蠓科
蠓科 | |
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Culicoides sonorensis,一只雌性的蠓 | |
科学分类 | |
界: | 动物界 Animalia |
门: | 节肢动物门 Arthropoda |
纲: | 昆虫纲 Insecta |
目: | 双翅目 Diptera |
亚目: | 长角亚目 Nematocera |
下目: | 蚊下目 Culicomorpha |
总科: | 摇蚊总科 Chironomoidea |
科: | 蠓科 Ceratopogonidae Grassi, 1900 |
亚科 | |
异名 | |
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蠓科几乎所有物种的雌虫均会以某种动物为宿主来吸血,其中的一些物种被认为是捕食其他小昆虫的,特别是蚊子的幼虫已经调查确认是 Bezzia 的通常的猎物。例如,已经对 Bezzia nobilis 进行了实验,表明它们依赖蚊子幼虫作为一种猎物来源。[4][5]它们也可以是无脊椎动物的吸血寄生虫,这取决于吸血攻击是否致命。[6]
与其它吸血飞虫一样,这些叮咬的蠓,如库蠓,不仅可以通过叮咬对人畜等产生直接骚扰性危害,还可以通过吸血活动传播多种病原体,传播的疾病包括寄生线虫的曼氏菌病、蓝舌病、非洲马病、流行性出血病、虫媒病毒[7]和非病毒性动物病原体。[8]。因此,蠓虫是与人畜疾病关系密切的重要的媒介昆虫。[9]
蠓的幼虫需要水分才能发育,还需要空气和食物。它们不是严格意义上的水生或陆生动物。[10]他们在世界各地几乎任何水栖或半水栖及山区都找到牠们的栖息地。蠓科物种的若虫总可在一些潮湿的地方发现的,例如:在树皮底下、在腐烂的木头里、堆肥里、泥里、小溪旁、树洞内或保水植物内(例如:phytotelmata)。
库蠓属(Culicoides)、铗蠓属(Forcipomyia)及细蠓属(Leptoconops)主要吸有脊椎动物的血。有些Atrichopogon及铗蠓属物种是其他较大型的昆虫的外寄生虫(ectoparasite)。 毛蠓属(Dasyhelea)几乎只吃花蜜。其他属的物种则捕食较小型的昆虫。
外貌
[编辑]本科物种体形细小,身长一般在1–4 mm之间,褐色或黑色,有复眼、单眼各一对,触角丝状,长而有毛,分为13~15节。下颚须有5节,喙短,雌虫为刺吸式。胸分三节,中胸背板最发达,前部两侧各有一肩坑。翅短而宽,翅端钝圆,接近前缘的二条翅脉甚粗大。翅上有细毛及粗毛,有的翅膀上有暗斑与白斑。翅在静止时重叠覆盖于腹部之上。足三对,较发达,腹部有10节,雌虫末端有尾须一对。[11]
头部近球形,复眼发达,触角丝状,吸血种类口器发达,但比蚊虫的口器短。胸部背面略隆起,前翅大,库蠓属前翅多淡色或有暗斑,后翅成平衡棒,腹部10节。雌蠓有受精囊和一对卵巢。仅雌蠓吸血。雄蠓尾端的外生殖器的构造因种类而异。蛹为裸蛹,分头、胸、腹3部,体前方背面有一对呼吸管。幼虫呈蠕虫状,上颚和咽发达,在水中作螺旋运动,行体壁呼吸。卵呈长纺锤形,长为宽的4倍以上。[12]
蠓为完全变态昆虫,这意味着它们的发育包括四个生命阶段:卵、幼虫、蛹和成虫。[13]在温暖气候中,常见的物种需要大约两到六周才能完成一个生命周期。成年雄性和雌性都以花蜜为食。大多数雌性还以脊椎动物(包括人类)的血液为食,以获取产卵所需的蛋白质。它们的叮咬很痛,并且会引起强烈的瘙痒损伤。[10] 它们的口器非常发达,可以切割宿主的皮肤。有些物种捕食其他昆虫。
分类
[编辑]常见种类
[编辑]全球已知蠓类5360种,其中库蠓1224~1530种、细蠓133种、铗蠓属蠛蠓亚属124种、澳蠓1种。吸血蠓虫(blood-suckingmidges)是蠓科昆虫中能叮刺人和畜等动物蠓虫的总称。吸血蠓虫是重要的医学昆虫。全世界已知吸血蠓虫为4属1503种,包括库蠓属(Culicoides)、蠛蠓属(Lasiohelea)、细蠓属(Leptoconops)和澳蠓属(Austroconops)。现知库蠓有1247种,占世界已知蠓科昆虫种类的23.3%,占已知4个吸血蠓属的83%。[14]
常见物种
[编辑]- Culicoides impunctatus—known as the Scottish midge, or Highland midge
- Culicoides imicola
亚科
[编辑]蠓科物种大致可以分类为四个亚科[15]:
- 蠓亚科(Ceratopogoninae Kieffer, 1906)——细长的幼虫,没有前腿或钩子。这个亚科的大多数幼虫都是捕食性的,成虫一般会吸食脊椎动物的血液或攻击其他昆虫,大多数雌性以大小相似的昆虫为食。[6]
- 库蠓属(Culicoides Latreille, 1809)
- 毛蠓亚科(Dasyheleinae Kieffer, 1911)——幼虫的特征是肛门节段具有可伸缩的后肢。幼虫是水生的,成虫不以脊椎动物的血液为食,也不捕食其他昆虫。它们只吃花蜜,这是角蠹科的一种不寻常的摄食行为。
- 铗蠓亚科(Forcipomyiinae (Lenz, 1934) Enderlein, 1936)——幼虫身上有前肢和后肢,幼虫是陆生和水生的,主要以藻类和真菌为食。一些物种是热带作物如可可豆的重要传粉者。
- 铗蠓属(Forcipomyia Meigen, 1818
- 细蠓亚科(Leptoconopinae Enderlein, 1936)——幼虫通过它们独特的头部巩膜和口器来识别。[16]
属
[编辑]本科包括以下属:[17]
- Acanthohelea Kieffer, 1917
- Adelohelea Borkent, 1995
- Afrohelea Wirth, 1965
- Afrostilobezzia Szadziewski & Dominiak, 2015
- Agilihelea Yu, 2005
- Alautunmyia Borkent, 1996
- Allohelea Kieffer, 1917
- Alloimyia Yu, 2005
- Alluaudomyia Kieffer, 1913
- Amerohelea Grogan & Wirth, 1981
- Anebomyia Borkent, 2014
- Ankylohelea de Meillon & Wirth, 1987
- Archiaustroconops Szadziewski, 1996
- Archiculicoides Szadziewski, 1996
- Atrichopogon Kieffer, 1906
- Atriculicoides Remm, 1976
- Atyphohelea Borkent, 1998
- Austroconops Wirth & Lee, 1958
- Austrohelea Wirth & Grogan, 1988
- Austrosphaeromias Spinelli, 1997
- Avaritia Fox, 1955
- Baeodasymyia Clastrier & Raccurt, 1979
- Baeohelea Wirth & Blanton, 1970
- Bahiahelea Wirth, 1992
- Bezzia Kieffer, 1899
- Boreohelea
- Borkenthelea Spinelli & Grogan, 1993
- Bothahelea Grogan & Wirth, 1983
- Bothamia Meiswinkel, 1987
- Brachycretacea Szadziewski, 1996
- Brachypogon
- Burmahelea Szadziewski & Sontag, 2019
- Cacaohelea Wirth & Grogan, 1988
- Calcarhelea Wirth & Grogan, 1988
- Calyptopogon Kieffer, 1910
- Camptopterohelea Wirth & Hubert, 1960
- Capehelea de Meillon & Wirth, 1987
- Ceratoculicoides Wirth & Ratanaworabhan, 1971
- Ceratohelea Wirth & Grogan, 1988
- Ceratopalpomyia Szadziewski, 1988
- Ceratopogon Meigen, 1803
- Chairopogon Yu, 2005
- Changania
- Chelohelea Giles & Wirth, 1985
- Clastrieromyia Spinelli & Grogan, 1985
- Clinohelea Kieffer, 1917
- Congohelea Wirth & Grogan, 1988
- Cotocripus Brethes, 1912
- Crispomyia Debenham, 1974
- Culicoides
- 毛蠓属 Dasyhelea Kieffer, 1911
- Devalquia
- Diaphanobezzia Ingram & Macfie, 1931
- Dibezzia Kieffer, 1911
- Diplosella Kieffer, 1921
- Downeshelea Wirth & Grogan, 1988
- Echinohelea Macfie, 1940
- Eohelea Petrunkevitch, 1957
- Euprojoannisia Brèthes, 1914
- Fanthamia De Meillon, 1939
- Fittkauhelea Wirth & Blanton, 1970
- 铗蠓属 Forcipomyia Meigen, 1818
- Fossihelea Szadziewski, 1988
- Fossileptoconops Szadziewski, 1996
- Gedanohelea Szadziewski, 1988
- Gerontodactus Borkent, 2019
- Gerontodacus Borkent, 2019
- Groganhelea Spinelli & Dippolito, 1995
- Guihelea Yu & Qian, 2005
- Haematomyidium Goeldi
- Hebetula Wirth & Debenham, 1977
- Heleageron Borkent, 1995
- Heteroceratopogon Wirth & Grogan, 1988
- Heterohelea Clastrier, 1985
- Heteromyia Say, 1825
- Homohelea Kieffer, 1917
- Hypsimyia Yu, 2005
- Indobezzia Dasgupta & Saha, 1995
- Isthmohelea Ingram & MacFie, 1931
- Jenkinshelea Macfie, 1934
- Johannsenomyia Malloch, 1915
- Jordanoconops Szadziewski, 2000
- Kolenohelea de Meillon & Wirth, 1981
- Lanatomyia Debenham, 1974
- Lanehelea Wirth & Blanton, 1972
- Lasiohelea Kieffer, 1921
- Lebanoculicoides Szadziewski, 1996
- Leehelea Debenham, 1974
- Leptoconops
- Leptohelea Wirth & Blanton, 1970
- Luciamyia De Meillon, 1937
- Mackerrasomyia Debenham, 1970
- Macropeza Meigen, 1818
- Macrurohelea Ingram & MacFie, 1931
- Mallochohelea Wirth, 1962
- Mantohelea Szadziewski, 1988
- Metacanthohelea Wirth & Grogan, 1988
- Metahelea Edwards, 1929
- Meunierohelea Szadziewski, 1988
- Minyohelea Borkent, 1995
- Monogedania Szadziewski, Grogan, Sontag & Bojarski, 2022
- Monohelea Kieffer, 1917
- Nannohelea Grogan & Wirth, 1980
- Nelohelea Szadziewski & Sontag, 2019
- Neobezzia Wirth & Ratanaworabhan, 1972
- Neoculicoides Pierce, 1966
- Neohelea Clastrier, 1988
- Neosphaeromias Das Gupta & Wirth, 1970
- Neurobezzia Wirth & Ratanaworabhan, 1973
- Neurohelea Kieffer, 1925
- Nilobezzia Kieffer, 1921
- Niphanohelea Grogan & Wirth, 1981
- Notiohelea Grogan & Wirth, 1979
- Notoceratopogon de Meillon & Downes, 1986
- Oculudentavis Xing, O'Connor, Schmitz, Chiappe, McKellar, Yi & Li, 2020
- Oecacta Poey
- Oxyria Yu, 2005
- Pachyhelea Wirth, 1959
- Palaeobrachypogon Borkent, 1995
- 须蠓属 Palpomyia Meigen, 1818
- Parabezzia Malloch, 1915
- Paraculicoides Pierce, 1966
- Paradasyhelea Macfie, 1940
- Paralluaudomyia Clastrier, 1960
- Parastilobezzia Wirth & Blanton, 1970
- Paryphoconus Enderlein, 1912
- Pellucidomyia Macfie, 1939
- Peronehelea Borkent, 1995
- Phaenobezzia Haeselbarth, 1965
- Physohelea Grogan & Wirth, 1979
- Probezzia Kieffer, 1906
- Prokempia
- Protoculicoides Boesel, 1937
- Pseudostilobezzia Wirth & Ratanaworabhan, 1973
- Rhynchohelea Wirth & Blanton, 1970
- Schizohelea Kieffer, 1917
- Schizonxyhelea
- Schizonyxhelea Clastrier, 1984
- Serromyia Meigen, 1818
- Sinhalohelea Grogan & Borkent, 1992
- Sinicohelea Yu, Wang & Tan, 2012
- Sphaerohelea Spinelli & Felippe-Bauer, 1990
- Sphaeromias Curtis, 1829
- Sphaeromias Stephens, 1829
- Spinellihelea Borkent, Grogan & Picado, 2008
- Stenoxenus Coquillett, 1899
- Stilobezzia
- Stiloculicoides Wirth & Grogan, 1988
- Tetrabezzia Kieffer, 1917
- Thyridomyia Saunders, 1925
- Trishelea
- Wannohelea Yu, 2005
- Washingtonhelea Wirth & Grogan, 1988
- Wirthohelea Szadziewski, 1988
- Wirthomyia Vargas, 1973
- Xenohelea Kieffer, 1917
- Yungahelea Spinelli & Ronderos, 2018
生活习性
[编辑]蠓虫为刺吸式口器,雄蠓吸食植物汁液,雌蠓具有吸血习性。因此雌蠓可作为传播疾病的重要媒介。一般蠓虫的吸血活动是在白天、黎明或黄昏进行。成虫多栖息于树丛、竹林、杂草、洞穴等避风、避光处。当温度、光照适合且无风时,成虫即成群飞出,蠓虫的飞行能力不强,一般不超过0.5km,因此蠓虫的活动范围一般在栖息地周围的300m内,吸血蠓类交配时常有群舞现象,交配后吸血,约3~4d后卵巢发育成熟而产卵。通常雌蠓一生产卵2~3次,一次产卵量约50~150粒,蠓生活史所需的时间与温度关系密切,在夏季约需一个月,通常一年可繁殖2~4代,视种类与地区不同而异,雄蠓交配后1~2d便死亡,雌蠓的寿命约一个月,一般以幼虫或卵越冬。[9]
出现季节
[编辑]中国北方通常出现在5~8月;南方出现在4~10月。
繁殖方式
[编辑]蠓主要孳生在潮湿、松软、富有腐殖质的土壤内及水塘、树洞、沼泽和住区附近的粪坑、污水沟等处。
病毒传播
[编辑]吸血蠓虫可以通过吸血传播多种虫媒病毒,目前已知的从吸血蠓虫分离鉴定的病毒有5科7属,包括弹状病毒科、水泡性病毒属及暂时热病毒属、呼肠孤病毒科、环状病毒属、布尼亚病毒科正布尼亚病毒属及内罗毕病毒属、披膜病毒科甲病毒属、黄病毒科黄病毒属。[9]
吸血蠓作为传播媒介可以在人和(或)动物之间传播多种病毒性疾病,蓝舌病是蠓虫传播的最重要的动物性疾病,以库蠓为主要传播媒介,主要侵害绵羊,该病的发生具有明显的地区性和季节性,以绵羊感染后临床症状最明显,表现为发热、面部水肿、颊粘膜和胃肠道粘膜严重的卡他性炎症为特征,严重病例舌头呈蓝紫色、发绀并出现溃疡和糜烂症状。孕羊患病可经胎盘传播给胎儿,引起母畜的流产、死胎或先天性异常的发生。目前在我国已经从库蠓中分离到蓝舌病毒,但尚未发现动物间有蓝舌病的流行。[9]
弹状病毒科中的新泽西水泡性口炎病毒所引起的水泡性口炎,又名鼻疮、口疮、伪口疮、“烂舌症”、“牛及马的口腔溃疡”等,是一种可引起马、牛、猪和鹿等多种动物发病的一种病毒性传染病,大量流涎是家畜感染最重要的症状,其特征表现为口腔黏膜、蹄冠状带及乳头皮肤出现水泡、糜烂及溃疡,临床症状与口蹄疫相似。人可因接触病畜而偶发感染,但常不显症状或仅轻微发烧或寒战,有的患者表现头疼、恶心、呕吐等。约有1/4的病人在口部、舌或鼻部出现疱疹样水疱病变,持续2~7天后完全恢复。VS最早于1926年和1927年在美国被报道,VS可以使马和牛的生产能力下降而造成严重的经济损失,并对国际贸易产生严重影响。[9]
此外,布尼亚病毒科的赤羽病毒所造成的赤羽病是引起牛、羊繁殖障碍的一种重要病毒性疫病,是我国从国外进口牛、羊必须检测的七种疫病之一,其临床特征表现为流产、早产、死胎、胎儿畸形、木乃伊胎、新生胎儿发生关节弯曲积水性无脑综合症等,该病最早在意大利暴发,随后在日本、美国、韩国、台湾、沙特阿拉伯等地区分离到该病毒,对养牛业和养羊业危害严重,是较为严重的一种疫病,布尼亚病毒科奥柔普西病毒也可引起蠓媒性人兽共患病等。总之,蠓媒性疾病给畜牧业造成了巨大的疾病负担,并带来了巨大的经济损失。[9]
参考文献
[编辑]- Blanton, F.S. and W.W. Wirth. 1979. The sand flies (Culicoides) of Florida (Ceratopogonidae). Arthropods of Florida and Neighboring Land Areas Volume 10. Florida Department of Agriculture and Consumer Services.
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- Downes, J.A. and W.W. Wirth. 1981. Chapter 28: Ceratopogonidae. Pp. 393–421. In: McAlpine, J.F., B.V. Peterson, G.E. Shewell, H.J. Teskey, J.R. Vockeroth, and D.M. Wood. Manual of Nearctic Diptera, Volume 1. Agriculture Canada Monograph 27.
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- ^ 10.0 10.1 common name: biting midges, no-see-ums, scientific name: Culicoides spp. (Insecta: Diptera: Ceratopogonidae). Featured Creatures. University of Florida. [20 September 2018]. (原始内容存档于2016-12-28).
- ^ 《消毒杀虫灭鼠手册》编写组.消毒杀虫灭鼠手册.北京:人民卫生出版社,1980.3:342 (中文).
- ^ 《消毒杀虫灭鼠手册》编写组.消毒杀虫灭鼠手册.北京:人民卫生出版社,1980.3:342.
- ^ Beckenbach, Andrew T.; Borkent, Art. Molecular analysis of the biting midges (Diptera: Ceratopogonidae), based on mitochondrial cytochrome oxidase subunit 2. Molecular Phylogenetics and Evolution. 2003-04-01, 27 (1): 21–35. ISSN 1055-7903. PMID 12679068. doi:10.1016/S1055-7903(02)00395-0 (英语).
- ^ 蠓虫及其传播的虫媒病毒 - 中国知网. cnki.cn-ki.net. [2021-07-23]. (原始内容存档于2021-07-23).
- ^ 虞以新 (编). 中国蠓科昆虫名录及其检索表. 军医医学科学出版社. 005-09 [2016-05-08]. (原始内容存档于2016-06-03) (中文(简体)).
- ^ Borkent, Art; Craig, Douglas A. Austroconops Wirth and Lee, a Lower Cretaceous genus of biting midges yet living in Western Australia: a new species, first description of the immatures and discussion of their biology and phylogeny (Diptera: Ceratopogonidae). American Museum Novitates. 23 August 2004, (3449): 1–2 [2022-08-03]. doi:10.1206/0003-0082(2004)449<0001:AWALAL>2.0.CO;2. hdl:2246/2814. (原始内容存档于2022-08-03).
- ^ Ceratopogonidae. GBIF. [2023-03-14]. (原始内容存档于2023-03-14).
延伸阅读
[编辑][在维基数据编辑]
外部链接
[编辑]- British insects: the families of Diptera (页面存档备份,存于互联网档案馆) – Delta guides, Biodiversity and Biological Collections
- Ceratopogonidae – AAFC, Government of Canada
- The Ceratopogonidae – Inbio Site, Instituto Nacional de Biodiversidad
- Ceratopogonid Web Page – Belmont University
- Biting midges (页面存档备份,存于互联网档案馆) on the UF / IFAS Featured Creatures Web site
- Flying Teeth (页面存档备份,存于互联网档案馆) Spain Buddy Website