长江口盐沼潮沟大型浮游动物群落生态学研究
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摘要
盐沼湿地的初级生产力通过潮汐涨退和生物输移运送到河口与近海生态系统,从而可能成为维持这些生态系统食物网中次级生产力的重要的能量基础。目前,许多科学研究已经证实大型浮游动物是河口浮游动物中主要的生物量组成部分,而且也是初级生产者和较高营养级消费者之间非常关键的中间联系。因此,认识河口盐沼大型浮游动物的群落特征以及与其它生物间的营养联系,对于理解盐沼湿地生态系统功能及其与邻近水体间物质与能量联系具有重要意义。本研究选择长江口崇明东滩盐沼湿地和九段沙盐沼湿地作为研究地点,以大型浮游动物的群落结构和输入输出估算为切入点,结合稳定同位素和胃含物分析等技术手段分析揭示大型浮游动物与其它盐沼生物间的取食关系,初步探讨了大型浮游动物在长江口盐沼湿地与河口生态系统间物质联系及交换中的作用。主要研究结果如下：
1)为了解长江口盐沼湿地大型浮游动物种类组成、多样性及时空分布格局,2010年5月和10月,使用插网作为采样工具,选择崇明东滩盐沼中盐度和浊度较高的小南港潮沟以及盐度和浊度较低的团结沙潮沟,进行了大型浮游动物的群落结构调查。共采集到大型浮游动物9目、18科、28个分类单元。在数量上占优势的是哲水蚤Calanoida、蟹类蚤状幼体Zoea larvae of crabs和短额刺糠虾Acanthomysis brevirostris,约占总捕获个体数得98%。方差分析结果表明大型浮游动物的分布具有明显的季节格局和空间差异。哲水蚤、蟹类幼体和仔稚鱼主要出现在5月,短额刺糠虾主要出现在10月。团结沙潮沟的优势类群/种类是仔稚鱼,而小南港潮沟优势类群/种类为哲水蚤、蟹类幼体和短额刺糠虾。
2)为了解大型浮游动物在盐沼湿地内的输入输出,2010年3月至2011年2月,使用帆式张网作为采样工具,开展了为期一周年的九段沙盐沼湿地潮沟大型浮游动物逐月研究,计算大型浮游动物相对流通量。周年总体而言,九段沙盐沼是大型浮游动物的源,盐沼向邻近水域有净输出。蟹类蚤状幼体、寄居蟹蚤状幼体Zoea larvae of Diogenidae、中国周眼钩虾Perioculodes meridichinensis、日I本沼虾幼体Juvenile of Macrobrachium nipponense、短额刺糠虾和尖叶大狐钩虾Grandifoxus cuspis为主要的生物输出,其中盐沼地蟹类在繁殖季节释放大量幼体对于盐沼向河口输出浮游动物生物量具有重要意义。部分类群、种类类如哲水蚤、仔稚鱼、天津厚蟹大眼幼体Megalopae of Helice tientsinensis、中华蜾赢蜚Corophium sinensis和谭氏泥蟹幼蟹Juvenile of Ilyoplax deschampsi则出现输入大于输出的现象,除了一些幼蟹或大眼幼体到盐沼生态系统完成其成体的生活史以外,哲水蚤、中华蜾赢蜚等可能为盐沼捕食者所食。
3)为了解进入盐沼生态系统的大型浮游动物的食源和互花米草入侵对其食源的影响,使用稳定同位素分析技术比较研究了崇明东滩盐沼北部互花米草入侵区域5条潮沟和南部土著植被区5条潮沟内大型浮游动物的食源。结果表明,北部潮沟中的有机颗粒物碳稳定同位素值显著高于南部潮沟,说明互花米草的存在影响了潮沟中的颗粒有机物,表明潮沟中具有来自盐沼高等植物包括互花米草分解形成的颗粒有机碎屑,是浮游动物的可能食源。北部潮沟中桡足类、仔稚鱼和虾类幼体的碳稳定同位素值显著高于南部潮沟,说明它们摄食了潮沟中来自盐沼初级生产力的颗粒有机物,而北部潮沟中的这些动物摄取了部分来自互花米草的有机质。
4)为了解盐沼潮沟大型浮游动物中的优势类群——蟹类幼体的能量基础,于2007年秋季和冬季和2008年春季和夏季,采集了互花米草和芦苇单一植被斑块内的盐沼蟹类,对优势种类进行了胃含物解剖分析。结果表明优势蟹类天津厚蟹Helice tientsinensis和无齿螳臂相手蟹Chiromantes dehaani胃含物的主要组成成分是植物性材料,分别占其食物组成的37.64-83.99%(天津厚蟹)和52.77-84.16%(无齿螳臂相手蟹)。互花米草入侵并不改变蟹类的取食特性,植物性材料仍然在这两种蟹类的胃含物中占据了最大比重。这说明蟹类通过其成体取食盐沼维管束植物,并于繁殖季节释放幼体,在利用并向河口输出盐沼有机质方面起了一定作用。
可见,有些盐沼大型浮游动物可通过摄食和阶段性的幼体释放把盐沼植物(包括外来植物)初级生产力输出到河口及外海生态系统,但也有一部分大型浮游动物类群由河口向盐沼净输入,从而将外源有机质带进盐沼生态系统。综上所述,大型浮游动物通过其自身的活动,以及与其它生物的食物网联系,在盐沼与河口/外海生态系统间的物质输移过程中扮演着重要的角色。
It is a general accepted view that the primary productions of saltmarshes could delivered to the adjacent estuary and offshore ecosystem through tide dynamic an biological migration and might become main energy resource maintaining the secondary productivity of the food web in these ecosystems. At present, a lot of previous studies have confirmed that the most biomass of zooplankton was contributed by macrozooplankton. Also many studies have revealed the intermediate linking role of macrozooplankton between primary producers and higher level consumers. Therefore, learning the community structure of salt marsh macrozooplanktons and the trophic connection between them and other biomes is very important for us to understanding the wetland ecosystem function and the linking of material and energy with adjacent water ecosystem. In this study, saltmarshes in the Chongming Island and Jiuduansha Islands of the Yangtze River estuary are selected as study sites. We estimated the input and output of the macrozooplankton in the salt marsh creeks. Meanwhile, the technology of stable isotopic analysis and stomach content analysis were used to reveal the predator and prey relationships between macrozooplankton and others organisms. Also the role of macrozooplankton in the material connection and exchange between the salt marsh and the estuarine ecosystem was preliminarily explored. The main results are listed as follows:
1) To understand the composition, diversity and the spatial and temporal pattern of macrozooplankton in saltmarshes of the Yangtze River estuary, using fyke-net, I researched the community structures of macrozooplankton in Xiaonangang creek which had higher salinity and turbidity and Tuanjiesha creek which had lower salinity and turbidity, of the Chongming Dongtan salt marsh. During the study, a total of 28 taxa belonging to 9 orders and 18 families were found from the salt marsh creeks of the Yangtze River estuary over two seasons. The most numerically abundant taxa were Calanoida, zoea larvae of crabs and Acanthomysis brevirostris, which contributing about 98% of the total abundance. The results of variance analysis showed that macrozooplankton distribution has obvious seasonal pattern and spatial difference. Calanoida, zoea larvae of crabs and fish larva mainly occurred in May. However A. brevirostris were found mainly in October. The dominant taxon of Tuanjiesha creek was fish larva. But the dominant taxa of Xiaonangang creek were Calanoida, zoea larvae of crabs and A. brevirostris.
2) To access the input and output of the macrozooplankton in the salt marsh wetland, an annual research about macrozooplankton fluxes was carried out monthly using stow net in Jiuduansha saltmarshes from March 2010 to February 2011. The relative flux of macrozooplankton was calculated through the formula of'Ferrari index'(Ⅰ). The calculation result suggested Jiuduansha salt marsh was the resource of macrozooplankton. There is net flux from Jiuduansha salt marsh to adjacent water ecosystem. Zoea larvae of crabs, zoea larvae of Diogenidae, Perioculodes meridichinensis, juvenile of Macrobrachium nipponense, A. brevirostris and Grandifoxus cuspis were the main biological output. Among them, a mass of larvae of salt marsh crabs releasing in breeding season had important significance for the total output. On the contrary, the input of other taxa, e.g. Calanoida, fish larva, Megalopae of Helice tientsinensis, Corophium sinensis and juvenile of Ilyoplax deschampsi, was more than output. I think some of juvenile crabs or megalopae of crabs coming into the salt marsh to finish its adult life cycle. And also the Calanoida and Corophium sinensis might be preyed in the salt marsh.
3) To understand the food source of macrozooplankton and the effects of the Spartina alterniflora invasion to its diet, I compared the food source of macrozooplankton collected from creeks lying in the north of Chongming Dongtan salt marsh having S. alterniflora distribution with macrozooplankton capture from creeks lying in the south of Chongming Dongtan salt marsh just having native plant distributing area through stable isotopic analysis technology. The results showed that carbon stable isotope values of POM, particulate organic matter, in the north creeks were significantly higher than in the south creeks. The reason was the invasion of S. alterniflora affected the POM in the creeks. It can be explained that there were POM coming decomposition from S. alterniflora and higher vegetation in the creeks. And the POM might be food resource of macrozooplankton. The carbon stable isotope values of Calanoida, fish larva and juvenile shrimps from the north creeks were significantly higher than that from the south creeks. The result suggested that macrozooplankton feeding POM from salt marsh primary productivity. Moreover macrozooplankton in the north creeks ingested organic matter from Sp. alterniflora.
4) In order to understand the energy basis of crab larvae, which is the dominant macrozooplankton species in salt marsh creeks, we sampled crabs from.S. alterniflora and Phragmites anstralis monoculture patches in autumn and winter of 2007 as well as spring and summer of 2008, and made stomach content analysis of the dominant species. The results indicate that the main components of the stomach content of the dominant species, Helice tientsinensis and Chiromanles dehaani arc plant material. The plant material accounts for 37.64-83.99% and 52.77-84.16% in the food composition of H. tientsinensis and C. dehaani, respectively. S. alterniflora invasion does not alter the feeding characteristics of crabs. Plant material still accounts for the largest proportion in the stomach contents of the two dominant crabs. It indicates that salt marsh crabs play a role in transport organic matters from salt marsh to estuary through the adult crab feeding on the vascular plants in salt marsh and release larvae in breeding season.
All the results shows that some salt marsh macrozooplankton could transport the primary productivity of the salt marsh plants (including exotic plant) through feeding and releasing larvae to the estuarine and offshore ecosystem. But also there were part of macrozooplankton taxa having net input from estuary to the salt marsh, thus exogenous organic matter were brought into the salt marsh ecosystem. In summary, macrozooplankton play an important role between the salt marsh and estuary offshore ecosystem material transporting processes through its own activities, as well as other biological food web links
1)为了解长江口盐沼湿地大型浮游动物种类组成、多样性及时空分布格局,2010年5月和10月,使用插网作为采样工具,选择崇明东滩盐沼中盐度和浊度较高的小南港潮沟以及盐度和浊度较低的团结沙潮沟,进行了大型浮游动物的群落结构调查。共采集到大型浮游动物9目、18科、28个分类单元。在数量上占优势的是哲水蚤Calanoida、蟹类蚤状幼体Zoea larvae of crabs和短额刺糠虾Acanthomysis brevirostris,约占总捕获个体数得98%。方差分析结果表明大型浮游动物的分布具有明显的季节格局和空间差异。哲水蚤、蟹类幼体和仔稚鱼主要出现在5月,短额刺糠虾主要出现在10月。团结沙潮沟的优势类群/种类是仔稚鱼,而小南港潮沟优势类群/种类为哲水蚤、蟹类幼体和短额刺糠虾。
2)为了解大型浮游动物在盐沼湿地内的输入输出,2010年3月至2011年2月,使用帆式张网作为采样工具,开展了为期一周年的九段沙盐沼湿地潮沟大型浮游动物逐月研究,计算大型浮游动物相对流通量。周年总体而言,九段沙盐沼是大型浮游动物的源,盐沼向邻近水域有净输出。蟹类蚤状幼体、寄居蟹蚤状幼体Zoea larvae of Diogenidae、中国周眼钩虾Perioculodes meridichinensis、日I本沼虾幼体Juvenile of Macrobrachium nipponense、短额刺糠虾和尖叶大狐钩虾Grandifoxus cuspis为主要的生物输出,其中盐沼地蟹类在繁殖季节释放大量幼体对于盐沼向河口输出浮游动物生物量具有重要意义。部分类群、种类类如哲水蚤、仔稚鱼、天津厚蟹大眼幼体Megalopae of Helice tientsinensis、中华蜾赢蜚Corophium sinensis和谭氏泥蟹幼蟹Juvenile of Ilyoplax deschampsi则出现输入大于输出的现象,除了一些幼蟹或大眼幼体到盐沼生态系统完成其成体的生活史以外,哲水蚤、中华蜾赢蜚等可能为盐沼捕食者所食。
3)为了解进入盐沼生态系统的大型浮游动物的食源和互花米草入侵对其食源的影响,使用稳定同位素分析技术比较研究了崇明东滩盐沼北部互花米草入侵区域5条潮沟和南部土著植被区5条潮沟内大型浮游动物的食源。结果表明,北部潮沟中的有机颗粒物碳稳定同位素值显著高于南部潮沟,说明互花米草的存在影响了潮沟中的颗粒有机物,表明潮沟中具有来自盐沼高等植物包括互花米草分解形成的颗粒有机碎屑,是浮游动物的可能食源。北部潮沟中桡足类、仔稚鱼和虾类幼体的碳稳定同位素值显著高于南部潮沟,说明它们摄食了潮沟中来自盐沼初级生产力的颗粒有机物,而北部潮沟中的这些动物摄取了部分来自互花米草的有机质。
4)为了解盐沼潮沟大型浮游动物中的优势类群——蟹类幼体的能量基础,于2007年秋季和冬季和2008年春季和夏季,采集了互花米草和芦苇单一植被斑块内的盐沼蟹类,对优势种类进行了胃含物解剖分析。结果表明优势蟹类天津厚蟹Helice tientsinensis和无齿螳臂相手蟹Chiromantes dehaani胃含物的主要组成成分是植物性材料,分别占其食物组成的37.64-83.99%(天津厚蟹)和52.77-84.16%(无齿螳臂相手蟹)。互花米草入侵并不改变蟹类的取食特性,植物性材料仍然在这两种蟹类的胃含物中占据了最大比重。这说明蟹类通过其成体取食盐沼维管束植物,并于繁殖季节释放幼体,在利用并向河口输出盐沼有机质方面起了一定作用。
可见,有些盐沼大型浮游动物可通过摄食和阶段性的幼体释放把盐沼植物(包括外来植物)初级生产力输出到河口及外海生态系统,但也有一部分大型浮游动物类群由河口向盐沼净输入,从而将外源有机质带进盐沼生态系统。综上所述,大型浮游动物通过其自身的活动,以及与其它生物的食物网联系,在盐沼与河口/外海生态系统间的物质输移过程中扮演着重要的角色。
It is a general accepted view that the primary productions of saltmarshes could delivered to the adjacent estuary and offshore ecosystem through tide dynamic an biological migration and might become main energy resource maintaining the secondary productivity of the food web in these ecosystems. At present, a lot of previous studies have confirmed that the most biomass of zooplankton was contributed by macrozooplankton. Also many studies have revealed the intermediate linking role of macrozooplankton between primary producers and higher level consumers. Therefore, learning the community structure of salt marsh macrozooplanktons and the trophic connection between them and other biomes is very important for us to understanding the wetland ecosystem function and the linking of material and energy with adjacent water ecosystem. In this study, saltmarshes in the Chongming Island and Jiuduansha Islands of the Yangtze River estuary are selected as study sites. We estimated the input and output of the macrozooplankton in the salt marsh creeks. Meanwhile, the technology of stable isotopic analysis and stomach content analysis were used to reveal the predator and prey relationships between macrozooplankton and others organisms. Also the role of macrozooplankton in the material connection and exchange between the salt marsh and the estuarine ecosystem was preliminarily explored. The main results are listed as follows:
1) To understand the composition, diversity and the spatial and temporal pattern of macrozooplankton in saltmarshes of the Yangtze River estuary, using fyke-net, I researched the community structures of macrozooplankton in Xiaonangang creek which had higher salinity and turbidity and Tuanjiesha creek which had lower salinity and turbidity, of the Chongming Dongtan salt marsh. During the study, a total of 28 taxa belonging to 9 orders and 18 families were found from the salt marsh creeks of the Yangtze River estuary over two seasons. The most numerically abundant taxa were Calanoida, zoea larvae of crabs and Acanthomysis brevirostris, which contributing about 98% of the total abundance. The results of variance analysis showed that macrozooplankton distribution has obvious seasonal pattern and spatial difference. Calanoida, zoea larvae of crabs and fish larva mainly occurred in May. However A. brevirostris were found mainly in October. The dominant taxon of Tuanjiesha creek was fish larva. But the dominant taxa of Xiaonangang creek were Calanoida, zoea larvae of crabs and A. brevirostris.
2) To access the input and output of the macrozooplankton in the salt marsh wetland, an annual research about macrozooplankton fluxes was carried out monthly using stow net in Jiuduansha saltmarshes from March 2010 to February 2011. The relative flux of macrozooplankton was calculated through the formula of'Ferrari index'(Ⅰ). The calculation result suggested Jiuduansha salt marsh was the resource of macrozooplankton. There is net flux from Jiuduansha salt marsh to adjacent water ecosystem. Zoea larvae of crabs, zoea larvae of Diogenidae, Perioculodes meridichinensis, juvenile of Macrobrachium nipponense, A. brevirostris and Grandifoxus cuspis were the main biological output. Among them, a mass of larvae of salt marsh crabs releasing in breeding season had important significance for the total output. On the contrary, the input of other taxa, e.g. Calanoida, fish larva, Megalopae of Helice tientsinensis, Corophium sinensis and juvenile of Ilyoplax deschampsi, was more than output. I think some of juvenile crabs or megalopae of crabs coming into the salt marsh to finish its adult life cycle. And also the Calanoida and Corophium sinensis might be preyed in the salt marsh.
3) To understand the food source of macrozooplankton and the effects of the Spartina alterniflora invasion to its diet, I compared the food source of macrozooplankton collected from creeks lying in the north of Chongming Dongtan salt marsh having S. alterniflora distribution with macrozooplankton capture from creeks lying in the south of Chongming Dongtan salt marsh just having native plant distributing area through stable isotopic analysis technology. The results showed that carbon stable isotope values of POM, particulate organic matter, in the north creeks were significantly higher than in the south creeks. The reason was the invasion of S. alterniflora affected the POM in the creeks. It can be explained that there were POM coming decomposition from S. alterniflora and higher vegetation in the creeks. And the POM might be food resource of macrozooplankton. The carbon stable isotope values of Calanoida, fish larva and juvenile shrimps from the north creeks were significantly higher than that from the south creeks. The result suggested that macrozooplankton feeding POM from salt marsh primary productivity. Moreover macrozooplankton in the north creeks ingested organic matter from Sp. alterniflora.
4) In order to understand the energy basis of crab larvae, which is the dominant macrozooplankton species in salt marsh creeks, we sampled crabs from.S. alterniflora and Phragmites anstralis monoculture patches in autumn and winter of 2007 as well as spring and summer of 2008, and made stomach content analysis of the dominant species. The results indicate that the main components of the stomach content of the dominant species, Helice tientsinensis and Chiromanles dehaani arc plant material. The plant material accounts for 37.64-83.99% and 52.77-84.16% in the food composition of H. tientsinensis and C. dehaani, respectively. S. alterniflora invasion does not alter the feeding characteristics of crabs. Plant material still accounts for the largest proportion in the stomach contents of the two dominant crabs. It indicates that salt marsh crabs play a role in transport organic matters from salt marsh to estuary through the adult crab feeding on the vascular plants in salt marsh and release larvae in breeding season.
All the results shows that some salt marsh macrozooplankton could transport the primary productivity of the salt marsh plants (including exotic plant) through feeding and releasing larvae to the estuarine and offshore ecosystem. But also there were part of macrozooplankton taxa having net input from estuary to the salt marsh, thus exogenous organic matter were brought into the salt marsh ecosystem. In summary, macrozooplankton play an important role between the salt marsh and estuary offshore ecosystem material transporting processes through its own activities, as well as other biological food web links
引文
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