类 Copepoda	夏	67.2±16.1	13.1	125±29.9	19.2
桡足类 Copepoda	秋	20.4±3.3	5.8	38±6.1	6.4
轮虫类 Rotifera	夏	34.7±1.7	6.7	121.4±6.1	18.6
轮虫类 Rotifera	秋	10.7±0.5	3.1	37.5±1.8	6.4
枝角类 Cladocera	夏	4.5±2	0.9	115.9±53.2	17.8
枝角类 Cladocera	秋	11.6±0.5	3.3	301.9±13.7	51.2
涡虫类 Turbellaria	夏	5.7±1.2	1.1	19.9±4.1	3.1
涡虫类 Turbellaria	秋	3.3±0.6	0.9	11.4±2	1.9
多毛类 Polychaeta	夏	1.9±0.8	0.4	26.7±11.4	4.1
多毛类 Polychaeta	秋	3.6±1.6	1.0	50.6±22.7	8.6
海螨类 Halacaroidea	夏	3.4±0.6	0.7	5.2±0.9	0.8
海螨类 Halacaroidea	秋	0.1±0.2	0	0.2±0.3	0
动吻类 Kinorhyncha	夏	2.7±1.1	0.5	5.4±2.1	0.8
动吻类 Kinorhyncha	秋	0.9±0.1	0.3	1.7±0.3	0.3
双壳类 Bivalvia	夏	2.3±0.9	0.4	9.6±3.9	1.5
双壳类 Bivalvia	秋	0.9±0.4	0.3	3.9±1.8	0.7
介形类 Ostracoda	夏	1.9±1.2	0.4	48.6±31.6	7.5
介形类 Ostracoda	秋	0.3±0.1	0.1	8.1±1.7	1.4
等足类 Isopoda	夏	1.0±0.1	0.2	14.6±1.2	2.2
等足类 Isopoda	秋	0.5±0.1	0.1	7±1.7	1.2
涟虫 Cumacea	夏	0.2±0.2	0	0.8±0.7	0.1
涟虫 Cumacea	秋	0	0	0	0
端足类 Amphipoda	夏	0.2±0.3	0	2.8±4.9	0.4
端足类 Amphipoda	秋	0.1±0.1	0	1.7±1.7	0.3
原足类 Tanaidacea	夏	0	0	0	0
原足类 Tanaidacea	秋	0.3±0.4	0.1	4.1±5.6	0.7
其他	夏	0.1±0.1	0	0.2±0.4	0
其他	秋	1.2±0.6	0.3	4.3±1.9	0.7
合计	夏	514.9±32.1	100.0	651.7±98.0	100.0
合计	秋	350.8±30.7	100.0	589.2±37.1	100.0

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表 4 南黄海海域海洋线虫和桡足类丰度之比

Tab. 4 Ratios of abundance of nematodes to copepods in the southern Yellow Sea

	站位	海洋线虫N	桡足类C	N/C比值		站位	海洋线虫N	桡足类C	N/C比值
夏季	15	246.4	80.3	3.1	秋季	15	462.0	60.6	7.6
	17	209.5	37.4	5.6		17	251.4	11.1	22.6
	19	338.5	59.8	5.7		18	322.1	25.2	12.8
	22	200.4	45.4	4.4		19	217.1	17.2	12.6
	24	142.4	106.0	1.3		20	225.7	24.7	9.1
	27	441.5	50.0	8.8		22	320.1	6.1	52.8
	28	559.4	108.1	5.2		24	130.8	10.1	13.0
	31	597.3	74.7	8.0		26	122.7	13.6	9.0
	33	742.7	81.3	9.1		27	72.2	5.0	14.3
	43	404.4	26.3	15.4		28	166.6	3.5	47.1
						30	1 080.0	44.9	24.0
						33	346.9	20.7	16.8
						39	140.9	22.7	6.2

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表 5 南黄海夏、秋季小型底栖动物与环境因子的BIOENV分析

Tab. 5 Results of BIOENV analysis between meiofaunal assemblage and environmental factors in the southern Yellow Sea in summer and autumn

环境因子变量数	相关系数	环境因子变量组合
2	0.424	Md, W
3	0.419	Md, W, OM
4	0.378	YT, Md, W, OM
3	0.365	YT, Md, W
4	0.346	Md, W, OM, BWS
3	0.342	Md, W, BWS
5	0.331	YT, Md, W, OM, BWS
4	0.329	Md, W, OM, BWT
4	0.323	Md, W, OM, WD
3	0.320	YT, W, OM
注：WD. 水深（water depth）；BWT. 底层水温度（bottom water temperature）；BWS. 底层水盐度（bottom water salinity）；Md. 中值粒径（sediment median diameter）；YT. 粉砂黏土含量（silt-clay percentage）；W. 含水量（water content）；Chl a. 叶绿素a含量（chlorophyll a content）；Pha. 脱镁叶绿酸含量（phaeophorbide content）；OM. 有机质含量（organic matter content）。

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表 6 南黄海夏、秋季小型底栖动物丰度、生物量与环境因子的相关分析结果

Tab. 6 Results of correlation analysis between abundance and biomass of meiofauna and environmental factors in the southern Yellow Sea in summer and autumn

	WD/m	BWT/℃	BWS	Md/mm	YT/%	W/%	Chl a/（μg·g⁻¹）	Pha/（μg·g⁻¹）	OM/%
MA	−0.417*	0.313	−0.297	−0.332	0.266	−0.531**	−0.013	−0.06	−0.339
MB	−0.345	0.301	−0.336	−0.257	0.195	−0.523*	−0.074	−0.113	−0.309
MA'	−0.417	0.187	−0.162	−0.482*	0.428*	−0.590**	0.102	0.031	−0.294
MB'	−0.314	0.197	−0.237	−0.336	0.286	−0.539**	−0.001	−0.054	−0.257
注：*表示在0.01水平上显著相关；表示在0.05水平上显著相关；WD. 水深（water depth）；BWT. 底层水温度（bottom water temperature）；BWS. 底层水盐度（bottom water salinity）；Md. 中值粒径（sediment median diameter）；YT. 粉砂黏土含量（silt-clay content）；W. 含水量（water content）；Chl a. 叶绿素a含量（chlorophyll a content）；Pha. 脱镁叶绿酸含量（phaeophorbide content）；OM. 有机质含量（organic matter content）；MA. 小型底栖动物丰度（meiofauna abundance）；MB. 小型底栖动物生物量（meiofauna biomass）；MA'. 去除秋季30号站位的小型底栖动物丰度（meiofauna abundance without Site 30 in autumn）；MB'. 去除秋季30号站位的小型底栖动物生物量（meiofauna biomass without Site 30 in autumn）。

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表 7 南黄海小型底栖动物历史数据对比

Tab. 7 Comparison of historical data of meiofauna in the southern Yellow Sea

采样时间	小型底栖动物类群数	小型底栖动物丰度/（ind.·（10 cm²）⁻¹）	海洋线虫占比/%	桡足类占比/%	C/N比值	参考文献
2000年6月	18	809.3±407.4	73.8	18.7	3.9	文献[26]
2003年6月	18	1 584.0±686.0	88.7	5.8	16.2	文献[28]
2004年1月	20	1 186.1±486.1	89.77	4.2	21.4	文献[29]
2007年1−2月	15	348.8	77.6	13.3	13.1	文献[2]
2007年6月	18	1 529.0±1121.0	88.0	6.8	14.8	文献[30]
2011年6月	20	1 194.0±873.0	89.7	5.5	16.3	文献[31]
2013年6月	15	900.8±365.6	84.2	4.2	20.9	文献[32]
2013年11月	13	758.4±403.6	94.0	2.4	39.6	文献[32]
2020年8月	14	514.9±32.1	75.6	13.1	5.8	本研究
2020年11月	14	350.8±30.7	84.6	5.8	14.5	本研究

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