中国淡水管壳缝目硅藻的分类学研究
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摘要
硅藻是藻类植物中种类最丰富的类群之一,是海洋和淡水中主要的初级生产力,在全球的碳和硅循环中具重要作用,同时在生物监测、古生态重建和地层关系上得到广泛应用。管壳缝目(Aulonoraphidinales)是硅藻植物中较进化的类群,目前我国对该类群的研究比较薄弱,仅在西藏、湖南、黑龙江等少数地区有比较系统的调查工作。
本论文在查阅和分析了管壳缝目硅藻在我国记录的详细情况的基础上,利用光学显微镜(LM)和扫描电子显微镜(SEM)着重对采自我国管壳缝目硅藻研究较薄弱的15个省区(河北、内蒙古、辽宁、吉林、黑龙江、上海、江苏、浙江、安徽、福建、湖北、湖南、云南、甘肃和新疆)的600多号硅藻标本进行显微观察和拍照,依据目前国际上广泛使用的Simonsen(1979)分类系统,结合我国淡水硅藻的研究现状,对其中的管壳缝目硅藻进行了形态分类学研究。并运用SPSS 13.0软件,对世界范围内的窗纹藻科硅藻进行了数量分类学研究。同时分析了我国淡水管壳缝目硅藻的地理分布和生态特征。研究的主要结果如下:
1.通过对600多号硅藻标本的观察,查阅国内外大量分类文献的基础上,鉴定出管壳缝目(Aulonoraphidinales)硅藻3科11属130种28变种1变型,通过文献收录管壳缝目硅藻3种6变种1变型,总计研究管壳缝目硅藻3科11属133种34变种2变型:其中杆状藻科(Bacillariaceae)5属84种10变种1变型,包括杆状藻属(Bacillaria)1种1变种、菱形藻属(Nitzschia)51种5变种、菱板藻属(Hantzschia)17种1变种1变型、盘杆藻属(Tryblionella)10种1变种、细齿藻属(Denticula)5种2变种;窗纹藻科(Epithemiaceae)2属15种15变种1变型,包括窗纹藻属(Epithemia)7种12变种1变型、棒杆藻属(Rhopalodia)8种3变种;双菱藻科(Surirellaceae)4属34种9变种,包括波缘藻属(Cymatopleura)3种5变种、双菱藻属(Surirella)25种3变种、长羽藻属(Stenopterobia)1种1变种、马鞍藻属(Campylodiscus)5种。对每一个种进行详细的特征描述,并结合光镜和扫描电镜下新发现的分类性状,对种内或种间关系进行讨论。附图版78幅,照片965张,其中光镜(LM)照片775张,扫描电镜(SEM)照片190张。
2.发现1个新种:光亮型窗纹藻(Epithemia arguiformis You & Wang);拟定3个新种,均为菱板藻属种类(尚未定名);新组合1种2变种1变型,分别是狭窄盘杆藻(Tryb.angustatula(Lange-Bertalot)nov.comb.)、渐窄盘杆藻尖变种(Tryb.angustata var.acuta(Grunow)nov.comb.)、弯曲长羽藻头端变种(Sten.curvula var.capitata(Fontell)nov.comb.)、侧生窗纹藻顶生变种二齿变型(Epithemia adnata var.proboscidea f.bidens(A.Cleve)nov.comb.);中国新记录27种5变种,隶属于3科8属。
3.首次在我国提出建立盘杆藻属(Tryblionella),该属与菱形藻属(Nitzschia)的主要区别在于:壳面粗壮,宽阔,常具褶曲,线纹常被断开。确定该属在中国有10种1变种:尖锥盘杆藻(Tryb.acuminata)、细尖盘杆藻(Tryb.apiculata)、渐窄盘杆藻(Tryb.angustata)、渐窄盘杆藻尖变种(Tryb.angustata var.acuta)、狭窄盘杆藻(Tryb.angustatula)、暖温盘杆藻(Tryb.calida)、细长盘杆藻(Tryb.gracilis)、匈牙利盘杆藻(Tryb.hungarica)、莱维迪盘杆藻(Tryb.levidensis)、岸边盘杆藻(Tryb.littoralis)和维多利亚盘杆藻(Tryb.victoriae)。支持Round et al.(1990)将盘杆藻属(Tryblionella)从菱形藻属(Nitzschia)中调出的观点。
4.选取窗纹藻科29个种和5个外群种为分类运算单位,选择包括硅藻细胞壁形态、壳缝及表面纹饰等在内的25个性状,应用SPSS 13.0的最近邻居法(Nearest neighbor)和组间连接法(Between groups method)进行分层聚类分析:结果显示34个种类分成了6大类,与传统形态分类系统基本吻合,其中细齿藻属(Denticula)种类与菱形藻属(Nitzschia)种类聚在一起,而不是与窗纹藻属(gpithemia)和棒杆藻属(Rhopalodia)种类聚在一起,这与Grunow(1862)和Simonsen(1979)的分类系统相吻合。本文在我国淡水管壳缝目硅藻的形态分类中,也采用这种分类方法:将细齿藻属(Denticula)与菱形藻属(Nitzschia)一起放入杆状藻科(Bacillariaceae)中,而不是放入窗纹藻科(Epithemiaceae)中。
5.通过对管壳缝目硅藻种类在我国各省区分布状况的统计,并结合这些种在世界范围内的分布,得出结论如下:我国淡水管壳缝目硅藻绝大部分属于世界广布种,在我国的分布有以下三种情况,一是分布较广,如线形菱形藻(Nitz.linearis)、两栖菱形藻(Nitz.amphibia)、小头端菱形藻(Nitz.capitellata)、小片菱形藻(Nitz.frustulum)、细长菱形藻(Nitz.gracilis)等,约占世界广布种数的60%;二是零星分布在几个地区,如细端菱形藻(Nitz.dissipata)、洛伦菱形藻(Nitz.lorenziana)、细尖盘杆藻(Tryb.apiculata)等,约占世界广布种数的25%;三是仅分布在1-2个地区,如平庸菱形藻(Nitz.inconspicua)、直菱板藻(Hant.virgata)等,约占世界广布种数的15%。后两种情况可能与我国藻类资源调查不足有关系;存在一些区域性分布种类,多为温带及亚热带生种类;一些首次在我国报道的新种类,是否为中国特有种,还需要进一步考证。
6.通过对中国淡水管壳缝目硅藻生态特征分析得出:管壳缝目硅藻大多属于底栖和附着生活类型,在静水或缓流水体中的种类较丰富,同时,水体中的pH值和导电率也影响种类的生态分布:在pH值呈中性或碱性的水体及导电率呈中或中-高的水体中,管壳缝目种类丰富,生物量大。
Diatoms are the most species-rich group of algae and the main primary productivity in the marine and fresh water.They are ecologically widespread and have global significance in the carbon and silicon cycles,and are used increasingly in ecological monitoring, paleoecological reconstruction and stratigraphic correlation.Aulonoraphidinales is more evolutional diatom plant,while it is study deficient in our country,only a few areas,in Tibet, Hunan,Heilongjiang et al.,had been relatively systematic investigated.
In this thesis,more than 600 samples were observed in the light microscope(LM) and scanning electron microscope(SEM),which collected from 15 provinces where the investigations of algae are relatively deficient,including Hebei,Inner Mongolia,Liaoning, Jilin,Heilongjiang,Shanghai,Jiangsu,Zhejiang,Anhui,Fujian,Hubei,Hunan,Yunnan, Gansu and Xinjiang.On the basis of the currently research situation in China and the results of samples examination,the morphological classification of Aulonoraphidinales from China was studied,according to classification system by Simonsen(1979);The numerical taxonomy of worldwide Epithemiaceae was analyzed by SPSS 13.0;Furthermore,the geographical distribution and ecological characteristics of freshwater Aulonoraphidinales from China were also analyzed.The main results are as follows:
1.Based on more than 600 samples,3 families,11 genera,130 species,28 varieties,1 forma were identified,namely Bacillariaceae(5 genera 84 species 7 varieties 1 forma), Epithemiaceae(2 genera 15 species 13 varieties) and Surirellaceae(4 genera 31 species 8 varieties).Moreover,3 species 6 varieties 1 forma were extracted from literatures.Detail characteristics of all species were described,and according to the new characters observed in the LM and SEM,intraspecific and interspecific relationships were discussed.78 plates were presented,concluding 965 photos,775 of which were LM photos,190 were SEM photos.
2.One new species was found:Epithemia arguiformis You & Wang;Three drafted new species were described,which all belong to Hantzschia(no nomenclature);New combination of one species,two varieties and one forma were found,Tryb.angustatula(Lange-Bertalot) Q. M.You & Q.X.Wang,Tryb.angustata var.acuta(Grunow) Q.M.You & Q.X.Wang,Sten. curvula var.capitata(Fontell) Q.M.You & Q.X.Wang and Epit.adnata var.proboscidea f. bidens(A.Cleve) Q.M.You & Q.X.Wang;27 species 5 varieties were new records in China, which belong to 3 families 8 genera.
3.The genus Tryblionella was firstly reported in China,it differs from the Nitzschia in which valves are robust,broad,usually undulate,striae usually interrupted by one or more sterna.10 species 1 variety were identified:Tryb.acuminata,Tryb.apiculata,Tryb. angustata,Tryb.angustata var.acuta,Tryb.angustatula,Tryb.calida,Tryb.gracilis,Tryb. hungarica,Tryb.levidensis,Tryb.littoralis and Tryb.victoriae.We support the view of classification of Round et al.(1990):Tryblionella was separated from Nitzschia as a geuns.
4.By using software SPSS 13.0,hierarchical cluster analysis of twenty-five morphological characters extracted from twenty-nine species in Epithemiaceae and five outgroup species in Bacillariaceae.The results show that:hierarchical clustering tree can be divided into six branches.It basically agrees with the traditional morphological classification; Denticula and Nitzschia were nested in one group,but not nested with Epithemia and Rhopalodia.It inosculates with classification system of Grunow(1862) and Simonsen(1979). We also adopt this system in our morphological classification.
5.Based on identified specimens and literature,the distributions of Aulonoraphidinales in China were discussed.The results show that:(1) Most of the species are cosmopolitan,the distribution in our country has the following three types:ⅰ.the species are widely distributed, e.g.Nitz.linearis,Nitz.amphibia,Nitz.capitellata,Nitz.frustulum,Nitz.gracilis etc., accounting for around 60%of cosmopolitan species;ⅱ.the species are sporadicly distributed, which only occur in several provinces,e.g.Nitz.dissipata,Nitz.lorenziana,Tryb.Apiculata etc.,accounting for around 25%of cosmopolitan species;ⅲ.the species are only distributed in one or two provinces,e.g.Nitz.inconspicua,Hant.virgata etc,accounting for around 15% of cosmopolitan species.The results of the latter two situations may be related to lack of resources investigation;(2) there are some regional species,spreading at temperate or subtropical zone;(3) some new species those first found in China,it need for further research to conform whether they are endemic species in China.
6.Analysis of ecological characteristics shows that most of Aulonoraphidinales are the benthic and epiphytic types,they are abundant in static water and slow-moving water. Moreover,the ecological distributions of species are affected by the pH and conductivity:the Aulonoraphidinales species are diverse and high biomass in the water with neutral to alkaline pH and medium to high electrolyte.
本论文在查阅和分析了管壳缝目硅藻在我国记录的详细情况的基础上,利用光学显微镜(LM)和扫描电子显微镜(SEM)着重对采自我国管壳缝目硅藻研究较薄弱的15个省区(河北、内蒙古、辽宁、吉林、黑龙江、上海、江苏、浙江、安徽、福建、湖北、湖南、云南、甘肃和新疆)的600多号硅藻标本进行显微观察和拍照,依据目前国际上广泛使用的Simonsen(1979)分类系统,结合我国淡水硅藻的研究现状,对其中的管壳缝目硅藻进行了形态分类学研究。并运用SPSS 13.0软件,对世界范围内的窗纹藻科硅藻进行了数量分类学研究。同时分析了我国淡水管壳缝目硅藻的地理分布和生态特征。研究的主要结果如下:
1.通过对600多号硅藻标本的观察,查阅国内外大量分类文献的基础上,鉴定出管壳缝目(Aulonoraphidinales)硅藻3科11属130种28变种1变型,通过文献收录管壳缝目硅藻3种6变种1变型,总计研究管壳缝目硅藻3科11属133种34变种2变型:其中杆状藻科(Bacillariaceae)5属84种10变种1变型,包括杆状藻属(Bacillaria)1种1变种、菱形藻属(Nitzschia)51种5变种、菱板藻属(Hantzschia)17种1变种1变型、盘杆藻属(Tryblionella)10种1变种、细齿藻属(Denticula)5种2变种;窗纹藻科(Epithemiaceae)2属15种15变种1变型,包括窗纹藻属(Epithemia)7种12变种1变型、棒杆藻属(Rhopalodia)8种3变种;双菱藻科(Surirellaceae)4属34种9变种,包括波缘藻属(Cymatopleura)3种5变种、双菱藻属(Surirella)25种3变种、长羽藻属(Stenopterobia)1种1变种、马鞍藻属(Campylodiscus)5种。对每一个种进行详细的特征描述,并结合光镜和扫描电镜下新发现的分类性状,对种内或种间关系进行讨论。附图版78幅,照片965张,其中光镜(LM)照片775张,扫描电镜(SEM)照片190张。
2.发现1个新种:光亮型窗纹藻(Epithemia arguiformis You & Wang);拟定3个新种,均为菱板藻属种类(尚未定名);新组合1种2变种1变型,分别是狭窄盘杆藻(Tryb.angustatula(Lange-Bertalot)nov.comb.)、渐窄盘杆藻尖变种(Tryb.angustata var.acuta(Grunow)nov.comb.)、弯曲长羽藻头端变种(Sten.curvula var.capitata(Fontell)nov.comb.)、侧生窗纹藻顶生变种二齿变型(Epithemia adnata var.proboscidea f.bidens(A.Cleve)nov.comb.);中国新记录27种5变种,隶属于3科8属。
3.首次在我国提出建立盘杆藻属(Tryblionella),该属与菱形藻属(Nitzschia)的主要区别在于:壳面粗壮,宽阔,常具褶曲,线纹常被断开。确定该属在中国有10种1变种:尖锥盘杆藻(Tryb.acuminata)、细尖盘杆藻(Tryb.apiculata)、渐窄盘杆藻(Tryb.angustata)、渐窄盘杆藻尖变种(Tryb.angustata var.acuta)、狭窄盘杆藻(Tryb.angustatula)、暖温盘杆藻(Tryb.calida)、细长盘杆藻(Tryb.gracilis)、匈牙利盘杆藻(Tryb.hungarica)、莱维迪盘杆藻(Tryb.levidensis)、岸边盘杆藻(Tryb.littoralis)和维多利亚盘杆藻(Tryb.victoriae)。支持Round et al.(1990)将盘杆藻属(Tryblionella)从菱形藻属(Nitzschia)中调出的观点。
4.选取窗纹藻科29个种和5个外群种为分类运算单位,选择包括硅藻细胞壁形态、壳缝及表面纹饰等在内的25个性状,应用SPSS 13.0的最近邻居法(Nearest neighbor)和组间连接法(Between groups method)进行分层聚类分析:结果显示34个种类分成了6大类,与传统形态分类系统基本吻合,其中细齿藻属(Denticula)种类与菱形藻属(Nitzschia)种类聚在一起,而不是与窗纹藻属(gpithemia)和棒杆藻属(Rhopalodia)种类聚在一起,这与Grunow(1862)和Simonsen(1979)的分类系统相吻合。本文在我国淡水管壳缝目硅藻的形态分类中,也采用这种分类方法:将细齿藻属(Denticula)与菱形藻属(Nitzschia)一起放入杆状藻科(Bacillariaceae)中,而不是放入窗纹藻科(Epithemiaceae)中。
5.通过对管壳缝目硅藻种类在我国各省区分布状况的统计,并结合这些种在世界范围内的分布,得出结论如下:我国淡水管壳缝目硅藻绝大部分属于世界广布种,在我国的分布有以下三种情况,一是分布较广,如线形菱形藻(Nitz.linearis)、两栖菱形藻(Nitz.amphibia)、小头端菱形藻(Nitz.capitellata)、小片菱形藻(Nitz.frustulum)、细长菱形藻(Nitz.gracilis)等,约占世界广布种数的60%;二是零星分布在几个地区,如细端菱形藻(Nitz.dissipata)、洛伦菱形藻(Nitz.lorenziana)、细尖盘杆藻(Tryb.apiculata)等,约占世界广布种数的25%;三是仅分布在1-2个地区,如平庸菱形藻(Nitz.inconspicua)、直菱板藻(Hant.virgata)等,约占世界广布种数的15%。后两种情况可能与我国藻类资源调查不足有关系;存在一些区域性分布种类,多为温带及亚热带生种类;一些首次在我国报道的新种类,是否为中国特有种,还需要进一步考证。
6.通过对中国淡水管壳缝目硅藻生态特征分析得出:管壳缝目硅藻大多属于底栖和附着生活类型,在静水或缓流水体中的种类较丰富,同时,水体中的pH值和导电率也影响种类的生态分布:在pH值呈中性或碱性的水体及导电率呈中或中-高的水体中,管壳缝目种类丰富,生物量大。
Diatoms are the most species-rich group of algae and the main primary productivity in the marine and fresh water.They are ecologically widespread and have global significance in the carbon and silicon cycles,and are used increasingly in ecological monitoring, paleoecological reconstruction and stratigraphic correlation.Aulonoraphidinales is more evolutional diatom plant,while it is study deficient in our country,only a few areas,in Tibet, Hunan,Heilongjiang et al.,had been relatively systematic investigated.
In this thesis,more than 600 samples were observed in the light microscope(LM) and scanning electron microscope(SEM),which collected from 15 provinces where the investigations of algae are relatively deficient,including Hebei,Inner Mongolia,Liaoning, Jilin,Heilongjiang,Shanghai,Jiangsu,Zhejiang,Anhui,Fujian,Hubei,Hunan,Yunnan, Gansu and Xinjiang.On the basis of the currently research situation in China and the results of samples examination,the morphological classification of Aulonoraphidinales from China was studied,according to classification system by Simonsen(1979);The numerical taxonomy of worldwide Epithemiaceae was analyzed by SPSS 13.0;Furthermore,the geographical distribution and ecological characteristics of freshwater Aulonoraphidinales from China were also analyzed.The main results are as follows:
1.Based on more than 600 samples,3 families,11 genera,130 species,28 varieties,1 forma were identified,namely Bacillariaceae(5 genera 84 species 7 varieties 1 forma), Epithemiaceae(2 genera 15 species 13 varieties) and Surirellaceae(4 genera 31 species 8 varieties).Moreover,3 species 6 varieties 1 forma were extracted from literatures.Detail characteristics of all species were described,and according to the new characters observed in the LM and SEM,intraspecific and interspecific relationships were discussed.78 plates were presented,concluding 965 photos,775 of which were LM photos,190 were SEM photos.
2.One new species was found:Epithemia arguiformis You & Wang;Three drafted new species were described,which all belong to Hantzschia(no nomenclature);New combination of one species,two varieties and one forma were found,Tryb.angustatula(Lange-Bertalot) Q. M.You & Q.X.Wang,Tryb.angustata var.acuta(Grunow) Q.M.You & Q.X.Wang,Sten. curvula var.capitata(Fontell) Q.M.You & Q.X.Wang and Epit.adnata var.proboscidea f. bidens(A.Cleve) Q.M.You & Q.X.Wang;27 species 5 varieties were new records in China, which belong to 3 families 8 genera.
3.The genus Tryblionella was firstly reported in China,it differs from the Nitzschia in which valves are robust,broad,usually undulate,striae usually interrupted by one or more sterna.10 species 1 variety were identified:Tryb.acuminata,Tryb.apiculata,Tryb. angustata,Tryb.angustata var.acuta,Tryb.angustatula,Tryb.calida,Tryb.gracilis,Tryb. hungarica,Tryb.levidensis,Tryb.littoralis and Tryb.victoriae.We support the view of classification of Round et al.(1990):Tryblionella was separated from Nitzschia as a geuns.
4.By using software SPSS 13.0,hierarchical cluster analysis of twenty-five morphological characters extracted from twenty-nine species in Epithemiaceae and five outgroup species in Bacillariaceae.The results show that:hierarchical clustering tree can be divided into six branches.It basically agrees with the traditional morphological classification; Denticula and Nitzschia were nested in one group,but not nested with Epithemia and Rhopalodia.It inosculates with classification system of Grunow(1862) and Simonsen(1979). We also adopt this system in our morphological classification.
5.Based on identified specimens and literature,the distributions of Aulonoraphidinales in China were discussed.The results show that:(1) Most of the species are cosmopolitan,the distribution in our country has the following three types:ⅰ.the species are widely distributed, e.g.Nitz.linearis,Nitz.amphibia,Nitz.capitellata,Nitz.frustulum,Nitz.gracilis etc., accounting for around 60%of cosmopolitan species;ⅱ.the species are sporadicly distributed, which only occur in several provinces,e.g.Nitz.dissipata,Nitz.lorenziana,Tryb.Apiculata etc.,accounting for around 25%of cosmopolitan species;ⅲ.the species are only distributed in one or two provinces,e.g.Nitz.inconspicua,Hant.virgata etc,accounting for around 15% of cosmopolitan species.The results of the latter two situations may be related to lack of resources investigation;(2) there are some regional species,spreading at temperate or subtropical zone;(3) some new species those first found in China,it need for further research to conform whether they are endemic species in China.
6.Analysis of ecological characteristics shows that most of Aulonoraphidinales are the benthic and epiphytic types,they are abundant in static water and slow-moving water. Moreover,the ecological distributions of species are affected by the pH and conductivity:the Aulonoraphidinales species are diverse and high biomass in the water with neutral to alkaline pH and medium to high electrolyte.
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