摘要
由于高度的细胞结构分化、表型水平上的形态多样性、独特的核双态性以及(有性的)接合生殖方式、及在微食物网中的枢纽地位等因素,纤毛虫原生动物一直是细胞学、真核生物遗传学、生态学等基础生物学领域中重要的模型动物或研究对象。
在国家自然科学基金项目(30430090,40676076)及中-英国际合作达尔文启动计划(14-015)的资助下,作者于2006-2009年期间,对中国黄渤海和南海近岸不同生境的自由生纤毛虫的区系开展了调查。结合活体观察及蛋白银、氨银法等现代染色方法对所涉纤毛虫的活体形态、纤毛图式或银线系及统计学特征进行了详细的研究和报道。共发现、分离和鉴定自由生海洋纤毛虫46种,分别隶属于9个目(前口目、刺钩目、合膜目、篮管目、管口目、吸管目、异毛目、丽克虫目、腹毛目)。根据形态学所给出的重要指征建立了若干新的分类阶元,包括2个新属、11个新种和4个新组合。
主要成果和创新点包括:
1)榴弹虫科,因许多重要分类特征(口器数目、排布,口区结构等)在活体观察中难以了解,造成该类群的分类学长期存在混乱和错误。本工作基于活体观察及细胞学染色制片,首次揭示了1新属(偏榴弹虫属)、4新种(东方诺兰德虫,中华诺兰德虫,巨大偏榴弹虫,相似巨大偏榴弹虫)及1新组合(相似瓦榴弹虫)的形态学和纤毛图式特征,并首次给出了迄今不明的纺锤披巾虫的口器结构。其中,新属(偏榴弹虫)特征为:具棘刺,盔板8环;盔板具肾形单窗口,脊靠近波浪状左边缘,且呈翼状,盔板右边缘的齿状结构与窗口交替排列;口器3个。
2)吸管亚纲是纤毛门中高度特化的类群,大部分阶元尤其是海洋类群近半个世纪以来一直无人问津。本文借助银染技术及分子信息完成了对14种的研究并建立1新属(拟粘管虫属)与1新种(半球拟粘管虫):生芽斑吸管虫,冠状斑吸管虫,球形斑吸管虫,柱状斑吸管虫,结节壳吸管虫,扁平壳吸管虫,粗壮壳吸管虫,具边似壳吸管虫,宽口似壳吸管虫,钟状贝吸管虫,锯齿杯吸管虫,小甲吸管虫,长号甲吸管虫,灰色曲壳吸管虫。拟粘管虫新属的特征为:虫体半球形,无柄、无鞘,通过基板直接附着在基质上;细胞明显分成致密的内质和透明的外质,后者呈凝胶状。吸管触手状,不成簇,散布于细胞表面。
3)异毛目隶属于分类地位最高等的多膜纲,虫体通常巨大,常见于各类水体,其研究史几乎贯穿了纤毛虫的研究史。因其各种类间活体特征的重叠性,在300多年的历史中积累了大量混乱和错误。本工作基于活体观察及详尽的纤毛图式资料,激活长期存在争议的突喇叭虫属且给出新定义,并对其模式种(耳状突喇叭虫)与绿色爽口虫进行了混乱清理、重认证及缺失资料的补足,同时,对8个已知种(蓝菌丽克虫,简单瓶囊虫,延长类瓶囊虫,佛氏环须虫,盐蚕豆虫,小突口虫,叶状突口虫,弯曲突口虫)进行了必要性重描述及显微照片的提供。突喇叭虫新激活属的特征为:虫体较大并具一定伸缩性,自由游泳;深而阔大的口前庭,前庭腹面边缘有一裂口;口前庭动基列仅分布在口前庭内壁上,口区底部没有毛基粒或动基列;纤毛图示与喇叭虫相似:具腹面缝合线和体动基列条带反差区;口围带几乎闭合,在腹面形成一楔形缺口;口侧膜长而明显。
4)合膜目种类相对较少,虽仅有7属,但大部分种类缺少纤毛图式及明确的分类标准,各属种间分类与系统地位长期不明。借助银染技术及分子手段,对该类群前人遗留的混乱做了清理和厘定,建立3个新组合(椭圆带膜虫,贪食带膜虫,长形带膜虫),给出该目种类最多属(带膜虫属)的新定义:虫体肾形或卵圆形;合膜粗壮,口区左右两侧均有分布,延伸到左侧背面;大核椭球形或长棒状;伸缩泡通常多个,位置不确定。
5)对管口目类群做了补足性研究,建立3个新种(热带管口虫,喙突宽管虫,似梳状偏体虫),对国际间长期存疑,分类学资料不详的3个种做了纤毛图式资料的建立和重描述,分别给出新定义、建立新模标本(北欧林奇虫,少齿篷管虫,寄生下管虫)。
6)前口目隶属裸口亚纲,分类地位原始,彼此间亲缘关系不明或存严重争议。本工作在详尽活体观察基础上,借助蛋白银等银染技术对该类群进行了初步探讨。建立1个新种(拟纹中圆虫),并首次给出迄今不明的优美壶形虫的纤毛图式和详细的口区结构,对其分类地位的确定提供了重要证据,同时对已知种盐扁体虫进行了补足性重描述以及统计数据的提订。
7)同时,本工作对篮管目与腹毛目2个新种进行了现代方法的研究,包括活体观察、蛋白银显微照片、纤毛图式及分子信息(红色篮口虫,多核拟双棘虫),对长期分类地位不明的盾片异列虫在纤毛图式水平上进行了清理、重认证以及缺失资料的补足,并分别给出了新定义。
Ciliated protozoa is the most complex and highly differentiated among unicellularorganisms,of which some groups are closely related to occurrence of red-tide,bait-animalculture in aquaculture,disease of animal,water quality purification in polysaprobicenvironment,material cycle and energy flow in“micro-food-loop”.
From 2006 to 2009,faunistic and taxonomic studies on free living ciliates mainly fromcoastal waters off Qingdao and Guangzhou (including sand,mollusks-culturing and-breedingwaters,tidal zone,mouth of the Pearl River,mangroves near Shenzhen) have been carried out.The morphology and infraciliature of 46 species representing 9 orders (Prostomatida,Haptorida,Synhymeniida,Nassulida,Cyrophorida,Suctorida,Heterotrichida,Licophorida,Hypotrichida),23 families and 33 genera were investigated using living observation and silverimpregnation methods.Main achievements are summarized as follows:
Two new genera were established,Apocoleps nov.gen.and Paramucophrya nov.gen..
Diagnosis of Apocoleps nov.gen.:Colepidae with spiny armour composed of eight tiers;plate with reniform uni-windows,ridge near left plate margin serrated and wing-like,teeth ofright plate margin arranged alternately with windows;three adoral organelles.
Diagnosis of Paramucophrya nov gen.:body hemispherical,composed of inner and outerplasma,the outer of which is a transparent mucous coat;without stalk or lorica,attached tosubstrate;tentacles sparsely scattered.
The complicated genus Condylostentor Jankowski,1978 which has never been studiedusing modern methods since it is originally established,is reactivated and restudied basedon investigation on the type species of this genus.
Improved diagnosis of Condylostentor:Slightly contractile,free-swimming Stentoridae;vestibular cavity deep and prominent,apical border with conspicuous ventral groove;ciliaturein Stentor-like pattern,i.e.ventral suture and contrast zone of somatic kineties present;adoralzone almost closed,interrupted on ventral side by a deep cleft;vestibulum kineties andparoral membrane present.
Eleven new species were established based on living morphology and infraciliature:Nolandia orientalis Chen et al.,2009;Nolandia sinica Chen et al.,2009;Apocoleps magnusChen et al.,2009;Apocoleps paramagnus Chen et al.,2009;Metacystis parastriata nov.spec.;Nassula rubra nov.spec.;Cyrtophoron tropic nov.spec.;Aegyriana rostellum nov.spec.; Dysteria pseudopectinata nov.spec.;Paramucophrya semisphaerica Chen et al.,2005;Parabirojimia multinucleata Chen et al.,2009.
Four new combinations were suggested and their infraciliature was reveal for thefirst time which has never been studied using modern staining methods:Pinacocolepssimilis (Kahl,1933) Chen et al.,2009;Zosterodasys oblonga (Maupas,1883) nov.comb.;Zosterodasys vorax (Stokes,1887) nov.comb.;Zosterodasys elongata (Kahl,1931) nov.comb.
Infraciliature of 8 species was revealed for the first time and their neotypes havebeen deposited.These included:Tiarinafusus (Clapar(?)de & Lachmann,1857) Bergh,1881;Lagynus elegans (Engelmann,1862) Quennerstedt,1867;Lynchella nordica Jankowski,1968;Chlamydonys paucidentatus Deroux,1976;Hypocoma parasitica Gruber,1884;Condylostentor auriculatus (Kahl,1932) Jankowski,1978;Climacostomum virens (Ehrenberg,1933) Kahl,1932;Anteholosticha scutellum (Cohn,1866) Berger,2003.
Fifteen species of suctorid ciliates were recorded and described for the first time inChina:Ephelota gemmipara (Hertwig,1876) Bütschli,1889;E.mammillata Dons,1915;E.crustaceorum Hailer,1880;E.truncata Fraipont,1878;Acineta tuberosa Ehrenberg,1833;A.compressa Clapar(?)de & Lachmann,1859;A.foetida Maupas,1881;Paracineta limbata(Maupas,1881) Collin,1912;P.patula (Clapar(?)de & Lachmann,1861) Collin,1911;Conchacineta complatana (Gruber,1884) Curds,1985;Actinocyathula crenata (Fraipont,1878) Curds,1987;Loricophrya parva (Schulz,1932) Matthes,1956;L.tuba (Zelinka,1914)Curds,1987;Paramucophrya semisphaerica Chen et al.,2005;Flectacineta livadiana(Mereschkowsky,1881)Jankowski,1978.
Nine species were redescribed based on characteristics of living feature andinfraciliature from the Qingdao and Guangzhou populations:Placus salinus Dietz,1964;Licnophora lyngbycola Faur(?)-Fremiet,1937;Folliculina simplex (Dons,1917) Song et al.,2003;Folliculinopsisproducta (Wright,1859) Faur(?)-Fremiet,1936;Peritromusfaurei Kahl,1932;Fabrea salina Henneguy,1890;Condylostoma minutum Bullington,1940;Condylostoma spatiosum Ozaki & Yagiu in Yagiu,1944;Condylostoma curva Burkovsky,1970.
引文
顾福康(1991):原生动物学概论.高等教育出版社.
沈蕴芬等(1999):原生动物学.科学出版社.
宋微波等(1999):原生动物学专论.青岛海洋大学出版社.
宋微波等(2003):海水养殖中的危害性原生动物.科学出版社.
宋微波等(2009):黄渤海的自由生纤毛虫.印刷中.
Aescht,E.(2001).Catalogue of the generic names of ciliates (Protozoa,Ciliophora).Denisia.,1:1-350.
Andrews,E.A.(1923).Folliculina:case making,anatomy and transformation.J.Morphol.,38:207-278.
Andrews,E.A.(1953).Valletofolliculina bicornis,a unique new genus and species of folliculinid (Ciliata:Heterotricha) from California.J.Washington Acad.Sci.,43:189-194.
Batisse,A.(1994).Sous-Classe des Suctoria Clapar(?)de et Lachmann,1958.Trait(?) Zool.,2:493-563.
Berger,H.,Foissner,W.(1989).Morphology and biometry of some soil hypotrichs (Protozoa,Ciliophora) from Europe and Japan.Bull.Br.Mus.Nat.Hist.(Zool) 55:19-46.
Berger,H.(2003).Redefinition of Holosticha Wrzesniowski,1877 (Ciliophora,Hypotricha).Europ.J.Protistol.,39:373-379.
Berger,H.(2004).Uroleptopsis Kahl,1932 (Ciliophora,Hypotricha):morphology and cell division of type species,redefinition,and phylogenetic relationships.Acta.Protozool.,43:99-121.
Berger,H.(2006).Monograph of the Urostyloidea (Ciliophora,Hypotricha).Monogr.Biol.,85:1-1304.
Bock,K.J.(1952).Zur (O|¨)kologie der Ciliaten des marinen Sandgrundes der Kieler Bucht.Ⅰ.Ⅱ.Kieler Mecresforschungen.Bd.IX.
Borror,A.C.,Wicklow,B.J.(1983).The suborder Urostylina Jankowski,(Ciliophora,Hypotrichida):morphology,systematics and identification of species.Acta.Protozool.,22:97-126.
Borror,A.C.(1972).Tidal marsh ciliates (Protozoa):morphology,ecology,systematics.Acta Protozool.,10:29-71.
Bott,T.L.,Kaplan,L.A.(1990).Potential for protozoan grazing of bacteria in streambed sediments.J.N.Am.Benthol.Soc.,9:336-345.
Bullington,W.E.(1940).Some ciliates from Tortugas.Pap.Tortugas Lab.,32:179-221.
Burkovsky,I.(1970).The ciliates of the mesopsammon of the Kandalaksha gulf (White Sea) I.Acta Protozool.,7:475-499.
B(u|¨)tschli,O.(1887-1889).Protozoa.Abt Ⅲ.Infusoria und System der Radiolaria.In:Klassen und Ordnungen des Their-Reichs (Ed.H.G.Bronns),C.F.Winter,Leipzig,1098-2035.
Calkins,G.N.(1902).Marine protozoa from Woods Hole.Bull.U.S.Fish Comm.,21:413-468.
Carey,P.G.(1992).Marine interstitial ciliates.An illustrated key.Chapman and Hall,London.
Chatton,E.,Lwoff,A.(1930).lmpr6gnation,par diffusion argentique,de l'infraciliature des cili(?)s matins et d'eau douce,apr(?)s fixation cytologique et sans desiccation.Compt.Rend.Soc.Biol.,104:834-836.
Gelei,J.Von,Horv(?)th,P.(1931).Eine nasse Silber-bzw.Goldmethode f(u|¨)r die Herstellung der reizleitenden Elemente bei den Ciliaten. Z. Wiss. Mikros., 48: 9-29.
Chen, X., Song, W., Al-Rasheid, K. A. S. Warren, A., Hu, X., Wang, Y., Long, H. (2009). Morphological studies on three marine colepid ciliates form Qingdao, China, including one new genus, one new combination and two new species (Ciliophora, Colepidae). Eur. J. Protistol. In press.
Chen, X., Song, W., Hu, X. (2005). Morphology of suctorid ciliates from coastal waters off Qingdao, with description of a new genus and a new species (Protozoa, Ciliophora). Acta Zootax. Sin., 30:493-500. (in Chinese with English summary)
Chen, X., Song, W., Warren, A., Al-Rasheid, K. A. S., Al-Farraj, S. A., Al-Quraishy, S. A., Gong, J. (2008). Redefinition of two marine suctorian ciliates, Ephelota gemmipara (Hertwig, 1876) Butschli, 1889 and Ephelota crustaceorum Haller, 1880 (Ciliophora, Suctoria), with a brief description of the asexual reproduction process in E. gemmipara. Acta Protozool., 47: 113-124.
Chen, X., Song, W., Al-Rasheid, K. A. S., Warren, A., Long, H., Shao, C, Al-Farraj, S. A., Gong, J., Hu, X. (2007). The morphology of three marine heterotrichous ciliates, Condylostentor auriculatus (Kahl, 1932) Jankowski, 1978, Condylostoma minutum Bullington, 1940 and C. spatiosum Ozaki & Yagiu in Yagiu, 1944 (Ciliophora: Heterotrichida). Acta Protozool., 46: 289-309.
Chen, X., Xu, H., Wang Y, Liu W., Gao, S., Al-Rasheid, K. A. S., Warren, A., Song, W. (2009). Morphology and gene sequences of three marine colepid ciliates: Nolandia sinica spec. nov., Apocoleps paramagnus spec. nov. and Tiarina fusus (Claparede & Lachmann, 1859) Bergh, 1881 (Ciliophora, Colepidae). Eur. J. Protistol. Submitted.
Clapar(?)de, E., Lachmann, J. (1859). (?)tudes sur les infusoires et les rhizopodes. M(?)m. Inst. natn. Gen(?)v. 6 (yr 1858): 261-482.
Cohn, F. (1866). Neue Infusorien im Seeaquarium. Zeitsch. Wiss. Zool. Leipzig., 16: 253-302. (In German).
Corliss, J. O. (1979). The ciliated protozoa: characterization classification and guide to the literature. 2nd ed. Pergamon Press, Oxford, New York, Toronto, Sydney, Paris, Frankfurt.
Curds, C. R. (1985a) A revision of the Suctoria (Ciliophora, Kinetofragminophora) 1. Acineta and its morphological relatives. Bull. Br. Mus. nat. Hist. (Zool.) 48: 75-129.
Curds, C. R. (1985b). A revision of the Suctoria (Ciliophora, Kinetofragminophora) 2. An addendun to Acineta. Bull. Br. Mus. nat. Hist. (Zool.), 49: 163-165.
Curds, C. R. (1985c). A revision of the Suctoria (Ciliophora, Kinetofragminophora) 3. Tokophrya and its morphological relatives. Bull. Br. Mus. nat. Hist. (Zool.), 49: 167-193.
Curds, C. R. (1986). A revision of the Suctoria (Ciliophora, Kinetofragminophora) 4. Podophrya and its morphological relatives. Bull. Br. Mus. nat. Hist. (Zool.), 50: 59-91.
Curds, C. R. (1987). A revision of the Suctoria (Ciliophora, Kinetofragminophora) 5. The Paracineta and Corynophrya problem. Bull. Br. Mus. nat. Hist. (Zool.) 52: 71-106.
Das, S. M. (1952). Indian Folliculindae (Ciliata, Heterotricha). Ann. Magaz. Nat. Hist., 6: 235-240.
Deroux, G (1965). Origine des cineties anterieures, gauches et buccales dans le genre Dysteria Huxley. Compt. Rend. Acad. Sci. Paris. 260: 6689-6691.
Deroux, G (1970). La serie "chlamydonellienne" chez les Chlamydodontidae (Holotriches, Cyrtophorina Faur(?)-Fremiet). Protistologica 6: 155-182.
Deroux, G. (1974). Les dispositifs adhesifes ciliaires chez les Cyrtophorida et la famille des Hypocomidae. Protistologica 10: 379-396.
Deroux, G. (1976a). Le plan cortical des Cyrtophorida unite d'expression et marges de variabilite. I. Le cas des Plesiotrichopidae, Fam. Nov., dans la nouvelle systematique. Protistologica 12: 469-481.
Deroux, G (1976b). Le plan cortical des Cyrtophorida unite d'expression et marges de variabilite. ?. Cyrtophorida a thigmotactisme ventral g(?)n(?)ralise. Protistologica 12: 483-500.
Deroux, G. (1976c). Le plan cortical des Cyrtophorida. III. Les structures differenciatrices chez les Dysteriina. Protistologica 12: 505-538.
Deroux, G. (1994). Sous-Classe des Cyrtophorida Faur(?)-Fremiet in Corliss, 1956. Trait(?). de. Zoologie. 2: 401-431.
Dons, C. (1918) Neue marine Ciliaten und Suctorien. Troms(?) Mus. Aarshrifter 38-39 (year 1915-1916): 75-100.
Dons, C. (1942). Paragolliculina labiata n. sp., ein neues karaffentier. Kong. Norske Cidensk. Selsk. Skr., 14: 195-198.
Dragesco, J. (1960). Cilies mesopsammiques littoraux. Systematique, morphologie, ecologie. Trav. Stn. boil. Roscoff(N. S.)122: 1-356.
Dragesco, J. & Dragesco-Kerneis, A. (1986). Cilies libres de l'Afrique intertropicale. Faune. Tropicale., 26: 1-559.
Dragesco, J. (1960). Cilies mesopsammiques littoraux. Trav. Stat. Biol. Roscoff(N. S.) 12: 1-356.
Dragesco, J. (1966). Quelques Cilies libres du Gabon. Biol. Gabon., 2: 91-117.
Dragesco, J., Dragesco-Kerneis. (1991). Free-living ciliates from the coastal area of Lake Tanganyika (Africa). Eur. J. Protistol., 26: 216-235.
Ehrenberg, C. G. (1830). Beitr(?)ge zur Kenntniss der Organisation der Infusorien und Ihrer geographischen Verbreitung, besonders in Sibirien. Abh. Dt. Akad. Wiss. Berlin, year 1830: 1-88.
Ehrenberg, C. G. (1831). (?)ber die Entwickelung und Lebensdauer der Infusionsthiere; nebst ferneren Beitr(?)gen zu einer Vergleichung ihrer organischen Systeme. Abh. Dt. Akad. Wiss. Berlin, year 1831: 1-154.
Ehrenberg, C. G. (1833). Dritter Beitr(?)gen zur Erkenntniss der grosser Organisation in der Richtung des kleinsten Raumes. Abh. Akad. Wiss. Berlin (yr 1835): 145-336.
Ehrenberg, C. G. (1838) Die Infusionsthierchen als vollkommene Organismen. Ein Blick in das tiefere organische Leben der Natur. Voss, Leipzig
Entz, G. (1884). (?)ber Infusorien des Golfes von Neapel. Mitt. Zoo. Stn. Neapel., 5: 289-444. (In German)
Faur(?)-Fremiet, E. (1932). Division et morphogen(?)se chez Folliculina ampulla O. F. M(?)ller. Bull. Biol., 66: 77-110.
Faur(?)-Fremiet, E. (1936). Condylostoma (Stentor) auriculatus (Gruber). Bull. Soc. Zool. Fran., 61: 511-519.
Faur(?)-Fremiet, E. (1936). The Folliculinidae (Infusoria, Heterotricha) of the Breton coast. Biol. Bull., 70: 353-360.
Faure-Fremiet, E. (1961). Les cili(?) Cyrtophorina et leur diversification morphologique. C. R. Acad. Sci. Paris., 267: 954-957.
Faure-Fremiet, E. (1965). Morphologie des Dysteriidae (Ciliata Cyrtophorina). Compt. Rend. Acad. Sci. Paris., 260: 6679-6684.
Faure-Fremiet, E., Andre J., Grnier M. C. (1968). Structure fine de l'organite fixateur des Dysteriidae (Ciliata Cyrtophorida). C. R. Acad. Sci. Paris., 105: 355-360.
Fenchel, T. (1987). Ecology of Protozoa. Springer-Verlag, Berlin.
Fenchel, T. (1969). The ecology of marine microbenthos. IV. Structure and function of the benthic ecosystem, its chemical and physical factors and the microfauna communities with special regerence to the ciliated protozoa. Ophelia 6: 1-182.
Fernandez-Leborans, G, Arndt, C. E., Gabilondo, R. (2006). Protozoan epibionts and their distribution on the Arctic ice-amphipod Gammarus wilkitzkii from Spitsbergen, Norway. Arct. Anta. Alp. Res., 38: 343-356.
Finlay, B. J., Berninger, U. G, Clarke, K. J., Cowling, A. J., Hindle, R. M., Rogerson, A. (1988). On the abundance and distribution of protozoa and their food in a productive freshwater pond. Eur. J. Protistol., 23:205-217.
Foissner, W. (1982). (?)kologie und Taxonomie der Hypotrichida (Protozoa: Ciliophora) eineger (?)sterreichischer Boden. Arch. Protistenkd., 126: 19-143. (In German)
Foissner, W., W(?)lfl, S. (1994). Revision of the genus Stentor Oken (Protozoa, Ciliophora) and description of S. areucanns nov. spec, from South American lakes. J. Plankton Res., 16: 255-289.
Foissner, W. (1983). Taxonomische Studien (?)ber die Ciliaten des Gro(?)glocknergebietes (Hohe Tauern, (?)sterreich) I. Familien Holophryidae, Prorodontidae, Plagiocampidae, Colepidae, Enchelyidae und Lacrymariidae nov. fam. Annln. naturh. Mus. Wien. 84B, 49-85.
Foissner, W. (1984). Infraciliatur, Silberliniensystem und Biometrie einigerneuer und wenig bekannter terrestrischer, limnischer und mariner Ciliaten (Protozoa: Ciliophora) aus den Klassen Kinetofragminophora, Colpodea und Polyhymenophora. Stapfia., 12: 1-165.
Foissner, W. (1987). Soil protozoa: fundamental problems, ecological significance, adaptations in ciliates and testaceans, bioindicator and guide to the literature. Prog. Protistol., 2: 69-212.
Foissner, W., Berger, H., Kohmann, F. (1992). Taxonomische und okologische Revision der Ciliaten des Saprobiensystems-Band ?: Peritrichia, Heterotrichida, Odontostomatida. Informationsbe. Bayer. Landesam. Wasserwirts., 5/92: 1-502.
Foissner, W., Berger, H., Kohmann, F. (1994). Taxonomische und (?)kologische Revision der Ciliaten des Saprobiensystems-Band ?: Hymenostomata, Prostomatida, Nassulida. Informationsbriche des Bayer. Landesamtes fur Wasserwirtschaft., 1/94: 1-548.
Foissner, W., Berger, H., Schaumburg, J. (1999). Identification and ecology of limnetic plankton ciliates. Informationsber. Bayer. Landesamtes Wasserwirtschaft., 3/99: 1-793.
Foissner, W., Kusuoka, Y., Shimano, S. (2008). Morphology and gene sequence of Levicoleps biwae n. gen., n. sp. (Ciliophora, Prostomatida), a proposed endemic from the ancient Lake Biwa, Japan. J. Eukaryot. Microbiol., 55: 185-200.
Fraipont, J. (1877). Recherches sur les Acin(?)tiniens de la c(?)te d'Ostende. Bull. Acad. Roy. Belg., 44: 287-297.
Fraipont, J. (1878). Recherches sur les Acin(?)tiniens de la c(?)te d'Ostende. Bull. Acad. Roy. Belg., 45: 475-516.
Gong, J., Song, W. (2003). Morphology and infraciliature of two marine benthic ciliates, Dysteria procera Kahl, 1931 and Dysteria magna nov. spec. (Protozoa, Ciliophora, Cyrtophorida), from China. Eur. J. Protistoi., 39: 301-310.
Gong, J., Song, W. (2004). Morphology and infraciliature of two marine species of Hartmannula (Protozoa, Ciliophora, Cyrtophorida), from scallop farming waters off Qingdao (Tsingtao), China. J. Nat. Hist., 38: 1327-1337.
Gong, J., Lin, X., Song, W. (2003). Redescription of a poorly-known marine cyrtophorid ciliate Dysteria pusilla (Claparede & Lachmann, 1859) (Protozoa: Ciliophora: Cyrtophorida) from Qingdao, China. Acta. Protozool., 42: 215-221.
Gong, J., Song, W., Warren, A. (2002). Redescriptions of two marine cyrtophorid Ciliates, Dysteria cristata (Gourret and Roeser, 1888) and Dysteria monostyla (Ehrenberg, 1838) Kahl, 1931 (Protozoa, Ciliophora, Cyrtophorida), from China. Eur. J. Protistoi., 38: 213-222.
Gong, J., Song, W., Warren, A. (2005). Periphytic ciliate colonization: annual cycle and responses to environmental conditions. Aauat. Microb. Ecol., 39:159-170.
Grell, G., Meister, A. (1984). Beitrage zur Ultrastruktur der Konjugation von Ephelota gemmipara R. Hertwig (Suctoria). Protistologica 20: 65-86.
Grell, K, G., Benwith, G. (1984). Die Ultrastruckture von Ephelota gemmipara Hertwig und E. plana Wailes (Suctoria): ein Vergleich. I. Die adulte Form, 1984.?. Der schw(?)rmer. Protistologica 20: 205-233.
Griffiths, B. S. (1994). Soil nutrient flow. In: Darbyshire J. F. (ed), Soil protozoa, CAB, Oxford, 65-91.
Gruber, A. (1884). Die Protozoen des Hafens von Genua. Nova Acta Leopold., 46: 475-539.
Guilcher, Y. (1950). Morphogen(?)se et morphologie compar(?)e chez les cilies gemmipares: chonotriches et tentaculiferes. Ann. Biol., 26: 465-478.
Guilcher, Y. (1951). Contribution (?) 1'(?)tude des cili(?)s gemmipares, chonotriches et tentaculif(?)res. Ann. Sc. Nat. Zool., 13:33-132.
Hartwig, E. (1973). Die Ciliaten des Gezeiten-Sandstrandes der Nordseeinsel Sylt. I. Systematik. Akad. Wiss. Lit. (Mainz) Math. Naturwiss. Kl. Mikrofauna Meersbod., 18: 387-453.
Hausmann, K., H(?)lsmann, N., Radek, R. (2003). Protistology, 3rdcompletely revised edition. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Berlin.
Hertwig, R. (1876). Ueber Podophrya gemmipara nebst Bemerkungen zum Bau und zur systematischen Stellung der Acineten. Morph. Jb., 1: 20-82.
Hertwig, R. (1877). Ueber den Bau und die Entwicklung der Spirochona gemmipara. Jen. Z. Naturwiss., 11: 149-187.
Hu, X. (2008). Cortical structure in non-dividing and dividing Diophrys japanica spec. nov. (Ciliophora, Euplotida) with notes on morphological variation. Eur. J. Protistol., 44: 115-129.
Hu, X., Song W. & Warren A. (2002). Observations on the morphology and morphogenesis of a new marine urostylid ciliate, Parabirojimia similis nov. gen., nov. spec. (Protozoa, Ciliophora, Hypotrichida). Europ. J. Protistol., 38, 351-364.
Huttenlauch, I. (1987). Ultrastructural aspects of the somatic and buccal infraciliature of Coleps ampacanthus Ehrenberg 1833. Protoplasma., 136: 191-198.
Huttenlauch, I., Bardele, C. F. (1987). Light and electron microscopical observations on the stomatogenesis of the ciliate Coleps amphacanthus Ehrenberg, 1833. J. Protozool., 34: 183-192.
ICZN (The International Commission of Zoological Nomenclature) (1999) International Code of Zoological Nomenclature. 4th ed. Tipografia La Garangola, Padova
Jankowski, A. W. (1967). New genera of tentacled Infusoria (Suctoria). Mat. V Konf. Mold. Zool. 35-36.
Jankowski, A. W. (1978). Revision of the system of the class Polyhymenophora (Spirotricha) (in Russian). Dokl. Akad. Nauk. SSSR., 1978: 39-40.
Kahl, A. (1932). Urtiere oder Protozoa I: Wimpertiere oder Ciliata (Infusoria) 3. Spirotricha. Tierwilt. Dtl. 25: 399-650. (In German)
Kahl, A. (1930). Urtiere oder Protozoa I: Wimpertiere oder Ciliata (Infusoria) 1. Allgemeiner Teil und Prostomata. Tierwelt. Dtl. 18: 1-180.
Kahl, A. (1933). Ciliata Libera et Ectocommensalia. Tierwelt Nord- und Ostsee, 23(Teil ?, c_3): 29-146. (In German)
Kahl, A. (1934). Suctoria. In: Die Tierwelt der Nord- Ostee, (Bd I, Teil Hc5) (Eds. G. Grimpe, E. Wagler), Becker & Erler Kom.-Ges, Leipzig, 184-226.
Kent, W. S. (1880-1882) A manual of the infusoria: including a description of all known flagellate, ciliate, and tentaculiferous protozoa British and foreign, and an account of the organization and affinities of the sponges. David Bogue, London, ?-?
Klein, B. M. (1926). Ergebnisse mit einer Silbermethode bei Ciliaten. Arch. Protistenk., 56: 243-279.
Li, L, Song, W., Warren, A., Al-Rasheid, K. A., Roberts, D., Yi, Z., Al-Farraj, S.A., Hu, H. (2008). Morphology and morphogenesis of a new marine ciliate, Apokeronopsis bergeri nov. spec. (Ciliophora, Hypotrichida) from the Yellow Sea, China. Eur. J. Protistol., 44: 208-219.
Lin, X., Li, J., Gong, J., Warren, A., Song W. (2008). Taxonomic studies on three marine pleurostomatid ciliates, Litonotus bergeri nov. spec, L. blattereri nov. spec. and L. petzi nov. spec. (Ciliophora, Pleurostomatida) from north China sea. Europ. J. Protistol., 44: 91-102.
Lin, X., Song, W., Li J. (2007). Amphileptus aeschtae nov. spec, and Amphileptus eigneri nov. spec. (Ciliophora, Pleurostomatida), two new marine pleurostomatid ciliates from China. Europ. J. Protist., 43: 77-86.
Lobban, C. S., Schefter, M., Simpson, A. G. B., Pochon, X., Pawlowski, J., Foissner, W. (2002). Maristentor dinoferus n. gen., n. sp., a giant heterotrich ciliate (Spirotrichea: Heterotrichida) with zooxanthellae, from coral reefs on Guam, Mariana Islands. Mar. Biol., 140: 411-423.
Lynn, D. H., Small, E. B. (2002). Phylum Cililphora Doflein, 1901. In: An Illustrated Guide to the Protozoa, 2nd ed. (Eds. J. J. Lee, G. F. Leedale, P. C. Bradbury). Society of Protozoologists, Lawrence, Kansas 371-656.
Lynn, D. H. (2008). The Ciliated Protozoa. Characterization, Classification and Guide to the Literature. 3~(rd) ed. Springer.
Ma, H., Ki, J., Song, W. (2003). An improved silver carbonate impregnation for marine ciliated protozoa. Acta. Protozool., 42: 161-164.
Mansfeld, K. (1923). 16 neue oder wenig bekannte marine Infusorien. Arch. Protistenkd., 46: 97-140. (In German)
Matthes, D. (1988). Suctoria. In: Suctoria und Urceolariidae (Peritricha), (D. Matthes, W. Guhl, G. Haider).Protozoenfauna 7/1. Gustav Fischer Verlag. Stuttgart. New York, 1-226.
Maupas, E. (1881). Contribution (?) 1' etude des acinetiens. Arch. Zool. exp. Gen., 9: 299-368.
Maupas, E. (1883). Contribution (?) 1' etude morphologique et anatomique des infusoires cili(?)s. Arch. Zool. Exp. G(?)n. (s(?)r. 2)., 1: 427-664.
Mulisch, M., Heep. T., Sturm, W., Borcherding, J. (1998). Redescription of Ascobius lentu, a rare freshwater folliculinid (Ciliophora: Heterotrichida) from a pond in Germany. Acta Protozool., 37: 29-39.
M(?)ller, O. F. (1773). Vermium Terrestrium et Fluviatilium, seu Animalium Infusoriorum, Helminthicorum et Testaceorum, non Marinorum, Succincta Historia. Heineck & Faber, Havniae & Lipsiae
M(?)ller, O. F. (1786). Animalcula Infusoria Fluviatilia et Marina, quae Detexit, Systematice Descripsit et ad Vivum Delineari Curavit. N. M(?)lleri, Hauniae.
Noble, A. E. (1929). Two new species of the protozoan genus Ephelota from Monterey Bay. California. Univ. Calif. Publ. Zool., 33: 13-26.
Noland, L. E. (1925). A review of the genus Coleps with descriptions of two new species. Trans. Am. micros. Soc, 44: 3-13.
Noland, L. E. (1937). Observations on marine ciliates of the Gulf Coast of Florida. Trans. Am. micros. Soc, 56: 160-171.
Nylander, J. A. A. (2004). MrModeltest. 2. Evolutionary Biology Centre, Uppsala University.
Obolkina, L. A. (1995). New species of the family Colepidae (Prostomatida, Ciliophora) from Lake Baikal. Zool. Zh., 74: 3-19.
Ozaki, Y., Yagiu, R. (1941). Studies on the marine ciliates of Japan, mainly from the Setonaikai (the inland sea of Japan). I. J. Sci. Hirosima Univ., 8: 165-184.
Packroff, G., Wilbert, N. (1991). Taxonomische Studien (?)ber die Ciliatenfauna (Protozoa, Ciliophora) der Eifelmaare.Arch. Protistenkd., 140: 121-139.
Pallas, P. S. (1766) Elenchus Zoophytorum. Petrum van Cleef, Hagae-Comitum.
Petz, W., Song, W., Wilbert, N. (1995). Taxonomy and ecology of the ciliate fauna (Protozoa; Ciliophora) in the endopagial and pelagial of the Weddell Sea, Antarctica. Stapfia., 40: 1-223.
Ronquist, F. R.., Huelsenbeck J. P. (2003). MrBayes 3: Bayesian phylogenetic under mixed models. Bioinformatics 19: 1572-1574.
Sand, E. (1899-1901). (?)tude monographique sur le groupe des infusoires tentaculif(?)res. Ann. Soc. Beige Micro., 24: 57-189,25: 1-204.
Shao, C, Miao, M, Song, W., Warren, A., Al-Rasheid, K. A. S., Al-Quaishy, S. A., Al-Farraj, S. A. (2008a). Studies on two marine Metaurostylopsis spp. from China with notes on the morphogenesis of M. sinica nov. spec. (Ciliophora, Urostylida). Acta. Protozool., 47: 95-112.
Shao, C, Song, W., Hu, X., Ma, H., Zhu, M., Wang, M. (2006). Cell division and morphology of the marine ciliate, Condylostoma spatiosum Ozaki and Yagiu (Ciliophora; Heterotrichida) based on a Chinese population. Europ. J. Protistol., 42: 9-19.
Small, E. B., Lynn, D. H. (1985). Phylum Ciliophora Doflein, 1901. In: Lee, J. J., Hutner, S. H. & Bovee, E. C. (ed.), An Illustrated Guide to the Protozoa. Society of Protozoologists, Lawrence, KS. p. 393-575.
Song, W., Wilbert, N. (1997a). Morphological studies on some free living ciliates (Ciliophora: Heterotrichida, Hypotrichida) from marine biotopes in Qingdao, China, with descriptions of three new species: Holosticha warreni nov. spec, Tachysoma ovata nov. spec. and T. dragescoi nov. spec. Europ. J. Protistol., 33: 48-62.
Song, W., Wilbert N. (1997b). Morphological investigations on some free living ciliates (Protoza, Ciliophora) from China Sea with description of a new hypotrichous genus, Hemigastrostyla nov. gen. Arch. Protistenkd., 148: 413-444.
Song, W, Wilbert, N. (1995). Benthische Ciliaten des S(?)wassers. In: R(?)ttger, R. (Ed.), Praktikum der Protozoologie. Gustav Fischer Verlag, New York, p. 156-168.
Song, W, Wilbert, N. (1997). Morphological investigations on some free-living ciliates (Protozoa; Ciliophora) from China sea with description of a new hypotrichous genus, Hemigastrostyla nov. gen. Arch. Protistenkd., 148: 413-444.
Song, W., Wilbert, N. (2002). Faunistic studies on marine ciliates form the Antarctic benthic area, including descriptions of one epizoic form, 6 new species and 2 new genera (Protozoa: Ciliophora). Acta Protozool., 41:23-61.
Song, W., Warren, A., Ji, D., Wang, M., Al-Rasheid, K. A. S. (2003). New contributions to two heterotrichous ciliates, Folliculina simplex (Dons, 1917), Condylostoma curva Burkovsky, 1970 and one licnophorid, Licnophora lyngbycola Faur(?)-Fremiet, 1937 (Protozoa, Ciliophora): descriptions of morphology and infraciliature. J. Eukaryot. Microbiol., 50: 449-462.
Spiegel, A. (1926). Einige neue marine Ciliaten. Arch. Protistenkd., 55: 184-190.
Stankovic, A., Borsuk, P., Koper, M., Weglenski, P. (2002). Studies on Ephelota sp., an epizoic suctorian found on Antarctic krill. Polar. Biology., 25: 827-832.
Swofford, D. L. (2002). PAUP*. Phylogenetic analysis using parsimony (* and other methods). Sinauer Associates, Sunderland, MA.
Tartar, V. (1958). Stentor introversus n. sp. J. Protozool., 5: 93-95.
Tartar, V. (1961) The biology of Stentor. Pergamon Press, London.
Wailes, G. H. (1925). Some new or rare Protozoa from British Columbia. Ann. Mag. nat. Hist. London. 16: 40-48.
Wailes, G. H. (1943). Protozoa et Suctoria. Can. Pac. Fauna., 1: 1-46.
Wang, C. C. (1934). Notes on the marine infusoria of Amoy. Rep. Mar. Biol. Ass., 3: 50-70.
Wang, C. C, Nie, D. (1932). A survey of the marine protozoa of Amoy. Contr. Biol. Lab. Sci. Soc. Chi., 8: 285-385.
Wang, Y., Hu X., Long, H., Al-Rasheid, K. A. S., Al-Farraj, S. A., Song, W. (2008). Morphological studies indicate that Pleuronema grolierei nov. spe. and P. coronatum Kent, 1881 represent different sections of the genus Pleuronema (Ciliophora: Scuticociliatida). Eur. J. Protistol., 44: 131-140.
Wilbert, N., Song, W. (2005). New contributions to the marine benthic ciliates from the Antarctic area, including description of seven new species (Protozoa, Ciliophora). J. Nat. Hist., (London) 39: 935-973.
Wilbert, N. (1975). Eine verbesserte Technik der Protargolimpr(?)gnation f(?)r Ciliaten. Mikrokosmos 64: 171-179. (In German)
Wilbert, N., Kahan, D. (1981). Ciliates of Solar Lake on the Red Sea shore. Arch. Protistenkd., 124: 70-95.
Wilbert, N., Schmall, G. (1976). Morphologie und Infraciliatur von Coleps nolandi Kahl, 1930. Protistologica 12: 193-197.
Wilbert, N., Song, W. (2005). New contributions to the marine benthic ciliates from the Antarctic area, including description of seven new species (Protozoa; Ciliophora). J. Nat. Hist., 39: 935-973.
Wright, T. S. (1858). Description of new protozoa. Edinburgh NewPhilos. J. (N. S)., 7: 276-281.
Wright, T. S. (1859). Description of new protozoa. Edinburgh NewPhilos. J. (N. S)., 10: 97-104. Yagiu, R. (1944). Studies on the ciliates Condylostoma I: taxonomy. J. Sci. Hiroshima Univ., (Ser. B., Div.
1)10: 161-184.