碘泡虫属的修订及中国部分碘泡虫物种的分类学研究
|
摘要
粘体动物(Myxozoa)是一类世界性分布的后生动物寄生虫,主要寄生于鱼类,能导致鱼类重大病害的发生,并造成严重的经济损失和生态污染。碘泡虫(Myxobolus)是粘体动物的最大类群,全世界报道850多种,我国记录320多种。本文运用形态学、组织学及分子生物学方法,研究了碘泡虫属的分类地位,系统整理了我国的碘泡虫种类,并对采集到的12种碘泡虫进行了鉴定和补充描述。主要结果如下：
一、碘泡虫属与尾孢虫属的分类关系研究：(1)本文发现丑陋圆形碘泡虫(Myxobolus turpisrotundus)、苍梧碘泡虫(M. tsangwuensis)、贝壳碘泡虫(M. musseliusae)、口腔碘泡虫(M. oralis)的部分孢子具有与尾孢虫类似的尾突。孢子发生研究显示碘泡虫的尾突与尾孢虫的尾突均由壳瓣基部延伸形成,说明碘泡虫尾突与尾孢虫尾突具有共同的来源和发育过程。(2)利用碘泡虫、尾孢虫及相近种属的18S rDNA序列作为分子标记进行系统发育分析,发现碘泡虫与尾孢虫在系统发育树上交替聚在一起没有形成独立的分支,说明碘泡虫属和尾孢虫属均非单系类群。(3)进化分析发现碘泡虫与尾孢虫的祖先种类不具有尾突,尾突是在后来的进化过程中出现的,并且尾突分化事件在整个进化过程中独立发生了多次。同时,本文发现祖先种类分化出尾突之后还可以在后来的进化过程中再次失去尾突重新形成无尾的种类。综上所述,尾突并非尾孢虫特有的形态特征亦不能代表尾孢虫之间的亲缘关系,无法作为区分碘泡虫属和尾孢虫属的分类指标。经典分类学依据尾突建立的尾孢虫属是一个无效属,加之其与碘泡虫具有极近的亲缘关系又无其他形态差异,应将其并入碘泡虫属。
二、中国碘泡虫物种的整理：(1)完善我国碘泡虫物种名录：将我国记录的30种尾孢虫、8种粘体虫修订入碘泡虫属；通过对我国碘泡虫物种进行系统梳理,厘定11物种,其中同名异物9种,同物异名1种,无效种1种,取消41种中国无效记录。(2)明确每个物种的典型宿主和寄生部位：根据首次描述时的宿主为典型宿主的原则明确了我国每个碘泡虫种类的典型宿主和寄生部位。根据上述结果建立了中国碘泡虫名录,包括326个碘泡虫种类,寄生于中国115种淡水鱼类,1种海水鱼类,其中寄生鲫、鲤的碘泡虫种类最多,分别为52种和25种。
三、12种碘泡虫的鉴定与补充描述：(1)厘定寄生异育银鲫体表、鳃、鳍条的碘泡虫为丑陋圆形碘泡虫(M. turpisrotundus)而非圆形碘泡虫(M. rotundus)。(2)厘定寄生我国鲤鳃丝的碘泡虫为贝壳碘泡虫(M. musseliusae)而非异型碘泡虫(M. dispar)。(3)鉴定寄生异育银鲫咽部的碘泡虫为洪湖碘泡虫新种(M. honghuensis sp.n.)。(4)鉴定寄生异育银鲫口腔上颌的碘泡虫为口腔碘泡虫新种(M. or alis sp. n.)。(5)依据本文采集种群对山东碘泡虫(M. shantungesis)、野鲤碘泡虫(M. koi)、鲤肠碘泡虫(M. cyprinicola)、吴李碘泡虫(M. wulii)、瓶囊碘泡虫(M.ampullicapsulatus)、倪李碘泡虫(M. nielii)、多涅茨碘泡虫(M. doneci)、鳜碘泡虫(M. sinipercus)等8种碘泡虫的孢子形态、超微结构、寄生特性及18S rDNA序列进行了补足性重描述。
Myxozoans are an economically important group of metazoan endoparasites infecting various fish throughtout the world. The diseases caused by them have resulted in mass mortality or loss of economic value of the cultured and wild fish. Myxobolus Butschli,1882is the genera with the greatest number of described species, with over850species identified throughout the world and320species reported in China. In the present paper, we investigated the taxonomy of Myxobolus, made a systematic revisions on Myxobolus species in China, and identified12Myxobolus species by morphological, histological, and molecular methods. The main results are as follows.
Taxonamy of Myxobolus and Henneguya:(1) In the present study, the spores of Myxobolus turpisrotundus, M. tsangwuensis, M. musseliusae and M. oralis were observed intermixed with atypical spores bearing caudal appendages. Ultrastructural observation showed that the caudal appendages on both Myxobolus and Henneguya spores came from the elongation of the posterior of the valves.(2) Phylogenetic analysis based on all the available ribosomal DNA sequence data of Myxobolus and Henneguya species still did not support a separation of these two genera.(3) Tracing the history of caudal appendages evolution showed that the morphology of the ancestral species of Myxobolus and Henneguya did not possess the caudal appendages and that caudal appendages have arisen on multiple occasions independently during the evolution. It is worth noting that some Henneguya-type species could lose their caudal appendages to form Myxobolus-type species. The above results indicated that caudal appendages were not a valid character to separate Myxobolus-type and Henneguya-type species into two separate genera. Genus Henneguya is not valid and its species should be assigned to genus Myxobolus.
Systematic revisions on the Myxobolus species in China:(1) Revisons on the list of Myxobolus species in China:30Henneguya and8Myxosoma species described in China were assigned to Myxobolus;9homonyms,1synonym,1invalid species, and41invalid Chinese records were identified and removed from the Chinese Myxobolus list.(2) Identification of the type host and the site of infection:the type host and the site of infection of the Myxobolus species in China were identified according to the original descriptions. In conclusion, the list of Myxobolus species in China includes326normal species infecting115different freshwater fish,1marine fish. Among them,52and25species were identified from gold fish and common carp, respectively.
Identification and redescription of12Myxobolus species collected in China:(1) The species infecting the skin, gills and fins of gibel carp Carassius auratus gibelio was identified as M. turpisrotundus.(2) The species infecting the gills of common carp Cyprinus carpio was identified as M. musseliusae.(3) The species infecting the pharynx of gibel carp was identified as M. honghuensis sp. n.(4) The species infecting the palate in the mouth of gibel carp was identified as M. oralis sp. n.(5) M. shantungesis, M. koi, M. cyprinicola, M. wulii, M. ampullicapsulatus, M. nielii, M. doneci, and M. sinipercus were redescribed by their morphological, ultrastructural, histological and molecular characteristics.
一、碘泡虫属与尾孢虫属的分类关系研究：(1)本文发现丑陋圆形碘泡虫(Myxobolus turpisrotundus)、苍梧碘泡虫(M. tsangwuensis)、贝壳碘泡虫(M. musseliusae)、口腔碘泡虫(M. oralis)的部分孢子具有与尾孢虫类似的尾突。孢子发生研究显示碘泡虫的尾突与尾孢虫的尾突均由壳瓣基部延伸形成,说明碘泡虫尾突与尾孢虫尾突具有共同的来源和发育过程。(2)利用碘泡虫、尾孢虫及相近种属的18S rDNA序列作为分子标记进行系统发育分析,发现碘泡虫与尾孢虫在系统发育树上交替聚在一起没有形成独立的分支,说明碘泡虫属和尾孢虫属均非单系类群。(3)进化分析发现碘泡虫与尾孢虫的祖先种类不具有尾突,尾突是在后来的进化过程中出现的,并且尾突分化事件在整个进化过程中独立发生了多次。同时,本文发现祖先种类分化出尾突之后还可以在后来的进化过程中再次失去尾突重新形成无尾的种类。综上所述,尾突并非尾孢虫特有的形态特征亦不能代表尾孢虫之间的亲缘关系,无法作为区分碘泡虫属和尾孢虫属的分类指标。经典分类学依据尾突建立的尾孢虫属是一个无效属,加之其与碘泡虫具有极近的亲缘关系又无其他形态差异,应将其并入碘泡虫属。
二、中国碘泡虫物种的整理：(1)完善我国碘泡虫物种名录：将我国记录的30种尾孢虫、8种粘体虫修订入碘泡虫属；通过对我国碘泡虫物种进行系统梳理,厘定11物种,其中同名异物9种,同物异名1种,无效种1种,取消41种中国无效记录。(2)明确每个物种的典型宿主和寄生部位：根据首次描述时的宿主为典型宿主的原则明确了我国每个碘泡虫种类的典型宿主和寄生部位。根据上述结果建立了中国碘泡虫名录,包括326个碘泡虫种类,寄生于中国115种淡水鱼类,1种海水鱼类,其中寄生鲫、鲤的碘泡虫种类最多,分别为52种和25种。
三、12种碘泡虫的鉴定与补充描述：(1)厘定寄生异育银鲫体表、鳃、鳍条的碘泡虫为丑陋圆形碘泡虫(M. turpisrotundus)而非圆形碘泡虫(M. rotundus)。(2)厘定寄生我国鲤鳃丝的碘泡虫为贝壳碘泡虫(M. musseliusae)而非异型碘泡虫(M. dispar)。(3)鉴定寄生异育银鲫咽部的碘泡虫为洪湖碘泡虫新种(M. honghuensis sp.n.)。(4)鉴定寄生异育银鲫口腔上颌的碘泡虫为口腔碘泡虫新种(M. or alis sp. n.)。(5)依据本文采集种群对山东碘泡虫(M. shantungesis)、野鲤碘泡虫(M. koi)、鲤肠碘泡虫(M. cyprinicola)、吴李碘泡虫(M. wulii)、瓶囊碘泡虫(M.ampullicapsulatus)、倪李碘泡虫(M. nielii)、多涅茨碘泡虫(M. doneci)、鳜碘泡虫(M. sinipercus)等8种碘泡虫的孢子形态、超微结构、寄生特性及18S rDNA序列进行了补足性重描述。
Myxozoans are an economically important group of metazoan endoparasites infecting various fish throughtout the world. The diseases caused by them have resulted in mass mortality or loss of economic value of the cultured and wild fish. Myxobolus Butschli,1882is the genera with the greatest number of described species, with over850species identified throughout the world and320species reported in China. In the present paper, we investigated the taxonomy of Myxobolus, made a systematic revisions on Myxobolus species in China, and identified12Myxobolus species by morphological, histological, and molecular methods. The main results are as follows.
Taxonamy of Myxobolus and Henneguya:(1) In the present study, the spores of Myxobolus turpisrotundus, M. tsangwuensis, M. musseliusae and M. oralis were observed intermixed with atypical spores bearing caudal appendages. Ultrastructural observation showed that the caudal appendages on both Myxobolus and Henneguya spores came from the elongation of the posterior of the valves.(2) Phylogenetic analysis based on all the available ribosomal DNA sequence data of Myxobolus and Henneguya species still did not support a separation of these two genera.(3) Tracing the history of caudal appendages evolution showed that the morphology of the ancestral species of Myxobolus and Henneguya did not possess the caudal appendages and that caudal appendages have arisen on multiple occasions independently during the evolution. It is worth noting that some Henneguya-type species could lose their caudal appendages to form Myxobolus-type species. The above results indicated that caudal appendages were not a valid character to separate Myxobolus-type and Henneguya-type species into two separate genera. Genus Henneguya is not valid and its species should be assigned to genus Myxobolus.
Systematic revisions on the Myxobolus species in China:(1) Revisons on the list of Myxobolus species in China:30Henneguya and8Myxosoma species described in China were assigned to Myxobolus;9homonyms,1synonym,1invalid species, and41invalid Chinese records were identified and removed from the Chinese Myxobolus list.(2) Identification of the type host and the site of infection:the type host and the site of infection of the Myxobolus species in China were identified according to the original descriptions. In conclusion, the list of Myxobolus species in China includes326normal species infecting115different freshwater fish,1marine fish. Among them,52and25species were identified from gold fish and common carp, respectively.
Identification and redescription of12Myxobolus species collected in China:(1) The species infecting the skin, gills and fins of gibel carp Carassius auratus gibelio was identified as M. turpisrotundus.(2) The species infecting the gills of common carp Cyprinus carpio was identified as M. musseliusae.(3) The species infecting the pharynx of gibel carp was identified as M. honghuensis sp. n.(4) The species infecting the palate in the mouth of gibel carp was identified as M. oralis sp. n.(5) M. shantungesis, M. koi, M. cyprinicola, M. wulii, M. ampullicapsulatus, M. nielii, M. doneci, and M. sinipercus were redescribed by their morphological, ultrastructural, histological and molecular characteristics.
引文
[1]陈启鎏.湖北省鱼病病原区系图志.北京：科学出版社,1973.
[2]陈启鎏,马成伦.中国动物志：粘体动物门,粘孢子虫纲.北京：科学出版社,1998.
[3]陈启鎏,谢杏人.斑鳢和乌鳢寄生孢子虫的研究.水生生物学集刊,1960,2：171-196.
[4]陈信廉,邹为民.鱼类寄生粘孢子虫两新种(粘孢子目：粘体科,碘泡科).动物分类学报,1998,23：113-116.
[5]董江丽,赵元莙,唐发辉,索栋.黄颡单尾虫(粘体门,双壳目)的重描述及基于28S rDNA和ITS-5.8S序列的系统地位分析.动物分类学报,2011,36-84-92.
[6]冯淑娟,汪建国.丹江口水库鱼类寄生粘孢子虫四新种.水生生物学报,1990,14：68-74.
[7]龚小宁,鲁义善,汪建国.淡水鱼类寄生粘孢子虫两新种的记述.水生生物学报,2004,28：418-420.
[8]黄城.厦门沿岸海水鱼类寄生粘孢子虫的分类学研究.[硕士学位论文].重庆：重庆师范大学,2006.
[9]李文宽.异育银螂寄生粘体虫属一新种(粘抱子纲：双壳目：粘体科).动物分类学报,22.-115-117.
[10]李卓凡,吴宝华.杭州西湖鱼类寄生粘抱子虫区系.杭州大学学报,1983,10-207-220.
[11]鲁义善,聂品.淡水鱼类粘孢子虫的18S rDNA分子系统学研究.水生生物学报,28：587-591.
[12]罗芬,张其中.寄生鲤的粘孢子虫一新种(粘孢子纲：双壳目).动物分类学报,26：162-165.
[13]马成伦.长江上游鱼类寄生粘抱子虫Ⅰ.重庆师范学院学报(自然科学版),1993,10：9-15.
[14]马成伦,董新民,王慈生.四川省鱼类寄生粘抱子虫新属新种(丝孢子纲：粘孢子目).动物分类学报,1982,7：341-352.
[15]马成伦,董新民,王慈生.四川省鱼类寄生粘孢子虫Ⅵ尾孢虫和单极虫(粘孢子纲：双壳目).重庆师范学院学报(自然科学版),1999,16：12-19.
[16]马成伦,王慈生,蔡志华.云南省鱼类寄生粘孢子虫Ⅰ尾孢子虫和拟尾孢虫(粘孢子目：碘泡科).动物分类学报,1986,11：236-244.
[17]倪达书,李连祥.湖北省花马湖鱼类寄生粘孢子虫的研究Ⅱ粘孢子虫新种的描述.动物分类学报,1992,17：133-150.
[18]冉江波.嘉陵江下游鲶类鱼寄生虫区系.[硕士学位论文].重庆：西南大学,2006.
[19]索栋,赵元若.尼氏单极虫(粘体门,双壳目)重描述及基于18S rDNA系统地位分析.2010,35：90-95
[20]吴宝华,姜乃澄,王淑霞.浙江省淡水鱼类寄生粘抱子虫三新种(粘孢子目：粘体科、两极科).动物分类学报,1985,10：337-340.
[21]吴宝华,李卓凡.浙江省淡水鱼类寄生粘孢子虫六新种(粘孢子虫目).动物分类学报,1986,11：1-9.
[22]吴宝华,王淑霞.寄生在东北银鲫Carassius auratus gibelio (Bloch)的粘孢子虫三新种.杭州大学学报(自然科学版),1982,9：91-94.
[23]吴英松,汪建国.寄生异育银鲫的碘泡虫一新种(粘孢子纲：双壳目,碘泡科).水生生物学报,1997,21：268-270.
[24]吴英松,汪建国.圆形碘泡虫孢子发生的超微结构研究.水生生物学报,2003,25：61-69.
[25]吴英松,汪建国.圆形碘泡虫和关桥碘泡虫孢子的光镜及扫描电镜观察.水生生物学报,2003,27：264-268.
[26]吴灶和,陈启鎏.淡水鱼类寄生粘孢子虫九新种.水生生物学报,1987,11：161-170.
[27]吴灶和,吴建平,华鼎可.南海鱼类寄生粘抱子虫的研究Ⅰ角形虫属五新种.热带海洋,1993,12：79-87.
[28]吴灶和,吴建平,华鼎可.南海鱼类寄生粘抱子虫的研究Ⅱ篮子鱼两极虫与尖吻鲈尾孢虫两新种.热带海洋,1994,13：67-71.
[29]叶灵通.鲤、鲫寄生的几种粘孢子虫的形态、分子系统发育及季节动态研究.[博 士学位论文].武汉：中国科学院水生生物研究所,2012.
[30]章晋勇.圆形碘泡虫抗原性及其与宿主相互作用研究.[博士学位论文].武汉：中国科学院水生生物研究所,2006.
[31]赵元若,马成伦.中国西部淡水鱼类寄生粘孢子虫的记述.动物分类学报,1994,19：257-267.
[32]赵元若,宋微波.黄海鱼类寄生角形虫属粘孢子虫一新种及四新纪录(粘体门,粘孢子纲,双壳目).动物分类学报,2003,28：13-19.
[33]赵元著,宋微波.青岛近岸海洋鱼类4种寄生薄壳虫属粘孢子虫,包括1新种的研究(粘体门,双壳目).水生生物学报,2009,33：61-66.
[34]周杨.中国近海部分海水鱼类寄生粘孢子虫的分类学研究与地理分布.[硕士学位论文].重庆：重庆师范大学,2007.
[35]Anderson CL, Canning EU, Okamura B. A triploblast origin for myxozoa? Nature, 1998,392:346-347.
[36]Bahri S. Abnormal forms of Myxobolus bizerti and Myxobolus mulleri (Myxosporea: Bivalvulida) spores with caudal appendages. Bull Eur Assoc Fish Pathol,2008,28: 252-255.
[37]Bartosova P, Fiala I, Jirku M, Cinkova M, Caffara M, Fioravanti ML, Atkinson SD, Bartholomew JL, Holzer AS. Sphaerospora sensu stricto:Taxonomy, diversity and evolution of a unique lineage of myxosporeans (Myxozoa). Mol Phylogenet Evol, 2013,68:93-105.
[38]Camus AC, Griffin MJ. Molecular characterization and histopathology of Myxobolus koi infecting the gills of a koi, Cyprinus carpio, with an amended morphological description of the agent. J Parasitol,2010,96:116-124.
[39]Canning EU, Curry A, Feist SW, Longshaw M, Okamura B. A new class and order of myxozoans to accommodate parasites of bryozoans with ultrastructural observations on Tetracapsula bryosalmonae (pkx organism). J Eukaryot Microbiol, 2000,47:456-468.
[40]Canning EU, Okamura B. Biodiversity and evolution of the myxozoa. Adv Parasitol, 2004,56:43-131.
[41]Carriero MM, Adriano EA, Silva MR, Ceccarelli PS, Maia AA. Molecular phylogeny of the Myxobolus and Henneguya genera with several new south american species. PLoS One,2013,8:e73713.
[42]Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol Biol Evol,2000,17:540-552.
[43]Cavalier-Smith T, Allsopp M, Chao E, Boury-Esnault N, Vacelet J. Sponge phylogeny, animal monophyly and the origin of the nervous system:18S rRNA evidence. Can J Zool,1996,74:2031-2045.
[44]Crawshaw M, Sweeting R. Myxobolus koi Kudo,1919 a new record for britain. J Fish Dis,1986,95:465-467.
[45]Dayoub A, Molnar K, Salman H, Al-Samman A, Szekely C. Myxobolus infections of common carp (Cyprinus carpio) in syrian fish farms. Acta Vet Hung,2007,55: 501-509.
[46]Eiras JC. Synopsis of the species of the genus Henneguya Thelohan, 1892 (Myxozoa:Myxosporea:Myxobolidae). Syst Parasitol,2002,52:43-54.
[47]Eiras JC, Adriano EA. A checklist of new species of Henneguya Thelohan,1892 (Myxozoa: Myxosporea, myxobolidae) described between 2002 and 2012. Syst Parasitol,2012,83:95-104.
[48]Eiras JC, Molnar K, Lu YS. Synopsis of the species of Myxobolus Biitschli,1882 (Myxozoa: Myxosporea: Myxobolidae). Syst Parasitol,2005,61:1-46.
[49]El-Mansy A. Revision of Myxobolus heterosporus Baker,1963 (syn. Myxosoma heterospora) (Myxozoa: Myxosporea) in african records. Dis Aquat Organ, 2005, 63:205-214.
[50]El-Mansy A, Molnar K. Development of Myxobolus hungaricus (Myxosporea: Myxobolidae) in oligochaete alternate hosts. Dis Aquat Organ, 1997,31:227-232.
[51]El-Matbouli M, Hoffmann RW. Light and electron microscopic studies on the chronological development of Myxobols cerebralis to the actinosporean stage in Tubifex tubifex to the antinsoporean stage triactinomyxon. Int J Parasitol, 1998,28: 495-217.
[52]Eszterbauer E. Genetic relationship among gill-infecting Myxobolus species (Myxosporea) of cyprinids:Molecular evidence of importance of tissue-specificity. Dis Aquat Organ,2004,58:35-40.
[53]Evans NM, Holder MT, Barbeitos MS, Okamura B, Cartwright P. The phylogenetic position of myxozoa: Exploring conflicting signals in phylogenomic and ribosomal data sets. Mol Biol Evol,2010,27:2733-2746.
[54]Ferguson JA, Atkinson SD, Whipps CM, Kent ML. Molecular and morphological analysis of Myxobolus spp. Of salmonid fishes with the description of a new Myxobolus species. J Parasitol,2008,94:1322-1334.
[55]Fiala Ⅰ. The phylogeny of Myxosporea (Myxozoa) based on small subunit ribosomal RNA gene analysis. Int J Parasitol,2006,36:1521-1534.
[56]Fiala I, Bartosova P. History of myxozoan character evolution on the basis of rDNA and EF-2 data. BMC Evol Biol,2010,10:228.
[57]Gilbert MA, Granath WO, Jr. Whirling disease of salmonid fish: Life cycle, biology, and disease. J Parasitol,2003,89:658-667.
[58]Griffin MJ, Goodwin AE. Thelohanellus toyamai (syn. Myxobolus toyamai) infecting the gills of koi Cyprinus carpio in the Eastern united states. J Parasitol, 2011,97:493-502.
[59]Gui J, Zhou L. Genetic basis and breeding application of clonal diversity and dual reproduction modes in polyploid Carassius auratus gibelio. Sci China Life Sci, 2010,53:409-415.
[60]Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate Maximum-Likelihood phylogenies:Assessing the performance of phyml 3.0. Syst Biol,2010,59:307-321.
[61]Gunter N, Adlard R. The demise of Leptotheca Thelohan,1895 (Myxozoa: Myxosporea: Ceratomyxidae) and assignment of its species to Ceratomyxa Thelohan,1892 (Myxosporea: Ceratomyxidae), Ellipsomyxa Koie,2003 (Myxosporea: Ceratomyxidae), Myxobolus Butschli, 1882 and Sphaerospora Thelohan,1892 (Myxosporea: Sphaerosporidae). Syst Parasitol,2010,75:81-104.
[62]Hall TA. Bioedit: A user-friendly biological sequence alignment editor and analysis program for windows 95/98/nt. Nucleic Acids Symposium Series,1999,41:95-98.
[63]Hanelt B, Van Schyndel D, Adema CM, Lewis LA, Loker ES. The phylogenetic position of Rhopalura ophiocomae (Orthonectida) based on 18S ribosomal DNA sequence analysis. Mol Biol Evol,1996,13:1187-1191.
[64]Hoffman GL. Parasites of North American freshwater fishes, 2nd Edn. Comstock Publishing Associates, New York: Ithaca,1999.
[65]Holzer AS, Sommerville C, Wootten R. Molecular relationships and phylogeny in a community of myxosporeans and actinosporeans based on their 18S rDNA sequences. Int J Parasitol,2004,34:1099-1111.
[66]Honigberg B, Balamuth W, Bovee E, CORLISS J, Gojdics M, Hall R, Kudo R, Levine N, Loetblich A, J R, Weiser J, Wenrich D. A revised classification of the phylum protozoa. J Protozool,1964,11:7-20.
[67]Ivasik VM, Gevkan II, Vorona NJ. Gill myxosporeosis in carp. Zool Zh (In Russian), 1967,46:941-943.
[68]Jimenez-Guri E, Philippe H, Okamura B, Holland PW. Buddenbrockia is a cnidarian worm. Science,2007,317:116-118.
[69]Jurine LL. Histoire des poissons du lac leman. Mem. Soc. Phys. Hist. Nat.:Geneve, 1825.
[70]Kallert DM, Eszterbauer E, El-Matoubli M, Erseus C, Haas W. The life cycle of Henneguya nuessli.u Schuberg & Schroder, 1905 (Myxozoa) involves a triactinomyxon-type actinospore. J Fish Dis,2005,28:71-79.
[71]Katoh K, Misawa K, Kuma K, Miyata T. Mafft: A novel method for rapid multiple sequence alignment based on fast fourier transform. Nucleic Acids Res, 2002, 30: 3059-3066.
[72]Kaur H, Singh R. A synopsis of the species of Myxobolus Biitschli,1882 (Myxozoa: Bivalvulida) parasitising indian fishes and a revised dichotomous key to myxosporean genera. Syst Parasitol,2012,81:17-37.
[73]Kent ML, Andree KB, Bartholomew JL, El-Matbouli M, Desser SS, Devlin RH, Feist SW, Hedrick RP, Hoffmann RW, Khattra J, Hallett SL, Lester RJ, Longshaw M, Palenzeula O, Siddall ME, Xiao C. Recent advances in our knowledge of the Myxozoa. J Eukaryot Microbiol,2001,48:395-413.
[74]Koie M, Whipps CM, Kent ML. Ellipsomyxa gobii (Myxozoa: Ceratomyxidae) in the common goby Pomatoschistus microps (Teleostei: Gobiidae) uses Nereis spp. (Annelida: Polychaeta) as invertebrate hosts. Folia Parasitol,2004,51:14-18.
[75]Kudo R. Protozoology. 5th ed. Thomas Publisher:Springfield Illinois USA, 1977.
[76]Landsberg JH, Lom J. Taxonomy of the genera of the Myxobolus/Myxosoma group (Myxobolidae: Myxosporea), current listing of species and revision of synonyms. Syst Parasitol,1991,18:165-186.
[77]Levine N. Taxonomy of protozoa. J Parasitol,1970,56:208-209.
[78]Levine N, Corliss J, Cox F, Deroux G, Grain J, Honigberg BM, Leedale GF, Loeblich AR,3rd, Lom J, Lynn D, Merinfeld EG, Page FC, Poljansky G, Sprague V, Vavra J, Wallace FG. A newly revised classification of the protozoa. J Protozool, 1980,27:37-58.
[79]Liu Y, Gu Z, Zhang Y, Zeng L. Redescription and molecular analysis of Myxobolus shantungensis Hu, 1965 (Myxozoa: Myxosporea) infecting common carp Cyprinus carpio haematopterus. Parasitol Res,2011a,109:1619-1623.
[80]Liu Y, Gu ZM, Luo YL. Some additional data to the occurrence, morphology and validity of Myxobolus turpisrotundus Zhang, 2009 (Myxozoa: Myxosporea). Parasitol Res,2010a,107:67-73.
[81]Liu Y, Jia L, Huang MJ, Gu ZM. Thelohanellus testudineus n. sp. (Myxosporea: Bivalvulida) infecting the skin of allogynogenetic gibel carp Carassius auratus gibelio (Bloch) in China. J Fish Dis,2013a,
[82]Liu Y, Whipps CM, Gu ZM, Huang MJ, He C, Yang HL, Molnar K. Myxobolus musseliusae (Myxozoa: Myxobolidae) from the gills of common carp Cyprinus carpio and revision of Myxobolus dispar recorded in China. Parasitol Res,2013b, 112:289-296.
[83]Liu Y, Whipps CM, Gu ZM, Zeng C, Huang MJ. Myxobolus honghuensis n. sp. (Myxosporea: Bivalvulida) parasitizing the pharynx of allogynogenetic gibel carp Carassius auratus gibelio (bloch) from Honghu lake, China. Parasitol Res, 2012, 110:1331-1336.
[84]Liu Y, Whipps CM, Gu ZM, Zeng LB. Myxobolus turpisrotundus (Myxosporea: Bivalvulida) spores with caudal appendages: Investigating the validity of the genus Henneguya with morphological and molecular evidence. Parasitol Res,2010b,107: 699-706.
[85]Liu Y, Whipps CM, Liu WS, Zeng LB, Gu ZM. Supplemental diagnosis of a myxozoan parasite from common carp Cyprinus carpio:Synonymy of Thelohanellus xinyangensis with Thelohanellus kitauei. Vet Parasitol,2011b,178: 355-359.
[86]Liu Y, Yuan J, Jia L, Huang M, Zhou Z, Gu Z. Supplemental description of Thelohanellus wuhanensis Xiao & Chen,1993 (Myxozoa: Myxosporea) infecting the skin of Carassius auratus gibelio (bloch):Ultrastructural and histological data. Parasitol Int,2014,63:489-491.
[87]Lom J. On a new taxonomic character in myxosporidia as demonstrated in descriptions of two new species of Myxobolus. Folia Parasitol,1969,16:97-103.
[88]Lom J, Arthur JR. A guideline for preparation of species descriptions in myxosporea. J Fish Dis,1989,12:151-156.
[89]Lom J, Dykova I. Fine structure of triactinomyxon early stages and sporogony: Myxosporean and actinosporean features compared. J Protozool,1992,39:16-27.
[90]Lom J, Dykova I. Myxozoan genera: Definition and notes on taxonomy, life-cycle terminology and pathogenic species. Folia Parasitol,2006,53:1-36.
[91]Lom J, Dykova I. Ultrastructural features of the actinosporean phase of Myxosporea (Phylum Myxozoa):a comparative study. Acta Protozool,1997,36:83-103
[92]Lom J, Noble E. Revised classification of the myxosporea Butschli, 1881. Folia Parasitol,1984,31:193-205.
[93]Molnar K. Comments on the host, organ and tissue specificity of fish myxosporeans and on the types of their intrapiscine development. Parasitol Hung, 1994, 27:5-20.
[94]Molnar K. Remarks to the validity of genbank sequences of Myxobolus spp. (Myxozoa, Myxosporidae) infecting Eurasian fishes. Acta Parasitol, 2011,56: 263-269.
[95]Molnar K. Site preference of fish myxosporeans in the gill. Dis Aquat Organ, 2002, 48:197-207.
[96]Molnar K, El-Mansy A, Szekely C, Baska F. Development of Myxobolus dispar (Myxosporea: Myxobolidae) in an oligochaete alternate host, Tubifex tubifex. Folia Parasitol,1999,46:15-21.
[97]Molnar K, Marton S, Eszterbauer E, Szekely C. Comparative morphological and molecular studies on Myxobolus spp. infecting chub from the river Danube, hungary, and description of Myxobolus muellericus sp. n. Dis Aquat Organ,2006,73:49-61.
[98]Molnar K, Marton S, Szekely C, Eszterbauer E. Differentiation of Myxobolus spp. (Myxozoa:Myxobolidae) infecting roach (Rutilus rutilus) in Hungary. Parasitol Res, 2010,107:1137-1150.
[99]Molnar K, Szakolczai J. Halbetegsegek., Budapest: Mezogazdasagi Kiado,1980 (in Hungarian).
[100]Molnar K, Szekely C, Hallett SL, Atkinson SD. Some remarks on the occurrence, host-specificity and validity of Myxobolus rotundus Nemeczek,1911 (Myxozoa: Myxosporea). Syst Parasitol,2009,72:71-79.
[101]Monteiro AS, Okamura B, Holland PW. Orphan worm finds a home: Buddenbrockia is a myxozoan. Mol Biol Evol,2002,19:968-971.
[102]Nemeczek A. Beitrage zur kenntnis der myxo-und microsporidien der fische. Archiv fur Protistenkunde,1911,22:143-169.
[103]Nesnidal MP, Helmkampf M, Bruchhaus Ⅰ, El-Matbouli M, Hausdorf B. Agent of whirling disease meets orphan worm: Phylogenomic analyses firmly place Myxozoa in cnidaria. PLoS One, 2013,8:e54576.
[104]Okamura B, Anderson C, Canning E. An ancient clade of myxozoans parasitic in freshwater bryozoans. American Zoologist,2000,40:1157.
[105]Okamura B, Canning EU. Orphan worms and homeless parasites enhance bilaterian diversity. Trends in Ecology and Evolution.2003,18:633-639.
[106]Okamura B, Curry A, Wood TS, Canning EU. Ultrastructure of Buddenbrockia identifies it as a myxozoan and verifies the bilaterian origin of the Myxozoa. .'arasitology,2002,124:215-223.
[107]Oumouna M, Halett SL, Hoffmann RW, El-Matbouli M. Seasonal occurrence of actinsporeans (Myxozoa) and oligochaetes (Annelida) at a trout hatchery in Bavaria, Germany. Parasitol Res,2002,89:170-184
[108]Posada D. Jmodeltest: Phylogenetic model averaging. Mol Biol Evol,2008,25: 1253-1256.
[109]Ronquist F, Huelsenbeck JP. Mrbayes 3:Bayesian phylogenetic inference under mixed models. Bioinformatics,2003,19:1572-1574.
[110]Schlegel M, Lom J, Stechmann A, Bernhard D, Leipe D, Dykova Ⅰ, Sogin ML. Phylogenetic analysis of complete small subunit ribosomal RNA coding region of Myxidium lieberkuehni:Evidence that Myxozoa are metazoa and related to the Bilateria. Archiv fur Protistenkunde,1996,147:1-9.
[111]Shulman SS. Myxosporidia of the fauna of the ussr. Moscow: Nauka,1966.
[112]Siddall M, Whiting M. Long-branch abstractions. Cladistics,1999, 15:9-24.
[113]Siddall ME, Martin DS, Bridge D, Desser SS, Cone DK. The demise of a phylum of protists:Phylogeny of Myxozoa and other parasitic cnidaria. J Parasitol, 1995,81: 961-967.
[114]Sikama Y. On a new species of Henneguya found in a freshwater fish in China. Journal of the Shangai Science Institute,1938,4:135-138.
[115]Smothers JF, von Dohlen CD, Smith LH, Jr., Spall RD. Molecular evidence that the myxozoan protists are metazoans. Science,1994,265:1719-1721.
[116]Stamatakis A. Raxml-vi-hpc:Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics,2006,22:2688-2690.
[117]Stole A. Actinomyxidies, nouveau groupe de mesozoaires parent des myxosporidies. Bull Intl Acad Sci Boheme,1899,22:1-12.
[118]Swofford DL (2003) PAUP*:phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer, Sunderland, MA.
[119]Szekely C, Hallett SL, Atkinson SD, Molnar K. Complete life cycle of Myxobolus rotundus (Myxosporea: Myxobolidae), a gill myxozoan of common bream Abramis brama. Dis Aquat Organ,2009,85:147-155.
[120]Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. Mega5:Molecular evolutionary genetics analysis using Maximum Likelihood, evolutionary distance, and Maximum Parsimony methods. Mol Biol Evol,2011,28:2731-2739.
[121]Thelohan P. Observations sur les myxosporidies et essai de classification de ces organismes. Bulletin de la Societe Philomatique de Paris,1892,4:167-178.
[122]Thelohan P. Recherches sur les myxosporidies. Bull Sci Fr Belg,1895,26:100-394.
[123]Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG. The clustal-X windows interface:Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res,1997,25:4876-4882.
[124]Walliker D. The nature of the iodinophilous vacuole of myxosporidian spores, and a proposal to synonymize the genus Myxosoma Thelohan,1892 with the genus Myxobolus Butschli,1882. J Protozool,1968,55:571-575.
[125]Wang GT, Yao WJ, Wang JG, Lu YS. Occurrence of Thelohanellosis caused by Thelohanellus wuhanensis (Myxosporea) in juvenile allogynogenetic silver crucian carp Carassius auratus gibelio (bloch), with an observation on the efficacy of fumagillin as a therapeutan. J Fish Dis,2001,24:57-60.
[126]Wang ZW, Zhu HP, Wang D, Jiang FF, Guo W, Zhou L, Gui JF. A novel nucleo-cytoplasmic hybrid clone formed via androgenesis in polyploid gibel carp. BMC Res Notes,2011,4:82.
[127]Weill R. L'interpretation des cnidosporidies et la valeur taxonomique de ieur cnidome Ieur cycle compare a la phase larvaire des narcomeduses cuninides. Travaux de la Station Zoologique de Wimereaux,1938,13:727-744.
[128]Whipps CM, Diggles BK. Kudoa alliaria in flesh of Argentinian hoki Macruronus magellanicus (Gadiformes; Merlucciidae). Dis Aquat Organ,2006,69:259-263.
[129]Whipps CM, El-Matbouli M, Hedrick RP, Blazer V, Kent ML. Myxobolus cerebralis internal transcribed spacer 1 (ITS-1) sequences support recent spread of the parasite to North America and within Europe. Dis Aquat Organ,2004a, 60: 105-108.
[130]Whipps CM, Grossel G, Adlard RD, Yokoyama H, Bryant MS, Munday BL, Kent ML. Phylogeny of the multivalvulidae (Myxozoa: Myxosporea) based on comparative ribosomal DNA sequence analysis. J Parasitol, 2004b, 90: 618-622.
[131]Whipps CM, Kent ML. Phylogeography of the cosmopolitan marine parasite Kudoa thyrsites (Myxozoa: Myxosporea). J Eukaryot Microbiol, 2006, 53:364-373.
[132]Wolf K, Markiw ME. Biology contravenes taxonomy in the Myxozoa: New discoveries show alternation of invertebrate and vertebrate hosts. Science, 1984, 225:1449-1452.
[133]Xi BW, Xie J, Zhou QL, Pan LK, Ge XP. Mass mortality of pond-reared Carassius gibelio caused by Myxobolus ampullicapsulatus in China. Dis Aquat Organ, 2011, 93:257-260.
[134]Ye LT, Li WX, Wu SG, Wang GT. Supplementary studies on Henneguya doneci Schulman,1962 (Myxozoa: Myxosporea) infecting the gill filaments of Carassius auratus gibelio (bloch) in China: Histologic, ultrastructural, and molecular data. Parasitol Res,2012,110:1509-1516.
[135]Yokoyama H, Freeman MA, Yoshnaga T, Ogawa K. Myxobolus buri, the myxosporean parasite causing scoliosis of yellowtail, is synonymous with Myxobolus acanthogobii infecting the brain of the yellowfin goby. Fish Sci, 2004, 70:1036-1042.
[136]Yokoyama H, Inoue D, Kumamaru A, Wakabayashi H. Myxobolus koi (Myxozoa: Myxosporea) forms large-and small-type "cysts" in the gills of common carp. Fish Pathol,1997,32:211-217.
[137]Yokoyama H, Ogawa K, Wakabayashi H. Myxobolus cultus n. sp. (Myxosporea: Myxobolidae) in thegoldfish Carassius auratus transformed from the actinsporean stage in the oligochaete Branchiura sowerbyi. J Parasitol,1995,81:446-451.
[138]Zhang JY, Wang JG, Li AH, Gong XL, Cai TZ. Redescription of Myxobolus pyramidis Chen,1958 (Myxosporea: Bivalvulida). Parasitol Res, 2006, 99: 65-69.
[139]Zhang JY, Wang JG, Li AH, Gong XN. Infection of Myxobolus turpisrotundus sp. n. in allogynogenetic gibel carp, Carassius auratus gibelio (Bloch), with revision of Myxobolus rotundus (s. L.) Nemeczek reported from C. auratus auratus (1.). J Fish Dis,2010a,33:625-638.
[140]Zhang JY, Yokoyama H, Wang JG, Li AH, Gong XN, Ryu-Hasegawa A, Iwashita M, Ogawa K. Utilization of tissue habitats by Myxobolus wulii Landsberg & Lom, 1991 in different carp hosts and disease resistance in allogynogenetic gibel carp: Redescription of M. wulii from China and Japan. J Fish Dis,2010b,33:57-68.
[141]Zhao Y, Sun C, Kent ML, Deng J, Whipps CM. Description of a new species of Myxobolus (Myxozoa: Myxobolidae) based on morphological and molecular data. J Parasitol,2008,94: 737-742.
[142]Zhao YJ, Li NN, Tang FH, Dong JL. Remarks on the validity of Myxobolus ampullicapsulatus and Myxobolus honghuensis (Myxozoa:Myxosporea) based on ssu rDNA sequences. Parasitol Res,2013,112:3817-3823.
[2]陈启鎏,马成伦.中国动物志：粘体动物门,粘孢子虫纲.北京：科学出版社,1998.
[3]陈启鎏,谢杏人.斑鳢和乌鳢寄生孢子虫的研究.水生生物学集刊,1960,2：171-196.
[4]陈信廉,邹为民.鱼类寄生粘孢子虫两新种(粘孢子目：粘体科,碘泡科).动物分类学报,1998,23：113-116.
[5]董江丽,赵元莙,唐发辉,索栋.黄颡单尾虫(粘体门,双壳目)的重描述及基于28S rDNA和ITS-5.8S序列的系统地位分析.动物分类学报,2011,36-84-92.
[6]冯淑娟,汪建国.丹江口水库鱼类寄生粘孢子虫四新种.水生生物学报,1990,14：68-74.
[7]龚小宁,鲁义善,汪建国.淡水鱼类寄生粘孢子虫两新种的记述.水生生物学报,2004,28：418-420.
[8]黄城.厦门沿岸海水鱼类寄生粘孢子虫的分类学研究.[硕士学位论文].重庆：重庆师范大学,2006.
[9]李文宽.异育银螂寄生粘体虫属一新种(粘抱子纲：双壳目：粘体科).动物分类学报,22.-115-117.
[10]李卓凡,吴宝华.杭州西湖鱼类寄生粘抱子虫区系.杭州大学学报,1983,10-207-220.
[11]鲁义善,聂品.淡水鱼类粘孢子虫的18S rDNA分子系统学研究.水生生物学报,28：587-591.
[12]罗芬,张其中.寄生鲤的粘孢子虫一新种(粘孢子纲：双壳目).动物分类学报,26：162-165.
[13]马成伦.长江上游鱼类寄生粘抱子虫Ⅰ.重庆师范学院学报(自然科学版),1993,10：9-15.
[14]马成伦,董新民,王慈生.四川省鱼类寄生粘抱子虫新属新种(丝孢子纲：粘孢子目).动物分类学报,1982,7：341-352.
[15]马成伦,董新民,王慈生.四川省鱼类寄生粘孢子虫Ⅵ尾孢虫和单极虫(粘孢子纲：双壳目).重庆师范学院学报(自然科学版),1999,16：12-19.
[16]马成伦,王慈生,蔡志华.云南省鱼类寄生粘孢子虫Ⅰ尾孢子虫和拟尾孢虫(粘孢子目：碘泡科).动物分类学报,1986,11：236-244.
[17]倪达书,李连祥.湖北省花马湖鱼类寄生粘孢子虫的研究Ⅱ粘孢子虫新种的描述.动物分类学报,1992,17：133-150.
[18]冉江波.嘉陵江下游鲶类鱼寄生虫区系.[硕士学位论文].重庆：西南大学,2006.
[19]索栋,赵元若.尼氏单极虫(粘体门,双壳目)重描述及基于18S rDNA系统地位分析.2010,35：90-95
[20]吴宝华,姜乃澄,王淑霞.浙江省淡水鱼类寄生粘抱子虫三新种(粘孢子目：粘体科、两极科).动物分类学报,1985,10：337-340.
[21]吴宝华,李卓凡.浙江省淡水鱼类寄生粘孢子虫六新种(粘孢子虫目).动物分类学报,1986,11：1-9.
[22]吴宝华,王淑霞.寄生在东北银鲫Carassius auratus gibelio (Bloch)的粘孢子虫三新种.杭州大学学报(自然科学版),1982,9：91-94.
[23]吴英松,汪建国.寄生异育银鲫的碘泡虫一新种(粘孢子纲：双壳目,碘泡科).水生生物学报,1997,21：268-270.
[24]吴英松,汪建国.圆形碘泡虫孢子发生的超微结构研究.水生生物学报,2003,25：61-69.
[25]吴英松,汪建国.圆形碘泡虫和关桥碘泡虫孢子的光镜及扫描电镜观察.水生生物学报,2003,27：264-268.
[26]吴灶和,陈启鎏.淡水鱼类寄生粘孢子虫九新种.水生生物学报,1987,11：161-170.
[27]吴灶和,吴建平,华鼎可.南海鱼类寄生粘抱子虫的研究Ⅰ角形虫属五新种.热带海洋,1993,12：79-87.
[28]吴灶和,吴建平,华鼎可.南海鱼类寄生粘抱子虫的研究Ⅱ篮子鱼两极虫与尖吻鲈尾孢虫两新种.热带海洋,1994,13：67-71.
[29]叶灵通.鲤、鲫寄生的几种粘孢子虫的形态、分子系统发育及季节动态研究.[博 士学位论文].武汉：中国科学院水生生物研究所,2012.
[30]章晋勇.圆形碘泡虫抗原性及其与宿主相互作用研究.[博士学位论文].武汉：中国科学院水生生物研究所,2006.
[31]赵元若,马成伦.中国西部淡水鱼类寄生粘孢子虫的记述.动物分类学报,1994,19：257-267.
[32]赵元若,宋微波.黄海鱼类寄生角形虫属粘孢子虫一新种及四新纪录(粘体门,粘孢子纲,双壳目).动物分类学报,2003,28：13-19.
[33]赵元著,宋微波.青岛近岸海洋鱼类4种寄生薄壳虫属粘孢子虫,包括1新种的研究(粘体门,双壳目).水生生物学报,2009,33：61-66.
[34]周杨.中国近海部分海水鱼类寄生粘孢子虫的分类学研究与地理分布.[硕士学位论文].重庆：重庆师范大学,2007.
[35]Anderson CL, Canning EU, Okamura B. A triploblast origin for myxozoa? Nature, 1998,392:346-347.
[36]Bahri S. Abnormal forms of Myxobolus bizerti and Myxobolus mulleri (Myxosporea: Bivalvulida) spores with caudal appendages. Bull Eur Assoc Fish Pathol,2008,28: 252-255.
[37]Bartosova P, Fiala I, Jirku M, Cinkova M, Caffara M, Fioravanti ML, Atkinson SD, Bartholomew JL, Holzer AS. Sphaerospora sensu stricto:Taxonomy, diversity and evolution of a unique lineage of myxosporeans (Myxozoa). Mol Phylogenet Evol, 2013,68:93-105.
[38]Camus AC, Griffin MJ. Molecular characterization and histopathology of Myxobolus koi infecting the gills of a koi, Cyprinus carpio, with an amended morphological description of the agent. J Parasitol,2010,96:116-124.
[39]Canning EU, Curry A, Feist SW, Longshaw M, Okamura B. A new class and order of myxozoans to accommodate parasites of bryozoans with ultrastructural observations on Tetracapsula bryosalmonae (pkx organism). J Eukaryot Microbiol, 2000,47:456-468.
[40]Canning EU, Okamura B. Biodiversity and evolution of the myxozoa. Adv Parasitol, 2004,56:43-131.
[41]Carriero MM, Adriano EA, Silva MR, Ceccarelli PS, Maia AA. Molecular phylogeny of the Myxobolus and Henneguya genera with several new south american species. PLoS One,2013,8:e73713.
[42]Castresana J. Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Mol Biol Evol,2000,17:540-552.
[43]Cavalier-Smith T, Allsopp M, Chao E, Boury-Esnault N, Vacelet J. Sponge phylogeny, animal monophyly and the origin of the nervous system:18S rRNA evidence. Can J Zool,1996,74:2031-2045.
[44]Crawshaw M, Sweeting R. Myxobolus koi Kudo,1919 a new record for britain. J Fish Dis,1986,95:465-467.
[45]Dayoub A, Molnar K, Salman H, Al-Samman A, Szekely C. Myxobolus infections of common carp (Cyprinus carpio) in syrian fish farms. Acta Vet Hung,2007,55: 501-509.
[46]Eiras JC. Synopsis of the species of the genus Henneguya Thelohan, 1892 (Myxozoa:Myxosporea:Myxobolidae). Syst Parasitol,2002,52:43-54.
[47]Eiras JC, Adriano EA. A checklist of new species of Henneguya Thelohan,1892 (Myxozoa: Myxosporea, myxobolidae) described between 2002 and 2012. Syst Parasitol,2012,83:95-104.
[48]Eiras JC, Molnar K, Lu YS. Synopsis of the species of Myxobolus Biitschli,1882 (Myxozoa: Myxosporea: Myxobolidae). Syst Parasitol,2005,61:1-46.
[49]El-Mansy A. Revision of Myxobolus heterosporus Baker,1963 (syn. Myxosoma heterospora) (Myxozoa: Myxosporea) in african records. Dis Aquat Organ, 2005, 63:205-214.
[50]El-Mansy A, Molnar K. Development of Myxobolus hungaricus (Myxosporea: Myxobolidae) in oligochaete alternate hosts. Dis Aquat Organ, 1997,31:227-232.
[51]El-Matbouli M, Hoffmann RW. Light and electron microscopic studies on the chronological development of Myxobols cerebralis to the actinosporean stage in Tubifex tubifex to the antinsoporean stage triactinomyxon. Int J Parasitol, 1998,28: 495-217.
[52]Eszterbauer E. Genetic relationship among gill-infecting Myxobolus species (Myxosporea) of cyprinids:Molecular evidence of importance of tissue-specificity. Dis Aquat Organ,2004,58:35-40.
[53]Evans NM, Holder MT, Barbeitos MS, Okamura B, Cartwright P. The phylogenetic position of myxozoa: Exploring conflicting signals in phylogenomic and ribosomal data sets. Mol Biol Evol,2010,27:2733-2746.
[54]Ferguson JA, Atkinson SD, Whipps CM, Kent ML. Molecular and morphological analysis of Myxobolus spp. Of salmonid fishes with the description of a new Myxobolus species. J Parasitol,2008,94:1322-1334.
[55]Fiala Ⅰ. The phylogeny of Myxosporea (Myxozoa) based on small subunit ribosomal RNA gene analysis. Int J Parasitol,2006,36:1521-1534.
[56]Fiala I, Bartosova P. History of myxozoan character evolution on the basis of rDNA and EF-2 data. BMC Evol Biol,2010,10:228.
[57]Gilbert MA, Granath WO, Jr. Whirling disease of salmonid fish: Life cycle, biology, and disease. J Parasitol,2003,89:658-667.
[58]Griffin MJ, Goodwin AE. Thelohanellus toyamai (syn. Myxobolus toyamai) infecting the gills of koi Cyprinus carpio in the Eastern united states. J Parasitol, 2011,97:493-502.
[59]Gui J, Zhou L. Genetic basis and breeding application of clonal diversity and dual reproduction modes in polyploid Carassius auratus gibelio. Sci China Life Sci, 2010,53:409-415.
[60]Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O. New algorithms and methods to estimate Maximum-Likelihood phylogenies:Assessing the performance of phyml 3.0. Syst Biol,2010,59:307-321.
[61]Gunter N, Adlard R. The demise of Leptotheca Thelohan,1895 (Myxozoa: Myxosporea: Ceratomyxidae) and assignment of its species to Ceratomyxa Thelohan,1892 (Myxosporea: Ceratomyxidae), Ellipsomyxa Koie,2003 (Myxosporea: Ceratomyxidae), Myxobolus Butschli, 1882 and Sphaerospora Thelohan,1892 (Myxosporea: Sphaerosporidae). Syst Parasitol,2010,75:81-104.
[62]Hall TA. Bioedit: A user-friendly biological sequence alignment editor and analysis program for windows 95/98/nt. Nucleic Acids Symposium Series,1999,41:95-98.
[63]Hanelt B, Van Schyndel D, Adema CM, Lewis LA, Loker ES. The phylogenetic position of Rhopalura ophiocomae (Orthonectida) based on 18S ribosomal DNA sequence analysis. Mol Biol Evol,1996,13:1187-1191.
[64]Hoffman GL. Parasites of North American freshwater fishes, 2nd Edn. Comstock Publishing Associates, New York: Ithaca,1999.
[65]Holzer AS, Sommerville C, Wootten R. Molecular relationships and phylogeny in a community of myxosporeans and actinosporeans based on their 18S rDNA sequences. Int J Parasitol,2004,34:1099-1111.
[66]Honigberg B, Balamuth W, Bovee E, CORLISS J, Gojdics M, Hall R, Kudo R, Levine N, Loetblich A, J R, Weiser J, Wenrich D. A revised classification of the phylum protozoa. J Protozool,1964,11:7-20.
[67]Ivasik VM, Gevkan II, Vorona NJ. Gill myxosporeosis in carp. Zool Zh (In Russian), 1967,46:941-943.
[68]Jimenez-Guri E, Philippe H, Okamura B, Holland PW. Buddenbrockia is a cnidarian worm. Science,2007,317:116-118.
[69]Jurine LL. Histoire des poissons du lac leman. Mem. Soc. Phys. Hist. Nat.:Geneve, 1825.
[70]Kallert DM, Eszterbauer E, El-Matoubli M, Erseus C, Haas W. The life cycle of Henneguya nuessli.u Schuberg & Schroder, 1905 (Myxozoa) involves a triactinomyxon-type actinospore. J Fish Dis,2005,28:71-79.
[71]Katoh K, Misawa K, Kuma K, Miyata T. Mafft: A novel method for rapid multiple sequence alignment based on fast fourier transform. Nucleic Acids Res, 2002, 30: 3059-3066.
[72]Kaur H, Singh R. A synopsis of the species of Myxobolus Biitschli,1882 (Myxozoa: Bivalvulida) parasitising indian fishes and a revised dichotomous key to myxosporean genera. Syst Parasitol,2012,81:17-37.
[73]Kent ML, Andree KB, Bartholomew JL, El-Matbouli M, Desser SS, Devlin RH, Feist SW, Hedrick RP, Hoffmann RW, Khattra J, Hallett SL, Lester RJ, Longshaw M, Palenzeula O, Siddall ME, Xiao C. Recent advances in our knowledge of the Myxozoa. J Eukaryot Microbiol,2001,48:395-413.
[74]Koie M, Whipps CM, Kent ML. Ellipsomyxa gobii (Myxozoa: Ceratomyxidae) in the common goby Pomatoschistus microps (Teleostei: Gobiidae) uses Nereis spp. (Annelida: Polychaeta) as invertebrate hosts. Folia Parasitol,2004,51:14-18.
[75]Kudo R. Protozoology. 5th ed. Thomas Publisher:Springfield Illinois USA, 1977.
[76]Landsberg JH, Lom J. Taxonomy of the genera of the Myxobolus/Myxosoma group (Myxobolidae: Myxosporea), current listing of species and revision of synonyms. Syst Parasitol,1991,18:165-186.
[77]Levine N. Taxonomy of protozoa. J Parasitol,1970,56:208-209.
[78]Levine N, Corliss J, Cox F, Deroux G, Grain J, Honigberg BM, Leedale GF, Loeblich AR,3rd, Lom J, Lynn D, Merinfeld EG, Page FC, Poljansky G, Sprague V, Vavra J, Wallace FG. A newly revised classification of the protozoa. J Protozool, 1980,27:37-58.
[79]Liu Y, Gu Z, Zhang Y, Zeng L. Redescription and molecular analysis of Myxobolus shantungensis Hu, 1965 (Myxozoa: Myxosporea) infecting common carp Cyprinus carpio haematopterus. Parasitol Res,2011a,109:1619-1623.
[80]Liu Y, Gu ZM, Luo YL. Some additional data to the occurrence, morphology and validity of Myxobolus turpisrotundus Zhang, 2009 (Myxozoa: Myxosporea). Parasitol Res,2010a,107:67-73.
[81]Liu Y, Jia L, Huang MJ, Gu ZM. Thelohanellus testudineus n. sp. (Myxosporea: Bivalvulida) infecting the skin of allogynogenetic gibel carp Carassius auratus gibelio (Bloch) in China. J Fish Dis,2013a,
[82]Liu Y, Whipps CM, Gu ZM, Huang MJ, He C, Yang HL, Molnar K. Myxobolus musseliusae (Myxozoa: Myxobolidae) from the gills of common carp Cyprinus carpio and revision of Myxobolus dispar recorded in China. Parasitol Res,2013b, 112:289-296.
[83]Liu Y, Whipps CM, Gu ZM, Zeng C, Huang MJ. Myxobolus honghuensis n. sp. (Myxosporea: Bivalvulida) parasitizing the pharynx of allogynogenetic gibel carp Carassius auratus gibelio (bloch) from Honghu lake, China. Parasitol Res, 2012, 110:1331-1336.
[84]Liu Y, Whipps CM, Gu ZM, Zeng LB. Myxobolus turpisrotundus (Myxosporea: Bivalvulida) spores with caudal appendages: Investigating the validity of the genus Henneguya with morphological and molecular evidence. Parasitol Res,2010b,107: 699-706.
[85]Liu Y, Whipps CM, Liu WS, Zeng LB, Gu ZM. Supplemental diagnosis of a myxozoan parasite from common carp Cyprinus carpio:Synonymy of Thelohanellus xinyangensis with Thelohanellus kitauei. Vet Parasitol,2011b,178: 355-359.
[86]Liu Y, Yuan J, Jia L, Huang M, Zhou Z, Gu Z. Supplemental description of Thelohanellus wuhanensis Xiao & Chen,1993 (Myxozoa: Myxosporea) infecting the skin of Carassius auratus gibelio (bloch):Ultrastructural and histological data. Parasitol Int,2014,63:489-491.
[87]Lom J. On a new taxonomic character in myxosporidia as demonstrated in descriptions of two new species of Myxobolus. Folia Parasitol,1969,16:97-103.
[88]Lom J, Arthur JR. A guideline for preparation of species descriptions in myxosporea. J Fish Dis,1989,12:151-156.
[89]Lom J, Dykova I. Fine structure of triactinomyxon early stages and sporogony: Myxosporean and actinosporean features compared. J Protozool,1992,39:16-27.
[90]Lom J, Dykova I. Myxozoan genera: Definition and notes on taxonomy, life-cycle terminology and pathogenic species. Folia Parasitol,2006,53:1-36.
[91]Lom J, Dykova I. Ultrastructural features of the actinosporean phase of Myxosporea (Phylum Myxozoa):a comparative study. Acta Protozool,1997,36:83-103
[92]Lom J, Noble E. Revised classification of the myxosporea Butschli, 1881. Folia Parasitol,1984,31:193-205.
[93]Molnar K. Comments on the host, organ and tissue specificity of fish myxosporeans and on the types of their intrapiscine development. Parasitol Hung, 1994, 27:5-20.
[94]Molnar K. Remarks to the validity of genbank sequences of Myxobolus spp. (Myxozoa, Myxosporidae) infecting Eurasian fishes. Acta Parasitol, 2011,56: 263-269.
[95]Molnar K. Site preference of fish myxosporeans in the gill. Dis Aquat Organ, 2002, 48:197-207.
[96]Molnar K, El-Mansy A, Szekely C, Baska F. Development of Myxobolus dispar (Myxosporea: Myxobolidae) in an oligochaete alternate host, Tubifex tubifex. Folia Parasitol,1999,46:15-21.
[97]Molnar K, Marton S, Eszterbauer E, Szekely C. Comparative morphological and molecular studies on Myxobolus spp. infecting chub from the river Danube, hungary, and description of Myxobolus muellericus sp. n. Dis Aquat Organ,2006,73:49-61.
[98]Molnar K, Marton S, Szekely C, Eszterbauer E. Differentiation of Myxobolus spp. (Myxozoa:Myxobolidae) infecting roach (Rutilus rutilus) in Hungary. Parasitol Res, 2010,107:1137-1150.
[99]Molnar K, Szakolczai J. Halbetegsegek., Budapest: Mezogazdasagi Kiado,1980 (in Hungarian).
[100]Molnar K, Szekely C, Hallett SL, Atkinson SD. Some remarks on the occurrence, host-specificity and validity of Myxobolus rotundus Nemeczek,1911 (Myxozoa: Myxosporea). Syst Parasitol,2009,72:71-79.
[101]Monteiro AS, Okamura B, Holland PW. Orphan worm finds a home: Buddenbrockia is a myxozoan. Mol Biol Evol,2002,19:968-971.
[102]Nemeczek A. Beitrage zur kenntnis der myxo-und microsporidien der fische. Archiv fur Protistenkunde,1911,22:143-169.
[103]Nesnidal MP, Helmkampf M, Bruchhaus Ⅰ, El-Matbouli M, Hausdorf B. Agent of whirling disease meets orphan worm: Phylogenomic analyses firmly place Myxozoa in cnidaria. PLoS One, 2013,8:e54576.
[104]Okamura B, Anderson C, Canning E. An ancient clade of myxozoans parasitic in freshwater bryozoans. American Zoologist,2000,40:1157.
[105]Okamura B, Canning EU. Orphan worms and homeless parasites enhance bilaterian diversity. Trends in Ecology and Evolution.2003,18:633-639.
[106]Okamura B, Curry A, Wood TS, Canning EU. Ultrastructure of Buddenbrockia identifies it as a myxozoan and verifies the bilaterian origin of the Myxozoa. .'arasitology,2002,124:215-223.
[107]Oumouna M, Halett SL, Hoffmann RW, El-Matbouli M. Seasonal occurrence of actinsporeans (Myxozoa) and oligochaetes (Annelida) at a trout hatchery in Bavaria, Germany. Parasitol Res,2002,89:170-184
[108]Posada D. Jmodeltest: Phylogenetic model averaging. Mol Biol Evol,2008,25: 1253-1256.
[109]Ronquist F, Huelsenbeck JP. Mrbayes 3:Bayesian phylogenetic inference under mixed models. Bioinformatics,2003,19:1572-1574.
[110]Schlegel M, Lom J, Stechmann A, Bernhard D, Leipe D, Dykova Ⅰ, Sogin ML. Phylogenetic analysis of complete small subunit ribosomal RNA coding region of Myxidium lieberkuehni:Evidence that Myxozoa are metazoa and related to the Bilateria. Archiv fur Protistenkunde,1996,147:1-9.
[111]Shulman SS. Myxosporidia of the fauna of the ussr. Moscow: Nauka,1966.
[112]Siddall M, Whiting M. Long-branch abstractions. Cladistics,1999, 15:9-24.
[113]Siddall ME, Martin DS, Bridge D, Desser SS, Cone DK. The demise of a phylum of protists:Phylogeny of Myxozoa and other parasitic cnidaria. J Parasitol, 1995,81: 961-967.
[114]Sikama Y. On a new species of Henneguya found in a freshwater fish in China. Journal of the Shangai Science Institute,1938,4:135-138.
[115]Smothers JF, von Dohlen CD, Smith LH, Jr., Spall RD. Molecular evidence that the myxozoan protists are metazoans. Science,1994,265:1719-1721.
[116]Stamatakis A. Raxml-vi-hpc:Maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics,2006,22:2688-2690.
[117]Stole A. Actinomyxidies, nouveau groupe de mesozoaires parent des myxosporidies. Bull Intl Acad Sci Boheme,1899,22:1-12.
[118]Swofford DL (2003) PAUP*:phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer, Sunderland, MA.
[119]Szekely C, Hallett SL, Atkinson SD, Molnar K. Complete life cycle of Myxobolus rotundus (Myxosporea: Myxobolidae), a gill myxozoan of common bream Abramis brama. Dis Aquat Organ,2009,85:147-155.
[120]Tamura K, Peterson D, Peterson N, Stecher G, Nei M, Kumar S. Mega5:Molecular evolutionary genetics analysis using Maximum Likelihood, evolutionary distance, and Maximum Parsimony methods. Mol Biol Evol,2011,28:2731-2739.
[121]Thelohan P. Observations sur les myxosporidies et essai de classification de ces organismes. Bulletin de la Societe Philomatique de Paris,1892,4:167-178.
[122]Thelohan P. Recherches sur les myxosporidies. Bull Sci Fr Belg,1895,26:100-394.
[123]Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG. The clustal-X windows interface:Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res,1997,25:4876-4882.
[124]Walliker D. The nature of the iodinophilous vacuole of myxosporidian spores, and a proposal to synonymize the genus Myxosoma Thelohan,1892 with the genus Myxobolus Butschli,1882. J Protozool,1968,55:571-575.
[125]Wang GT, Yao WJ, Wang JG, Lu YS. Occurrence of Thelohanellosis caused by Thelohanellus wuhanensis (Myxosporea) in juvenile allogynogenetic silver crucian carp Carassius auratus gibelio (bloch), with an observation on the efficacy of fumagillin as a therapeutan. J Fish Dis,2001,24:57-60.
[126]Wang ZW, Zhu HP, Wang D, Jiang FF, Guo W, Zhou L, Gui JF. A novel nucleo-cytoplasmic hybrid clone formed via androgenesis in polyploid gibel carp. BMC Res Notes,2011,4:82.
[127]Weill R. L'interpretation des cnidosporidies et la valeur taxonomique de ieur cnidome Ieur cycle compare a la phase larvaire des narcomeduses cuninides. Travaux de la Station Zoologique de Wimereaux,1938,13:727-744.
[128]Whipps CM, Diggles BK. Kudoa alliaria in flesh of Argentinian hoki Macruronus magellanicus (Gadiformes; Merlucciidae). Dis Aquat Organ,2006,69:259-263.
[129]Whipps CM, El-Matbouli M, Hedrick RP, Blazer V, Kent ML. Myxobolus cerebralis internal transcribed spacer 1 (ITS-1) sequences support recent spread of the parasite to North America and within Europe. Dis Aquat Organ,2004a, 60: 105-108.
[130]Whipps CM, Grossel G, Adlard RD, Yokoyama H, Bryant MS, Munday BL, Kent ML. Phylogeny of the multivalvulidae (Myxozoa: Myxosporea) based on comparative ribosomal DNA sequence analysis. J Parasitol, 2004b, 90: 618-622.
[131]Whipps CM, Kent ML. Phylogeography of the cosmopolitan marine parasite Kudoa thyrsites (Myxozoa: Myxosporea). J Eukaryot Microbiol, 2006, 53:364-373.
[132]Wolf K, Markiw ME. Biology contravenes taxonomy in the Myxozoa: New discoveries show alternation of invertebrate and vertebrate hosts. Science, 1984, 225:1449-1452.
[133]Xi BW, Xie J, Zhou QL, Pan LK, Ge XP. Mass mortality of pond-reared Carassius gibelio caused by Myxobolus ampullicapsulatus in China. Dis Aquat Organ, 2011, 93:257-260.
[134]Ye LT, Li WX, Wu SG, Wang GT. Supplementary studies on Henneguya doneci Schulman,1962 (Myxozoa: Myxosporea) infecting the gill filaments of Carassius auratus gibelio (bloch) in China: Histologic, ultrastructural, and molecular data. Parasitol Res,2012,110:1509-1516.
[135]Yokoyama H, Freeman MA, Yoshnaga T, Ogawa K. Myxobolus buri, the myxosporean parasite causing scoliosis of yellowtail, is synonymous with Myxobolus acanthogobii infecting the brain of the yellowfin goby. Fish Sci, 2004, 70:1036-1042.
[136]Yokoyama H, Inoue D, Kumamaru A, Wakabayashi H. Myxobolus koi (Myxozoa: Myxosporea) forms large-and small-type "cysts" in the gills of common carp. Fish Pathol,1997,32:211-217.
[137]Yokoyama H, Ogawa K, Wakabayashi H. Myxobolus cultus n. sp. (Myxosporea: Myxobolidae) in thegoldfish Carassius auratus transformed from the actinsporean stage in the oligochaete Branchiura sowerbyi. J Parasitol,1995,81:446-451.
[138]Zhang JY, Wang JG, Li AH, Gong XL, Cai TZ. Redescription of Myxobolus pyramidis Chen,1958 (Myxosporea: Bivalvulida). Parasitol Res, 2006, 99: 65-69.
[139]Zhang JY, Wang JG, Li AH, Gong XN. Infection of Myxobolus turpisrotundus sp. n. in allogynogenetic gibel carp, Carassius auratus gibelio (Bloch), with revision of Myxobolus rotundus (s. L.) Nemeczek reported from C. auratus auratus (1.). J Fish Dis,2010a,33:625-638.
[140]Zhang JY, Yokoyama H, Wang JG, Li AH, Gong XN, Ryu-Hasegawa A, Iwashita M, Ogawa K. Utilization of tissue habitats by Myxobolus wulii Landsberg & Lom, 1991 in different carp hosts and disease resistance in allogynogenetic gibel carp: Redescription of M. wulii from China and Japan. J Fish Dis,2010b,33:57-68.
[141]Zhao Y, Sun C, Kent ML, Deng J, Whipps CM. Description of a new species of Myxobolus (Myxozoa: Myxobolidae) based on morphological and molecular data. J Parasitol,2008,94: 737-742.
[142]Zhao YJ, Li NN, Tang FH, Dong JL. Remarks on the validity of Myxobolus ampullicapsulatus and Myxobolus honghuensis (Myxozoa:Myxosporea) based on ssu rDNA sequences. Parasitol Res,2013,112:3817-3823.
© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号 地址：北京市海淀区学院路29号 邮编：100083 电话：办公室：(+86 10)66554848；文献借阅、咨询服务、科技查新：66554700 |