蘋果豬籠草
蘋果豬籠草（學名：Nepenthes ampullaria）是一種分佈廣泛的豬籠草。其存在於在婆羅洲、摩鹿加群島、新幾內亞、西馬來西亞、新加坡、蘇門答臘和泰國。[2][3][4][5][6][7]其種加詞「ampullaria」來源於拉丁文「ampulla」，意為「燒瓶」。
蘋果豬籠草
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| 蘋果豬籠草的地面籠，可以看到作為遮雨的籠蓋葉子是外放的，開口隨時用來接取掉落的落葉，攝於婆羅洲巴哥國家公園 | |
科學分類 
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| 界： | 植物界 Plantae |
| 演化支： | 維管束植物 Tracheophyta |
| 演化支： | 被子植物 Angiosperms |
| 演化支： | 真雙子葉植物 Eudicots |
| 目： | 石竹目 Caryophyllales |
| 科： | 豬籠草科 Nepenthaceae |
| 屬： | 豬籠草屬 Nepenthes |
| 種： | 蘋果豬籠草 N. ampullaria
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| 二名法 | |
| Nepenthes ampullaria Jack, 1835
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| 異名 | |
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與其他物種的區別在於其食蟲性已退化，並已具有部分的食腐性，轉為以捕蟲籠收集和消化落葉。[8][9][10]因此素食性使得相當多的生物居住其中，形成特殊的共生關係，而這些生物有時能提供氮源。
植物學史
蘋果豬籠草的捕蟲籠形態和生長習性均十分獨特，與其餘同屬植物具有顯著的不同。弗朗西斯·歐內斯特·勞埃德翻譯了特羅爾（Troll）在1932年對蘋果豬籠草的描述：[注 1][11]
| “ | 在蘇門答臘以西的西比路島上，我在一個沼澤森林的大量植被中發現了蘋果豬籠草。這真是一個令人難以置信又難忘的景象。在藤蔓織成的大網中，獨特形態的捕蟲籠散發着微光，它們一簇簇地生長着；而更令人驚嘆的是，在長滿苔蘚的淤泥中，點綴着一個又一個地面籠，油然一片廣闊的地毯。 | ” |
形態特徵
蘋果豬籠草的莖呈淡棕色，可以攀爬至15米的高處。[12]植株生長到一定程度後倒伏，由植株基本再生長出側芽，使其呈叢狀。[2]
蘋果豬籠草的葉片呈淺綠色，可長達25厘米，寬至6厘米。籠蔓可長達15厘米。 [12]
蘋果豬籠草水罐狀的捕蟲籠一般都很小，很少超過10厘米高，7厘米寬。唇大大內趨，內緣部分約佔截面表面總長度的85%。[13]上位籠十分罕見，且其尺寸遠小於下位籠。籠蓋後翻，不具遮雨功能。[2]捕蟲籠的顏色為淺綠色至深紅色；據記錄。蘇門答臘和西馬來西亞的蘋果豬籠草幾乎都為全綠色或為綠底紅斑；全紅的蘋果豬籠草大多只存在於婆羅洲。在新幾內亞曾記錄到具巨大捕蟲籠的蘋果豬籠草。[12][14]
蘋果豬籠草的花序為密集的圓錐花序。它是在蘇門答臘和西馬來西亞發現的唯一有着圓錐花序的豬籠草屬植物。[14]
蘋果豬籠草幼株的各個部位均長有密集的棕色短毛，但成株的毛被除了花序部分都很稀疏。[12]
原生地
蘋果豬籠草是豬籠草屬內分佈較為廣泛的物種之一，其存在於婆羅洲、摩鹿加群島、新幾內亞、馬來西亞半島、新加坡、蘇門答臘、泰國。[3]其也存在於周圍許多小型島嶼中，包括邦加島、望加麗島、蘭塔島、達魯島、浮羅交怡、孟德爾島、明打威群島（西比路島）、美蘭地群島（巴東島、朗桑島、直名丁宜島）、尼亞斯島、檳榔嶼、廖內群島（林加群島、納土納群島[15]、廖內島）、魯帕島[16]
蘋果豬籠草一般生長在海拔1200米[2]或2100米[17]以下的潮濕陰涼的森林中。在婆羅洲，蘋果豬籠草主要分佈在海拔0至1000米相對平坦的巽他荒原森林、泥炭沼澤森林和退化沼澤林（degraded swamp forest）中。[12]
在蘇門答臘和西馬來西亞，蘋果豬籠草生長於海拔1100米以下荒原森林的平坦地帶、巴丹疏林（「巴丹」即「padang」，在馬來語中意為「平原」）、貝魯卡（矮灌木林）、泥炭沼澤林、退化沼澤森林以及淺水區域中。[14]
食蟲性
蘋果豬籠草與其他豬籠草相比，其捕獲獵物的數量極少，捕蟲籠內多為其他植物的落葉。因此，其被認為是靠吸收捕蟲籠內腐敗有機質釋放的營養素而獲取營養，[2]由此看來，蘋果豬籠草已擁有部分的食腐性。
蘋果豬籠草已進化出了多種適應獲得落葉的特徵：
- 在豬籠草屬中，蘋果豬籠草是少數幾種捕蟲籠中缺少「新月形」細胞的物種。[18][19]這些經過形態改變的氣孔保衛細胞，被認為可以讓昆蟲更加容易地滑入籠中。[11]
- 不同於其他豬籠草，蘋果豬籠草的籠蓋較小，且是向內翻折的，有助於落葉直接落入籠中。[14]
- 在捕食昆蟲時起到重要作用的蜜腺已經極為罕見，某些變型的籠蓋上甚至已經完全消失。[14]
- 與其他物種相比，唇上的蜜腺也大量減少。[14]
- 在地面籠中，腺體區幾乎延伸到了唇，而唇上只有少量或幾乎沒有蠟質區，[12][19][20][21][22]而蠟質區的作用是使昆蟲滑落入籠中的消化液里。[14]
- 蘋果豬籠草具有豬籠草屬中是十分少見的地下莖，讓捕蟲籠可以如同地毯一般覆蓋住土壤，擴大了接住落葉的區域範圍。[14]
- 蘋果豬籠草的捕蟲籠可以維持相對較長時間不枯萎，因為該物種有賴於逐步積累吸收到的營養素。[14]
- 一些豬籠草底內動物，如孑孓，被認為能夠促進落葉的分解，並通過排泄銨離子的方式為植物轉化出可利用的氮。一些分解落葉的細菌也可以幫助產生銨離子。[14]
- 蘋果豬籠草的捕蟲籠也是一部分水生生物的繁殖場所。
這些葉片為它的生長提供了大量的營養。實驗表明，生長於森林樹冠下（有落葉供給）的蘋果豬籠草，其葉內氮-15的豐度要遠低於沒有落葉供給的同類。然而，前者總體的氮濃度卻高於生長於空曠地帶的後者。2003年的研究估算，生長於森林樹冠下的蘋果豬籠草，其葉內有35.7%（±0.1%）的氮素來自落葉。[8]2011年的研究發現，蘋果豬籠草葉片42.7%（±5.5%）和捕蟲籠54.8%（±7.0%）的氮素來源於落葉，這表明食腐可增加葉片的淨光合速率。[9]
底內生物
在蘋果豬籠草的捕蟲籠內已記錄到了59種底內生物。[23]已知最小的舊大陸蛙——豬籠草姬蛙（Microhyla nepenthicola）和一種蟹蛛——豬籠草花蛛（Misumenops nepenthicola），生存於蘋果豬籠草的捕蟲籠中。[24][25]
種下類群
記載中最常見的變種是發現於沙勞越的總狀蘋果豬籠草（N. ampullaria var. racemosa），其具有總狀花序。B·H·丹瑟將其餘視為非重要的變種。[26]已發表的品種包括：[27]
- N. ampullacea var. picta Hort.Parker ex Rafarin (1869)
- N. ampullacea var. vittata Hort.Van Houtte ex Rafarin (1869)
- N. ampullaria var. geelvinkiana Becc. (1886)
- N. ampullaria var. guttata D.Moore (1872)[28]
- N. ampullaria var. longicarpa Becc. (1886)
- N. ampullaria var. microsepala Macfarl. (1911)
- N. ampullaria var. papuana Becc. in sched. nom.nud.
- N. ampullaria var. racemosa J.H.Adam & Wilcock (1990)
- N. ampullaria var. viridis Hort. ex Teijsm. (1859)[29]nom.nud.
- N. ampullaria var. vittata-major Mast. (1872)[30]
自然雜交種
蘋果豬籠草每年開花一至兩次，花期數周。它的花期與其他許多豬籠草屬植物吻合，因此它很容易和其他種形成自然雜交種。以下為已發現的自然雜交種。
- N. albomarginata × N. ampullaria[12]
- N. ampullaria × N. bicalcarata[12]
- N. ampullaria × N. eustachya[3]
- N. ampullaria × N. gracilis [=N. × trichocarpa][12]
- (N. ampullaria × N. gracilis) × N. bicalcarata [=N. × trichocarpa × N. bicalcarata]
- N. ampullaria × N. hemsleyana[31]
- N. ampullaria × N. hirsuta[12]
- N. ampullaria × N. mirabilis [=N. × kuchingensis, Nepenthes cutinensis][12]
- N. ampullaria × N. neoguineensis[3]
- N. ampullaria × N. rafflesiana [=N. × hookeriana][12][32]
- ? (N. ampullaria × N. rafflesiana) × N. mirabilis [=N. × hookeriana × N. mirabilis][33]
- N. ampullaria × N. reinwardtiana[14]
- N. ampullaria × N. spathulata[34]
- N. ampullaria × N. tobaica[14]
-
蘋果豬籠草與真穗豬籠草的自然雜交種 -
毛果豬籠草 -
蘋果豬籠草與新幾內亞豬籠草的自然雜交種 -
虎克豬籠草
註釋
- ^ F·E·勞埃德翻譯特羅爾1932年對蘋果豬籠草的描述原文：[11]
“ I came across N. ampullaria among the massive vegetations of a swamp-forest on the island of Siberut off the west coast of Sumatra. It was a fabulous, unforgettable sight. Everywhere, through the network of lianas the peculiarly-formed pitchers of this species gleamed forth, often in tight clusters and, most remarkably, the muddy moss-overgrown soil was spotted with the pitchers of this plant, so that one got the impression of a carpet. ”
參考文獻
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