长江口及邻近海域大型底栖生物群落生态学研究
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摘要
根据2006～2011年长江口及邻近海域范围内的大型底栖生物调查资料,运用方差分析、Bray-Curtis相似性聚类、多维尺度排序、典范对应分析、丰度/生物量比较曲线、大型底栖生物污染指数等方法从时间、空间、沉积物类型和溶解氧等几个不同方面定量分析了长江口及邻近海域的大型底栖生物群落结构的分布格局以及多样性特征,探讨了不同影响因素对大型底栖生物的分布和群落结构的影响,并将大型底栖生物做为目标生物,使用三种不同的评价方法评价了长江口及邻近海域的环境质量,主要研究结果如下：
从时间对大型底栖生物的影响来看：长江口海域大型底栖生物的种类组成没有显著的季节和年际变化,种类数的高值区也一直在长江口锋面附近。而优势种则随着时间的推移不断变化,群落初期占据绝对优势的竹节虫到后期逐渐消亡,被毛头犁体星虫所取代。生物量无显著的时间差异,但总丰度的时间差异显著。次级生产力也存在显著的时间差异。对于群落结构的分析可知,各个季节群落都可划分为河口群落和近岸群落两类,而夏季和冬季河口群落又可细分为内河群落和河口群落。研究的主要结果表明,长江口大型底栖生物在近年没有显著的年际变化和季节变化,但是优势种有所变化。
从空间对大型底栖生物的影响来看：将长江口划分为河口区、近岸区和远岸区进行分析,结果发现,不同区域之间的种类数、生物量和密度均存在差异显著,河口区的种类数、密度和生物量均为最低。SIMPER分析结果则表明,不同区域之间的大型底栖生物的优势种不同,其中河口区群落结构相对简单,优势种明显,各个季节优势种不同,近岸区群落结构相对复杂,优势种优势度相对不明显,其中小头虫在各个季节均占优势。远岸区各个季节优势种相对明显,且季节间不同。研究的主要结果表明,人类活动已经对长江口河口区域产生了极大影响,对近岸区也有影响,但影响的程度不大,对远岸区的影响较小。
从沉积物类型对大型底栖生物的影响来看：四种不同类型的沉积物中总共鉴定出293种大型底栖生物,各不同类型沉积物中,多毛类、软体动物均为其中的主要类群。双因素方差分析表明多毛类、软体动物、甲壳类和棘皮动物和总的物种数均存在显著的底质类型间差异,但除了多毛类外,其他类群的物种数均无显著的季节差异。根据SIMPER分析结果,不同底质类型的沉积物中优势种也有较大区别：粉砂型底质中的主要优势种为小型多毛类和甲壳动物,而砂质泥类型底质的主要优势种则为较大体型的多毛类或大型的棘皮动物和甲壳类等。双因素方差分析表明,无论生物量、密度还是多样性指数,均存在显著的底质类型间差异,生物量从低到高排序依次是粉砂<粉砂质砂<砂质泥<砂质粉砂,而丰度从低到高排序则依次是粉砂<粉砂质砂<砂质粉砂<砂质泥。群落相似性分析则表明,除了秋季以外,其余各个李节均无显著的群落差异。CCA分析表明,不同底质类型的沉积物环境所占主导因子的环境因子不同。粉砂底质的主要环境影响因素是营养盐、盐度、水深和溶解氧(负相关)；粉砂质砂区主要环境影响因素是溶解氧和pH值；砂质粉砂区主要环境影响因素是温度；而砂质泥区的主要环境影响因素是磷酸盐、硅酸盐、溶解氧和pH值。研究的主要结果表明,不同沉积物类型中食物可得性的差异、生境异质性和沉积速率是影响大型底栖生物分布的重要因素。
从溶解氧对大型底栖生物的影响来看：低氧区的多毛类和软体动物物种数、丰度和生物量均显著高于非低氧区,低氧区多毛类和软体动物物种所占百分比高于非低氧区,而甲壳类物种所占百分比则低于非低氧区。根据SIMPER分析结果,低氧区与非低氧区的主要优势种也有很大差别,特别在夏季,低氧区主要优势种为小头虫、中华异稚虫等小型多毛类；而非低氧区的主要优势种不但包括多毛类,纹尾长眼虾也成为主要优势种。聚类分析和多维尺度排序则表明,低氧区群落和非低氧区群落无显著差异,群落相似性分析结果也证明了这一点。典范对应分析结果表明低氧区多毛类的主要影响因子是盐度,甲壳类的主要影响因子是水深、温度和总有机碳等,而非低氧区多毛类的主要影响因子是盐度和水深,甲壳类的主要影响因子则是悬浮物和无机氮。年际变化的分析表明,低氧区生物量、丰度等参数无显著年际变化,但主要优势种发生变化。研究的主要结果表明,短期中度低氧不一定引起大型底栖生物总丰度的下降,但可能引起大型底栖生物群落重建,类群组成发生明显变化。
利用大型底栖生物,通过多样性指数、MPI指数和ABC曲线三种方法对长江口的环境质量进行评价,结果表明：多样性指数评价情况下,各季节重度污染的站位均在1/3以上,而属于轻度污染和清洁的站位各季节均未超过20个。MPI指数评价的情况下,各季节重度污染的站位则超过了80%,而轻度污染和清洁的站位则不超过10个,根据ABC曲线则表明,只有春季和冬季近岸区的群落处于扰动之中。研究的主要结果表明,三种方法对环境质量的评价各有优缺点,但对于低多样性的长江口海域,使用MPI指数方法可能会过高的估计污染程度。与同期调查的国内其他河口港湾相比,长江口的环境质量在国内仅好于杭州湾和椒江口,因此对于长江口海域的环境保护刻不容缓。
A quantitative analysis on the distribution pattern and diversity of macrobenthos communities in the Yangtze Estuary (YE) and adjacent waters (YEAW) was performed based on survey data from2006to2011, in terms of spatio-temporal properties, sediment types, and dissolved oxygen (DO). Several multivariate methods were applied to explore the macrobenthos distribution and community structure in relation to the environmental factors, such as the analysis of variance, Bray-Curtis similarity-based cluster, multidimensional scaling (MDS), analysis of similarity (ANOSIM), and canonical correspondence analysis (CCA). Besides, all the methods of curve for abundance and biomass comparison (ABC), macrobenthos pollution index (MPI) and diversity index were used to evaluate the environmental quality of the YEAW on the basis of the present macrobenthos data.The main results described as follows.
Regarding the temporal influence on macrobenthos, no significant seasonal and annual variations in the species composition in the YEAW were observed. The high-value region of species number located around the front in the YE. However, the dominants varied with temporal change, and Maldane sp. were found to be the dominant species at the initial stage of community development. These Polychaeta group then gradually died out at the later stage and were finally replaced by Apionsoma trichocephala. No apparent difference was observed in the biomass, whereas the total abundance and secondary productivity show remarkable temporal differences. Community structure analysis reveals that macrobenthos in each season could be divided into estuarine and inshore assemblages. The estuarine community in summer and winter can be further subdivided into riverine and estuarine groups. The results indicated no evident interannual and seasonal variations in macrobenthos in the YE in recent years.
Regarding the spatial influence on macrobenthos, the Yangtze estuary is divided into estuarine, inshore, and offshore subregion. We found that these three subregions have significant different species number, biomass, and density and the former presenting the lowest values for these community parameters. According to SIMPER analysis results, the dominant species of macrobenthos varied geographically. The community structure in the estuarine region was relatively simple with obvious dominant species, whereas the inshore region showed complicated community structure without absolute dominantsand Capitella capitata dominated in all the seasons. In the offshore region, the dominants were evidently differ in each season. The results indicate thai human activities have a huge impact on the estuanme area, some effects on the offshore area, and minimal influence on the far shore area. Regarding the influence of the sediment types on macrobenthos, a total of293species are identified in four different types of sediment. Polychaeta and mollusca were the main groups in each type of sediment. Two-way analysis of variance indicates that polychaeta, mollusca, crustacea, echinodermata significantly differ from all other groups in terms of the substrate type of sediment. However, the species of other groups except for polychaeta did not show apparent seasonal differences. SIMPER results reveal that different types of sediment are dominated bydifferent species. The dominant species in silt-substrate-typed sediment were small-sized polychaeta and crustacea, whereas the dominants in silt-mud substrate-type sediment were large-sized polychaeta, echinodermata, crustacea, and so on. Two-way analysis of variance also indicates that the biomass, density, and diversity index evidently vary with different substrate types of sediment. The biomass from low to high is silt, silt sand, silt mud, and sandy silt, in turn, whereas the abundance is silt, silt sand, sandy silt, and silt mud, in turn. Community ANOSIM indicates no evident differencein different seasons, except in autumn. CCA shows that themacrobenthos in different sediment types are affected by different environmental factors. The main factors influencing the macrobenthos in silt substrate include nutrients, salinity, water depth, and DO (negative correlation); in silt sand are DO and pH; in sandy silt is temperature; and in sandy mud are phosphate, silicate, DO, and pH. Results demonstrate that variations in food availability in different sediment types, habitat heterogeneity, and deposition rate significantly effect the macrobenthos distribution.
Regarding the influence of DO on macrobenthos, the species number, abundance, and biomass of polychaeta and mollusca in hypoxic zones are much higher than in non-hypoxic zones. A higher percentage of polychaeta and mollusca exists in hypoxic zones, whereas higher crustacea exists in non-hypoxic zones. SIMPER analysis results show that the dominant species are different between these two zones. Especially in summer, the dominant species in hypoxic zones were small-sized polychaeta (e.g., C. capitata and Heterospio sinica). However, dominant species in non-hypoxic include not only polychaeta but also the Ogyrides striaticauda. Both the analysis of clusterand MDS indicate that communities in these two zones have no significant difference, which is also confirmed through the ANOSIM. CCA results indicate that the polychaeta in hypoxic zones is principally affected by salinity, whereas crustacea is depth, temperature, total organic carbon, and so on. In non-hypoxic, the polychaeta is mainly influenced by salinity and depth, whereas crustacea is suspended matter and inorganic nitrogen. Annual variation analysis shows no, and so on. Variations in the dominant species are observed, although there was no evident annual variation in biomass and abundance. These findings indicate that a short-term, moderately low-oxygen supply does not necessarily lead to decreased total abundance of macrobenthos but can result in the community reconstruction and evident variation in assemblage composition.
Three methods (diversity index, MPI, and ABC curve) were used to study the effects of pollution on macrobenthos. Diversity index evaluation shows that the stations presented heavy pollution in each season is above one third, whereas the slight pollution and cleanness is below20stations. MPI evaluation indicates that the heavy-pollution stations in each season is>80%, whereas the influence of slight-pollution and cleanness is below10stations. ABC curve reveals that the community in the offshore area is disturbed only in spring and winter. These results indicate that the three methods used to evaluate environmental quality have both merits and demerits. However, the MPI method may overestimate the pollution degree in the YE with low diversity. Compared with other domestic estuaries and harbors investigated at the same period, environmental quality of the YE is only better than that of Hangzhou Bay and Jiaojiang Estuary. Therefore, the environmental protection in the YE allows of no delay.
从时间对大型底栖生物的影响来看：长江口海域大型底栖生物的种类组成没有显著的季节和年际变化,种类数的高值区也一直在长江口锋面附近。而优势种则随着时间的推移不断变化,群落初期占据绝对优势的竹节虫到后期逐渐消亡,被毛头犁体星虫所取代。生物量无显著的时间差异,但总丰度的时间差异显著。次级生产力也存在显著的时间差异。对于群落结构的分析可知,各个季节群落都可划分为河口群落和近岸群落两类,而夏季和冬季河口群落又可细分为内河群落和河口群落。研究的主要结果表明,长江口大型底栖生物在近年没有显著的年际变化和季节变化,但是优势种有所变化。
从空间对大型底栖生物的影响来看：将长江口划分为河口区、近岸区和远岸区进行分析,结果发现,不同区域之间的种类数、生物量和密度均存在差异显著,河口区的种类数、密度和生物量均为最低。SIMPER分析结果则表明,不同区域之间的大型底栖生物的优势种不同,其中河口区群落结构相对简单,优势种明显,各个季节优势种不同,近岸区群落结构相对复杂,优势种优势度相对不明显,其中小头虫在各个季节均占优势。远岸区各个季节优势种相对明显,且季节间不同。研究的主要结果表明,人类活动已经对长江口河口区域产生了极大影响,对近岸区也有影响,但影响的程度不大,对远岸区的影响较小。
从沉积物类型对大型底栖生物的影响来看：四种不同类型的沉积物中总共鉴定出293种大型底栖生物,各不同类型沉积物中,多毛类、软体动物均为其中的主要类群。双因素方差分析表明多毛类、软体动物、甲壳类和棘皮动物和总的物种数均存在显著的底质类型间差异,但除了多毛类外,其他类群的物种数均无显著的季节差异。根据SIMPER分析结果,不同底质类型的沉积物中优势种也有较大区别：粉砂型底质中的主要优势种为小型多毛类和甲壳动物,而砂质泥类型底质的主要优势种则为较大体型的多毛类或大型的棘皮动物和甲壳类等。双因素方差分析表明,无论生物量、密度还是多样性指数,均存在显著的底质类型间差异,生物量从低到高排序依次是粉砂<粉砂质砂<砂质泥<砂质粉砂,而丰度从低到高排序则依次是粉砂<粉砂质砂<砂质粉砂<砂质泥。群落相似性分析则表明,除了秋季以外,其余各个李节均无显著的群落差异。CCA分析表明,不同底质类型的沉积物环境所占主导因子的环境因子不同。粉砂底质的主要环境影响因素是营养盐、盐度、水深和溶解氧(负相关)；粉砂质砂区主要环境影响因素是溶解氧和pH值；砂质粉砂区主要环境影响因素是温度；而砂质泥区的主要环境影响因素是磷酸盐、硅酸盐、溶解氧和pH值。研究的主要结果表明,不同沉积物类型中食物可得性的差异、生境异质性和沉积速率是影响大型底栖生物分布的重要因素。
从溶解氧对大型底栖生物的影响来看：低氧区的多毛类和软体动物物种数、丰度和生物量均显著高于非低氧区,低氧区多毛类和软体动物物种所占百分比高于非低氧区,而甲壳类物种所占百分比则低于非低氧区。根据SIMPER分析结果,低氧区与非低氧区的主要优势种也有很大差别,特别在夏季,低氧区主要优势种为小头虫、中华异稚虫等小型多毛类；而非低氧区的主要优势种不但包括多毛类,纹尾长眼虾也成为主要优势种。聚类分析和多维尺度排序则表明,低氧区群落和非低氧区群落无显著差异,群落相似性分析结果也证明了这一点。典范对应分析结果表明低氧区多毛类的主要影响因子是盐度,甲壳类的主要影响因子是水深、温度和总有机碳等,而非低氧区多毛类的主要影响因子是盐度和水深,甲壳类的主要影响因子则是悬浮物和无机氮。年际变化的分析表明,低氧区生物量、丰度等参数无显著年际变化,但主要优势种发生变化。研究的主要结果表明,短期中度低氧不一定引起大型底栖生物总丰度的下降,但可能引起大型底栖生物群落重建,类群组成发生明显变化。
利用大型底栖生物,通过多样性指数、MPI指数和ABC曲线三种方法对长江口的环境质量进行评价,结果表明：多样性指数评价情况下,各季节重度污染的站位均在1/3以上,而属于轻度污染和清洁的站位各季节均未超过20个。MPI指数评价的情况下,各季节重度污染的站位则超过了80%,而轻度污染和清洁的站位则不超过10个,根据ABC曲线则表明,只有春季和冬季近岸区的群落处于扰动之中。研究的主要结果表明,三种方法对环境质量的评价各有优缺点,但对于低多样性的长江口海域,使用MPI指数方法可能会过高的估计污染程度。与同期调查的国内其他河口港湾相比,长江口的环境质量在国内仅好于杭州湾和椒江口,因此对于长江口海域的环境保护刻不容缓。
A quantitative analysis on the distribution pattern and diversity of macrobenthos communities in the Yangtze Estuary (YE) and adjacent waters (YEAW) was performed based on survey data from2006to2011, in terms of spatio-temporal properties, sediment types, and dissolved oxygen (DO). Several multivariate methods were applied to explore the macrobenthos distribution and community structure in relation to the environmental factors, such as the analysis of variance, Bray-Curtis similarity-based cluster, multidimensional scaling (MDS), analysis of similarity (ANOSIM), and canonical correspondence analysis (CCA). Besides, all the methods of curve for abundance and biomass comparison (ABC), macrobenthos pollution index (MPI) and diversity index were used to evaluate the environmental quality of the YEAW on the basis of the present macrobenthos data.The main results described as follows.
Regarding the temporal influence on macrobenthos, no significant seasonal and annual variations in the species composition in the YEAW were observed. The high-value region of species number located around the front in the YE. However, the dominants varied with temporal change, and Maldane sp. were found to be the dominant species at the initial stage of community development. These Polychaeta group then gradually died out at the later stage and were finally replaced by Apionsoma trichocephala. No apparent difference was observed in the biomass, whereas the total abundance and secondary productivity show remarkable temporal differences. Community structure analysis reveals that macrobenthos in each season could be divided into estuarine and inshore assemblages. The estuarine community in summer and winter can be further subdivided into riverine and estuarine groups. The results indicated no evident interannual and seasonal variations in macrobenthos in the YE in recent years.
Regarding the spatial influence on macrobenthos, the Yangtze estuary is divided into estuarine, inshore, and offshore subregion. We found that these three subregions have significant different species number, biomass, and density and the former presenting the lowest values for these community parameters. According to SIMPER analysis results, the dominant species of macrobenthos varied geographically. The community structure in the estuarine region was relatively simple with obvious dominant species, whereas the inshore region showed complicated community structure without absolute dominantsand Capitella capitata dominated in all the seasons. In the offshore region, the dominants were evidently differ in each season. The results indicate thai human activities have a huge impact on the estuanme area, some effects on the offshore area, and minimal influence on the far shore area. Regarding the influence of the sediment types on macrobenthos, a total of293species are identified in four different types of sediment. Polychaeta and mollusca were the main groups in each type of sediment. Two-way analysis of variance indicates that polychaeta, mollusca, crustacea, echinodermata significantly differ from all other groups in terms of the substrate type of sediment. However, the species of other groups except for polychaeta did not show apparent seasonal differences. SIMPER results reveal that different types of sediment are dominated bydifferent species. The dominant species in silt-substrate-typed sediment were small-sized polychaeta and crustacea, whereas the dominants in silt-mud substrate-type sediment were large-sized polychaeta, echinodermata, crustacea, and so on. Two-way analysis of variance also indicates that the biomass, density, and diversity index evidently vary with different substrate types of sediment. The biomass from low to high is silt, silt sand, silt mud, and sandy silt, in turn, whereas the abundance is silt, silt sand, sandy silt, and silt mud, in turn. Community ANOSIM indicates no evident differencein different seasons, except in autumn. CCA shows that themacrobenthos in different sediment types are affected by different environmental factors. The main factors influencing the macrobenthos in silt substrate include nutrients, salinity, water depth, and DO (negative correlation); in silt sand are DO and pH; in sandy silt is temperature; and in sandy mud are phosphate, silicate, DO, and pH. Results demonstrate that variations in food availability in different sediment types, habitat heterogeneity, and deposition rate significantly effect the macrobenthos distribution.
Regarding the influence of DO on macrobenthos, the species number, abundance, and biomass of polychaeta and mollusca in hypoxic zones are much higher than in non-hypoxic zones. A higher percentage of polychaeta and mollusca exists in hypoxic zones, whereas higher crustacea exists in non-hypoxic zones. SIMPER analysis results show that the dominant species are different between these two zones. Especially in summer, the dominant species in hypoxic zones were small-sized polychaeta (e.g., C. capitata and Heterospio sinica). However, dominant species in non-hypoxic include not only polychaeta but also the Ogyrides striaticauda. Both the analysis of clusterand MDS indicate that communities in these two zones have no significant difference, which is also confirmed through the ANOSIM. CCA results indicate that the polychaeta in hypoxic zones is principally affected by salinity, whereas crustacea is depth, temperature, total organic carbon, and so on. In non-hypoxic, the polychaeta is mainly influenced by salinity and depth, whereas crustacea is suspended matter and inorganic nitrogen. Annual variation analysis shows no, and so on. Variations in the dominant species are observed, although there was no evident annual variation in biomass and abundance. These findings indicate that a short-term, moderately low-oxygen supply does not necessarily lead to decreased total abundance of macrobenthos but can result in the community reconstruction and evident variation in assemblage composition.
Three methods (diversity index, MPI, and ABC curve) were used to study the effects of pollution on macrobenthos. Diversity index evaluation shows that the stations presented heavy pollution in each season is above one third, whereas the slight pollution and cleanness is below20stations. MPI evaluation indicates that the heavy-pollution stations in each season is>80%, whereas the influence of slight-pollution and cleanness is below10stations. ABC curve reveals that the community in the offshore area is disturbed only in spring and winter. These results indicate that the three methods used to evaluate environmental quality have both merits and demerits. However, the MPI method may overestimate the pollution degree in the YE with low diversity. Compared with other domestic estuaries and harbors investigated at the same period, environmental quality of the YE is only better than that of Hangzhou Bay and Jiaojiang Estuary. Therefore, the environmental protection in the YE allows of no delay.
引文
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