菊虎科
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菊虎科![]() 化石时期:白垩纪早期-现代
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瘟疫丽艳菊虎(Chauliognathus lugubris) | |
科学分类 ![]() | |
界: | 动物界 Animalia |
门: | 节肢动物门 Arthropoda |
纲: | 昆虫纲 Insecta |
目: | 鞘翅目 Coleoptera |
亚目: | 多食亚目 Polyphaga |
下目: | 叩头虫下目 Elateriformia |
总科: | 叩甲总科 Elateroidea |
科: | 菊虎科 Cantharidae Imhoff, 1856 |
模式属 | |
菊虎属 Cantharis L., 1758
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亚科 | |
异名 | |
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花萤科(Cantharidae),又称菊虎科,为鞘翅目下的一科,属于叩头虫总科的一支。目前已知超过130属5000种,分属5个亚科[1]。该科成员一般体态修长、鞘翅柔软,且许多种类具有鲜艳的警戒色。菊虎主要栖息于森林中,广布于各大洲的可栖息地,常可见到其停栖于花朵上。成虫主要为肉食性,会捕食小型无脊椎动物,如蚜虫或蛾类幼虫等。但也会采食花蜜,为授粉昆虫之一。
从相关的化石记录来看,该科的多样性于中生代末期已相当丰富,且部分亚科可能在演化初期,就已具备了相当的物种多样性。目前该科成员最早可以追溯到白垩纪早期。
下属分类
[编辑]本科包括以下属:
- Absidiella Wittmer, 1972
- Afronycha Wittmer, 1949
- Allocotoma Gorham, 1895
- Ancistronycha Märkel, 1852
- 始源小菊虎属 Archaeomalthodes Hsiao, Slipinski & Pang, 2016
- Armidia Mulsant, 1862
- Arturmiles Fanti, 2021
- Asilis Broun, 1893
- Asiopodabrus Wittmer, 1982
- Asiosilis Wittmer, 1977
- Atalantycha Kazantsev, 2005
- Athemellus Wittmer, 1972
- Athemus Lewis, 1895
- Autosilis Kazantsev, 2011
- Bactrocantharis Barovskii, 1926
- Bactronycha Kazantsev, 2001
- Belotus Gorham, 1881
- Bisadia Wittmer, 1972
- Boveycantharis Wittmer, 1969
- Brachysilidius Pic, 1949
- Brevipterus Yang, Liu & Zhao, 2022
- Burmomiles Fanti, Damgaard & Ellenberger, 2018
- Caccodes Sharp, 1885
- Cacomorphocerus Schaufuss, 1892
- Callosonotatum Pic, 1945
- Cantharis Linnaeus, 1758
- Cantharomorphus Fiori, 1914
- Cephalomalthinus Pic, 1921
- Chauliognathus Hentz, 1830
- Cnathrion Kazantsev & Perkovsky, 2019
- Compsonycha Wittmer, 1949
- Cordicantharis Svihla, 1999
- Cordylocera Guérin, 1830
- Cordylocerellus Wittmer, 1969
- Cratosilis Motschulsky, 1860
- Cretocantharis Hsiao, Li, Ren & Pang, 2021
- Crudosilis Kazantsev, 1994
- Cultellunguis McKey-Fender, 1950
- Curche Alekseev & Kazantsev, 2014
- Curticantharis Zhang, 1989
- Cyrebion Fairmaire, 1891
- Cyrtomoptera Motschulsky, 1860
- Cyrtomoptila Motschoulsky, 1859
- Daiphron Gorham, 1881
- Delkeskampia Wittmer, 1969
- Dichelotarsus Motschulsky, 1860
- Discodon Gorham, 1881
- Ditemnomorphus Champion, 1915
- Ditemnus LeConte, 1861
- Dysmorphocerus Solier, 1849
- Electronycha Kazantsev, 2013
- Electrosilis Kazantsev, 2013
- Elektrokleinia Ellenberger & Fanti, 2019
- Eridanula Fanti & Damgaard, 2018
- Eusilis Reitter, 1887
- Falsomalthinus Pic, 1924
- Falsomalthodes Pic, 1924
- Falsopodabrus Pic, 1927
- Fissocantharis Pic, 1921
- Flabelloontelus Pic, 1911
- Frostia Fender, 1951
- Gallerucocantharis Pic, 1913
- Geigyella Wittmer, 1972
- Grandesilis Pic, 1955
- Guineapolemius Wittmer, 1969
- Gymnocantharis Wittmer, 1979
- Habronychus Wittmer, 1981
- Habronychus (Monohabronychus) gen. Monohabronychus
- Hansasilis Pic, 1936
- Haplous Fairmaire, 1885
- Hatchiana Fender, 1966
- Hemipodistra Ganglbauer, 1922
- Hemipodistra Heikertinger, 1922
- Heteromastix Boheman, 1858
- Hoffeinsensia Kuska & Kania, 2010
- Hukawngichthyurus Fanti & Ellenberger, 2018
- Hyponotum Wittmer, 1949
- Ichthyurini Champion, 1915
- Ichthyurus Westwood, 1848
- Incisosilis Pic, 1908
- Indopolemius Wittmer, 1969
- Inmalthodes Pic, 1938
- Islamocantharis Wittmer & Magis, 1978
- Juratelacrima Fanti & Damgaard, 2018
- Kandyosilis Pic, 1929
- Katyacantharis Kazantsev & Perkovsky, 2019
- Kuskaella Fanti & Kupryjanowicz, 2017
- Laemoglyptus Fairmaire, 1886
- Leiothorax Wittmer, 1978
- Lithocantharis Linnaeus, 1997
- Lobetus Kiesenwetter, 1852
- Lycocerus Gorham
- Lycopolemius Pic, 1921
- Macrocerus Motschulsky, 1845
- Macromalthinus Pic, 1919
- Macrosilis Pic, 1911
- Malchinomorphus Pic, 1922
- Malthaster Gorham, 1885
- Malthesis Motschulsky, 1853
- Malthinellus Kiesenwetter, 1874
- Malthinocantharis Pic, 1914
- Malthinus Latreille, 1806
- Malthodes Kiesenwetter, 1852
- Malthoichthyurus Pic, 1919
- Malthopterus Motschulsky, 1853
- Maltypus Motschoulsky, 1860
- Manizalesum Pic, 1955
- Mantimalthinus Fanti & Castiglione, 2017
- Markus Fanti & Pankowski, 2018
- Maroniodes Brancucci, 1981
- Maronius Gorham, 1881
- Metacantharis Bourgeois, 1886
- Michalskantharis Fanti, 2017
- Microdaiphron Pic, 1926
- Microichthyurus Pic, 1919
- Micronotum Wittmer, 1949
- Micropodabrus Pic, 1920
- Mikadocantharis Wittmer & Magis, 1978
- Mimomalthinus Pic, 1931
- Mimoplatycis Kazantsev, 2013
- Mimopodabrus Wittmer, 1997
- Mimopolemius Pic, 1921
- Molliberus Peris & Fanti, 2018
- Myamalycocerus Fanti, 2016
- Nastonycha Motschulsky, 1853
- Neogressittia Wittmer, 1969
- Neoontelus Wittmer, 1972
- Noergaardia Fanti & Damgaard, 2018
- Nothotytthonyx Li, Biffi, Kundrata & Cai, 2022
- Occathemus Svihla, 1999
- Onychotelusia Wittmer, 1969
- Oontelus Solier, 1849
- Ornatomalthinus Poinar & Fanti, 2016
- Pachymesia Westwood, 1849
- Pachymesiosilis Pic, 1911
- Pacificanthia Kazantsev, 2002
- Pakabsidia Wittmer, 1972
- Palaeocantharis Hsiao, Li, Ren & Pang, 2021
- Palmnickeneoceras Fanti & Damgaard, 2018
- Paracantharis Wittmer, 1969
- Paradiscodon Wittmer, 1969
- Paramalthinus Brancucci, 1984
- Paramaronius Wittmer, 1963
- Parasilis Gorham, 1885
- Peltariosilis Wittmer, 1952
- Photinomorpha Champion, 1915
- Photurocantharis Pic, 1914
- Plectocephalon Pic, 1928
- Plectonotum Gorham, 1891
- Podabrinus Fairmaire, 1896
- Podabrus Dejean, 1833
- Podabrus Leconte, 1850
- Podabrus Westwood, 1838
- Podistra Motschulsky, 1839
- Podistra Victor, 1839
- Podosilis Wittmer, 1978
- Poinarelektronmiles Fanti & Damgaard, 2020
- Polemiosilis Pic, 1921
- Polemius LeConte, 1851
- Porostenus Motschulsky, 1853
- Prosthaptus Gorham, 1900
- Prothemellus Svihla, 1992
- Prothemus Champion, 1926
- Prothemus blankae
- Prothemus chinensis Wittmer, 1987
- Prothemus hisamatsui Wittmer, 1984
- 鹿野氏圆胸菊虎 Prothemus kanoi Wittmer, 1984
- Prothemus kubani
- Prothemus laticornis Y.Yang & X.Yang, 2011[2]
- Prothemus limbolarius (Fairmaire, 1900)
- Prothemus lycoceroides Kazantsev, 2011[2]
- Prothemus minor Wittmer, 1984
- Prothemus notsui Wittmer, 1984
- Prothemus piluensis Wittmer, 1987
- Prothemus shibatai Okushima, 2008
- Prothemus svihlai Wittmer, 1987
- Prothemus varicolor Wittmer, 1954
- 渡边氏圆胸菊虎 Prothemus watanabei Okushima & M.Satô, 2002
- Pseudoabsidia Wittmer, 1969
- Pseudocerocoma Pic, 1919
- Pseudocratosilis Moscardini & Sassi, 1970
- Pseudodiscodon Wittmer, 1969
- Pseudolobetus Champion, 1915
- Pseudopachymesia Pic, 1911
- Pseudopodabrus Pic, 1906
- Pseudosilis Pic, 1911
- Psilorrhynchus Gemminger & Harold, 1869
- Pygodiscodon Wittmer, 1952
- Rambesilis Pic, 1911
- Rhagonycha Eschscholtz, 1830
- Rhaxonycha Motschulsky, 1860
- Sanaungulus Fanti, Damgaard & Ellenberger, 2018
- Satoichthyurus Wittmer, 1978
- Sidabia Svihla, 1994
- Silicantharis Svihla, 1992
- Silidiscodon Leng & Mutchler, 1922
- Silidius Gorham, 1883
- Silis Charpentier, 1825
- Silis Dejean, 1825
- Silisonycha Wittmer, 1949
- Silvanotelus Wittmer, 1969
- Sinometa Wittmer, 1969
- Sogdocantharis Kasantsev, 1993
- Sphaerarthrum Waterhouse, 1884
- Stenopodabrus Nakane, 1992
- 狭胸菊虎属 Stenothemus Bourgeois, 1907
- Sucinocantharis Kuska & Kania, 2010
- Sucinorhagonycha Kuska, 1996
- Taiwanocantharis Wittmer, 1984
- Taocantharis Švihla, 2011
- 丽菊虎属 Themus Motschulsky, 1857
- Tibiopodabrus Svihla, 2004
- Trypheridium Brancucci, 1985
- Trypherus LeConte, 1851
- Tylocerus Dalman, 1833
- Tytthonyx LeConte, 1851
- Varcalium Li, Ruta, Tihelka & Cai, 2022
- Vitalfranzius Fanti & Müller, 2022
- Walteriella Kazantsev, 2001
- Wittmercantharis Sato, 1986
- Wittmerus Magis, 1957
- Wongyekokia Linnaeus, 1997
- Xenismus Waterhouse, 1879
- Yukikoa Sato, 1976
形态描述
[编辑]菊虎科成员的成虫长约1.2至18毫米[3],一般体态修长、鞘翅柔软、色彩多变,且有金属光泽[1],许多物种呈现鲜艳的警戒色[4]。腹部7至8节[5],胸部及腹部两侧外缘具有腺体,可外翻分泌忌食物质,以驱离掠食者[5]。通常两性形态有些差异[3]。足部长,所有腿皆为5个跗节[3][4]。
头节突出前胸背板,触角11节,大多成长丝状,但也有部分物种成锯齿状、梳状,或扇状[3]。上唇膜状,常位于头楯下方[3]。髋节位于中胸,且彼此相邻[3]。前胸背板边缘平整,可能为正方形或横扁形,少数成狭长形[3]。
雄虫的阳茎成三叶状(trilobed),多数由一根硬质的管状被盖(tegmen)包覆,内部则为一膜质柱状构造[3]。被盖由一阳茎基(phallobase)和两根侧突所构成[3]。阳茎基大多膜质,形态多样,且许多成员有减化的现象[3]。
生态学
[编辑]目前我们对于该科成员的生态学所知甚微[3]。本科成员主要栖息于森林中,广布于各大洲的可栖息地,常可见到其停栖于花朵上[1]。成虫主要为肉食性,会捕食小型无脊椎动物,如蚜虫或蛾类幼虫等[1][5]。此外也会采食花蜜及花粉,为重要的授粉甲虫[5]。
菊虎、拟天牛、拟金花虫、天牛、耀夜萤科、细花萤等访花性甲虫在色彩和外型上相似,其中有不少物种具有分泌忌食物质的能力[5]。这样的群集里可能同时牵涉不同的拟态策略,包含无分泌忌食物质的物种,模仿有分泌的物种,使捕食过难吃个体的掠食者会避免捕食类似物种,这种策略称为贝氏拟态。又或是多种不同会分泌忌食物种的种类,产生类似的花纹或警戒色,这种策略则称为穆氏拟态[5]。
生活史
[编辑]菊虎为完全变态昆虫[3]。幼虫一般住在土中,喜好潮湿土壤,常栖息于较松软的土壤,或是叶片下、石块下,且常出现于朽木附近[3]。菊虎幼虫为肉食性,会掠食其他节肢动物的幼虫及卵,并吸食其体液[3]。幼虫期最多可达十个龄期,幼虫期快结束时,会于土壤中建造蛹室化蛹[3]。
分类学
[编辑]菊虎科在亲缘谱系上与萤科的关系最为接近[6],然而已知所有的菊虎种类都不会发光。本种相当高的物种多样性,目前全世界已知种超过5,000种[1],且持续增加中[5]。但仅有北美和欧洲的基础分类学研究较为透彻,其他地区尚有许多种类未被描述[5]。且由于该科大多数种类发表时,是根据博物馆的搜藏。在材料不足的情形下,常仅发表雄虫的交尾器形态依据,雌虫的相关形态则付之阙如,使雌虫鉴定成为一个相当困难的工作[5]。
本科的共有衍征包含翅脉减化、膜状上唇、幼虫体表被覆角质层短绒,且成虫腹部第1–8节、幼虫胸腹1–8节或1–9节背片两侧具有成对腺状孔洞[1]。过去分类学家曾将萤科、红萤科、拟萤科、角唇萤科、平萤科、捕蜈萤科,以及雌光萤科统称为菊虎总科(Cantharoidea),但后来以幼生期形态为主的比较形态学研究和系统发育分析则支持菊虎总科为叩头虫总科的一个支序,为稳定叩头虫总科的单系性,便将其划入叩头虫总科,而分子系统发育学也证实这样的处理[6]。目前本科已知有菊虎亚科、丽艳菊虎亚科、异荧菊虎亚科、尖须菊虎亚科、荧菊虎亚科等五个亚科[1]。
古生物学
[编辑]目前该科最早的纪录则来自黎巴嫩琥珀(1.25~1.35亿年前,白垩纪早期),然而身体后方完全损毁,以致无法确认物种分类。澳大利亚则于库恩瓦拉(Koonwarra)化石层有白垩纪早期(1.15亿年前)有菊虎化石的纪录[7],然而保存状况亦不佳。目前除了三例白垩纪缅甸琥珀的种类(均来自缅甸),大多数琥珀化石菊虎皆描述自新生代的琥珀,绝大多数来自波罗的海琥珀[3]。从相关的化石记录来看,该科的多样性于中生代末期已相当丰富,且尖须菊虎亚科可能在菊虎科演化初期,就已具备了相当的物种多样性。
相关文化
[编辑]中文俗名“菊虎”的命名缘由,至今仍不清楚[5]。日文名“浄海坊”(ジョウカイボン),得名于日本平安时代后期的权臣平清盛出家后的法名。平氏生前为热病所苦,又由于斑蝥素这种化学物质会使皮肤红肿、发疱发炎。但其实菊虎并不会产生斑蝥素,而是拟天牛科(Oedemeridae)的物种,可见当时人们尚无法区分此两科物种[5]。英文的俗名为“兵甲虫”(Soldier beetle),是由于欧陆最早被描述的种类体色与早期英国士兵的红色外套类似而得名[5]。
相关研究
[编辑]瘟疫丽艳菊虎(Chauliognathus lugubris)是一种澳洲特有的菊虎[8],在遭遇掠食者攻击时,会分泌一种白色粘稠液体驱敌[9]。澳洲联邦科学与工业研究组织(CSIRO)的研究团队在分析液体成分后,发现含有一种称为8Z-二氢母菊酸(8Z-dihydromatricaria acid,DHMA)的脂肪酸[9],具有抗肿瘤、微生物能力[5]。该物种也是目前唯一已知可分泌这种化学物质的真核生物[9]。
参考文献
[编辑]- ^ 1.0 1.1 1.2 1.3 1.4 1.5 1.6 Hsiao, Yun; Ślipiński, Adam; Deng, Congshuang; Pang, Hong. A new genus and species of soldier beetle from Upper Cretaceous Burmese amber (Coleoptera, Cantharidae, Malthininae). Cretaceous Research: 119–123. [2017-08-04]. doi:10.1016/j.cretres.2016.09.002. (原始内容存档于2017-08-02).
- ^ 2.0 2.1 Yang Y, Kazantsev SV, Yang X (2011) Two remarkable new species of Prothemus Champion from China and Thailand, with comments on their systematic status (Coleoptera, Cantharidae). ZooKeys 119: 53–61. doi: 10.3897/zookeys.119.1621
- ^ 3.00 3.01 3.02 3.03 3.04 3.05 3.06 3.07 3.08 3.09 3.10 3.11 3.12 3.13 3.14 3.15 Ramsdale, Alistair S. 64. CANTHARIDAE Imhoff 1856. American Beetles, Volume II: Polyphaga: Scarabaeoidea through Curculionoidea. Boca Raton, Florida: CRC Press. 2002-06-19: 202–218 [2017-08-05]. ISBN 9781420041231. (原始内容存档于2019-09-01) (英语).
- ^ 4.0 4.1 Family Cantharidae - Soldier Beetles - BugGuide.Net. bugguide.net. [2017-08-04]. (原始内容存档于2021-04-22).
- ^ 5.00 5.01 5.02 5.03 5.04 5.05 5.06 5.07 5.08 5.09 5.10 5.11 5.12 萧昀. 常見卻鮮少人知的菊虎科昆蟲. 自然保育季刊. 2015-12-01, (92) [2017-08-04]. ISSN 1607-2928. (原始内容存档于2017-08-07).
- ^ 6.0 6.1 Kundrata, Robin; Bocakova, Milada; Bocak, Ladislav. The comprehensive phylogeny of the superfamily Elateroidea (Coleoptera: Elateriformia). Molecular Phylogenetics and Evolution: 162–171. [2017-08-04]. doi:10.1016/j.ympev.2014.03.012. (原始内容存档于2020-06-03).
- ^ Jell, Peter A. "The fossil insects of Australia.". Memoirs of the Queensland Museum. 2004, (50): pt 1.
- ^ 3. Chauliognathus lugubris (Fabricius). www.ces.csiro.au. [2017-08-06]. (原始内容存档于2015-09-23).
- ^ 9.0 9.1 9.2 Haritos, Victoria S.; Horne, Irene; Damcevski, Katherine; Glover, Karen; Gibb, Nerida; Okada, Shoko; Hamberg, Mats. The convergent evolution of defensive polyacetylenic fatty acid biosynthesis genes in soldier beetles. Nature Communications. 2012-10-23, 3: ncomms2147 [2017-08-06]. doi:10.1038/ncomms2147. (原始内容存档于2022-04-07) (英语).
外部链接
[编辑]- Delta key (页面存档备份,存于互联网档案馆)
- Key to the British genera of Cantharidae
- Key to the British species of genus Cantharis
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