菊虎科
菊虎科(Cantharidae),又稱花螢科,為鞘翅目下的一科,屬於叩頭蟲總科的一支。目前已知超過130屬5000種,分屬5個亞科[1]。該科成員一般體態修長、鞘翅柔軟,且許多種類具有鮮豔的警戒色。菊虎主要棲息於森林中,廣布於各大洲的可棲息地,常可見到其停棲於花朵上。成蟲主要為肉食性,會捕食小型無脊椎動物,如蚜蟲或蛾類幼蟲等。但也會採食花蜜,為授粉昆蟲之一。
菊虎科 化石時期:白堊紀早期-現代
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瘟疫麗艷菊虎(Chauliognathus lugubris) | |
科學分類 | |
界: | 動物界 Animalia |
門: | 節肢動物門 Arthropoda |
綱: | 昆蟲綱 Insecta |
目: | 鞘翅目 Coleoptera |
亞目: | 多食亞目 Polyphaga |
下目: | 叩頭蟲下目 Elateriformia |
總科: | 叩甲總科 Elateroidea |
科: | 菊虎科 Cantharidae Imhoff, 1856 |
模式屬 | |
菊虎屬 Cantharis L., 1758
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亞科 | |
異名 | |
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從相關的化石記錄來看,該科的多樣性於中生代末期已相當豐富,且部分亞科可能在演化初期,就已具備了相當的物種多樣性。目前該科成員最早可以追溯到白堊紀早期。
下屬分類
本科包括以下屬:
- Absidiella Wittmer, 1972
- Afronycha Wittmer, 1949
- Allocotoma Gorham, 1895
- Ancistronycha Märkel, 1852
- 始源小菊虎屬 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
- 鹿野氏圓胸菊虎 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
- 渡邊氏圓胸菊虎 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
- 狹胸菊虎屬 Stenothemus Bourgeois, 1907
- Sucinocantharis Kuska & Kania, 2010
- Sucinorhagonycha Kuska, 1996
- Taiwanocantharis Wittmer, 1984
- Taocantharis Švihla, 2011
- 麗菊虎屬 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
形態描述
菊虎科成員的成蟲長約1.2至18毫米[3],一般體態修長、鞘翅柔軟、色彩多變,且有金屬光澤[1],許多物種呈現鮮豔的警戒色[4]。腹部7至8節[5],胸部及腹部兩側外緣具有腺體,可外翻分泌忌食物質,以驅離掠食者[5]。通常兩性形態有些差異[3]。足部長,所有腿皆為5個跗節[3][4]。
頭節突出前胸背板,觸角11節,大多成長絲狀,但也有部分物種成鋸齒狀、梳狀,或扇狀[3]。上唇膜狀,常位於頭楯下方[3]。髖節位於中胸,且彼此相鄰[3]。前胸背板邊緣平整,可能為正方形或橫扁形,少數成狹長形[3]。
雄蟲的陽莖成三葉狀(trilobed),多數由一根硬質的管狀被蓋(tegmen)包覆,內部則為一膜質柱狀構造[3]。被蓋由一陽莖基(phallobase)和兩根側突所構成[3]。陽莖基大多膜質,形態多樣,且許多成員有減化的現象[3]。
生態學
目前我們對於該科成員的生態學所知甚微[3]。本科成員主要棲息於森林中,廣布於各大洲的可棲息地,常可見到其停棲於花朵上[1]。成蟲主要為肉食性,會捕食小型無脊椎動物,如蚜蟲或蛾類幼蟲等[1][5]。此外也會採食花蜜及花粉,為重要的授粉甲蟲[5]。
菊虎、擬天牛、擬金花蟲、天牛、耀夜螢科、細花螢等訪花性甲蟲在色彩和外型上相似,其中有不少物種具有分泌忌食物質的能力[5]。這樣的群集裡可能同時牽涉不同的擬態策略,包含無分泌忌食物質的物種,模仿有分泌的物種,使捕食過難吃個體的掠食者會避免捕食類似物種,這種策略稱為貝氏擬態。又或是多種不同會分泌忌食物種的種類,產生類似的花紋或警戒色,這種策略則稱為穆氏擬態[5]。
生活史
菊虎為完全變態昆蟲[3]。幼蟲一般住在土中,喜好潮濕土壤,常棲息於較鬆軟的土壤,或是葉片下、石塊下,且常出現於朽木附近[3]。菊虎幼蟲為肉食性,會掠食其他節肢動物的幼蟲及卵,並吸食其體液[3]。幼蟲期最多可達十個齡期,幼蟲期快結束時,會於土壤中建造蛹室化蛹[3]。
分類學
菊虎科在親緣譜系上與螢科的關係最為接近[6],然而已知所有的菊虎種類都不會發光。本種相當高的物種多樣性,目前全世界已知種超過5,000種[1],且持續增加中[5]。但僅有北美和歐洲的基礎分類學研究較為透徹,其他地區尚有許多種類未被描述[5]。且由於該科大多數種類發表時,是根據博物館的蒐藏。在材料不足的情形下,常僅發表雄蟲的交尾器形態依據,雌蟲的相關形態則付之闕如,使雌蟲鑑定成為一個相當困難的工作[5]。
本科的共有衍徵包含翅脈減化、膜狀上唇、幼蟲體表被覆角質層短絨,且成蟲腹部第1–8節、幼蟲胸腹1–8節或1–9節背片兩側具有成對腺狀孔洞[1]。過去分類學家曾將螢科、紅螢科、擬螢科、角唇螢科、平螢科、捕蜈螢科,以及雌光螢科統稱為菊虎總科(Cantharoidea),但後來以幼生期形態為主的比較形態學研究和系統發育分析則支持菊虎總科為叩頭蟲總科的一個支序,為穩定叩頭蟲總科的單系性,便將其劃入叩頭蟲總科,而分子系統發育學也證實這樣的處理[6]。目前本科已知有菊虎亞科、麗艷菊虎亞科、異熒菊虎亞科、尖鬚菊虎亞科、熒菊虎亞科等五個亞科[1]。
古生物學
目前該科最早的紀錄則來自黎巴嫩琥珀(1.25~1.35億年前,白堊紀早期),然而身體後方完全損毀,以致無法確認物種分類。澳大利亞則於庫恩瓦拉(Koonwarra)化石層有白堊紀早期(1.15億年前)有菊虎化石的紀錄[7],然而保存狀況亦不佳。目前除了三例白堊紀緬甸琥珀的種類(均來自緬甸),大多數琥珀化石菊虎皆描述自新生代的琥珀,絕大多數來自波羅的海琥珀[3]。從相關的化石記錄來看,該科的多樣性於中生代末期已相當豐富,且尖鬚菊虎亞科可能在菊虎科演化初期,就已具備了相當的物種多樣性。
相關文化
中文俗名「菊虎」的命名緣由,至今仍不清楚[5]。日文名「浄海坊」(ジョウカイボン),得名於日本平安時代後期的權臣平清盛出家後的法名。平氏生前為熱病所苦,又由於斑蝥素這種化學物質會使皮膚紅腫、發皰發炎。但其實菊虎並不會產生斑蝥素,而是擬天牛科(Oedemeridae)的物種,可見當時人們尚無法區分此兩科物種[5]。英文的俗名為「兵甲蟲」(Soldier beetle),是由於歐陸最早被描述的種類體色與早期英國士兵的紅色外套類似而得名[5]。
相關研究
瘟疫麗艷菊虎(Chauliognathus lugubris)是一種澳洲特有的菊虎[8],在遭遇掠食者攻擊時,會分泌一種白色粘稠液體驅敵[9]。澳洲聯邦科學與工業研究組織(CSIRO)的研究團隊在分析液體成分後,發現含有一種稱為8Z-二氫母菊酸(8Z-dihydromatricaria acid,DHMA)的脂肪酸[9],具有抗腫瘤、微生物能力[5]。該物種也是目前唯一已知可分泌這種化學物質的真核生物[9]。
參考文獻
- ^ 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. (原始內容存檔於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. (原始內容存檔於2019-09-01) (英語).
- ^ 4.0 4.1 Family Cantharidae - Soldier Beetles - BugGuide.Net. bugguide.net. [2017-08-04]. (原始內容存檔於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 蕭昀. 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. (原始內容存檔於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. (原始內容存檔於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]. (原始內容存檔於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. (原始內容存檔於2022-04-07) (英語).
外部連結
- Delta key (頁面存檔備份,存於網際網路檔案館)
- Key to the British genera of Cantharidae
- Key to the British species of genus Cantharis