海洋纤毛虫的多样性研究：盾纤类、膜口类及腹毛类
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摘要
海洋纤毛虫作为异养原生动物,是海洋微食物网(marine microbial food web)中的重要组成部分,在海洋初-次级能量流动和碳循环(pico-、nano-级向macro-级转移)过程中起着重要的枢纽作用。相关领域的研究(水体能量动力学与碳循环、近岸生产力评估、养殖生态学、环境保护等)目前所面临的突出和焦点问题在于:处于核心地位的纤毛虫原生动物,由于鉴定资料的严重缺乏和残缺而不得不被忽略。
自二十世纪七十年代引入能够揭示其现代分类学特征(纤毛图式和银线系等)的银染方法以来,迄今关于海洋种类的现代研究资料极度匮乏,大部分已命名的“已知种类”等待确认,大量未明阶元有待鉴定和描述。
鉴于以上原因,在国家自然科学基金重点项目的资助下,作者于2005-2008年间对青岛、烟台沿海及深圳红树林自然保护区等多类生境中的盾纤类、膜口类及腹毛类纤毛虫生物多样性开展了现代标准的分类学研究,共发现、鉴定纤毛虫38种隶属20属。其中新种12种,国内新纪录15种。主要成果包括:
1.新属
●阔口虫属Eurystoma nov. gen.,根据形态学的特征将该新属归入到盾纤目,阔口虫科,其定义为:口区硕大,三片小膜紧密排列于口器顶端;口侧膜几近围口一周,胞口位于口区前半部。
2.十二新种
在详尽的活体形态学与纤毛图式研究及与已知种的充分比较的基础上,建立十二个新种,丰富了海洋纤毛虫的生物多样性:盾纤类9种(相似裂纱虫,中华裂纱虫,查皮克帆口虫,维亚可夫帆口虫,中华帆口虫,维尔伯特帆口虫,格罗利帆口虫,中华阔口虫,异指状污栖虫),腹毛类2种(浅黄后尾柱虫,中华尾刺虫),低等动基片纲1种(细长尾毛虫)。
3.十五种国内新纪录
利用现代研究方法对以下十五种纤毛虫的中国种群进行了首次报道,补充了活体及纤毛图式信息:盐鬃毛虫,神圣柔页虫,加州环须虫,砂栖突口虫,纤细具刺虫,强健纤口虫,朦胧长吻虫,蠕状心口虫,坡捞膜袋虫,双小核草履虫,埃氏盘头虫,寡毛腹针虫,阿拉腹针虫,无色腹针虫,迪惕腹梭虫。
4.对地位不明种类的廓清
对国际间分类学资料不详且长期存疑的7个种的活体信息或纤毛图式做了首次报道,对前人遗留的混乱和错误进行了修订和澄清,给出了种的新定义并指定了新模:盐鬃毛虫,神圣柔页虫,加州环须虫,砂栖突口虫,纤细具刺虫,强健纤口虫和朦胧长吻虫。
5.典型的沙隙类群的描述
国内首次对典型的沙隙类群(纤细具刺虫,寡毛腹针虫,阿拉腹针虫,无色腹针虫和迪惕腹梭虫)做了完整的形态学描述并提供了其小核糖体亚基基因序列,为这一特殊类群的系统起源分析提供了证据。
6.心口虫属的定义重修订
鉴于心口虫属在定义上迄今缺乏完整的活体和纤毛图式水平的刻画,本工作在总结前人及本研究的基础上,给出了基于现代研究的新定义:心口虫属Cardiostomatella:虫体呈长柱状,体纤毛整齐;口器与四膜虫相似,口前缝合线明显;具口后动基列;尾纤毛一至数根,伸缩泡一至多枚。
7.对12个已知种重描述
基于青岛种群对以下种类的活体及纤毛图式做了补足性描述:蠕状心口虫,坡捞膜袋虫,冠帆口虫,双小核草履虫,特氏前口虫,缩原克鲁虫,维尔伯特泡毛虫,埃氏盘头虫,寡毛腹针虫,阿拉腹针虫,无色腹针虫,迪惕腹梭虫,恩茨伸颈虫。
Marine ciliate as heterotrophic protozoa is a key component of marine microbial food web and plays an important role in the carbon cycle between the producers - secondary consumers. The current focal question on related research (kinetic energy of water and carbon cycle, coastal productivity assessment, ecology, environmental protection etc.) is: the ciliated protozoa (key groups) has to be ignored because of the deficiency and incomplete identification information.
Although the morphology study of marine ciliates has carried out for more than a century, but most study remain at the living obversation level. Since the silver-staining method was used to reveal the infraciliature and silverline system at the twenty century, lots of known forms need to be restudied using morden methods and many unknown species are waiting to be identified and described. At the same time, the development of industry and aquaculture results in the deterioration of the marine environment and the loss of coastal zone which leads to the reduction of biodiversity.
Because of the reasons above, funded by the Natural Science Foundation of China (Project No. 30430090), morphological and faunistic studies on free living ciliates (Protozoa: Ciliophora) in the coastal waters off Qingdao and Shenzhen Mangrove area (sandy beach, open sea, tidal area, maricultural ponds etc.) were carried out during the last four years. In the study, altogether 38 species belonging to 20 genera (including twelve new species and fifiteen new records) were collected and then studied using modern methods (living observations, protargol impregnation and silver carbonate impregnation methods). The achievements are summarized as follows:
●One New Genus: Eurystoma nov. gen.
According to the infraciliature, it was classified to the Order Scuticociliatida. The diagnose for this genus is as followings: Eurystomatidae with closely arranged oral membranelles including small M1, bipartite M2, and well-developed M3. Body dorso-ventrally flattened. Surface of cell faintly sculptured with silverline system in quadrangular mesh-pattern. Cytostome located at anterior 1/3 of large buccal field.
●Twelve new species were described:
Based on the detailed living morphology, infraciliature and comparion with known species, twelve new forms were founded and described:
Schizocalyptra similis, Schizocalyptra sinica, Pleuronema czapikae, Pleuronema wiackowskii, Pleuronema sinica, Pleuronema wilberti, Pleuronema grolierei, Eurystoma sinicum, Philaster apodigitiformis, Metaurostylopsis flavicans, Uronychia sinica, Urotricha elongata.
●Fifteen new records:
As the following species have never been founded in China, based on Qingdao population the redescription were made according to the complete morphology and infraciliature:
Cardistomatella vermiformis, Chaenea robusta, Condylostoma arenarium, Cyclidium ploumeuri, Discocephalus ehrenbergi, Hippocomos salinus, Kentrophoros graciles, Lacrymaria maurea, Paramecium aurelia, Peritromus californicus, Sathrophilus agitatus, Trachelocerca ditis, Tracheloraphis aragoi, Tracheloraphis discolor, Tracheloraphis oligotriata.
●Descrition of free living ciliates which could be found only in the sandy beach.
This is the first time in China to describe the psammophilic ciliates with complete morphology. And also the SSU rRNA gene, which is helpful to evaluate their sysmatic position, was sequenced: Trachelocerca ditis, Tracheloraphis aragoi, Tracheloraphis discolor, Tracheloraphis oligotriata.
●Improved diagmosis for one genus:
Since no clear definition for the genus Cardistomatella is available, a new diagnosis based on all data obtained is given: cell elongated, with uniform somatic ciliation; both pre- and postoral sutures present; buccal cavity inconspicuous; three membranelles and one C-shaped paroral membrane arranged in Tetrahymena-pattern; postoral kineties present.
●The living information or infraciliature of seven species has been revealed for the first time and their neotypes have been deposited.
Hippocomos salinus, Sathrophilus agitatus, Peritromus californicus, Condylostoma arenarium, Kentrophoros graciles, Chaenea robusta, Lacrymaria maurea.
●Redescription of twelve species:
Based on the Qingdao population, addiontial living and infraciliature information of the following twelve species was afforded:
Cardistomatella vermiformis, Cyclidium ploumeuri, Pleuronema coronatum, Paramecium aurelia, Frontonia tchibisovae, Protocruzia contrax, Tunicothrix wilberti, Discocephalus ehrenbergi, Tracheloraphis oligotriata, Tracheloraphis aragoi, Tracheloraphis discolor, Trachelocerca ditis, Trachelotractus entzi.
自二十世纪七十年代引入能够揭示其现代分类学特征(纤毛图式和银线系等)的银染方法以来,迄今关于海洋种类的现代研究资料极度匮乏,大部分已命名的“已知种类”等待确认,大量未明阶元有待鉴定和描述。
鉴于以上原因,在国家自然科学基金重点项目的资助下,作者于2005-2008年间对青岛、烟台沿海及深圳红树林自然保护区等多类生境中的盾纤类、膜口类及腹毛类纤毛虫生物多样性开展了现代标准的分类学研究,共发现、鉴定纤毛虫38种隶属20属。其中新种12种,国内新纪录15种。主要成果包括:
1.新属
●阔口虫属Eurystoma nov. gen.,根据形态学的特征将该新属归入到盾纤目,阔口虫科,其定义为:口区硕大,三片小膜紧密排列于口器顶端;口侧膜几近围口一周,胞口位于口区前半部。
2.十二新种
在详尽的活体形态学与纤毛图式研究及与已知种的充分比较的基础上,建立十二个新种,丰富了海洋纤毛虫的生物多样性:盾纤类9种(相似裂纱虫,中华裂纱虫,查皮克帆口虫,维亚可夫帆口虫,中华帆口虫,维尔伯特帆口虫,格罗利帆口虫,中华阔口虫,异指状污栖虫),腹毛类2种(浅黄后尾柱虫,中华尾刺虫),低等动基片纲1种(细长尾毛虫)。
3.十五种国内新纪录
利用现代研究方法对以下十五种纤毛虫的中国种群进行了首次报道,补充了活体及纤毛图式信息:盐鬃毛虫,神圣柔页虫,加州环须虫,砂栖突口虫,纤细具刺虫,强健纤口虫,朦胧长吻虫,蠕状心口虫,坡捞膜袋虫,双小核草履虫,埃氏盘头虫,寡毛腹针虫,阿拉腹针虫,无色腹针虫,迪惕腹梭虫。
4.对地位不明种类的廓清
对国际间分类学资料不详且长期存疑的7个种的活体信息或纤毛图式做了首次报道,对前人遗留的混乱和错误进行了修订和澄清,给出了种的新定义并指定了新模:盐鬃毛虫,神圣柔页虫,加州环须虫,砂栖突口虫,纤细具刺虫,强健纤口虫和朦胧长吻虫。
5.典型的沙隙类群的描述
国内首次对典型的沙隙类群(纤细具刺虫,寡毛腹针虫,阿拉腹针虫,无色腹针虫和迪惕腹梭虫)做了完整的形态学描述并提供了其小核糖体亚基基因序列,为这一特殊类群的系统起源分析提供了证据。
6.心口虫属的定义重修订
鉴于心口虫属在定义上迄今缺乏完整的活体和纤毛图式水平的刻画,本工作在总结前人及本研究的基础上,给出了基于现代研究的新定义:心口虫属Cardiostomatella:虫体呈长柱状,体纤毛整齐;口器与四膜虫相似,口前缝合线明显;具口后动基列;尾纤毛一至数根,伸缩泡一至多枚。
7.对12个已知种重描述
基于青岛种群对以下种类的活体及纤毛图式做了补足性描述:蠕状心口虫,坡捞膜袋虫,冠帆口虫,双小核草履虫,特氏前口虫,缩原克鲁虫,维尔伯特泡毛虫,埃氏盘头虫,寡毛腹针虫,阿拉腹针虫,无色腹针虫,迪惕腹梭虫,恩茨伸颈虫。
Marine ciliate as heterotrophic protozoa is a key component of marine microbial food web and plays an important role in the carbon cycle between the producers - secondary consumers. The current focal question on related research (kinetic energy of water and carbon cycle, coastal productivity assessment, ecology, environmental protection etc.) is: the ciliated protozoa (key groups) has to be ignored because of the deficiency and incomplete identification information.
Although the morphology study of marine ciliates has carried out for more than a century, but most study remain at the living obversation level. Since the silver-staining method was used to reveal the infraciliature and silverline system at the twenty century, lots of known forms need to be restudied using morden methods and many unknown species are waiting to be identified and described. At the same time, the development of industry and aquaculture results in the deterioration of the marine environment and the loss of coastal zone which leads to the reduction of biodiversity.
Because of the reasons above, funded by the Natural Science Foundation of China (Project No. 30430090), morphological and faunistic studies on free living ciliates (Protozoa: Ciliophora) in the coastal waters off Qingdao and Shenzhen Mangrove area (sandy beach, open sea, tidal area, maricultural ponds etc.) were carried out during the last four years. In the study, altogether 38 species belonging to 20 genera (including twelve new species and fifiteen new records) were collected and then studied using modern methods (living observations, protargol impregnation and silver carbonate impregnation methods). The achievements are summarized as follows:
●One New Genus: Eurystoma nov. gen.
According to the infraciliature, it was classified to the Order Scuticociliatida. The diagnose for this genus is as followings: Eurystomatidae with closely arranged oral membranelles including small M1, bipartite M2, and well-developed M3. Body dorso-ventrally flattened. Surface of cell faintly sculptured with silverline system in quadrangular mesh-pattern. Cytostome located at anterior 1/3 of large buccal field.
●Twelve new species were described:
Based on the detailed living morphology, infraciliature and comparion with known species, twelve new forms were founded and described:
Schizocalyptra similis, Schizocalyptra sinica, Pleuronema czapikae, Pleuronema wiackowskii, Pleuronema sinica, Pleuronema wilberti, Pleuronema grolierei, Eurystoma sinicum, Philaster apodigitiformis, Metaurostylopsis flavicans, Uronychia sinica, Urotricha elongata.
●Fifteen new records:
As the following species have never been founded in China, based on Qingdao population the redescription were made according to the complete morphology and infraciliature:
Cardistomatella vermiformis, Chaenea robusta, Condylostoma arenarium, Cyclidium ploumeuri, Discocephalus ehrenbergi, Hippocomos salinus, Kentrophoros graciles, Lacrymaria maurea, Paramecium aurelia, Peritromus californicus, Sathrophilus agitatus, Trachelocerca ditis, Tracheloraphis aragoi, Tracheloraphis discolor, Tracheloraphis oligotriata.
●Descrition of free living ciliates which could be found only in the sandy beach.
This is the first time in China to describe the psammophilic ciliates with complete morphology. And also the SSU rRNA gene, which is helpful to evaluate their sysmatic position, was sequenced: Trachelocerca ditis, Tracheloraphis aragoi, Tracheloraphis discolor, Tracheloraphis oligotriata.
●Improved diagmosis for one genus:
Since no clear definition for the genus Cardistomatella is available, a new diagnosis based on all data obtained is given: cell elongated, with uniform somatic ciliation; both pre- and postoral sutures present; buccal cavity inconspicuous; three membranelles and one C-shaped paroral membrane arranged in Tetrahymena-pattern; postoral kineties present.
●The living information or infraciliature of seven species has been revealed for the first time and their neotypes have been deposited.
Hippocomos salinus, Sathrophilus agitatus, Peritromus californicus, Condylostoma arenarium, Kentrophoros graciles, Chaenea robusta, Lacrymaria maurea.
●Redescription of twelve species:
Based on the Qingdao population, addiontial living and infraciliature information of the following twelve species was afforded:
Cardistomatella vermiformis, Cyclidium ploumeuri, Pleuronema coronatum, Paramecium aurelia, Frontonia tchibisovae, Protocruzia contrax, Tunicothrix wilberti, Discocephalus ehrenbergi, Tracheloraphis oligotriata, Tracheloraphis aragoi, Tracheloraphis discolor, Trachelocerca ditis, Trachelotractus entzi.
引文
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