Comparison of community structure and diversity of free-living marine nematodes in the sandy intertidal zone of Dalian in winter
-
摘要: 本研究于2015年12月在濒临渤海的大连夏家河子沙滩和濒临黄海的大连金沙滩采集海洋线虫样品,对两处砂质潮间带海洋线虫群落结构及多样性特征进行了研究。研究结果表明,两处砂质潮间带海洋线虫丰度、优势种、摄食结构及多样性差异显著。夏家河子海洋线虫平均丰度为(592.0±318.9) ind./(10 cm2),金沙滩海洋线虫平均丰度为(54.6±53.8) ind./(10 cm2),显著低于夏家河子。共鉴定海洋线虫43种,共有种13个。夏家河子沙滩海洋线虫优势种为Daptonema sp.1、Daptonema sp.2、Theristus sp.1、Setosabatieria sp.1、Prochromadorella sp.1、Daptonema sp.3、Paracyatholaimus sp.1、Sabatieria breviseta,以非选择性沉积食性线虫为主;金沙滩海洋线虫优势种为Oncholaimus sp.1、Chromadorita sp.1、Theristus sp.2, Neochromadora sp.1、Enoplus sp.1、Prochromadorella sp.1,以杂食性或捕食性线虫为主。两处砂质潮间带海洋线虫种数S、丰富度指数d和香农−威纳多样性指数H'存在极显著的差异,其值均表现为夏家河子显著高于金沙滩。相似性分析结果显示,两处沙滩的海洋线虫群落结构存在显著差异。间隙水pH、沉积物有机质含量及中值粒径是引起海洋线虫群落结构及多样性差异的主要因子,其中,沉积物中值粒径对线虫群落特征的影响最为突出。就两处沙滩潮区差异而言,受沉积物粒径特征及pH的影响,夏家河子海洋线虫群落特征在不同潮区之间也存在差异,其低潮带海洋线虫多样性高于高、中潮带,杂食者和捕食者丰度高,反映其海洋线虫群落结构较复杂。金沙滩不同潮区的环境因子无显著差异,线虫群落结构及多样性潮区差异不显著,较为均一。此外,水动力对海洋线虫群落结构沙滩差异和潮区差异也产生重要影响。一方面,水动力通过影响不同粒径沉积物的沉积影响海洋线虫群落特征;另一方面,水动力可影响线虫的分散性,进而对其潮区分布产生影响。Abstract: In this study, marine nematode samples were collected from Dalian Xiajiahezi Beach near the Bohai Sea and Dalian Golden Beach near the Yellow Sea in December 2015, and the community structure and diversity characteristics of marine nematodes in two sandy intertidal zones were studied. The results showed that there were significant differences in the abundance, dominant species, trophic structure and diversity of marine nematodes at the two beaches. The average abundance of marine nematodes were (592.0±318.9) ind./(10 cm2) in the Xiajiahezi Beach, and (54.6±53.8) ind./(10 cm2 ) in the Golden Beach, which was significantly lower than that in the Xiajiahezi Beach. A total of 43 species of marine nematodes were identified, and 13 common species. The dominant species of marine nematodes in the Xiajiahezi Beach were Daptonema sp.1, Daptonema sp.2, Theristus sp.1, Setosabatieria sp.1, Prochromadorella sp.1, Daptonema sp.3, Paracyatholaimus sp.1, Sabatieria breviseta, which were mainly non-selective sedimenous nematodes. The dominant species in the Golden Beach were Oncholaimus sp.1 Chromadorita sp.1, Theristus sp.2, Neochromadora sp.1, Enoplus sp.1, Prochromadorella sp.1, which were mainly omnivorous or predatory nematodes. There were significant differences in species number (S), Margalef’s species richness (d) and Shannon–Wiener index(H′) between the two sandy beaches, and the values were significantly higher in the Xijiahezi Beach than the Golden Beach. The Analysis of Similarity tests (ANOSIM) results showed that there were significant differences in marine nematodes community structure between the two beaches. Interstitial water temperature and median diameter (MDΦ) were the main environmental factors that cause differences in marine nematode community structure and diversity. The characteristics of the marine nematodes community in the Xijiahezi Beach were also different among different tidal zones due to the variations of sediment granularity and pH. The diversity of marine nematodes in low tidal zone was higher than that in high and mid tidal zones, and the abundance of omnivores and predators was higher, which reflected the complex structure of the marine nematodes community. In the Golden Beach, there was no significant difference in environmental factors in different tidal zones, and the difference in nematodes community structure and diversity of tidal areas was not significant and relatively uniform. In addition, hydrodynamic force function was also an important factor that causes differences in marine nematode community structures at the studied beaches. On the one hand, hydrodynamic force affected the community characteristics of marine nematodes by influencing the deposition of sediments with different particle sizes. On the other hand, hydrodynamic force could affect the dispersity of nematodes which in turn affects their tidal distribution.
-
Key words:
- Dalian; beach /
- marine nematodes /
- community structure /
- feeding type
-
图 2 两处沙滩环境因子主成分分析
X为夏家河子;J为金沙滩;H、M、L分别代表高、中、低潮带;T为间隙水温度;Sal为 间隙水盐度;DO为间隙水溶解氧;Chl a为叶绿素 a;OM为有机质,MDΦ为中值粒径
Fig. 2 Principal component analysis of environmental factors in two beaches
X represents Xiajiahezi Beach; J represents Golden Beach; H, M, and L represent high, mid, and low tidal zones respectively; T represents interstitial water temperature; Sal represents interstitial water salinity; DO represents interstitial water dissolved oxygen; Chl a represents chlorophyll a; OM represents organic matter; MDΦ represents medium grain size
表 1 两处沙滩各潮间带环境因子
Tab. 1 Environmental factors in different tidal zones of two beaches
采样地点 潮区 间隙水温度/℃ pH 间隙水溶解氧浓度/(mg·L−1) 间隙水盐度 Chl a 质量比/(μg·g−1) 有机质含量/% 中值粒径/(Φ) 夏家河子 高潮带 2.45 8.33 5.53 28.80 1.07 0.10 2.59 中潮带 1.19 8.23 5.53 28.80 1.76 0.04 2.72 低潮带 0.74 8.47 3.86 30.20 0.73 0.04 3.14 金沙滩 高潮带 7.00 8.12 9.21 34.00 0.09 0.84 0.47 中潮带 8.10 7.82 7.07 33.95 0.19 0.80 −0.32 低潮带 7.79 7.91 5.22 35.00 0.44 0.75 1.30 表 2 两处沙滩海洋线虫优势种贡献率及其摄食类型(SIMPER分析,优势度≥5%)
Tab. 2 The contributions and feeding types of dominant nematodes species in two beaches (SIMPER analysis, dominance ≥5%)
优势种 摄食类型 夏家河子 金沙滩 高潮带优势种
贡献率/%中潮带势种
贡献率/%低潮带势种
贡献率/%高潮带势种
贡献率/%中潮带势种
贡献率/%低潮带势种
贡献率/%Axonolaimus sp.1 2A 5.22 Bathylaimus sp.1 1B 6.02 Chromadorita sp.1 2A 26.67 26.83 Daptonema sp.1 1B 11.68 34.75 14.39 5.92 Daptonema sp.2 1B 16.63 22.47 14.18 Daptonema sp.3 1B 16.35 Enoplolaimus sp.1 2B 6.32 Enoplus sp.1 2B 12.15 5.00 Neochromadora sp.1 2A 7.20 6.53 Oncholaimus sp.1 2B 36.39 57.62 59.18 Oncholaimus sp.2 2B 10.52 Paracyatholaimus sp.1 2A 9.83 Prochromadorella sp.1 2A 13.11 13.44 7.85 Sabatieria breviseta 1B 12.14 Setosabatieria sp.1 1B 19.67 16.17 Thalamonhystera sp.1 1B 6.32 Theristus sp.1 1B 13.31 11.55 12.23 Theristus sp.2 1B 12.15 8.35 5.00 Viscosia sp.1 2B 12.8 表 3 研究沙滩海洋线虫多样性指数
Tab. 3 Diversity indices of marine nematodes in study beaches
潮间带 潮区 物种数 物种丰富度指数 均匀度指数 香农−威纳多样性指数 优势度指数 夏家河子 H 19 4.049 0.698 5 2.967 0.812 4 M 20 4.353 0.504 7 2.181 0.645 2 L 26 6.147 0.705 8 3.318 0.872 7 金沙滩 H 11 3.453 0.703 8 2.435 0.807 6 M 6 3.200 0.909 1 2.350 0.986 9 L 15 4.776 0.602 1 2.352 0.728 0 表 4 大连沙滩海洋线虫丰度及多样性指数与环境因子的相关性分析结果
Tab. 4 Results of correlation analysis of marine nematode abundance and diversity indices with environmental factors in Dalian beaches
多样性指数 间隙水温度 pH 间隙水溶解氧浓度 间隙水盐度 Chl a 质量比 有机质含量 中值粒径 丰度 −0.882** 0.648* −0.485 −0.812** 0.705* −0.687* 0.799** 物种数 −0.822** 0.778** −0.570 −0.544 0.623* −0.683* 0.855** 物种丰富度指数 −0.505 0.507 −0.415 −0.168 0.314 −0.506 0.621* 均匀度指数 0.260 −0.279 0.187 0.178 −0.471 0.088 −0.296 香农−威纳多样性指数 −0.601* 0.589* −0.379 −0.406 0.210 −0.583* 0.670* 优势度指数 0.040 −0.113 −0.039 0.019 −0.244 −0.093 −0.038 注:*代表差异显著(p<0.05);**代表差异极显著(p<0.01)。 表 5 不同沙滩海洋线虫丰度及多样性指数与环境因子的相关性分析
Tab. 5 Correlation analysis of marine nematode abundance and diversity indices with environmental factors in different beaches
潮间带 环境因子 间隙水温度 间隙水pH 间隙水溶解氧浓度 间隙水盐度 Chl a质量比 有机质含量 中值粒径 夏家河子 丰度 0.615 −0.136 0.300 0.307 0.025 0.493 −0.568 物种数 −0.234 0.646 −0.474 0.630 −0.039 −0.590 0.475 物种丰富度指数 −0.358 0.703 −0.499 0.625 −0.139 −0.614 0.600 均匀度指数 −0.028 0.854* −0.367 0.189 −0.836* 0.173 0.546 香农−威纳多样性指数 −0.134 0.895* −0.485 0.413 −0.613 −0.131 0.604 优势度指数 −0.045 0.820* −0.350 0.234 −0.652 −0.079 0.548 金沙滩 丰度 0.006 −0.392 0.043 −0.634 −0.170 0.775 −0.228 物种数 −0.350 0.090 −0.293 0.372 0.327 0.295 0.657 物种丰富度指数 0.070 −0.212 −0.082 0.460 0.172 −0.251 0.680 均匀度指数 0.692 −0.651 0.259 −0.389 −0.307 −0.250 −0.449 香农−威纳多样性指数 0.310 −0.498 0.155 0.031 −0.101 −0.161 0.345 优势度指数 0.658 −0.713 0.080 −0.248 −0.185 −0.195 −0.286 注:*代表差异显著(p<0.05);**代表差异极显著(p<0.01)。 -
[1] McLachlan A, Turner I. The interstitial environment of sandy beaches[J]. Marine Ecology, 1994, 15(3/4): 177−212. [2] McLachlan A, Dorvlo A. Global patterns in sandy beach macrobenthic communities[J]. Journal of Coastal Research, 2005, 2005(214): 674−687. [3] Vassallo P, Paoli C, Fabiano M. Ecosystem level analysis of sandy beaches using thermodynamic and network analyses: a study case in the NW Mediterranean Sea[J]. Ecological Indicators, 2012, 15(1): 10−17. doi: 10.1016/j.ecolind.2011.09.016 [4] Gheskiere T, Vincx M, Weslawski J M, et al. Meiofauna as descriptor of tourism-induced changes at sandy beaches[J]. Marine Environmental Research, 2005, 60(2): 245−265. doi: 10.1016/j.marenvres.2004.10.006 [5] Heip C H R, Vincx M, Vranken G. The ecology of marine nematodes[J]. Oceanography and Marine Biology, 1985, 23: 399−489. [6] Schratzberger M, Gee J M, Rees H L, et al. The structure and taxonomic composition of sublittoral meiofauna assemblages as an indicator of the status of marine environments[J]. Journal of the Marine Biological Association of the UK, 2000, 80(6): 969−980. doi: 10.1017/S0025315400003039 [7] Coull B C, Chandler G T. Pollution and meiofauna: field, laboratory, and mesocosm studies[J]. Oceanography Marine Biology, 1992, 30: 191−271. [8] Santos G H C, Cardoso R S, Maria T F. Bioindicators or sediment relationships: evaluating ecological responses from sandy beach nematodes[J]. Estuarine, Coastal and Shelf Science, 2019, 224: 217−227. doi: 10.1016/j.ecss.2019.04.035 [9] Pavlyuk O N, Trebukhova Y A. Intertidal meiofauna of Jeju Island, Korea[J]. Ocean Science Journal, 2011, 46(1): 1−11. doi: 10.1007/s12601-011-0001-3 [10] Mundo-Ocampo M, Lambshead P J D, Debenham N, et al. Biodiversity of littoral nematodes from two sites in the gulf of california[J]. Hydrobiologia, 2007, 586(1): 179−189. doi: 10.1007/s10750-006-0624-z [11] Delgado J D, Riera R, Monterroso Ó, et al. Distribution and abundance of meiofauna in intertidal sand substrata around Iceland[J]. Aquatic Ecology, 2009, 43(2): 221−233. doi: 10.1007/s10452-008-9200-0 [12] Armenteros M, Ruiz-Abierno A, Fernández-Garcés R, et al. Biodiversity patterns of free-living marine nematodes in a tropical bay: cienfuegos, Caribbean Sea[J]. Estuarine, Coastal and Shelf Science, 2009, 85(2): 179−189. doi: 10.1016/j.ecss.2009.08.002 [13] Kotwicki L, Szymelfenig M, Troch M D, et al. Latitudinal biodiversity patterns of meiofauna from sandy littoral beaches[J]. Biodiversity & Conservation, 2005, 14(2): 461−474. [14] Lee M R, Riveros M. Latitudinal trends in the species richness of free-living marine nematode assemblages from exposed sandy beaches along the coast of Chile (18°–42°S)[J]. Marine Ecology, 2012, 33(3): 317−325. doi: 10.1111/j.1439-0485.2011.00497.x [15] 张志南, 钱国珍. 小型底栖生物取样方法的研究[J]. 海洋湖沼通报, 1990(4): 37−42.Zhang Zhinan, Qian Guozhen. A study on sampling methods for meiofauna[J]. Transactions of Oceanology and Limnology, 1990(4): 37−42. [16] 张志南. 秦皇岛砂滩海洋线虫的数量研究[J]. 青岛海洋大学学报, 1991, 21(1): 63−75.Zhang Zhinan. A study on the abundance of free-living marine nematodes on four intertidal sandy beaches at the Qin Huang Dao Bay, north China[J]. Journal of Ocean University of Qingdao, 1991, 21(1): 63−75. [17] 张志南, 党宏月, 于子山. 青岛湾有机质污染带小型底栖生物群落的研究[J]. 中国海洋大学学报(自然科学版), 1993, 23(1): 83−91.Zhang Zhinan, Dang Hongyue, Yu Zishan. A study on the meiobenthic community in an organically polluted beach of Qingdao Bay[J]. Journal of Ocean University of China, 1993, 23(1): 83−91. [18] 张志南, 林霞, 于子山. 大连石槽岩滩附植小型动物的初步研究[J]. 中国海洋大学学报(自然科学版), 1994, 24(3): 373−383.Zhang Zhinan, Lin Xia, Yu Zishan. Preliminary study on the phytal meiofauna from the rocky beach at Shicao, Dalian[J]. Periodical of Ocean University of China, 1994, 24(3): 373−383. [19] 张志南, 周红, 华尔, 等. 中国小型底栖生物研究的40年——进展与展望[J]. 海洋与湖沼, 2017, 48(4): 657−671.Zhang Zhinan, Zhou Hong, Hua Er, et al. Meiofauna study for the forty years in China—Progress and prospect[J]. Oceanologia et Limnologia Sinica, 2017, 48(4): 657−671. [20] Liu Haibin, Zhang Zhinan, Fan Shiliang, et al. Seasonal variability in free-living marine nematode community structure in a sandy beach of the Taiping Bay of Qingdao, China[J]. Acta Oceanologica Sinica, 2008, 27(2): 102−115. [21] 蔡立哲, 李复雪. 厦门潮间带泥滩和虾池小型底栖动物类群的丰度[J]. 台湾海峡, 1998, 17(1): 91−95.Cai Lizhe, Li Fuxue. Abundance of meiofauna on intertidal mudflat and shrimp ponds in Xiamen[J]. Journal of Oceanography in Taiwan Strait, 1998, 17(1): 91−95. [22] 蔡立哲, 厉红梅, 邹朝中. 厦门钟宅泥滩海洋线虫群落的种类组成及其多样性[J]. 厦门大学学报(自然科学版), 2000, 39(5): 669−675.Cai Lizhe, Li Hongmei, Zou Chaozhong. Species composition and diversity of marine nematode community on intertidal mudflat in Zhongzhai, Xiamen[J]. Journal of Xiamen University (Natural Science), 2000, 39(5): 669−675. [23] 黄德铭, 刘晓收, 林明仙, 等. 污水排海对小型底栖生物丰度和生物量的影响[J]. 应用生态学报, 2014, 25(10): 3023−3031.Huang Deming, Liu Xiaoshou, Lin Mingxian, et al. Effects of sewage discharge on abundance and biomass of meiofauna[J]. Chinese Journal of Applied Ecology, 2014, 25(10): 3023−3031. [24] 范士亮, 刘海滨, 张志南, 等. 青岛太平湾砂质潮间带小型底栖生物丰度和生物量的研究[J]. 中国海洋大学学报(自然科学版), 2006, 36(S1): 98−104.Fan Shiliang, Liu Haibin, Zhang Zhinan, et al. Study on the abundance and biomass of meiofauna in the sandy beach of Taiping Bay, Qingdao[J]. Periodical of Ocean University of China, 2006, 36(S1): 98−104. [25] 李佳, 华尔, 张志南. 青岛砂质潮间带小型底栖动物分布及季节动态[J]. 应用生态学报, 2012, 23(12): 3458−3466.Li Jia, Hua Er, Zhang Zhinan. Distribution and seasonal dynamics of meiofauna in intertidal zone of Qingdao sandy beaches, Shandong Province of East China[J]. Chinese Journal of Applied Ecology, 2012, 23(12): 3458−3466. [26] 慕芳红, 张婷, 李佳, 等. 厦门大德记沙滩小型底栖动物的时空分布及影响因素[J]. 中国海洋大学学报(自然科学版), 2020, 50(9): 34−45.Mu Fanghong, Zhang Ting, Li Jia, et al. Spatiotemporal distribution of meiofauna and its influencing factors at the Dadeji Beach, Xiamen[J]. Periodical of Ocean University of China, 2020, 50(9): 34−45. [27] Hua Er, Mu Fanghong, Zhang Zhinan, et al. Nematode community structure and diversity pattern in sandy beaches of Qingdao, China[J]. Journal of Ocean University of China, 2016, 15(1): 33−40. doi: 10.1007/s11802-016-2686-5 [28] 陈玉珍, 郭玉清, 刘爱原. 福建海坛岛长江澳沙滩自由生活海洋线虫群落研究[J]. 生态学报, 2019, 39(7): 2573−2582.Chen Yuzhen, Guo Yuqing, Liu Aiyuan. Community structures of free-living marine nematodes on sandy beach of Changjiang’ao (Haitan Island, Fujian Province)[J]. Acta Ecologica Sinica, 2019, 39(7): 2573−2582. [29] 华尔, 李佳, 董洁, 等. 砂质潮间带自由生活海洋线虫对缺氧的响应—微型受控生态系研究[J]. 生态学报, 2012, 32(13): 3975−3986. doi: 10.5846/stxb201106080765Hua Er, Li Jia, Dong Jie, et al. Responses of sandy beach nematodes to oxygen deficiency: microcosm experiments[J]. Acta Ecologica Sinica, 2012, 32(13): 3975−3986. doi: 10.5846/stxb201106080765 [30] Hua Er, Zhang Zhinan, Zhou Hong, et al. Meiofauna distribution in intertidal sandy beaches along China shoreline (18°–40°N)[J]. Journal of Ocean University of China, 2016, 15(1): 19−27. doi: 10.1007/s11802-016-2740-3 [31] 杨丹. 大连湾局部海域石油污染对潮间带海洋线虫群落结构影响的研究[D]. 大连: 辽宁师范大学, 2014.Yang Dan. Studies on responds of marine nematode community to crude oil contamination in intertidal zone of Bathing Beach, Dalian Bay[D]. Dalian: Liaoning Normal University, 2014. [32] 张冲, 杨同军. 我国东部沿海平均潮差分布特征及变化规律[J]. 水运工程, 2013(1): 60−65.Zhang Chong, Yang Tongjun. Distribution and variation law of mean tidal range in eastern coastal China[J]. Port & Waterway Engineering, 2013(1): 60−65. [33] 张江泉, 郑崇伟, 李荣川, 等. 黄渤海风、浪、流等海洋水文要素特征分析[J]. 科技资讯, 2013(31): 112−115.Zhang Jiangquan, Zheng Chongwei, Li Rongchuan, et al. Analysis of sea surface wind speed, wave and current in the Bohai Sea and Yellow Sea[J]. Science & Technology Information, 2013(31): 112−115. [34] Zhu Longhai, Hu Rijun, Zhu Haijun, et al. Modeling studies of tidal dynamics and the associated responses to coastline changes in the Bohai Sea, China[J]. Ocean Dynamics, 2018, 68(12): 1625−1648. doi: 10.1007/s10236-018-1212-2 [35] Platt H M, Warwick R M. Freeliving Marine Nematodes: II. British Chromadorids: Pictorial Key to World Genera and Notes for the Identification of British Species[M]. Great Britain: The Bath Press, 1988. [36] Wieser W. Die Beziehung zwischen Mundhöhlengestalt, Ernährungsweise und Vorkommen bei freilebenden marinen Nematoden[J]. Arkiv für Zoologi, 1953, 4: 439−484. [37] 国家海洋局. GB/T 17378.5−2007, 海洋监测规范第5部分: 沉积物分析[S]. 北京: 中国标准出版社, 2007.State Oceanic Administration. GB/T 17378.5−2007, The specification for marine monitoring—Part 5: sediment analysis[S]. Beijing: China Standard Press, 2007. [38] 国家海洋局. GB/T 127633−2007, 海洋调查规范 第3部分: 海洋气象观测[S]. 北京: 中国标准出版社, 2008.State Oceanic Administration. GB/T 12763.3−2007, Specifications for oceanographic survey—Part 3: marine meteorolgical observations[S]. Beijing: China Standard Press, 2008. [39] Hulings N C, Gray J W. Physical factors controlling abundance of meiofauna on tidal and atidal beaches[J]. Marine Biology, 1976, 34(1): 77−83. doi: 10.1007/BF00390790 [40] 刘海滨. 青岛太平湾砂质潮间带小型底栖生物群落结构与多样性的研究[D]. 青岛: 中国海洋大学, 2007.Liu Haibin. Studies on community structure and biodiversity of meiofauna in Taiping Bay, Qingdao, China[D]. Qingdao: Ocean University of China, 2007. [41] Giere O. Meiobenthology: The Microscopic Motile Fauna of Aquatic Sediments[M]. Berlin, Heidelberg: Springer, 2009. [42] Fenchel T M. The ecology of micro-and meiobenthos[J]. Annual Review of Ecology and Systematics, 1978, 9: 99−121. doi: 10.1146/annurev.es.09.110178.000531 [43] Warwick R M. Nematode associations in the Exe estuary[J]. Journal of the Marine Biological Association of the United Kingdom, 1971, 51(2): 439−454. doi: 10.1017/S0025315400031908 [44] Moens T, Vincx M. Observations on the feeding ecology of estuarine nematodes[J]. Journal of the Marine Biological Association of the United Kingdom, 1997, 77(1): 211−227. doi: 10.1017/S0025315400033889 [45] Urban-Malinga B, Hedtkamp S I C, van Beusekom J E E, et al. Comparison of nematode communities in baltic and North Sea sublittoral, permeable sands–diversity and environmental control[J]. Estuarine, Coastal and Shelf Science, 2006, 70(1/2): 224−238. [46] 刘晓收, 赵瑞, 华尔, 等. 莱州湾夏季大型底栖动物群落结构特征及其与历史资料的比较[J]. 海洋通报, 2014, 33(3): 283−292.Liu Xiaoshou, Zhao Rui, Hua Er, et al. Macrofaunal community structure in the Laizhou Bay in summer and the comparison with historical data[J]. Marine Science Bulletin, 2014, 33(3): 283−292. [47] 吴金浩, 宋广军, 韩家波, 等. 渤海老铁山海域沉积物粒度分布及其对大型底栖生物群落的影响[J]. 水产科学, 2019, 38(5): 624−635.Wu Jinhao, Song Guangjun, Han Jiabo, et al. Granularity distribution of sediments and effect on macrobenthos community in Laotie Mountain area of Bohai Sea[J]. Fisheries Science, 2019, 38(5): 624−635. [48] Heininger P, Höss S, Claus E, et al. Nematode communities in contaminated river sediments[J]. Environmental Pollution, 2006, 146(1): 64−76. [49] 周艳芝. 波浪作用下海滩沉积物的动力学行为——以青岛仰口湾海水浴场为例[D]. 青岛: 中国海洋大学, 2010.Zhou Yanzhi. Dynamics behavior of beach sediment under wave action—a case study of Yangkou Bay[D]. Qingdao: Ocean University of China, 2010. [50] Gray J S, Rieger R M. A quantitative study of the meiofauna of an exposed sandy beach, at Robin Hood’s Bay, Yorkshire[J]. Journal of the Marine Biological Association of the United Kingdom, 1971, 51(1): 1−19. doi: 10.1017/S0025315400006408 [51] Barquín J, Death R G. Spatial patterns of macroinvertebrate diversity in New Zealand springbrooks and rhithral streams[J]. Journal of the North American Benthological Society, 2006, 25(4): 768−786. doi: 10.1899/0887-3593(2006)025[0768:SPOMDI]2.0.CO;2 [52] Lü Ying, Zhang Weidong, Gao Yan, et al. Preliminary study on responses of marine nematode community to crude oil contamination in intertidal zone of Bathing Beach, Dalian[J]. Marine Pollution Bulletin, 2011, 62(12): 2700−2706. doi: 10.1016/j.marpolbul.2011.09.018