虹琉璃蟻屬
虹琉璃蟻屬(Iridomyrmex),俗名彩虹蟻(rainbow ant, 因其帶有藍綠色彩虹光澤),隸屬於蟻科琉璃蟻亞科,由澳洲昆蟲學家古斯塔夫·邁爾 (Gustav Mayr) 於1862年首次描述,現生有79種,已滅絕有5種,大部分物種原生於澳洲,其餘的原生於亞洲及大洋洲,它們也被人為引入巴西、紐西蘭及阿拉伯地區。已滅絕的物種分布於中國、法國及美國。
虹琉璃蟻屬![]() 化石時期:
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肉蟻(I. purpureus) | |
科學分類 ![]() | |
界: | 動物界 Animalia |
門: | 節肢動物門 Arthropoda |
綱: | 昆蟲綱 Insecta |
目: | 膜翅目 Hymenoptera |
科: | 蟻科 Formicidae |
亞科: | 琉璃蟻亞科 Dolichoderinae |
族: | 細臭蟻族 Leptomyrmecini |
屬: | 虹琉璃蟻屬 Iridomyrmex Mayr, 1862[1] |
模式種 | |
肉蟻 Formica detecta Smith, 1858
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物種 | |
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多樣性[2] | |
現生約79種 已滅絕5種 |
它們是優勢種,它們會擾亂土壤組成或進入屋中,所以有時候被當成害蟲。澳洲的農夫會利用肉蟻(I. purpureus)來清除動物屍體,將屍體放在蟻丘上,數週內就會只剩下骨頭。肉蟻會進行儀式性(ritualised)的打鬥,以解決不同群體間的領地衝突,這種方式可以減少傷亡。本屬最大的物種是 I. purpureus 物種複合群(species group),體長約8毫米。
蟻后在婚飛後會建立新的群體,可以是單獨進行,或是透過分巢,也就是一些母群體的工蟻偕同蟻后一起遷徙到新的地點,有時候數隻蟻后會合作築巢,但在工蟻羽化後便會互相攻擊,最後只會剩下一隻蟻后。卵羽化成工蟻需花費44到61天。虹琉璃蟻習性及棲地多樣,因物種而異,有些物種築巢於土壤中,群體大小從數百個體到超過三十萬個體,巢體樣式可以是被卵石覆蓋,有好幾個出入口的大型蟻丘,有些物種在枝條中築巢,有些物種則會建立數個蟻巢,並以通道相連,蟻巢群的長度甚至可以達到650公尺;有些物種會和其他生物共生,像是會提供螞蟻分泌物或蜜露的鱗翅目幼蟲;I. bicknelli 會替蘭花授粉;虹琉璃蟻是掠食者也是清除者,會捕食獵物餵養幼蟲;虹琉璃蟻免疫海蟾蜍的毒素,能夠獵食未成熟的海蟾蜍;虹琉璃蟻的獵食者有蜘蛛、鳥類、蜥蜴或其他螞蟻。
系統發生學
虹琉璃蟻屬與最近的祖先約在1200萬年前分化[3],雖然虹琉璃蟻屬與其姊妹群 Froggattella 屬所歷經的時間相同[4],但 Froggattella 屬只有2個物種,相較於其他印澳地區的屬,該支系較為年輕,琉璃蟻亞科約在2300萬年前分化,虹琉璃蟻屬的化石分布於使新世及寡新世[5]。以下為虹琉璃蟻屬在印澳地區琉璃蟻亞科的屬的系統發生地位[3]。
Indo‑Australian genera |
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分類學
虹琉璃蟻屬由澳洲昆蟲學家 Gustav Mayr 於1862年首次描述[6],但直到1903年才指定模式種[4],模式種為 Formica detecta,Iridomyrmex purpureus 的同物異名[7][8],1878年琉璃蟻亞科建立之後虹琉璃蟻屬被分類到其下[9],虹琉璃蟻屬的分類早期錯綜複雜,因為早期的研究者沒有辨認出虹琉璃蟻屬的關鍵特徵,因此把很多物種錯置於虹琉璃蟻屬下[4][10]。
螞蟻學家 William Brown 二世是第一個質疑虹琉璃蟻屬並非單系群的人[4],他指出阿根廷蟻(舊時隸屬虹琉璃蟻屬)和其他虹琉璃蟻屬物種的相異之處[11]。虹琉璃蟻屬首次重整是在1990年代,其下的91個物種被重新分配到了6個屬,分別是Anonychomyrma 屬、Doleromyrma 屬、Linepithema 屬、Ochetellus 屬、Papyrius 屬和 Philidris 屬 ,重整後只有62個物種仍隸屬虹琉璃蟻屬[4],2011年,虹琉璃蟻屬再次重整,第二次重整後,虹琉璃蟻屬現在包含79個現生物種及5個已滅絕物種,其中有31個是新發表的物種[4],4個物種被分配到其他屬之下,25個物種及亞種被歸類為同物異名。根據單篇資料來源指出,澳洲虹琉璃蟻屬的物種可能有350種[12]。
虹琉璃蟻屬的學名:Iridomyrmex ,意思是彩虹般的螞蟻,因其藍綠色的彩虹光澤而得名[13],Irido 為古希臘文,意思是彩虹[14];myrmex ,也是古希臘文,意思是螞蟻[15]。
物種
(已滅絕物種以 † 標示)
- Iridomyrmex adstringatus Heterick & Shattuck, 2011
- Iridomyrmex agilis Forel, 1907
- Iridomyrmex alpinus Heterick & Shattuck, 2011
- Iridomyrmex anceps (Roger, 1863)
- Iridomyrmex anderseni Shattuck, 1993
- Iridomyrmex angusticeps Forel, 1901
- Iridomyrmex anteroinclinus Shattuck, 1993
- Iridomyrmex atypicus Heterick & Shattuck, 2011
- Iridomyrmex azureus Viehmeyer, 1914
- Iridomyrmex bicknelli Emery, 1898
- Iridomyrmex bigi Shattuck, 1993
- Iridomyrmex brennani Heterick & Shattuck, 2011
- †Iridomyrmex breviantennis Théobald, 1937
- Iridomyrmex brunneus Forel, 1902
- Iridomyrmex calvus Emery, 1914
- Iridomyrmex cappoinclinus Shattuck, 1993
- Iridomyrmex cephaloinclinus Shattuck, 1993
- Iridomyrmex chasei Forel, 1902
- Iridomyrmex coeruleus Heterick & Shattuck, 2011
- Iridomyrmex conifer Forel, 1902
- Iridomyrmex continentis Forel, 1907
- Iridomyrmex cuneiceps Heterick & Shattuck, 2011
- Iridomyrmex cupreus Heterick & Shattuck, 2011
- Iridomyrmex curvifrons Heterick & Shattuck, 2011
- Iridomyrmex cyaneus Wheeler, 1915
- Iridomyrmex difficilis Heterick & Shattuck, 2011
- Iridomyrmex discors Forel, 1902
- Iridomyrmex dromus Clark, 1938
- Iridomyrmex elongatus Heterick & Shattuck, 2011
- Iridomyrmex exsanguis Forel, 1907
- †Iridomyrmex florissantius Carpenter, 1930
- Iridomyrmex fulgens Heterick & Shattuck, 2011
- Iridomyrmex galbanus Shattuck, 1993
- Iridomyrmex gibbus Heterick & Shattuck, 2011
- Iridomyrmex gumnos Heterick & Shattuck, 2011
- Iridomyrmex hartmeyeri Forel, 1907
- Iridomyrmex hertogi Heterick & Shattuck, 2011
- Iridomyrmex hesperus Shattuck, 1993
- Iridomyrmex infuscus Heterick & Shattuck, 2011
- Iridomyrmex innocens Forel, 1907
- Iridomyrmex lividus Shattuck, 1993
- Iridomyrmex longisoma Heterick & Shattuck, 2011
- Iridomyrmex luteoclypeatus Heterick & Shattuck, 2011
- Iridomyrmex macrops Heterick & Shattuck, 2011
- †Iridomyrmex mapesi Wilson, 1985
- Iridomyrmex mattiroloi Emery, 1898
- Iridomyrmex mayri Forel, 1915
- Iridomyrmex meridianus Heterick & Shattuck, 2011
- Iridomyrmex minor Forel, 1915
- Iridomyrmex mirabilis Heterick & Shattuck, 2011
- Iridomyrmex mjobergi Forel, 1915
- Iridomyrmex neocaledonica Heterick & Shattuck, 2011
- Iridomyrmex niger Heterick & Shattuck, 2011
- Iridomyrmex nudipes Heterick & Shattuck, 2011
- †Iridomyrmex obscurans Carpenter, 1930
- Iridomyrmex obscurior Forel, 1902
- Iridomyrmex obsidianus Emery, 1914
- Iridomyrmex omalonotus Heterick & Shattuck, 2011
- Iridomyrmex pallidus Forel, 1901
- Iridomyrmex phillipensis Heterick & Shattuck, 2011
- Iridomyrmex prismatis Shattuck, 1993
- Iridomyrmex purpureus (Smith, 1858)
- Iridomyrmex reburrus Shattuck, 1993
- Iridomyrmex roseatus Heterick & Shattuck, 2011
- Iridomyrmex rubriceps Forel, 1902
- Iridomyrmex rufoinclinus Shattuck, 1993
- Iridomyrmex rufoniger (Lowne, 1865)
- Iridomyrmex sanguineus Forel, 1910
- Iridomyrmex setoconus Shattuck & McMillan, 1998
- †Iridomyrmex shandongicus Zhang, 1989
- Iridomyrmex spadius Shattuck, 1993
- Iridomyrmex splendens Forel, 1907
- Iridomyrmex spodipilus Shattuck, 1993
- Iridomyrmex spurcus Wheeler, 1915
- Iridomyrmex suchieri Forel, 1907
- Iridomyrmex suchieroides Heterick & Shattuck, 2011
- Iridomyrmex tenebrans Heterick & Shattuck, 2011
- Iridomyrmex tenuiceps Heterick & Shattuck, 2011
- Iridomyrmex trigonoceps Heterick & Shattuck, 2011
- Iridomyrmex turbineus Shattuck & McMillan, 1998
- Iridomyrmex victorianus Forel, 1902
- Iridomyrmex viridiaeneus Viehmeyer, 1914
- Iridomyrmex viridigaster Clark, 1941
- Iridomyrmex xanthocoxa Heterick & Shattuck, 2011
描述
虹琉璃蟻屬不同於其他琉璃蟻亞科的物種,虹琉璃蟻屬的頭楯前緣高於大顎;眼睛位於頭部的最上方,遠離大顎[4][16],虹琉璃蟻屬為中小型物種:I. conifer 物種複合群的物種體長約為4毫米,體色紅棕色,前伸腹板明顯突出;I. purpureus 物種複合群的物種體型最大,體長約為8毫米[17]。
虹琉璃蟻屬物種的工蟻罕有多形性[4]。
觀察結果顯示,畸形的工蟻存在於自然環境中,例如以下案例,有隻工蟻的前伸腹節與腹垂節癒合,雖然腹柄節存在,但卻異常增大,儘管畸形導致一點阻礙,該工蟻仍有活動能力[18]。
分布地區及棲地
虹琉璃蟻屬的物種分布在許多國家和島嶼,範圍橫跨好幾個大陸,在大洋洲地區分布於斐濟、印尼、新多里喀尼亞、巴布亞紐幾內亞和索羅門群島[4][19][20];澳洲的每個州都有分布[21];紐西蘭則是被人為引入,分布於北島及南島[22];在亞洲則分布於緬甸、東帝汶、印度、馬來西亞和菲律賓[4][23]。唯一分布於中東地區的物種是 I. anceps ,被人為引入阿拉伯聯合大公國;I. rufoniger 被人為引入巴西[4][24];大洋洲的諾福克島和菲律普島也有族群[4]。已滅絕物種的分布範圍大於現生的分布範圍,中國、法國和美國都曾出土過虹琉璃蟻的化石[25][26][27]。
虹琉璃蟻棲息在市郊,乾燥和半乾燥地區, 棲息於相思樹、貝殼杉、班庫樹、木麻黃、桉樹等林地中,也棲息於牧場、澳洲柏森林、都市公園、農地、草原、石楠樹林地、沙丘、稀樹草原、沼澤、都市花園、潮濕雨林、硬葉林,或是人造建物。
虹琉璃蟻大部分會在土壤中築巢,但有些也會住在石頭下、朽木中,甚至是沙灘、人行道中[4],某些物種會避免在沙質土壤中築巢[28],巢體外觀多變,大部分是具有單個出入口的小型蟻丘,有些物種的巢體則是被卵石覆蓋且具有多個出入口的大型蟻丘[4],某些物種會築出超級巢體,數個巢體彼此透過通道相連,長度可達650公尺[16],曾經有個案例,有個群體佔據了超過10公頃的地,總共有85個巢體和1500個出入口[29]。肉蟻不會攻擊巢友,但是會攻擊居住在不同巢體但同一群體的個體[30]。巢體可以是高於地表或低於地表,有些物種如 I. conifer 高於或低於地表的巢都有,冬天時它們會築巢於樹枝上,天氣回暖時棲息在地下[16],地下巢築在食物充足的地區,冬天時群體會移居到陽光充足的地方[31]。
行為和生態學
大部分的虹琉璃蟻習性兇悍,會主動攻擊騷擾巢穴或是覓食時遇到的生物,特別是 I. purpureus 物種複合群的物種,它們會湧出巢穴殺死入侵者[4];小型的物種也很兇悍,它們會噴灑虹琉璃蟻素(iridomyrmecin),一種虹琉璃蟻屬專有的防禦物質[4],虹琉璃蟻數量大,生性兇悍,活動力強,在澳洲是優勢種,某些物種會佔據資源,阻止其他昆蟲前來取食[32],它們數量多又活躍,甚至能影響其他螞蟻的覓食行為[4]。
雖然虹琉璃蟻以兇悍著名,有些物種生性膽小,遭到騷擾只會逃離或躲藏,也會避免和其他螞蟻接觸[4];如果巢穴受到攻擊,它們不會反擊,而是把幼蟲搬走,然後什麼也不做,它們會躲在落葉裡或植物裡,有些物種如 I. victorianus 遭受威脅時會靜止不動[4]。
當鄰近的肉蟻群體發生領地衝突時,它們會透過儀式性的打鬥來解決紛爭[33][34],不同群體的個體相遇時,它們會快速地揮動觸角,張開大顎,盡可能的張開身體讓自己看起來更高更大,這行為暗示著肉蟻的打鬥是透過比較體型進行[34],打鬥結束後,它們會清潔自己的身體,繼續尋找下一隻螞蟻[33],有些群體間的衝突可以持續數月甚至數年,但是真正會導致嚴重傷亡的打鬥極為罕見。
覓食
虹琉璃蟻是獵食者和清除者,它們也會和半翅目及鱗翅目幼蟲互動以取得蜜露[4],它們會獵食小型昆蟲,取食花粉[4],它們通常在地面上覓食,但大多數物種還會爬到植株上覓食;虹琉璃蟻大多數活動於晝夜,但有些是夜行性[4],它們白天時會堵住出入口,如果天氣合適,有些夜行性的物種也會在白天活動,但是有些物種,如 I. bigi 無論如何都不會在白天活動;有些物種會築巢在沒有陰影的地方,日出之後他們就可以曬太陽暖活身子,然後開始覓食[35]。
食性
虹琉璃蟻主要取食甜食,如花粉及蜜露[4],工蟻會獵食昆蟲或其他螞蟻來餵食幼蟲[16],工蟻會被長有油質體的種子吸引,它們會收集種子,剝下油質體後丟棄種子,這些留在螞蟻附近的種子擁有較高的存活率[5]。
它們有時候會在白蟻丘裡築巢[36],如此它們就可以就近捕食白蟻,除了捕食以外,目前沒有任何證據顯示白蟻和螞蟻有任何交互作用[4],有些免疫海蟾蜍毒素的物種會捕食未成年的海蟾蜍[37],如 I. purpureus 和 I. ruburrus[38][39],由於此特性,有人提議在海蟾蜍棲地釋放虹琉璃蟻巢,以達到生物防治的目的。
肉蟻是澳洲唯一會取食海鳥糞的螞蟻[40]。
獵食者
有些無脊椎動物專食虹琉璃蟻: 遊走蛛(Habronestes bradleyi) 會追蹤它們的緊戒費洛蒙[41][42][43][44];澳洲多刺魔蜥(Moloch horridus)是伏擊型的獵食者,專食虹琉璃蟻屬的物種,它們甚至會專食特定單一物種,拒食同屬其他物種[45];盲蛇 Ramphotyphlops nigrescens 會追蹤它們的行經的路徑,取食部分物種的幼蟲和卵[46];有些地甲蟲會躲在蟻巢附近的土中,捕食經過的工蟻[47]。
短吻針鼴(Tachyglossus aculeatus)會在婚飛期間捕食高脂肪含量的處女蟻后[48],它們也會在八月到十月攻擊蟻巢,捕食巢中的繁殖型個體[49],蟻后在婚飛期後仍然處於危險之中,因為有些鳥,如澳洲喜鵲、渡鴉,會被剛翻過的土壤吸引,它們會翻開土壤捕食那些剛建立新群體的蟻后[50];寄生性的絛蟲也會感染虹琉璃蟻[51]。
生命週期與繁殖
在溫帶潮濕氣候區,它們全年都會婚飛[4],婚飛進行時,繁殖蟻會離開巢穴爬到高處,準備起飛和交配[4],蟻后會和單隻或多隻雄蟻交配,因物種而異[52];根據觀察結果,肉蟻是雄蟻先起飛再來才是蟻后[50];20至40隻蟻后會聚集到巢穴的頂端,並在身體暖活後起飛,這個週期一天內會持續數次,並且維持數天,直到氣候改變或是巢中沒有任何蟻后[50],雄蟻在交配之後就會死去,蟻后則會尋找適合的地方建立群體,尋找的過程中它們會暴露在鳥類或其他獵食者的威脅之中[53],也會因為各種原因,如疾病或飢餓導致死亡[50];大多數的蟻后獨自築巢,但也可以是多個蟻后合作築巢、被其他群體納入巢中,或是「分巢」,也就是母群體的一部分離開舊巢建立新巢[54]。10%的蟻后會和其他蟻后合作築巢[55]。
蟻后掘出巢室後會產下大約20顆卵,它們會在一個月內孵化,卵羽化成成蟲需花費44到61天[54][56];成熟巢穴個體數各異,從數百到30萬以上不等[5][57],大多數的群體是單後型,但也有多達四隻蟻后的群體[50][58],寡後型(Oligogynous)的群體也存在,巢中存在多個蟻后,蟻后間彼此爭奪產權,但不同蟻后產下的工蟻不會攻擊不同的蟻后[54][59],蟻后只會照顧自己的卵幼[58][60],群體成長到一定規模後,蟻后就不會繼續聚集[54][61]。
共生生物
虹琉璃蟻會和鱗翅目幼蟲、蚜蟲及介殼蟲共生[16][62][63],工蟻會保護鱗翅目幼蟲,有時候還會把它們搬入巢中[16]。會和虹琉璃蟻共生的鱗翅目包含佳灰蝶(Jalmenus clementi、Jalmenus daemeli、Jalmenus eichhorni、Jalmenus evagoras、Jalmenus icilius、Jalmenus inous、Jalmenus lithochroa)[64]:246–248 [65]、澳灰蝶(Ogyris amaryllis、Ogyris olane、Ogyris oroetes)[64]:234–236、灰尖角灰蝶(Anthene lycaenoides)[64]:270、希特坎灰蝶(Candalides heathi)[64]:266、Candalides margarita (trident pencil-blue), Deudorix diovis (bright cornelian),[64]:256–258 Euchrysops cnejus (gram blue),[64]:290 Lampides boeticus (pea blue),[64]:290 Leptotes plinius (plumbago blue),[64]:288 Lucia limbaria (chequered copper),[64]:214 Nesolycaena caesia (Kimberly spotted opal),[64]:214 Neolucia agricola (fringed heath-blue)[64]:280 and Theclinesthes serpentata (saltbush blue).[64]:288 。I. bicknelli 會替 Microtis parviflora 授粉[66],且研究指出螞蟻不會傷害花粉[67]。
肉蟻會在清晨時用卵石和沙土堵住 Camponotus consobrinus 的巢穴出入口,阻擋它們離開巢穴,遭到這種對待的螞蟻也會回擊,用植物纖維堵住肉蟻的巢穴出入口[68][69];如果肉蟻的巢穴被樹木或其他陰影蓋住,由於照曬不到陽光,群體的健康狀況會惡化[70],Camponotus consobrinus 就會入侵肉蟻的巢穴,用一種黑色的樹脂類物質堵住通道,受攻擊的肉蟻群體會搬到其他的地方[71]。
與人類的互動
在澳洲郊外,肉蟻是農夫不可或缺的幫手,農夫會把動物屍體放到肉蟻巢上,數周內,屍體會被吃到剩骨頭[72];虹琉璃蟻有時候也是害蟲,它們會進到屋中,取食人類的食物[73],還會翻動土壤擾亂土質[74],清除蟻巢很困難,因為如果沒有徹底清除,來自鄰近群體的個體會補充待清除蟻巢的個體數[75];有些物種在都市地區適應良好;在坎培拉早期,人為活動影響環境,改變後的植被與花園提供了食物來源[76],於是肉蟻數量快速成長。肉蟻會造成阻礙生物防治半翅目昆蟲,特別是那些會分泌蜜露的,進而造成柑橘種植者的困擾[77]。
延伸閱讀
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引用文本
- Hölldobler, Bert; Wilson, Edward O. The Ants. Cambridge, Mass.: Belknap Press of Harvard University Press. 1990. ISBN 978-0-674-04075-5.
外部連結
- 維基共享資源上的相關多媒體資源:Iridomyrmex
- 維基物種上的相關資訊:Iridomyrmex
- Iridomyrmex at the AntWiki – Bringing Ants to the World (頁面存檔備份,存於網際網路檔案館)