【作者】 丰美萍；

【导师】 肖天； 张武昌；

【作者基本信息】 中国科学院研究生院（海洋研究所） ， 海洋生态学， 2014， 博士

【摘要】 微型浮游动物是海洋浮游生态系统的重要部分，砂壳纤毛虫是其中的一个重要类群，可作为海洋浮游生态研究的模式种，其生态学的研究具有重要意义。本论文以海洋砂壳纤毛虫为研究对象，汇总了以往所有砂壳纤毛虫资料，厘清并理顺分类系统，汇集图谱资料形成《砂壳纤毛虫图谱》一书。以此为基础，研究了多个海区(北极王湾、吐噶喇海峡、西里伯斯海附近、哥斯达黎加近海、南海、胶州湾)的砂壳纤毛虫群落的生态分布，并在胶州湾进行了砂壳纤毛虫群落长期变化(2003–2012年)的监测和研究。砂壳纤毛虫种名录和图谱砂壳纤毛虫是纤毛虫中种类最多的目(Lynn,2008)，迄今为止已累计大量资料，但没有一份详细系统的名录，且分类系统存在分歧和混乱。本论文总结了自Kofoid&Campbell (1929)以来的分类学资料，参照Lynn (2008)的分类系统，整理完成世界砂壳纤毛虫种名录，并整理图谱资料，出版了《砂壳纤毛虫图谱》，此后建设了网站Planktonic Ciliate Project。厘清世界今生砂壳纤毛虫15科67属931种，世界海洋种共计15科67属900种，淡水种共计4科6属31种。我国水体共记录有13科36属163种，海洋种13科36属143种，淡水种2科3属20种。除9个种(详见第二章)外，其他淡水种在我国都有记录。在所有属中，拟铃虫属(Tintinnopsis)包含种类数最多，为141种。该名录中，除了已有中文名的159种外，其余的种我们给出了中文译名，并将对有分歧的中文译名进行了统一。我们在长江口海域发现砂壳纤毛虫一新种，定名为河口拟铃虫(Tintinnopsisestuariensis)。该种壳体为不透明黏着壳，通体覆盖矿物质颗粒，两端开口，壳体上半部分呈圆筒状，下半部分侧扁，壳体长度120–180μm，口径50–75μm，属于拟铃虫属。该种区别于同属其他相似种类的特点是壳体下半部分侧扁、下部开口为垂直开口，口径较大。该种于2005年的5月、9月和11月出现，2月未见出现，9月丰度最高。该种出现时，水体表层温度范围为14–27°C，表层盐度范围为3.9–29.8。该种的纤毛图式、细胞及基因特征尚需进一步研究。砂壳纤毛虫生态调查北极王湾北极黄河站附近的王湾(Kongsfjorden)是重要的生态系统监测点。我们首次报道了王湾砂壳纤毛虫群落的空间分布与时间变化，并且细化到种。王湾共发现6属8种砂壳纤毛虫：挪威棘口虫、水晶薄铃虫、贫齿拟网纹虫、巨拟网纹虫、钝笛杯虫、Stenosemellaspp.、细小拟铃虫和Tintinnopsisspp.。砂壳纤毛虫总丰度变化范围为0–2435ind L-1，总生物量变化范围为0–6.33μgC L-1。有4种砂壳纤毛虫是优势种，占总丰度的93.1%。不同的优势种垂直分布类型不同：细小拟铃虫出现在采样全深度(0–200m)，丰度高值出现在表层；Tintinnopsisspp.只在30m以浅的水层出现；挪威棘口虫出现在采样全深度但丰度整体均低；Stenosemellaspp.从不在表层出现。砂壳纤毛虫垂直分布可能受到冰川融水入湾的影响。对连续站的砂壳纤毛虫群落变化研究，并未发现其与潮时有明确的相关关系。聚类分析结果表明，王湾内砂壳纤毛虫属于同一群落，且在连续采样期间群落结构没有明显改变。砂壳纤毛虫可以作为王湾海洋浮游生态监测的指示种。环太平洋3个海区对环太平洋3个海区的砂壳纤毛虫进行了初步调查，分别为西太平洋的吐噶喇海峡(TokaraStrait)、热带太平洋西部的西里伯斯海(CelebesSea)附近以及热带太平洋东部的哥斯达黎加(CostaRica)近海。发现的砂壳纤毛虫种类数为：吐噶喇海峡样品中10科24属36种，西里伯斯海附近样品中10科25属39种，哥斯达黎加近海样品中12科27属40种。3个海区发现砂壳纤毛虫多为透明壳，种类结构较为一致。砂壳纤毛虫总丰度与水深呈极显著的负相关关系。在垂直分布上，哥斯达黎加近海的砂壳纤毛虫群落分布较浅，西里伯斯海附近的砂壳纤毛虫最深。南海2007年10月在南海北部调查了大型砂壳纤毛虫(>76m)的生态分布。发现砂壳纤毛虫7属22种，其中拟铃虫属最多，共10种。斯氏拟铃虫和根状拟铃虫为优势种。砂壳纤毛虫总丰度为0–41768ind m-3，生物量为0–609.92g Cm-3。76.83%站位的砂壳纤毛虫丰度低于1000ind m-3。砂壳纤毛虫生物量与丰度分布的总体趋势为近岸高且站位间差异大，远洋低且站位间差异小。砂壳纤毛虫集中于雷州半岛以东的近岸浅水区，此区水文状况为高温低盐、高Chl a浓度；不同种的分布规律不同，多呈斑块状分布。砂壳纤毛虫的丰度、生物量以及种丰富度与Chl a浓度均为正相关关系。2007年8–9月调查南海西部海域砂壳纤毛虫的生态分布，共发现7属12种。砂壳纤毛虫总水体丰度为0–11622indm-2。优势种有5种：尖形钟形虫、宽口钟形虫、费瓦拟平顶虫、南方类铃虫、圆锥条纹虫。圆锥条纹虫聚集强度最高，属于适盐适温范围较窄的浮游动物；尖形钟形虫与宽口钟形虫的聚集性最弱，分布较为均匀。南海西部砂壳纤毛虫群落受南海流系、水团和季风的影响，整体稳定性均较低。砂壳纤毛虫总丰度、种丰富度与环境因子没有明显相关性。圆锥条纹虫与环境因子的相关性系数高。与南海北部相比，南海西部砂壳纤毛虫类群与南海北部为同一区系，但是种类组成中远洋种占主要部分，丰度大大低于南海北部。圆锥条纹虫的丰度高，且聚集强度高，与温盐因子的相关性较好。砂壳纤毛虫群落长期调查胶州湾是我国近海长期观测点，积累了大量长时间序列数据，对近海生态系统的长期变化有着重要意义。我们对胶州湾砂壳纤毛虫群落进行了长时间监测和研究，调查时间为2003年5月至2012年12月。共发现砂壳纤毛虫10属27种，拟铃虫属种类最多，共15种。砂壳纤毛虫丰度范围为0–3970ind L-1。每年砂壳纤毛虫丰度在6月最高，7–8月继续保持高值，9月开始下降，1月最低。8月份砂壳纤毛虫种类数最高。聚类分析表明，砂壳纤毛虫在年周期内可以分为3个群落：春夏群落(2–6月)、夏秋群落(7–10月)以及冬季群落(11–1月)。冬季群落的聚集程度最紧密，说明冬季群落最为稳定。砂壳纤毛虫群落在7月与11月出现剧烈的结构变动。砂壳纤毛虫丰度的年内变化要远远大于年际变化。就年际变化而言，砂壳纤毛虫丰度从2004年至2007年呈升高趋势，往后呈明显下降趋势，这主要是6–8月的丰度变化引起。相关性分析表明，温度对砂壳纤毛虫丰度有明显影响，而物种数则受盐度负相关影响和温度的正相关影响。温度与盐度协同作用于砂壳纤毛虫群落的变化。百乐拟铃虫与白领细壳虫是胶州湾砂壳纤毛虫群落变动的关键种。更多还原

【Abstract】 Microzooplankton is a key trophic group in pelagic ecosystems, as theseorganisms consume most of the primary production. Tintinnids are one of the bestknown groups of planktonic ciliates because of their loricae, which make themconspicuous in plankton samples and permit species identification. Although usuallynot the dominant component of the microzooplankton, tintinnids are regarded as modelsfor marine plankton research, as more is known about tintinnids than most any othergroup of microzooplankters (Dolan et al.,2013). In this study, a check list ofcontemporary living tintinnids around the world was compiled according toreferences available since Kofoid&Campbell (1929). Based on this work, we studiedthe distribution of tintinnid communities in South China Sea, Jiaozhou Bay,Kongsfjorden (Svalbard, Arctic), Tokara Strait, near Celebes Sea and coast off CostaRica. Besides, a long-term monitoring was conducted on tintinnid communities inJiaozhou Bay from May2003to December2012.Check list of contemporary tintinnid speciesA check list of contemporary living tintinnid (Ciliophora, Spirotrichea, Tintinnida)ciliates around the world was compiled according to available references since Kofoid&Campbell (1929). A total of931species belonging to67genera and15families werelisted in marine and fresh waters all over the world. Among them,900species live inmarine habitats and31species live in fresh water biotopes. Tintinnopsis is the mostabundant genus (141species). In China,163species including20species in freshwaters, belonging to36genera and13families, have been recorded. There were9species, namely Codonella crater, Stenosemella lacustris, Tintinnidium ephemeridum,Tintinnidium ranunculi, Tintinnidium semiciliatum, Tintinnopsis fusiformis,Tintinnopsis illinoisensis, Tintinnopsis ovalis, and Tintinnopsis cylindrata, living infresh waters in the list were not recorded in China. In the list, Chinese species name were given, of which159were adopted from the present literatures.We found a new tintinnid ciliate species, Tintinnopsis estuariensis in the YangtzeRiver estuary, China in2005. Its arenaceous lorica is cylindro-conical in shape,(120–180)×(50–75) μm in size, agglutinated with mineral particles, and open at both ends.Therefore, we assigned it to the genus Tintinnopsis (Phylum Ciliophora, ClassSpirotrichea, Subclass Choreotrichia, Order Tintinnida, Family Codonellidae). Twocharacters distinguish it from its congeners: the unique lateral flattened posteriorportion with a vertical aboral opening and wider oral diameter. It occurred in water witha surface temperature of14–27°C and surface salinity of3.9–29.8.Ecological studies of tintinnid communities in different seasKongsfjordenWe studied the tintinnid communities in Kongsfjorden, Svalbard (Arctic) duringJuly2012, which is the first to provide quantitative, species-specific data about thetintinnids there. Eight morphotypes were found: Acanthostomella norvegica,Leprotintinnus pellucidus, Parafavella edentata, P. gigantea, Ptychocylis obtusa,Stenosemella spp., Tintinnopsis parvula, and Tintinnopsis spp. Total tintinnidabundance and biomass ranged from undetectable (<1ind L–1) to2435ind L–1and0to6.33μg C L–1, respectively. Tintinnid community composition, in terms of speciespresent, appeared uniform throughout the fjord. However, four species showed distinctvertical distributions: Tintinnopsis parvula occurred throughout the water column butshowed a peak at the surface; Tintinnopsis spp. occurred only in the upper30m; A.norvegica occurred in low abundance throughout the water column; and Stenosemellaspp. was not detected in surface waters. The vertical distribution of tintinnid speciesmay be influenced by the presence of glacier melt water. At the coastal station, thetintinnid community showed considerable temporal variation in total abundance andthe identity of the most abundant species. These changes were not related to tide-andwind-induced salinity shifts. The observed temporal and spatial variations suggest thatthe marine plankton community of the Kongsfjorden is highly dynamic and thattintinnids can be used to assess the variation of marine planktonic communities.Three sites around Pacific Ocean Primary investigation of tintinnid communities around the Pacific Ocean in TokaraStrait, near Celebes Sea and off the coast of Costa Rica. Twelve families,27genera,40species were found off coast of Costa Rica,10families,25genera,39species werefound in Celebes Sea, and10families24genera36species were found in Tokara Strait.Most of the tintinnids had hyaline loricae.South China SeaThe abundance, biomass and species richness of large tintinnids (>76m) in thenorthern South China Sea (in82stations in13sections) were investigated in October,2007. Totally22species in7genera were found, in which genus Tintinnopsis wasdominant in abundance. Seven species were new recorded for the South China Sea. Theabundance of tintinnids ranged from0to41768ind m-3, on average (28517244) indm-3, and their biomass ranged from0to609.92g C m-3, on average (42.75110.80)g C m-3. Compared with those in pelagic area, the abundance and biomass of tintinnidswere higher in coastal shallow waters in the east of Leizhou Peninsula, where the waterwas warmer, less salty and with more Chl a. Most species were distributed in patches.The species richness of all stations was from0to12. The abundance, biomass andspecies richness of tintinnids were closely related to Chl a concentration. Tintinnopsisschotti and T. radix were dominant species.A total of12species of tintinnids in7genera were found in the western SouthChina Sea in summer2007and2species (Eutintinnus elegans and Xystonellopsisbrandti) were recorded for the first time in South China Sea. The range of tintinnidsabundance was from0to11622ind m-2with an average of2218±2254ind m-2, andthe species richness was from0to6. Five dominant species were Cyttarocylisacutiformis, Cyttarocylis eucecryphalus, Xystonellopsis favata, Codonellopsismeridionalis and Rhabdonella conica, while the dominance index of R. conica washighest. R. conica was observed for the highest aggregation intensity, which meant thatit was more closely related to environmental changes. Cyttarocylis acutiformis and C.eucecryphalus had the lowest aggregation intensity and more evenly distributed. Boththe species diversity and species evenness in the western South China Sea were lowdue to the effect of a variety of monsoon, ocean currents and water masses, which indicated that tintinnids community were under an unstable state. Correlation analysisby SPSS software revealed no significant correlation among tintinnids abundance,species abundance and environment factors. The community structure wascharacterized by more pelagic species and more unstable in the western South ChinaSea than in the northern part of South China Sea. R. conica was considered as biologicalindicator species of water masses for its high average abundance and maximumabundance, high aggregation intensity and fine correlation with environmental factors.Long-term monitoring of tintinnid communities in Jiaozhou BaySeveral studies on tintinnids in Jiaozhou Bay have been conducted since1952, yetstudies focusing on temporal variation of tintinnid community are still lack. Long-termvariations in tintinnid abundance and species composition were investigated monthlyfor more than9years at4sites in Jiaozhou Bay (Yellow Sea) from May2003toDecember2012. A total of27species belonging to10genera were recorded. Amongthem genus Tintinnopsis has more species (15) than other genus, and the remaininggenera were represented by one or two species. Species richness at every sampling dateranged from0to21. At the monthly scale, tintinnid abundance reached maximum inJune from the minimum of January. Abundance stayed high in July and August, andthen went down to December. Species richness reached its maximum in August.Distinct clusters of tintinnid communities were revealed associated with differentmonths and seasonal patterns: Spring-summer cluster (February-June), Summer-autumn cluster (July-October), and winter cluster (November-January). Winter clusterswas tightest among the three clusters, which showed that the winter communities wererelatively stable. The structure of tintinnid communities changed dramatically duringJune-July and October-November.Temporally, tintinnid abundances increased from2004to2007, and thendecreased to2012. Spatially, tintinnid abundances were higher in inner bay than in outerbay. Tintinnid abundance was found positively correlated to temperature (p<0.01),while species richness was positively correlated to temperature (p<0.01) and negativelyto salinity (p<0.01). No correlation was found between Chl a and tintinnid abundanceor species richness. It was supposed there was a combined effect on the tintinnid communities from temperature and salinity. Tintinnopsis beroidea and Stenosemellanivalis are identified as the key species of tintinnid communities in Jiaozhou Bay andthe key factor for temporal variations of tintinnid abundance during2003–2012.更多还原