Article(id=1292187252554223825, tenantId=1146029695717560320, journalId=1291416733694918677, issueId=1292187163098112845, articleNumber=null, orderNo=null, doi=10.11693/hyhz20250300089, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1743350400000, receivedDateStr=2025-03-31, revisedDate=1748880000000, revisedDateStr=2025-06-03, acceptedDate=null, acceptedDateStr=null, onlineDate=1786011048687, onlineDateStr=2026-08-06, pubDate=1780070400000, pubDateStr=2026-05-30, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1786011048687, onlineIssueDateStr=2026-08-06, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1786011048687, creator=13701087609, updateTime=1786011048687, updator=13701087609, issue=Issue{id=1292187163098112845, tenantId=1146029695717560320, journalId=1291416733694918677, year='2026', volume='57', issue='3', pageStart='579', pageEnd='830', issueExtLink='null', onlineDate='null', pubDate='1780070400000', pubDateStr='2026-05-30', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1786011027358, creator='13701087609', updateTime=1786013993148, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1292199602611056777, tenantId=1146029695717560320, journalId=1291416733694918677, issueId=1292187163098112845, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1292199602611056778, tenantId=1146029695717560320, journalId=1291416733694918677, issueId=1292187163098112845, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=777, endPage=784, ext={EN=ArticleExt(id=1292187253120454866, articleId=1292187252554223825, tenantId=1146029695717560320, journalId=1291416733694918677, language=EN, title=DOMINANT SPECIES AND COMMUNITY CHARACTERISTICS OF MACROBENTHOS IN THE SHELLFISH AQUACULTURE AREA OF XIAOQING RIVER ESTUARY——ECOLOGICAL NICHE AND FUNCTIONAL GROUP ANALYSIS, columnId=1292187198095384612, journalTitle=Oceanologia et Limnologia Sinica, columnName=ARTICLES, runingTitle=null, highlight=null, articleAbstract=

To understand the impact of rainfall on the functional group characteristics of macrobenthic communities in the shellfish farming area of Xiaoqing River estuary in Laizhou Bay, Bohai Sea, survey data of rainfall were collected in July (pre-rainfall) and August (post-rainfall) 2019 and analyzed. By integrating Shannon-Wiener niche breadth, Pianka niche overlap, and canonical correspondence analysis (CCA), the variation in dominant microbenthic species and their environmental drivers were examined. Results are as follows. (1) Eight dominant species were identified, of which Moerella iridescens and Nephthys oligobranchia were consistently dominant in the two months. (2) Significant shifts in life-form and feeding functional groups were observed: July was dominated by endobenthic-planktivores (mean density 265 ind./m2, 71%), while August shifted to carnivores (mean density 103 ind./m2, 58%), with the GS/GSB ratio increasing from 0 to 2/3. (3) Dominant species exhibited marked niche breadth variation (0.494~2.945), featuring coexistence of broad-niche species (e.g., Nephthys oligobranchia) and narrow-niche specialists (e.g., Potamocorbula laevis), with weak interspecific competition (60% of species pairs showed a narrow niche overlap value (<0.3). (4) CCA revealed that the salinity (32.7% explained variance, P<0.01) was the primary driver of functional group succession (GS/GSB ratio) and niche differentiation. The stenohaline bivalve Potamocorbula laevis (optimal salinity 22~34) served as a bioindicator for salinity thresholds. This study shows that rainfall-induced environmental changes (salinity, dissolved oxygen, pH, chlorophyll a) critically drive functional group succession and niche partitioning in macrobenthos, providing a scientific basis for conservation and sustainable utilization of benthic resources in the Xiaoqing River estuary.

, authors=Yu-Ze TANG1, Xiao-Wei XU2, Jing-Kun DING3, Su-Yan XUE1, Jia-Qi LI1, Lu-Lei LIU1, Ang LI1, Yu-Ze MAO1, , authorsList=Yu-Ze TANG, Xiao-Wei XU, Jing-Kun DING, Su-Yan XUE, Jia-Qi LI, Lu-Lei LIU, Ang LI, Yu-Ze MAO, authorCompany=null, correspAuthors=Yu-Ze MAO, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1292187255028863197, articleId=1292187252554223825, tenantId=1146029695717560320, journalId=1291416733694918677, language=CN, title=小清河口贝类养殖区大型底栖动物优势种及群落特征——生态位与功能群分析, columnId=1292187198254768166, journalTitle=海洋与湖沼, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

为深入分析降雨对小清河口贝类养殖区大型底栖动物群落功能群特征的影响, 基于2019年7月(降雨前)和8月(降雨后)调查数据, 结合Shannon-Wiener生态位宽度、Pianka生态位重叠及典范对应分析(CCA), 研究了大型底栖动物优势种群落变化特征及环境驱动因素。结果表明: (1) 共鉴定8个优势种, 彩虹明樱蛤(Moerella iridescens)、寡鳃齿吻沙蚕(Nephthys oligobranchia)为两个月份共有优势种。(2) 降雨前后大型底栖动物生活型和摄食功能群结构呈现显著差异, 7月以底内型—浮游生物食者(平均密度265 ind./m2, 占比71%)为主导, 8月以肉食者为主(平均密度103 ind./m2, 占比58%), GS/GSB值由0增至2/3。(3) 优势种生态位宽度差异显著(0.494~2.945), 广生态位种(如寡鳃齿吻沙蚕Nephthys oligobranchia)与窄生态位种(如光滑河蓝蛤Potamocorbula laevis)共存, 物种间竞争较弱(60%的种对生态位重叠Qik<0.3)。(4) 典范对应分析表明, 盐度(解释率32.7%, P<0.01)是调控功能群演替(GS/GSB值)及生态位分化的首要驱动因子。(5) 光滑河蓝蛤作为狭盐种(适宜盐度22~34), 可作为盐度阈值指示种。研究结果揭示了降雨引起的环境变化(盐度、溶解氧、pH、叶绿素a)是驱动底栖动物GS/GSB值及生态位分化的重要因素, 为小清河口底栖生物资源保护与可持续利用提供科学依据。

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唐玉泽, 硕士研究生, E-mail:

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毛玉泽, 博士生导师, 研究员, E-mail:
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注: 盐度(SAL); 水温(WT); 溶氧(DO); 水深(WD); 叶绿素a (chl a)

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Dominant species of macrofauna in July and August in the Xiaoqing River estuary

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月份编号种名优势度(Y)
2019.07光滑河蓝蛤Potamocorbula laevis0.11
彩虹明樱蛤Moerella iridescens0.08
寡鳃齿吻沙蚕 Nephthys oligobranchia0.03
2019.08彩虹明樱蛤Moerella iridescens0.11
托氏䗉螺Umbonium thomasi0.02
寡鳃齿吻沙蚕 Nephthys oligobranchia0.06
尖齿拳蟹Philyra acutidens0.02
东方长眼虾Ogyrides orientalis0.03
), ArticleFig(id=1292187259575488808, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, language=CN, label=表1, caption=

小清河口近岸7~8月大型底栖动物优势种

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月份编号种名优势度(Y)
2019.07光滑河蓝蛤Potamocorbula laevis0.11
彩虹明樱蛤Moerella iridescens0.08
寡鳃齿吻沙蚕 Nephthys oligobranchia0.03
2019.08彩虹明樱蛤Moerella iridescens0.11
托氏䗉螺Umbonium thomasi0.02
寡鳃齿吻沙蚕 Nephthys oligobranchia0.06
尖齿拳蟹Philyra acutidens0.02
东方长眼虾Ogyrides orientalis0.03
), ArticleFig(id=1292187259642597673, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, language=EN, label=Tab.2, caption=

Life forms and habitat groups of dominant macrofaunal species in different months

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优势种生活型生活类群月份
光滑河蓝蛤Potamocorbula laevis底内型GSB
彩虹明樱蛤Moerella iridescens底内型GSB▲●
寡鳃齿吻沙蚕Nephthys oligobranchia底内型GSB▲●
托氏䗉螺Umbonium thomasi底上型GS
尖齿拳蟹Philyra acutidens穴居型GSB
东方长眼虾Ogyrides orientalis游泳型GS
), ArticleFig(id=1292187259713900842, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, language=CN, label=表2, caption=

不同月份大型底栖动物优势种生活型及生活类群

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优势种生活型生活类群月份
光滑河蓝蛤Potamocorbula laevis底内型GSB
彩虹明樱蛤Moerella iridescens底内型GSB▲●
寡鳃齿吻沙蚕Nephthys oligobranchia底内型GSB▲●
托氏䗉螺Umbonium thomasi底上型GS
尖齿拳蟹Philyra acutidens穴居型GSB
东方长眼虾Ogyrides orientalis游泳型GS
), ArticleFig(id=1292187259806175531, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, language=EN, label=Tab.3, caption=

Functional group and density of dominant macrofauna species in different months

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月份优势物种栖息密度/(ind./m2)功能群
7月光滑河蓝蛤Potamocorbula laevis93Pl
彩虹明樱蛤Moerella iridescens26Pl
寡鳃齿吻沙蚕Nephthys oligobranchia11C
8月彩虹明樱蛤Moerella iridescens34Pl
托氏䗉螺Umbonium thomasi23C
寡鳃齿吻沙蚕Nephthys oligobranchia18C
尖齿拳蟹Philyra acutidens12O
东方长眼虾Ogyrides orientalis12C
), ArticleFig(id=1292187259881673004, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, language=CN, label=表3, caption=

不同月份大型底栖动物群落优势种功能群及栖息密度

, figureFileSmall=null, figureFileBig=null, tableContent=
月份优势物种栖息密度/(ind./m2)功能群
7月光滑河蓝蛤Potamocorbula laevis93Pl
彩虹明樱蛤Moerella iridescens26Pl
寡鳃齿吻沙蚕Nephthys oligobranchia11C
8月彩虹明樱蛤Moerella iridescens34Pl
托氏䗉螺Umbonium thomasi23C
寡鳃齿吻沙蚕Nephthys oligobranchia18C
尖齿拳蟹Philyra acutidens12O
东方长眼虾Ogyrides orientalis12C
), ArticleFig(id=1292187259948781869, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, language=EN, label=Tab.4, caption=

The niche breadth and overlap of dominant species in macrofaunal communities

, figureFileSmall=null, figureFileBig=null, tableContent=
编号种类BiQik
S1S2S3S4S5S6
S1光滑河蓝蛤0.4941
S2寡鳃齿吻沙蚕2.9450.0041
S3彩虹明樱蛤2.9430.0120.7091
S4托氏䗉螺1.3500.0240.2030.2761
S5尖齿拳蟹2.0550.1590.5630.6130.4511
S6东方长眼虾2.1940.0010.4500.3340.0630.1801
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大型底栖动物群落优势种的生态位宽度和生态位重叠

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编号种类BiQik
S1S2S3S4S5S6
S1光滑河蓝蛤0.4941
S2寡鳃齿吻沙蚕2.9450.0041
S3彩虹明樱蛤2.9430.0120.7091
S4托氏䗉螺1.3500.0240.2030.2761
S5尖齿拳蟹2.0550.1590.5630.6130.4511
S6东方长眼虾2.1940.0010.4500.3340.0630.1801
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小清河口贝类养殖区大型底栖动物优势种及群落特征——生态位与功能群分析
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唐玉泽 1 , 徐晓玮 2 , 丁敬坤 3 , 薛素燕 1 , 李加琦 1 , 刘鲁雷 1 , 李昂 1 , 毛玉泽 1,
海洋与湖沼 | 研究论文 2026,57(3): 777-784
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海洋与湖沼 |研究论文 2026 , 57 (3) : 777 -784
小清河口贝类养殖区大型底栖动物优势种及群落特征——生态位与功能群分析
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266071, China), AuthorCompanyExt(id=1292187255251161312, tenantId=1146029695717560320, journalId=1291416733694918677, articleId=1292187252554223825, companyId=1292187255234384094, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1海水养殖生物育种与可持续产出全国重点实验室(中国水产科学研究院黄海水产研究所) 山东青岛 266071)])])]
唐玉泽1 , 徐晓玮2, 丁敬坤3, 薛素燕1, 李加琦1, 刘鲁雷1, 李昂1, 毛玉泽1,
作者信息
  • 1海水养殖生物育种与可持续产出全国重点实验室(中国水产科学研究院黄海水产研究所) 山东青岛 266071
  • 2自然资源部北海海域海岛中心 山东青岛 266061
  • 3中交水运规划设计院有限公司青岛分公司 山东青岛 266000
通讯作者:
毛玉泽, 博士生导师, 研究员, E-mail:
作者简介:

唐玉泽, 硕士研究生, E-mail:

DOMINANT SPECIES AND COMMUNITY CHARACTERISTICS OF MACROBENTHOS IN THE SHELLFISH AQUACULTURE AREA OF XIAOQING RIVER ESTUARY——ECOLOGICAL NICHE AND FUNCTIONAL GROUP ANALYSIS
Yu-Ze TANG1 , Xiao-Wei XU2, Jing-Kun DING3, Su-Yan XUE1, Jia-Qi LI1, Lu-Lei LIU1, Ang LI1, Yu-Ze MAO1,
Affiliations
  • 1State Key Laboratory of Mariculture Biobreeding and Sustainable Goods, Yellow Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences, Qingdao 266071, China
  • 2North Sea Marine and Island Affairs Center, Ministry of Natural Resources, Qingdao 266061, China
  • 3Qingdao Branch of China Communications Water Transport Planning and Design Institute Co., Ltd., Qingdao 266000, China
出版时间: 2026-05-30 doi: 10.11693/hyhz20250300089
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为深入分析降雨对小清河口贝类养殖区大型底栖动物群落功能群特征的影响, 基于2019年7月(降雨前)和8月(降雨后)调查数据, 结合Shannon-Wiener生态位宽度、Pianka生态位重叠及典范对应分析(CCA), 研究了大型底栖动物优势种群落变化特征及环境驱动因素。结果表明: (1) 共鉴定8个优势种, 彩虹明樱蛤(Moerella iridescens)、寡鳃齿吻沙蚕(Nephthys oligobranchia)为两个月份共有优势种。(2) 降雨前后大型底栖动物生活型和摄食功能群结构呈现显著差异, 7月以底内型—浮游生物食者(平均密度265 ind./m2, 占比71%)为主导, 8月以肉食者为主(平均密度103 ind./m2, 占比58%), GS/GSB值由0增至2/3。(3) 优势种生态位宽度差异显著(0.494~2.945), 广生态位种(如寡鳃齿吻沙蚕Nephthys oligobranchia)与窄生态位种(如光滑河蓝蛤Potamocorbula laevis)共存, 物种间竞争较弱(60%的种对生态位重叠Qik<0.3)。(4) 典范对应分析表明, 盐度(解释率32.7%, P<0.01)是调控功能群演替(GS/GSB值)及生态位分化的首要驱动因子。(5) 光滑河蓝蛤作为狭盐种(适宜盐度22~34), 可作为盐度阈值指示种。研究结果揭示了降雨引起的环境变化(盐度、溶解氧、pH、叶绿素a)是驱动底栖动物GS/GSB值及生态位分化的重要因素, 为小清河口底栖生物资源保护与可持续利用提供科学依据。

大型底栖动物  /  盐度  /  生态位宽度  /  GS/GSB  /  CCA

To understand the impact of rainfall on the functional group characteristics of macrobenthic communities in the shellfish farming area of Xiaoqing River estuary in Laizhou Bay, Bohai Sea, survey data of rainfall were collected in July (pre-rainfall) and August (post-rainfall) 2019 and analyzed. By integrating Shannon-Wiener niche breadth, Pianka niche overlap, and canonical correspondence analysis (CCA), the variation in dominant microbenthic species and their environmental drivers were examined. Results are as follows. (1) Eight dominant species were identified, of which Moerella iridescens and Nephthys oligobranchia were consistently dominant in the two months. (2) Significant shifts in life-form and feeding functional groups were observed: July was dominated by endobenthic-planktivores (mean density 265 ind./m2, 71%), while August shifted to carnivores (mean density 103 ind./m2, 58%), with the GS/GSB ratio increasing from 0 to 2/3. (3) Dominant species exhibited marked niche breadth variation (0.494~2.945), featuring coexistence of broad-niche species (e.g., Nephthys oligobranchia) and narrow-niche specialists (e.g., Potamocorbula laevis), with weak interspecific competition (60% of species pairs showed a narrow niche overlap value (<0.3). (4) CCA revealed that the salinity (32.7% explained variance, P<0.01) was the primary driver of functional group succession (GS/GSB ratio) and niche differentiation. The stenohaline bivalve Potamocorbula laevis (optimal salinity 22~34) served as a bioindicator for salinity thresholds. This study shows that rainfall-induced environmental changes (salinity, dissolved oxygen, pH, chlorophyll a) critically drive functional group succession and niche partitioning in macrobenthos, providing a scientific basis for conservation and sustainable utilization of benthic resources in the Xiaoqing River estuary.

macrofauna  /  salinity  /  niche breadth  /  GS/GSB  /  CCA
唐玉泽, 徐晓玮, 丁敬坤, 薛素燕, 李加琦, 刘鲁雷, 李昂, 毛玉泽. 小清河口贝类养殖区大型底栖动物优势种及群落特征——生态位与功能群分析. 海洋与湖沼, 2026 , 57 (3) : 777 -784 . DOI: 10.11693/hyhz20250300089
Yu-Ze TANG, Xiao-Wei XU, Jing-Kun DING, Su-Yan XUE, Jia-Qi LI, Lu-Lei LIU, Ang LI, Yu-Ze MAO. DOMINANT SPECIES AND COMMUNITY CHARACTERISTICS OF MACROBENTHOS IN THE SHELLFISH AQUACULTURE AREA OF XIAOQING RIVER ESTUARY——ECOLOGICAL NICHE AND FUNCTIONAL GROUP ANALYSIS[J]. Oceanologia et Limnologia Sinica, 2026 , 57 (3) : 777 -784 . DOI: 10.11693/hyhz20250300089
河口生态系统作为海洋与陆地的交错地带, 是物质循环与能量流动的关键界面, 也是全球生态系统中生物多样性最为丰富的区域之一(吴斌等, 2014)。其独特的地理位置和复杂的水动力条件, 使得河口生态系统在调节区域气候、维持生物多样性、促进营养物质交换以及支撑渔业资源等方面发挥着不可替代的作用。然而, 河口生态系统也是生态敏感性极高的区域, 其生物群落结构和功能极易受到自然和人为因素的干扰, 如盐度变化、营养盐输入、极端气候事件以及人类活动的影响。
大型底栖动物作为河口生态系统的关键功能物种, 通过生物扰动(摄食、避敌、排泄)以调控沉积物-水体界面的物质交换与能量流动, 其群落结构和功能的变化不仅反映生态系统对环境胁迫的敏感性, 还可作为生态健康的重要指示物(彭欣等, 2011; 詹启鹏等, 2023)。近年来, 随着全球气候变化和人类活动的加剧, 极端降水事件频发, 导致河口水动力条件和盐度梯度的剧烈变化, 进而对大型底栖动物群落的功能和结构产生深远影响(Grilo et al, 2011)。生态位理论作为解析群落构建机制的核心工具, 通过量化物种资源利用策略与生境适应能力, 能够有效揭示群落的竞争-共存机制(赵永强等, 2009; 张合烨等, 2024)。近年来, 基于多变量功能群与生态位研究逐渐成为生态学领域的研究热点。相比较于传统的单变量多样性研究, 多变量分析能够更全面地反映生态系统功能的稳定性(Kraft et al, 2008; Hawkins et al, 2015)。
小清河口作为莱州湾重要的河口生态系统, 其喇叭状河口地貌特征及多变的环境特征, 加剧了淡水输入后的盐度梯度效应, 成为大型底栖动物群落结构分异的主导因子(Liu et al, 2023)。尽管前期研究已揭示了该区域大型底栖动物群落的组成特征及空间分异规律(袁伟等, 2020; 丁敬坤等, 2021; 董建宇等, 2022), 但针对极端降水事件驱动下生态位和功能群演替的生态位分化机制, 仍缺乏系统研究。本研究选取2019年7月和8月(降水后盐度大幅度下降)的数据, 结合功能群分析、生态位宽度、生态位重叠及CCA分析, 探讨降雨扰动下大型底栖动物生态位与功能群的动态响应机制。研究旨在揭示极端降水事件对河口底栖生物群落结构、功能及生态位的影响, 为河口生态修复、底栖生物资源保护及经济贝类增养殖提供理论基础。
研究区域位于莱州湾小清河口附近海域(37°15′32.04″N~37°20′46.32″N, 119°2′36.66″E~119°8′54.66″E), 调查站位见图1。现场使用YSI Pro (Yellow Springs, OH, USA)测定盐度(Sal)、水温(T)、溶解氧(DO), 叶绿素a采样和分析方法按《海洋监测规范 第4部分: 海水分析》(GB 17378.4—2007)和《海洋监测规范 第7部分:近海污染生态调查和生物监测》(GB 17378.7—2007)进行。现场使用抓斗采泥器(型号为Van Veen grab 0.1 m2)采集3~4次合并为一个样品, 现场用0.5 mm网筛进行筛选, 仔细挑选大型底栖动物样品, 用5%的甲醛溶液固定, 带回实验室对样品进行鉴定分类、称重, 根据采样面积计算丰度(ind./m2)和生物量(g/m2)。样品处理等均按照《海洋调查规范 第6部分: 海洋生物调查》(中华人民共和国国家质量监督检验检疫总局等, 2008)进行。
根据优势度公式确定优势种指数(刘坤等, 2016); 优势种更替率计算公式见参考文献(徐晓群等, 2013); 采用以Shannon-Wiener多样性指数为基础的生态位宽度指数(Shannon, 1948); 采用Pianka重叠指数计算生态位重叠(Pianka, 1973)。
Y = (ni / N) × fi,
式中, Y为物种优势度; ni为第i种生物的个体数; N为研究区域内生物的总个体数; fi为该种生物在各站位出现的频率。
R=a+b-2ca+b-c×100%
式中, R为优势种更替率; ab分别为7、8月优势种的个数; c为7、8月共有优势种个数。
Bi=-j=1Rpijln Pij
式中, pij是物种ij站位的个体数与i种所有个体数的比值, 即:
pij = nij / Ni,
Ni=j=1Rnij
式中, R为环境资源点位的总数, 本文中R=38 (19个站点在2个月份, 每个取样位点相当于一个环境位点)。Bi为物种i的生态位宽度, 其阈值为[0, lnR]。Bi>2.0为广生态位种, 1.0<Bi≤2.0 为中生态位种, 0<Bi≤1.0 为窄生态位种, Bi=0单独划分为一组。
按生活型功能群特征将大型底栖动物分为底上生活功能群(GS)和底下生活功能群(GSB), 前者包括底上型和游泳型, 后者包括底内型和穴居型。GS/GSB的值可以反映生境状况对两类生活类群动物的数量影响(范航清等, 2000)。
按摄食功能群特征, 将大型底栖动物分为以下5类功能群: 浮游生物食者(planktophag-ous group, Pl)、植食者(phytophagous group, Ph)、肉食者(camivorous group, C)、杂食者(omnivorous group, O)、碎屑食者(detritivorous group, D)。
使用R 4.1.2软件中的“spaa”包及Primer 7对生态位宽度、生态位重叠等指数进行计算, 使用Canoco for Windows 5软件进行CCA分析。使用Excel计算各功能群7月和8月的平均栖息密度和平均生物量。站位分布图使用ArcGis 10.7软件进行绘制。
该海区环境呈现明显的时空分布特征, 水深范围在0.5~6 m, 因降雨的影响8月盐度出现了明显的下降, 距离河口近的站位较其他站位(S8、S9、S10、S14、S15、S16)盐度下降明显, 溶解氧亦表现出同样趋势。盐度、水温、pH总体表现出7月大于8月, 而溶解氧、叶绿素a总体表现出8月大于7月。见图2
本研究共获取8个优势种(7月3个优势种, 8月5个优势种), 其中, 彩虹明樱蛤、寡鳃齿吻沙蚕为2个月份共有优势种(表1)。
基于Bray-Curtis相似性聚类分析(图3), 研究区域大型底栖动物在26.3%相似性水平上形成两个显著分离的类群, 光滑河蓝蛤(Potamocorbula laevis)具有显著生态位特异性。7月和8月优势种更替明显(更替率为66.7%)。7月第一优势种是光滑河蓝蛤, 而8月第一优势种是彩虹明樱蛤。
该海域7月和8月优势种生活型分类见表2。7月优势种生活型均为底内型, 8月底内型、穴居型、底上型、游泳型均有分布。底内型优势种彩虹明樱蛤、寡鳃齿吻沙蚕在7月和8月均有分布。GS/GSB值表现为8月(2/3)>7月(0)。
该海域7月和8月大型底栖动物优势种的摄食功能群见表3。总物种数最多的是肉食者(3种), 占总优势种的50%, 浮游生物食者(2种)、杂食者(1种)。3种功能群的栖息密度和生物量组成见图4。按栖息密度排序依次为浮游生物食者(265 ind./m2)、肉食者(103 ind./m2)、杂食者(14 ind./m2), 按生物量排序依次为浮游生物食者(24.6 g/m2)、肉食者(5.6 g/m2)、杂食者(5.5 g/m2), 浮游生物食者的栖息密度和生物量均最高。7月浮游生物食者的种类数、生物量、栖息密度均大于8月, 而8月肉食者及杂食者的种类数、生物量、栖息密度则相反, 大于7月。
该海域7月和8月大型底栖动物优势种的生态位宽度值范围为0.494~2.945 (表4), 广生态位种4种, 为寡鳃齿吻沙蚕、彩虹明樱蛤、尖齿拳蟹、东方长眼虾; 中生态位种1种, 为托氏䗉螺; 窄生态位种1种, 为光滑河蓝蛤。
优势种种群生态位重叠值的范围为0.001~0.709 (表4)。当生态位重叠值大于0.6时, 种间生态位重叠较为显著, 大于0.6的有2组, 分别为彩虹明樱蛤和寡鳃齿吻沙蚕(0.709)、尖齿拳蟹(0.613)。光滑河蓝蛤与东方长眼虾生态位重叠值最小, 为0.001。
CCA分析了6种大型底栖动物优势种与环境因子(水深、水温、盐度、叶绿素a、pH和溶解氧)的生态位分化特征, 见图5。前两排序轴累计解释80.7%的物种变异(λ1=0.24, λ2=0.07), 能较好表征物种与环境互作关系(Ter Braak, 1987)。第一排序轴与溶解氧、叶绿素a和pH值呈正相关, 与其他环境因子呈负相关; 第二排序轴与水深、溶解氧和叶绿素a呈正相关, 与其他环境因子呈负相关。根据优势种在排序空间的分布格局(王宗兴等, 2012), 可将其划分为3个生态适应类群: Ⅰ类群(光滑河蓝蛤、彩虹明樱蛤、寡鳃齿吻沙蚕)聚集于高盐度附近, 对高盐度生境有较好的适应性; Ⅱ类群托氏䗉螺独立分布于高pH区域, 与pH呈正相关; Ⅲ类群(尖齿拳蟹与东方长眼虾)与溶解氧呈正相关(图5)。
大型底栖动物功能群特征与生境状况和物种生物学密切相关(Naeem et al, 1997)。本研究发现, 该海域7月以浮游生物食者为主(占比71%), 其中营底内生活的双壳贝类(光滑河蓝蛤、彩虹明樱蛤)优势显著, 而8月则转变为肉食者为主(占比58%), 且底内型双壳贝类的丰度大幅度下降。这一功能群更替现象可能与盐度骤降直接相关, 2019年8月10~12日台风“利奇马”过境导致小清河入海口区域降水量激增(寒亭区平均降水量达196.7 mm), 海域盐度显著降低(黄玉啟, 2022; 黄文彦等, 2024; 张端禹等, 2024)。光滑河蓝蛤作为狭盐种(适宜盐度22~34), 其渗透压调节能力在盐度低于22时急剧下降(滕炜鸣等, 2018), 导致其滤食效率降低而退出优势种行列。彩虹明樱蛤作为广盐种(适宜盐度7~42), 始终维持其优势地位(顾晓英等, 1998)。这种差异与两物种的生态位宽度特征高度吻合, 彩虹明樱蛤(Qik=2.943)与寡鳃齿吻沙蚕(Qik=2.945)作为广生态位种表现出更强的环境适应力, 而光滑河蓝蛤(Qik=0.494)的窄生态位特性使其成为盐度敏感指示种。
值得注意的是, 受“台风”利奇马过境影响, 类似水域温度、溶解氧、叶绿素a有所增加(卢龙飞等, 2020; 丰新倩等, 2022)。另外, 小清河等多条河流为该海域带来大量营养盐、有机物, 促进浮游生物的增殖, 适宜的生境和丰富的食物资源使得浮游生物食者在该海域占据优势地位(王勇等, 2002)。但盐度骤降导致滤食性贝类(如光滑河蓝蛤)生理受抑, 摄食能力下降, 其对浮游生物的捕食压力减弱, 可能间接丰富浮游生物资源, 为游泳型甲壳类(如东方长眼虾)提供更丰富的食物基础(朱根海等, 1998)。此外, 8月台风过境导致的咸淡水交换增强带来的高溶解氧环境进一步增强肉食者的代谢效率, 这些因素共同促使其在8月占据优势地位(叶丰等, 2012)。这种“功能群让渡”现象揭示了底栖动物通过生态位分化响应环境梯度的适应性策略(Taupp, 2018)。
生态位分化程度可通过重叠指数量化物种间资源利用的相似程度及潜在的竞争优势。本研究显示, 广生态位种间重叠值较高(如彩虹明樱蛤-寡鳃齿吻沙蚕Qik=0.709), 反映其资源利用方式的相似性。窄生态位种(如光滑河蓝蛤)与其他物种重叠值普遍低于0.3 (与东方长眼虾Qik≈0), 表明生态位分离显著(张零念等, 2022)。另外, 高重叠种对间仍可通过多维分化降低竞争, 彩虹明樱蛤(滤食性)与寡鳃齿吻沙蚕(杂食性)在食性上的分化, 前者为浮游生物摄食, 而后者可利用沉积物中的有机碎屑资源, 从而避免直接竞争。光滑河蓝蛤(底内型滤食者)与东方长眼虾(游泳型捕食者)则通过生活型和摄食功能的分离, 形成资源利用上的互补关系(李治霖等, 2021)。这种“食性-生境”多维分化模式(彭松耀等, 2015)有效缓解了种间竞争压力, 增强群落对环境扰动的缓冲能力(杨龙等, 2023)。
海洋环境的多变性和动力学过程使得影响大型底栖动物的环境因素极其复杂。CCA分析进一步揭示了环境因子的协同驱动作用, 光滑河蓝蛤、彩虹明樱蛤聚集于高盐生境, 尖齿拳蟹等游泳型甲壳类喜好高溶解氧生境, 托氏䗉螺则与pH显著正相关, pH<7.5时其钙化过程受抑制(王有基等, 2014)。环境因素的协同作用导致物种在资源位点上呈阶梯状分布, 形成“特化种-广适种”的特定分布格局。其中, 特化种(如光滑河蓝蛤)通过高盐依赖实现资源独占, 其丰度异常减少可作为盐度阈值突破的生物指示物。广适种(如寡鳃齿吻沙蚕)则通过生态位扩张维持群落功能稳定性。
(1) 盐度是小清河口贝类养殖区大型底栖动物功能群演变的关键驱动因子, 极端降水事件通过抑制滤食性贝类生理活动并释放浮游生物资源, 进而引发功能群更替。
(2) 广生态位种与窄生态位种通过盐度适应性、食性分化和生境分离实现共存, 群落稳定性依赖于功能群间的生态位互补。
(3) 光滑河蓝蛤作为特化种, 其丰度变化可预警盐度阈值。广适种通过生态位扩张维持群落功能稳定性, 为河口生态系统健康管理提供参考。

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doi: 10.11693/hyhz20250300089
  • 接收时间:2025-03-31
  • 首发时间:2026-08-06
  • 出版时间:2026-05-30
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  • 收稿日期:2025-03-31
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    1海水养殖生物育种与可持续产出全国重点实验室(中国水产科学研究院黄海水产研究所) 山东青岛 266071
    2自然资源部北海海域海岛中心 山东青岛 266061
    3中交水运规划设计院有限公司青岛分公司 山东青岛 266000

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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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