长江口鱼类群落多样性及基于多元排序方法群落动态的研究
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摘要
河口位于淡水和海洋生境交汇处，生产力高、环境复杂多变，是生态学重要的生态交错区，也是生态脆弱区。长江口是中国生产力最高的渔业生态系统，同时也是世界上最大的河口之一。鱼类群落是河口生态系统的重要组成部分，如鱼类通过捕食和被捕食行为直接或间接地影响着生态系统的健康。数十年来，长江口鱼类资源由于过度捕捞、环境污染、气候变化等外界胁迫作用的影响，已经受到很大程度的扰动，因而，探明这些胁迫作用下关于长江口鱼类群落多样性、鱼类主要环境驱动因子、群落结构时空动态变化及历史变动的研究有着重要的科学意义，其成果可以为科学管理和利用长江口鱼类资源并最终保持长江口生态系统健康发展提供理论依据。
本研究通过对2010年5~6月至2011年2月长江口4个航次底拖网鱼类资源及环境因子的季度调查，首次基于多元排序方法分析了长江口全局（从淡水至海洋水体）鱼类群落结构的季度与空间分布格局及群落与环境因子的关系，探明了种类组成、生态类群、群落多样性的季节及空间分布格局及群落主要环境驱动因子；通过2010年5月至2011年4月长江口12个月张网鱼类资源的月度，分析了鱼类群落的月度时间结构特点，并探讨了不同时间尺度鱼类群落多样性分析的差异；通过对近三十年（1985~1986年；2004年；2010~2011年）3个不同时期来长江口鱼类群落分类学多样性的变动，对分类学多样性的长期变动作了探讨。
首先分析了鱼类群落季度及空间动态。结果显示季度调查共记录62种，按照科包含种类数目的多少，鳀科（7种）、鰕虎鱼科（7种）、鲤科（7种）、石首鱼科（6种）、鲀科（5种）、舌鳎科（5种）为主要的科，有19个科为单种科（只包含1个种类）。而按照生态类群，无论是种类数目和丰富度均由海洋洄游性和河口性鱼类占据主导。根据相对重要性指数IRI，IRI排序前6种鱼类的累积比例就已达92.7%，这6种优势种按比例大小排列分别是棘头梅童鱼（Collichthyslucidus）（51.9%）、矛尾鰕虎鱼（Chaeturichthys stigmatias）（20.9%）、刀鲚（Coiliaectenes）（8.5%）、龙头鱼（Harpadon nehereus (Hamilton,1822)（4.2%））、凤鲚（Coilia mystus）（4.1%）和焦氏舌鳎（Cynoglossus joyneri（3.1%）。丰度或者相对重要性指数IRI的比例，均由小型鱼类占据主导地位。空间上，丰富度（由河段水域往外逐渐降低）、丰度由河段水域往外逐渐降低及△+（由河段水域往外逐渐升高）有明显的变化趋势，说明长江河段水域不但鱼类种类数目和数量少，而这些种类在形态学上的亲缘关系也更近。空间上群落结构体现出了一定的差异性，NMDS分析显示河段水域有明显的边界，可视作淡水鱼类群落，其特征种主要由淡水鱼类（Tachysurus nitidus）及洄游性和半洄游性鱼类（刀鲚、凤鲚）构成，但其他水域鱼类群落的空间边界并不明显。而季度上，多样性指数的差异并不明显，同时鱼类群落结构在春夏秋基本保持稳定，但到冬季有较大的变化；在全局尺度内，DCCA分析显示盐度、水温及甲壳类生物量是3个与鱼类群落关系最为密切的环境因子，而空间格局主要由盐度决定，时间格局主要由水温决定。G-独立样本检验显示群落中舌鳎属或鰕虎鱼科鱼类的丰度在空间上有错位分布现象。在全局尺度上，长江口同时还是重要的鱼类（河口性鱼类）产卵场。
其次分析了鱼类群落的月度结构，结果显示月度调查共记录56种鱼类（与底拖网共有种为38种），鱼类丰度也由少数几种鱼类占据主导，月度调查优势种为棘头梅童鱼、龙头鱼、刀鲚、凤鲚、尖海龙（Syngnathus acus）、康氏小公鱼（Stolephorus commersonii），经济鱼类的比例均占有重要地位，但鱼类小型化也很严重。海洋洄游性鱼类与河口性鱼类的种类数目相似，但前者的丰度明显高于后者。这也反映出张网采样所在水域扮演的是鱼类的索饵场或者育幼场的角色。群落多样性（种类数目、均匀度、△+、△）的月度结构变化明显。过一般相邻月份群落结构也较为相似，一年中基本可以分为4组（第Ⅰ组：1~4月，第Ⅱ组：5~8月上半月，第Ⅲ组：11月下半月~12月；第Ⅳ组：8月下半月~11月上半月），但12月与1月群落结构有较大的变动。在局域尺度内，水体温度的时间变化影响了丰度的变化。鳀科的主要种类在时间上有错位分布现象。通过分析珍稀鱼类刀鲚丰度的月度分布特征，认为应该增加11月为张网休渔期。
最后分析了近三十年来三段时期（1985~1986年、2004年、2010~2011年）鱼类群落分类多样性的变化，结果显示2004年所有4个航次及2010~2011年3个航次鱼类△+显著低于长江口总名录的△+，同时统计检验显示2004年及2010~2011年各航次△+的平均值低；而各个航次鱼类Λ+的分布都没有偏离长江口鱼类总名录Λ+。此外，群落△+的减低与种类数目的变化有关，但Λ+与种类数目无关。鱼类群落分类学多样性的下降反映出长江口鱼类群落分类学关系的范围变窄，鱼类群落由亲缘关系更近的物种组成。
Estuaries are critical ecotones where fresh waters and marine environment meet, theenvironmental gradients in estuaries are very steep, and biological productivity areespecially high, estuaries are also viewed as eco-crisis regions. The Yangtze Riverestuary is the most productive fishery ecological system in China and one of thebiggest estuaries in the globe. Estuarine fish assemblage takes a key role in structuringthe system, i.g. the stability of the system has been influenced directly and indirectlyby fishes via food chain and web. Recently decades, fish community of the systemchanges significantly because of human impacts and natural disturbances such asoverfishings, pollutants, climate changes and so on, which implying that it isnecessary to know community diversity, to test key structuring factors, to examine thetemporal and spatial variations of community structure and to analyze the historicalsuccessions of the community. So, our work could provide valuable information onassessing the fish resources and maintaining the sustainable development of theYangtze River estuary ecosystem.
Firstly, fish and environmental data using bottom trawling carried across the entiresalinity gradient in the Yangtze River estuary from spring (May)2010to winter2011(February)were used, and relations in environmental variables (seven abioticinfluences and three biotic factors) and fish community, seasonal and spatial changesin fish community were firstly analyzed based on multivariate analysis. The resultsindicated that62species belonging to29families were collected, the most speciosefamilies were Enguridae (7species), Gobiidae (7species), Cyprinidae (7species),Sciaenidae (6species), Tetraodontidae (5species) and Cynoglossidae (5species),while19single family species were also captured. It is clearly that marine migrantand estuarine species dominated the fish assemblages either in total abundances or innumbers of species. Total catches were dominated by few species, for example, interms of Index of Relative importance IRI, six species (Collichthys lucidus (51.9%),Chaeturichthys stigmatias (20.9%), Coilia ectenes (8.5%), Harpadon nehereus(4.2%), Coilia mystus (4.1%), and Cynoglossus joyneri (3.1%)) were defined asdominant taxa. The increasing trends in species richness and abundance coupling with decreasing patterns in△+from riverine zone to the adjacent sea reflect that thenumbers of species and individuals are smaller, also they tend to be related moreclosely than other zones. The community structures showed spatial variation, nonmulti-dimension scaling analysis NMDS highlighted that there is a obvious boundarybetween riverine zone and other zones, and fish assemblages in this zone could bedefined as freshwater community, SIIMPER analysis further indicated that diagnosticspecies were freshwater species (Tachysurus nitidus), anadromous (C. ectenes) andsemi-anadromous (C. mystus). However, there is no clear spatial patterns among otherzones. Both Community diversity and community structure were relatively stableduring the surveys, and community structure in winter versus that in both spring andsummer did differ significantly. Detrended canonical correspondence analysis DCCAshowed that among the seven abiotic factors (temperature, salinity, depth, silt, sand,ph, grain size) and three biotic (crustacean biomass, benthos biomass, and chlorophyll)influences analyzed, salinity, temperature, and crustacean biomass were the threemost important factors driving the community, thus, spatial and temporal variationswere mainly structured by salinity and temperature, respectively. Results of G-tests ofindependence tests indicated that spatial separations occurred among the Gobiidae orCynoglossus whatever season, suggesting that one species is abundant in some sites,while the other taxon may be abundant in other sites.
Secondly, fish and environmental factors using setnet surveyed monthly from May2010to April2011were analyzed. The results showed that56species were collectedduring the study period. In terms of species, marine migrants and estuarine speciesdominated the community, while, based on abundance, the marine migrant was themost abundant guild. Total IRI were totally represented by several small-size taxa,thus, six species (H. nehereus (34.7%), C. mystus (27.5%),C. lucidus (20.0%), C.ectenes (3.1%), Syngnathus acus (2.6%),and Stolephorus commersonii (2.2%))accounting for>90.0%of the IRI. Clearly temporal differences in communitydiversity were observed. NMDS and Cluster analysis showed community structures intwo continued months are similar, however, that in December2010differs from thatin January2011, these two techniques also illustrated a clear division into four distinctgroups of samples (GroupⅠ: samples capturing from January to April; GroupⅡ:samples capturing from May to the first half month of August; Group Ⅲ: samplescapturing from the second half month of November to December; Group Ⅳ: samplessampling between the second half month of August and the first half month of November), Results of G-tests of independence tests indicated that spatial separationsoccurred among the Enguridae during the study period, suggesting that one species isabundant in some months, while the other taxon may be abundant in other months. Itis clearly that abundance of dominant species differed among months. In order tofurther protect C. ectenes, we suggest that November should also be fishing closingtime. Multi regression demonstrated that temperature was the most importantinfluencing the monthly mean abundance across the study period. When comparedwith seasonal samplings via bottom trawlings, a stronger temporal patterns incommunity structure and diversity of the monthly samplings based setnet could befound.
Finally the long-term changes in taxonomic diversity (average taxnomicdistinctness and variations in taxonomic distinctness) of fish community of theYangtze River estuary are examined. Fish species lists recorded between threedifferent periods (1985~1986,2004,2010~2011) and12surveys were used toestimate the∧~+and△+of per survey, the results showed that△+in2004and2010~2011(except spring survey) fell below the95%probability funnels for△+caculating the full list of fish species from the Yangtze River estuary, morever, thedecreasing trends in△+were also observed. However, patterns in∧~+was differentfrom△+, as all∧~+distributed in the95%probability funnels for∧~+calculating thefull list of fish species, mean∧~+didn’t differ significance between different periods.Declines in△+of the Yangtze River estuary was related to the overfishing, pollutantsand hydroprojects, suggesting that relations among species were more closely inrecent years than that in1985~1986, which implying that the niche similarity betweenspecies increases, and may cause the loss of stability or resilience of the community.
本研究通过对2010年5~6月至2011年2月长江口4个航次底拖网鱼类资源及环境因子的季度调查，首次基于多元排序方法分析了长江口全局（从淡水至海洋水体）鱼类群落结构的季度与空间分布格局及群落与环境因子的关系，探明了种类组成、生态类群、群落多样性的季节及空间分布格局及群落主要环境驱动因子；通过2010年5月至2011年4月长江口12个月张网鱼类资源的月度，分析了鱼类群落的月度时间结构特点，并探讨了不同时间尺度鱼类群落多样性分析的差异；通过对近三十年（1985~1986年；2004年；2010~2011年）3个不同时期来长江口鱼类群落分类学多样性的变动，对分类学多样性的长期变动作了探讨。
首先分析了鱼类群落季度及空间动态。结果显示季度调查共记录62种，按照科包含种类数目的多少，鳀科（7种）、鰕虎鱼科（7种）、鲤科（7种）、石首鱼科（6种）、鲀科（5种）、舌鳎科（5种）为主要的科，有19个科为单种科（只包含1个种类）。而按照生态类群，无论是种类数目和丰富度均由海洋洄游性和河口性鱼类占据主导。根据相对重要性指数IRI，IRI排序前6种鱼类的累积比例就已达92.7%，这6种优势种按比例大小排列分别是棘头梅童鱼（Collichthyslucidus）（51.9%）、矛尾鰕虎鱼（Chaeturichthys stigmatias）（20.9%）、刀鲚（Coiliaectenes）（8.5%）、龙头鱼（Harpadon nehereus (Hamilton,1822)（4.2%））、凤鲚（Coilia mystus）（4.1%）和焦氏舌鳎（Cynoglossus joyneri（3.1%）。丰度或者相对重要性指数IRI的比例，均由小型鱼类占据主导地位。空间上，丰富度（由河段水域往外逐渐降低）、丰度由河段水域往外逐渐降低及△+（由河段水域往外逐渐升高）有明显的变化趋势，说明长江河段水域不但鱼类种类数目和数量少，而这些种类在形态学上的亲缘关系也更近。空间上群落结构体现出了一定的差异性，NMDS分析显示河段水域有明显的边界，可视作淡水鱼类群落，其特征种主要由淡水鱼类（Tachysurus nitidus）及洄游性和半洄游性鱼类（刀鲚、凤鲚）构成，但其他水域鱼类群落的空间边界并不明显。而季度上，多样性指数的差异并不明显，同时鱼类群落结构在春夏秋基本保持稳定，但到冬季有较大的变化；在全局尺度内，DCCA分析显示盐度、水温及甲壳类生物量是3个与鱼类群落关系最为密切的环境因子，而空间格局主要由盐度决定，时间格局主要由水温决定。G-独立样本检验显示群落中舌鳎属或鰕虎鱼科鱼类的丰度在空间上有错位分布现象。在全局尺度上，长江口同时还是重要的鱼类（河口性鱼类）产卵场。
其次分析了鱼类群落的月度结构，结果显示月度调查共记录56种鱼类（与底拖网共有种为38种），鱼类丰度也由少数几种鱼类占据主导，月度调查优势种为棘头梅童鱼、龙头鱼、刀鲚、凤鲚、尖海龙（Syngnathus acus）、康氏小公鱼（Stolephorus commersonii），经济鱼类的比例均占有重要地位，但鱼类小型化也很严重。海洋洄游性鱼类与河口性鱼类的种类数目相似，但前者的丰度明显高于后者。这也反映出张网采样所在水域扮演的是鱼类的索饵场或者育幼场的角色。群落多样性（种类数目、均匀度、△+、△）的月度结构变化明显。过一般相邻月份群落结构也较为相似，一年中基本可以分为4组（第Ⅰ组：1~4月，第Ⅱ组：5~8月上半月，第Ⅲ组：11月下半月~12月；第Ⅳ组：8月下半月~11月上半月），但12月与1月群落结构有较大的变动。在局域尺度内，水体温度的时间变化影响了丰度的变化。鳀科的主要种类在时间上有错位分布现象。通过分析珍稀鱼类刀鲚丰度的月度分布特征，认为应该增加11月为张网休渔期。
最后分析了近三十年来三段时期（1985~1986年、2004年、2010~2011年）鱼类群落分类多样性的变化，结果显示2004年所有4个航次及2010~2011年3个航次鱼类△+显著低于长江口总名录的△+，同时统计检验显示2004年及2010~2011年各航次△+的平均值低；而各个航次鱼类Λ+的分布都没有偏离长江口鱼类总名录Λ+。此外，群落△+的减低与种类数目的变化有关，但Λ+与种类数目无关。鱼类群落分类学多样性的下降反映出长江口鱼类群落分类学关系的范围变窄，鱼类群落由亲缘关系更近的物种组成。
Estuaries are critical ecotones where fresh waters and marine environment meet, theenvironmental gradients in estuaries are very steep, and biological productivity areespecially high, estuaries are also viewed as eco-crisis regions. The Yangtze Riverestuary is the most productive fishery ecological system in China and one of thebiggest estuaries in the globe. Estuarine fish assemblage takes a key role in structuringthe system, i.g. the stability of the system has been influenced directly and indirectlyby fishes via food chain and web. Recently decades, fish community of the systemchanges significantly because of human impacts and natural disturbances such asoverfishings, pollutants, climate changes and so on, which implying that it isnecessary to know community diversity, to test key structuring factors, to examine thetemporal and spatial variations of community structure and to analyze the historicalsuccessions of the community. So, our work could provide valuable information onassessing the fish resources and maintaining the sustainable development of theYangtze River estuary ecosystem.
Firstly, fish and environmental data using bottom trawling carried across the entiresalinity gradient in the Yangtze River estuary from spring (May)2010to winter2011(February)were used, and relations in environmental variables (seven abioticinfluences and three biotic factors) and fish community, seasonal and spatial changesin fish community were firstly analyzed based on multivariate analysis. The resultsindicated that62species belonging to29families were collected, the most speciosefamilies were Enguridae (7species), Gobiidae (7species), Cyprinidae (7species),Sciaenidae (6species), Tetraodontidae (5species) and Cynoglossidae (5species),while19single family species were also captured. It is clearly that marine migrantand estuarine species dominated the fish assemblages either in total abundances or innumbers of species. Total catches were dominated by few species, for example, interms of Index of Relative importance IRI, six species (Collichthys lucidus (51.9%),Chaeturichthys stigmatias (20.9%), Coilia ectenes (8.5%), Harpadon nehereus(4.2%), Coilia mystus (4.1%), and Cynoglossus joyneri (3.1%)) were defined asdominant taxa. The increasing trends in species richness and abundance coupling with decreasing patterns in△+from riverine zone to the adjacent sea reflect that thenumbers of species and individuals are smaller, also they tend to be related moreclosely than other zones. The community structures showed spatial variation, nonmulti-dimension scaling analysis NMDS highlighted that there is a obvious boundarybetween riverine zone and other zones, and fish assemblages in this zone could bedefined as freshwater community, SIIMPER analysis further indicated that diagnosticspecies were freshwater species (Tachysurus nitidus), anadromous (C. ectenes) andsemi-anadromous (C. mystus). However, there is no clear spatial patterns among otherzones. Both Community diversity and community structure were relatively stableduring the surveys, and community structure in winter versus that in both spring andsummer did differ significantly. Detrended canonical correspondence analysis DCCAshowed that among the seven abiotic factors (temperature, salinity, depth, silt, sand,ph, grain size) and three biotic (crustacean biomass, benthos biomass, and chlorophyll)influences analyzed, salinity, temperature, and crustacean biomass were the threemost important factors driving the community, thus, spatial and temporal variationswere mainly structured by salinity and temperature, respectively. Results of G-tests ofindependence tests indicated that spatial separations occurred among the Gobiidae orCynoglossus whatever season, suggesting that one species is abundant in some sites,while the other taxon may be abundant in other sites.
Secondly, fish and environmental factors using setnet surveyed monthly from May2010to April2011were analyzed. The results showed that56species were collectedduring the study period. In terms of species, marine migrants and estuarine speciesdominated the community, while, based on abundance, the marine migrant was themost abundant guild. Total IRI were totally represented by several small-size taxa,thus, six species (H. nehereus (34.7%), C. mystus (27.5%),C. lucidus (20.0%), C.ectenes (3.1%), Syngnathus acus (2.6%),and Stolephorus commersonii (2.2%))accounting for>90.0%of the IRI. Clearly temporal differences in communitydiversity were observed. NMDS and Cluster analysis showed community structures intwo continued months are similar, however, that in December2010differs from thatin January2011, these two techniques also illustrated a clear division into four distinctgroups of samples (GroupⅠ: samples capturing from January to April; GroupⅡ:samples capturing from May to the first half month of August; Group Ⅲ: samplescapturing from the second half month of November to December; Group Ⅳ: samplessampling between the second half month of August and the first half month of November), Results of G-tests of independence tests indicated that spatial separationsoccurred among the Enguridae during the study period, suggesting that one species isabundant in some months, while the other taxon may be abundant in other months. Itis clearly that abundance of dominant species differed among months. In order tofurther protect C. ectenes, we suggest that November should also be fishing closingtime. Multi regression demonstrated that temperature was the most importantinfluencing the monthly mean abundance across the study period. When comparedwith seasonal samplings via bottom trawlings, a stronger temporal patterns incommunity structure and diversity of the monthly samplings based setnet could befound.
Finally the long-term changes in taxonomic diversity (average taxnomicdistinctness and variations in taxonomic distinctness) of fish community of theYangtze River estuary are examined. Fish species lists recorded between threedifferent periods (1985~1986,2004,2010~2011) and12surveys were used toestimate the∧~+and△+of per survey, the results showed that△+in2004and2010~2011(except spring survey) fell below the95%probability funnels for△+caculating the full list of fish species from the Yangtze River estuary, morever, thedecreasing trends in△+were also observed. However, patterns in∧~+was differentfrom△+, as all∧~+distributed in the95%probability funnels for∧~+calculating thefull list of fish species, mean∧~+didn’t differ significance between different periods.Declines in△+of the Yangtze River estuary was related to the overfishing, pollutantsand hydroprojects, suggesting that relations among species were more closely inrecent years than that in1985~1986, which implying that the niche similarity betweenspecies increases, and may cause the loss of stability or resilience of the community.
引文
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