养殖和自然生境: 中国潮间带底栖纤毛虫多样性和群落结构的对比研究

doi:10.3969/j.issn.1000-5641.2026.03.008

摘要/Abstract

摘要：

近年来, 持续的沿海开发导致潮间带自然泥滩大规模转变为养殖滩涂, 造成底栖生物多样性丧失和群落结构改变. 然而, 当前对转变后的养殖泥滩和自然泥滩在底栖生物多样性、群落结构上的差异及其驱动因素, 尤其是在大尺度空间格局下的驱动效应仍缺乏系统认知. 本研究以中国潮间带养殖和自然泥滩为研究区域, 聚焦底栖纤毛虫群落, 通过单因素方差分析 (one-way ANOVA)、线性回归模型 (LR)、基于距离的冗余分析 (dbRDA) 和变异拆分分析 (VPA), 解析了两类泥滩底栖纤毛虫三方面的α多样性 (分类多样性、功能多样性和系统发育多样性) 和群落结构 (物种组成和功能组成) 的差异及其驱动因素. 结果显示: ① 两区域的本地环境变量无显著差异, 养殖活动未显著改变生境理化条件; ② 底栖纤毛虫的α多样性、物种组成和功能组成在两类泥滩间均存在显著差异; ③ 年平均气温 (MAT) 是导致纤毛虫α多样性和群落结构在两区域呈现显著差异的主要驱动因素. 研究表明, 相较于养殖活动的影响, 大尺度气候变量是导致两类泥滩底栖纤毛虫多样性和群落结构差异的关键驱动因素. 本研究为协调潮间带滩涂养殖开发与生物多样性保护提供了参考依据.

关键词: 潮间带泥滩, 底栖纤毛虫, α多样性, 群落结构

Abstract:

In recent years, continuous coastal development has led to large-scale conversion of natural mudflats to aquaculture mudflats in intertidal zones, resulting in the loss of benthic biodiversity and alterations in community structure. However, systematic understanding of the differences in benthic biodiversity and community structure between aquaculture and natural mudflats, as well as their driving mechanisms—particularly large-scale spatial drivers—remains limited. This study focused on benthic ciliate communities in China’s intertidal aquaculture and natural mudflats. Using one-way analysis of variance (ANOVA), linear regression (LR), distance-based redundancy analysis (dbRDA), and variation partitioning analysis (VPA), we examined the differences in three dimensions of α-diversity (taxonomic, functional, and phylogenetic diversity) and community structure (species composition and functional composition) of benthic ciliates between the two types of mudflat, along with their driving factors. The results revealed that: ① no significant differences exist in local environmental variables between the two regions, indicating that aquaculture activities did not significantly modify habitat physicochemical conditions; ② significant differences existed in α-diversity, species composition, and functional composition of benthic ciliates between aquaculture and natural mudflats; ③ mean annual temperature (MAT) emerged as the primary driver of the observed differences in ciliate α-diversity and community structure. The results of this study demonstrate that large-scale climatic variables, rather than aquaculture impacts, are the key drivers of divergent patterns in benthic ciliate diversity and community structure between the two mudflat types. These findings provide critical insights for balancing intertidal mudflat aquaculture development with biodiversity conservation.

Key words: intertidal mudflats, benthic ciliates, α-diversity, community structure

中图分类号:

Q958.1

李徐, 许媛. 养殖和自然生境: 中国潮间带底栖纤毛虫多样性和群落结构的对比研究[J]. 华东师范大学学报（自然科学版）, 2026, 2026(3): 98-109.

Xu LI, Yuan XU. Aquaculture and natural habitats: Comparison of diversity and community structure of benthic ciliates in China’s intertidal zones[J]. J* E* C* N* U* N* S*, 2026, 2026(3): 98-109.

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参考文献 66

1	LEVIN L A, BOESCH D F, COVICH A, et al.. The function of marine critical transition zones and the importance of sediment biodiversity. Ecosystems, 2001, 4 (5): 430- 451.
2	MARTÍNEZ M L, INTRALAWAN A, VÁZQUEZ G, et al.. The coasts of our world: Ecological, economic and social importance. Ecological Economics, 2007, 63 (2/3): 254- 272.
3	MURRAY N J, PHINN S R, DEWITT M, et al.. The global distribution and trajectory of tidal flats. Nature, 2019, 565 (7738): 222- 225.
4	HONG C P, BURNEY J A, PONGRATZ J, et al.. Global and regional drivers of land-use emissions in 1961–2017. Nature, 2021, 589 (7843): 554- 561.
5	SMALL C, NICHOLLS R J.. A global analysis of human settlement in coastal zones. Journal of Coastal Research, 2003, 19, 584- 599.
6	WANG X X, XIAO X M, XU X, et al.. Rebound in China’s coastal wetlands following conservation and restoration. Nature Sustainability, 2021, 4 (12): 1076- 1083.
7	FENG L H, MA Y J.. Evolution of tidal flats in China and ecological exploitation of tidal flat resources. Environmental Earth Sciences, 2012, 67 (6): 1639- 1649.
8	LONG X H, LIU L P, SHAO T Y, et al.. Developing and sustainably utilize the coastal mudflat areas in China. Science of The Total Environment, 2016, 569, 1077- 1086.
9	王丹, 吴反修. 中国渔业统计年鉴2023 [M]. 北京: 中国农业出版社, 2023.
10	DAVID C P C, STA MARIA Y Y, SIRINGAN F P, et al.. Coastal pollution due to increasing nutrient flux in aquaculture sites. Environmental Geology, 2009, 58 (2): 447- 454.
11	FARMAKI E G, THOMAIDIS N S, PASIAS I N, et al.. Environmental impact of intensive aquaculture: Investigation on the accumulation of metals and nutrients in marine sediments of Greece. Science of The Total Environment, 2014, 485, 554- 562.
12	SMITH V H, SCHINDLER D W.. Eutrophication science: Where do we go from here?. Trends in Ecology & Evolution, 2009, 24 (4): 201- 207.
13	GRALL J, CHAUVAUD L.. Marine eutrophication and benthos: The need for new approaches and concepts. Global Change Biology, 2002, 8 (9): 813- 830.
14	GAO D Z, LIU M, HOU L J, et al.. Effects of shrimp-aquaculture reclamation on sediment nitrate dissimilatory reduction processes in a coastal wetland of southeastern China. Environmental Pollution, 2019, 255, 113219.
15	HUANG Y Y, LI Y M, CHEN Q, et al.. Effects of reclamation methods and habitats on macrobenthic communities and ecological health in estuarine coastal wetlands. Marine Pollution Bulletin, 2021, 168, 112420.
16	BOUCHET V M P, SAURIAU P G.. Influence of oyster culture practices and environmental conditions on the ecological status of intertidal mudflats in the Pertuis Charentais (SW France): A multi-index approach. Marine Pollution Bulletin, 2008, 56 (11): 1898- 1912.
17	FINLAY B J, CORLISS J O, ESTEBAN G, et al.. Biodiversity at the microbial level: The number of free-living ciliates in the biosphere. The Quarterly Review of Biology, 1996, 71 (2): 221- 237.
18	ROSSI F, FORSTER R M, MONTSERRAT F, et al.. Human trampling as short-term disturbance on intertidal mudflats: Effects on macrofauna biodiversity and population dynamics of bivalves. Marine Biology, 2007, 151 (6): 2077- 2090.
19	WEISSE T.. Functional diversity of aquatic ciliates. European Journal of Protistology, 2017, 61, 331- 358.
20	AZOVSKY A, MAZEI Y.. Do microbes have macroecology? Large-scale patterns in the diversity and distribution of marine benthic ciliates. Global Ecology and Biogeography, 2013, 22 (2): 163- 172.
21	MIRONOVA E, TELESH I, SKARLATO S.. Diversity and seasonality in structure of ciliate communities in the Neva Estuary (Baltic Sea). Journal of Plankton Research, 2012, 34 (3): 208- 220.
22	STOECK T, KOCHEMS R, FORSTER D, et al.. Metabarcoding of benthic ciliate communities shows high potential for environmental monitoring in salmon aquaculture. Ecological Indicators, 2018, 85, 153- 164.
23	XU Y, FAN X P, WARREN A, et al.. Functional diversity of benthic ciliate communities in response to environmental gradients in a wetland of Yangtze Estuary, China. Marine Pollution Bulletin, 2018, 127, 726- 732.
24	ZHANG J W, CHEN X Y, SOETAERT K, et al.. The relative roles of multiple drivers on benthic ciliate communities in an intertidal zone. Marine Pollution Bulletin, 2023, 187, 114510.
25	XU Y, SOININEN J, ZHANG S K, et al.. Disentangling the relative roles of natural and anthropogenic-induced stressors in shaping benthic ciliate diversity in a heavily disturbed bay. Science of the Total Environment, 2021, 801, 149683.
26	MU H W, LI X C, WEN Y N, et al.. A global record of annual terrestrial Human Footprint dataset from 2000 to 2018. Scientific Data, 2022, 9, 176.
27	SANDERSON E W, JAITEH M, LEVY M A, et al.. The human footprint and the last of the wild: The human footprint is a global map of human influence on the land surface, which suggests that human beings are stewards of nature, whether we like it or not. BioScience, 2002, 52 (10): 891- 904.
28	VENTER O, SANDERSON E W, MAGRACH A, et al.. Global terrestrial human footprint maps for 1993 and 2009. Scientific Data, 2016, 3, 160067.
29	类彦立, 徐奎栋. 海洋微型底栖生物调查规范: HY/T140–2011 [S]. 北京: 中国标准出版社, 2011.
30	王苗勋, 许媛.. 甲醛与戊二醛固定剂对基于Ludox–QPS染色方法的研究结果对比. 华东师范大学学报(自然科学版), 2022 (3): 82- 89.
31	宋微波, 沃伦 A, 胡晓钟. 中国黄渤海的自由生纤毛虫 [M]. 北京: 科学出版社, 2009.
32	LYNN D H. The Ciliated Protozoa: Characterization, Classification and Guide to the Literature[M]. 3rd ed. Dordrecht, Netherlands: Springer, 2008.
33	范鑫鹏, 潘旭明. 中国盾纤亚纲和咽膜亚纲纤毛虫 [M]. 北京: 科学出版社, 2020.
34	XU Y, SOININEN J.. Spatial patterns of functional diversity and composition in marine benthic ciliates along the coast of China. Marine Ecology Progress Series, 2019, 627, 49- 60.
35	GARNIER E, CORTEZ J, BILLÈS G, et al.. Plant functional markers capture ecosystem properties during secondary succession. Ecology, 2004, 85 (9): 2630- 2637.
36	CLARKE K R, WARWICK R M.. A taxonomic distinctness index and its statistical properties. Journal of Applied Ecology, 1998, 35 (4): 523- 531.
37	LI Z F, HEINO J, LIU Z Y, et al.. The drivers of multiple dimensions of stream macroinvertebrate beta diversity across a large montane landscape. Limnology and Oceanography, 2021, 66 (1): 226- 236.
38	何书金, 王仰麟, 罗明, 等. 中国典型地区沿海滩涂资源开发 [M]. 北京: 科学出版社, 2005.
39	HU M J, SARDANS J, YAN R B, et al.. Substantial increase in P release following conversion of coastal wetlands to aquaculture ponds from altered kinetic exchange and resupply capacity. Water Research, 2023, 230, 119586.
40	LIN S Y, ZHOU Y X, WANG W Q, et al.. Losses and destabilization of soil organic carbon stocks in coastal wetlands converted into aquaculture ponds. Global Change Biology, 2024, 30 (9): e17480.
41	WU H, PENG R H, YANG Y, et al.. Mariculture pond influence on mangrove areas in South China: Significantly larger nitrogen and phosphorus loadings from sediment wash-out than from tidal water exchange. Aquaculture, 2014, 426, 204- 212.
42	SIDIK F, KUSUMA D W, PRIYONO B, et al. Managing sediment dynamics through reintroduction of tidal flow for mangrove restoration in abandoned aquaculture ponds [M]// Dynamic Sedimentary Environments of Mangrove Coasts. Amsterdam: Elsevier, 2021: 563-582.
43	MONTANI S, MAGNI P, SHIMAMOTO M, et al.. The effect of a tidal cycle on the dynamics of nutrients in a tidal estuary in the Seto Inland Sea, Japan. Journal of Oceanography, 1998, 54 (1): 65- 76.
44	FINDLAY S, FISCHER D.. Ecosystem attributes related to tidal wetland effects on water quality. Ecology, 2013, 94 (1): 117- 125.
45	WILSON A M, MORRIS J T.. The influence of tidal forcing on groundwater flow and nutrient exchange in a salt marsh-dominated estuary. Biogeochemistry, 2012, 108 (1): 27- 38.
46	GIMMLER A, KORN R, DE VARGAS C, et al.. The Tara Oceans voyage reveals global diversity and distribution patterns of marine planktonic ciliates. Scientific Reports, 2016, 6, 33555.
47	WEISSE T, KARSTENS N, MEYER V, et al.. Niche separation in common prostome freshwater ciliates: The effect of food and temperature. Aquatic Microbial Ecology, 2001, 26, 167- 179.
48	CLARKE A, FRASER K P P.. Why does metabolism scale with temperature?. Functional Ecology, 2004, 18 (2): 243- 251.
49	FUHRMAN J A, STEELE J A, HEWSON I, et al.. A latitudinal diversity gradient in planktonic marine bacteria. Proceedings of the National Academy of Sciences of the United States of America, 2008, 105 (22): 7774- 7778.
50	FINLAY B J.. The dependence of reproductive rate on cell size and temperature in freshwater ciliated protozoa. Oecologia, 1977, 30 (1): 75- 81.
51	WEISSE T.. Physiological mortality of planktonic ciliates: Estimates, causes, and consequences. Limnology and Oceanography, 2024, 69 (3): 524- 532.
52	BROWN J H.. Why are there so many species in the tropics?. Journal of Biogeography, 2014, 41 (1): 8- 22.
53	KISSLING W D, SEKERCIOGLU C H, JETZ W.. Bird dietary guild richness across latitudes, environments and biogeographic regions. Global Ecology and Biogeography, 2012, 21 (3): 328- 340.
54	SLATYER R A, HIRST M, SEXTON J P.. Niche breadth predicts geographical range size: A general ecological pattern. Ecology Letters, 2013, 16 (8): 1104- 1114.
55	SI X F, CADOTTE M W, ZENG D, et al.. Functional and phylogenetic structure of island bird communities. Journal of Animal Ecology, 2017, 86 (3): 532- 542.
56	王善, 沈卓, 王庆, 等.. 汕头南澳海域不同养殖区浮游纤毛虫群落结构的比较. 生态学杂志, 2015, 34 (8): 2215- 2221.
57	LI J Q, XU H L, LIN X F, et al.. Colonization of periphytic ciliated protozoa on an artificial substrate in mariculture waters with notes on responses to environmental factors. Progress in Natural Science, 2009, 19 (10): 1235- 1240.
58	JACOBS P, RIEGMAN R, VAN DER MEER J.. Impact of introduced juvenile mussel cultures on the pelagic ecosystem of the western Wadden Sea, The Netherlands. Aquaculture Environment Interactions, 2016, 8, 553- 566.
59	LANGENHEDER S, RAGNARSSON H.. The role of environmental and spatial factors for the composition of aquatic bacterial communities. Ecology, 2007, 88 (9): 2154- 2161.
60	VERLEYEN E, VYVERMAN W, STERKEN M, et al.. The importance of dispersal related and local factors in shaping the taxonomic structure of diatom metacommunities. Oikos, 2009, 118 (8): 1239- 1249.
61	JYRKÄNKALLIO-MIKKOLA J, MEIER S, HEINO J, et al.. Disentangling multi-scale environmental effects on stream microbial communities. Journal of Biogeography, 2017, 44 (7): 1512- 1523.
62	SAMUELSSON K, BERGLUND J, ANDERSSON A.. Factors structuring the heterotrophic flagellate and ciliate community along a brackish water primary production gradient. Journal of Plankton Research, 2006, 28 (4): 345- 359.
63	HEINO J, TOLKKINEN M, PIRTTILÄ A M, et al.. Microbial diversity and community–environment relationships in boreal streams. Journal of Biogeography, 2014, 41 (12): 2234- 2244.
64	GRIGOROPOULOU A, SCHMIDT-KLOIBER A, MÚRRIA C.. Incongruent latitudinal patterns of taxonomic, phylogenetic and functional diversity reveal different drivers of caddisfly community assembly across spatial scales. Global Ecology and Biogeography, 2022, 31 (5): 1006- 1020.
65	SOININEN J, JAMONEAU A, ROSEBERY J, et al.. Global patterns and drivers of species and trait composition in diatoms. Global Ecology and Biogeography, 2016, 25 (8): 940- 950.
66	ATKINSON D, CIOTTI B J, MONTAGNES D J S.. Protists decrease in size linearly with temperature: ca. 2.5%℃^–1. Proceedings of the Royal Society of London Series B: Biological Sciences, 2003, 270 (1533): 2605- 2611.

响应变量	驱动因子	系数	标准误差	95%置信区间	p值
丰度	PO₄^3–	–0.846	0.247	[–1.36, –0.33]	**
分类多样性指数TD	MAT	7.674	1.450	[4.62, 10.73]	***
	Chl a	5.468	1.481	[2.34, 8.59]	**
	NO₂^–	4.974	1.363	[2.10, 7.85]	**
功能多样性指数FD	MAT	1.595	0.411	[0.73, 2.46]	**
功能多样性指数FD	SiO₃^2–	–0.954	0.411	[–1.82, –0.09]	*
系统发育多样性指数PD	MAT	0.286	0.105	[0.07, 0.51]	*
系统发育多样性指数PD	SiO₃^2–	–0.266	0.105	[–0.49, –0.05]	*

组成类型	驱动因子	单因子解释率/%	AIC值	Pseudo-F值	p值
物种组成	MAT	8.1	42.093	2.754	***
	PO₄^3–	7.7	41.117	2.740	***
	SED	2.2	41.364	1.479	0.063
功能组成	MAT	6.7	10.488	2.428	*
	PO₄^3–	5.6	7.693	4.617	**
	SED	1.5	11.615	1.309	0.256

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