白环猪笼草

白环猪笼草（学名：）是婆罗洲、西马来西亚和苏门答腊特有的热带食虫植物。Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.Adam, J.H., C.C. Wilcock & M.D. Swaine 1989. Ecology and taxonomy of Bornean Nepenthes. University of Aberdeen Tropical Biology Newsletter 56: 2–4.其种加词“albomarginata”来源于拉丁文“albus”和“marginatus”，意为“白色”和“边”，指其唇下特有的白色环状绒毛。

 植物学史 
1848年，托马斯·洛布首次采集到了白环猪笼草。一年后，约翰·林德利在《园丁纪事》中正式描述了白环猪笼草。Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.

1856年，白环猪笼草被引种至英国。

 形态特征 
白环猪笼草为藤本植物。茎攀爬至4米的高处，直径可达5毫米，呈圆柱形。节间距可达15厘米。

白环猪笼草的叶片为革质，呈披针形，可长达25厘米，宽至2厘米。叶尖急尖，叶基渐尖，抱茎。白环猪笼草的叶片无叶柄。纵脉不明显。笼蔓可长达20厘米。

白环猪笼草下位笼的下三分之一为球形，上部为圆柱形。下位笼的体型较小，仅可高达15厘米，宽至4厘米。腹面具一对可宽达5毫米的笼翼。笼口呈圆形，基部拉長形成一個較短的唇頸。唇為圓柱形，可寬達2毫米，唇齿不明显。唇内缘部分约为截面表面总长度的34%。Bauer, U., C.J. Clemente, T. Renner & W. Federle 2012. Form follows function: morphological diversification and alternative trapping strategies in carnivorous Nepenthes pitcher plants. Journal of Evolutionary Biology 25(1): 90–102. 唇下有一层密集的白色绒毛，其可能具有引诱白蚁的功能。下位笼球形部分对应的内表面覆盖着消化腺。笼盖近圆形，无附属物。笼盖基部的后方具一根不分叉的笼蔓尾，可长达3毫米。

白环猪笼草的上位笼为圆柱形至漏斗形。笼翼缩小为一对隆起。其余部分与下位笼类似。

白环猪笼草的花序为总状花序，通常雄性花序较长。总花梗可长达25厘米，花序轴可达40厘米。花梗带1至2朵花，长约30毫米，无苞片。萼片为卵形至椭圆形，可长达4毫米。一项对来自同一副标本花粉大小的研究显示，其120粒花粉的平均直径为31.8微米。Adam, J.H. & C.C. Wilcock 1999.  Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.

白环猪笼草植株的大部分披被着密集的白色短毛，但在叶片下面的毛被通常较长。

白环猪笼草捕虫笼的颜色可为绿色、红色、黑色或具斑点。

 食虫性 
白环猪笼草最特别的是它只捕食白蚁；大部分猪笼草不会对猎物进行选择。据植物学家马利斯·A·梅尔巴赫和他同事的所述，白环猪笼草是一种食性专一的食虫植物。Merbach, M.A., D.J. Merbach, U. Maschwitz, W.E. Booth, B. Fiala & G. Zizka 2002. Carnivorous plants: Mass march of termites into the deadly trap. Nature 415: 36–37. Clarke, T. 2002. Plant has taste for termites . Nature News, January 3, 2002. Moran, J.A., M.A. Merbach, N.J. Livingston, C.M. Clarke & W.E. Booth 2001.  Annals of Botany 88: 307–311.Merbach, M.A., D.J. Merbach, W.E. Booth, U. Maschwitz, G. Zizka & B. Fiala 2000. A unique niche in plant carnivory: Nepenthes albomarginata feeds on epigaeically mass foraging termites. Tagungsband gtö 2000 13. Jahrestagung der Deutschen Gesellschaft für Tropenökologie 1–3. March 2000 in Würzburg Lehrstuhl für Tierökologie und Tropenbiologie Universität Würzburg. p. 105.

白环猪笼草具有独特的形态特征：在其唇下具一圈白色的绒毛。已捕获到白蚁的捕虫笼上的白色环状绒毛往往已消失。梅尔巴赫说：“有时会接连几日什么也没有发生，接着某个夜晚过后，捕虫笼内充满了白蚁，同时唇下的白毛也不见了。”

马利斯·A·梅尔巴赫为了研究这一现象，将一棵白毛完整的白环猪笼草放在正在寻找食物的白蚁附近。当白蚁发现了白环猪笼草的捕虫笼上的白毛后，便会啃食它。在吃了那些白毛后，不管是工蚁还是兵蚁都陆陆续续的掉入了捕虫笼内。

一旦它们掉入了捕虫笼内就再也无法爬出来了。马利斯·A·梅尔巴赫记录到的最高记录是每分钟有22只白蚁落入笼内。一个小时过后，白毛全部消失，无白毛的捕虫笼不能吸引白蚁，也就不再有白蚁落入捕虫笼。

尚未清楚这些白毛是如何吸引白蚁的。但马利斯·A·梅尔巴赫在实验中发现，这些白毛对于白蚁并没有远距离的吸引力。他说：“这一切看起来好像都是巧合，那些白蚁经常在离白毛只有不到1厘米的地方却没有发现它。”

马利斯·A·梅尔巴赫还指出，白环猪笼草是唯一一种将自身的一部分作为诱饵引诱猎物的食虫植物。

 生态关系 
白环猪笼草存在于婆罗洲、西马来西亚和苏门答腊。也存在于尼亞斯島及槟榔屿等较小型的岛屿。McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Peninsular Malaysia and Indochina. Redfern Natural History Productions, Poole.McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.其海拔分布范围为0至1200米。Adam, J.H., C.C. Wilcock & M.D. Swaine 1992.  Journal of Tropical Forest Science 5(1): 13–25.

白环猪笼草的原生地主要为巽他荒原森林，但也可能存在于低海拔山峰的山顶植被中。其陆生于泥炭和石灰石基质中。Anderson, J.A.R. 1965. Limestone habitat in Sarawak. Proceedings of the Symposium on Ecological Research in Humid Tropics Vegetation, July 1963, Kuching, Sarawak. pp. 49–57.

 相关物种 
2001年，查尔斯·克拉克对产自苏门答腊岛和西马来西亚的猪笼草进行了分支系统学分析，共利用了70个形态特征。以下为猪笼草进化树的“第6分支”，其相关性达50%。

 种下类群 
N. albomarginata f. sanguinea Toyoda ex Hinode-Kadan (1985) nom.nud.
N. albomarginata var. rubra (Hort. ex Macfarl.) Macfarl. (1908)Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler. Das Pflanzenreich IV, III, Heft 36: 1–91.
N. albomarginata var. tomentella (Miq.) Beck (1895) Beck, G. 1895. Die Gattung Nepenthes. Wiener Illustrirte Garten-Zeitung 20(3–6): 96–107, 141–150, 182–192, 217–229.
N. albomarginata var. typica Beck (1895)nom.illeg.
N. albomarginata var. villosa Hook.f. (1873) Hooker, J.D. 1873. Ordo CLXXV bis. Nepenthaceæ. In: A. de Candolle Prodromus Systematis Naturalis Regni Vegetabilis 17: 90–105.

 自然杂交种 

N. albomarginata × N. ampullariaClarke, C.M. 1997. Nepenthes of Borneo.  Natural History Publications (Borneo), Kota Kinabalu.
? N. albomarginata × N. chanianaMcPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
N. albomarginata × N. clipeata
N. albomarginata × N. eustachya
N. albomarginata × N. gracilis
N. albomarginata × N. hirsuta
N. albomarginata × N. macrovulgaris
N. albomarginata × N. northiana [=N. × cincta]Masters, M.T. 1884. New garden plants. Nepenthes cincta (Mast.), n. sp. . The Gardeners' Chronicle, new series, 21(540): 576–577.
N. albomarginata × N. rafflesianaLowrie, A. 1983.  Carnivorous Plant Newsletter 12(4): 88–95.
N. albomarginata × N. reinwardtiana [=N. × ferrugineomarginata]
? N. albomarginata × N. sanguineaShivas, R.G. 1985.  Carnivorous Plant Newsletter 14(1): 13–14.
N. albomarginata × N. veitchii

 白环猪笼草与诺斯猪笼草的自然杂交种 

由于诺斯猪笼草的分布局限，仅生长在沙捞越石龙门的石灰岩基质上，所以雪线猪笼草更为罕见。

白环猪笼草在杂交中很强势，这使得其与诺斯猪笼草杂交后代完全失去了宽大的唇。该杂交种的捕虫笼为窄漏斗形，笼身为奶油色至暗紫色，并具有红色或黑色的斑点。Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.

 白环猪笼草与两眼猪笼草的自然杂交种 

红脉猪笼草的分布范围涵盖了婆罗洲和苏门答腊。1981年，仓田重夫在西加里曼丹省采集到了红脉猪笼草的模式标本。次年，仓田重夫描述了这个自然杂交种。

 参考文献 

 [Anonymous] 1877. Reports of Societies. Royal Horticultural . The Gardeners' Chronicle 8(197): 441.
 [Anonymous] 1883. Mr. A. E. Ratcliff's Nepenthes . The Gardeners' Chronicle 20(497): 18–19.
 Adam, J.H. 1997.  Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.
 Beveridge, N.G.P., C. Rauch, P.J.A. Keßler, R.R. van Vugt & P.C. van Welzen 2013. A new way to identify living species of Nepenthes (Nepenthaceae): more data needed! Carnivorous Plant Newsletter 42(4): 122–128.
  Blondeau, G. 2001. Nepenthes albomarginata. In: Les Plantes Carnivores. De Vecchi, Paris. p. 69.
 Bonhomme, V., H. Pelloux-Prayer, E. Jousselin, Y. Forterre, J.-J. Labat & L. Gaume 2011. Slippery or sticky? Functional diversity in the trapping strategy of Nepenthes carnivorous plants. New Phytologist, published online on March 24, 2011. 

 Damayanti, F., M. Mansur & I. Roostika 2011. Diversity of Nepenthes spp. in West Kalimantan.  International Journal of Biodiversity and Conservation 3(13): 705–708. 
 Dixon, W.E. 1889. Nepenthes.  The Gardeners' Chronicle, series 3, 6(144): 354.
 Fashing, N.J. 2010.  In: M.W. Sabelis & J. Bruin (eds.) Trends in Acarology: Proceedings of the 12th International Congress . Springer Science, Dordrecht. pp. 81–84. 
 Hernawati & P. Akhriadi 2006. A Field Guide to the Nepenthes of Sumatra. PILI-NGO Movement, Bogor.
 Hooker, J.D. 1859. XXXV. On the origin and development of the pitchers of Nepenthes, with an account of some new Bornean plants of that genus . The Transactions of the Linnean Society of London 22(4): 415–424. 
 Kato, M., M. Hotta, R. Tamin & T. Itino 1993. Inter- and intra-specific variation in prey assemblages and inhabitant communities in Nepenthes pitchers in Sumatra. Tropical Zoology 6(1): 11–25. Abstract
 Kitching, R.L. 2000. Food Webs and Container Habitats: The natural history and ecology of phytotelmata . Cambridge University Press, Cambridge.
  Kurniawati, R. 2010. Serangga yang terdapat pada kantong Nepenthes albomarginata T. Lobb ex Lindl. dan Nepenthes eustachya Miq. di Kawasan Cagar Alam Lembah Harau Kabupaten Limapuluh Kota. Thesis, Andalas University, Padang. Abstract 
 Kurup, R., A.J. Johnson, S. Sankar, A.A. Hussain, C.S. Kumar & S. Baby 2013. Fluorescent prey traps in carnivorous plants. Plant Biology 15(3): 611–615. 
 Lecoufle, M. 1990. Nepenthes albo-marginata. In: Carnivorous Plants: Care and Cultivation. Blandford, London. pp. 128–129.
 Lee, C.C. 2000. Recent Nepenthes Discoveries . [video] The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.
  Macák, M. 1998. Portréty rostlin - Nepenthes albomarginata . Trifid 1998(3–4): 57–59. (page 2 , page 3 )
 Macfarlane, J.M. 1914. Family XCVI. Nepenthaceæ.  [pp. 279–288] In: J.S. Gamble. Materials for a flora of the Malayan Peninsula, No. 24.  Journal & Proceedings of the Asiatic Society of Bengal 75(3): 279–391.
  Mansur, M. 2001.  In: Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253.
  Mansur, M. 2007. Keanekaragaman jenis Nepenthes (kantong semar) dataran rendah di Kalimantan Tengah. [Diversity of lowland Nepenthes (kantong semar) in Central Kalimantan.] Berita Biologi 8(5): 335–341. Abstract
 Mansur, M. & F.Q. Brearley 2008. Ecological studies on Nepenthes at Barito Ulu, Central Kalimantan, Indonesia. Jurnal Teknologi Lingkungan 9(3): 271–276.
 Masters, M.T. 1872. The cultivated species of Nepenthes.  The Gardeners' Chronicle and Agricultural Gazette 1872(16): 540–542.
 McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
 Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. Plant Biology 3(2): 164–175. 
  Meimberg, H. 2002.  Ph.D. thesis, Ludwig Maximilian University of Munich, Munich.
 Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. Plant Biology 8(6): 831–840. 
 Meimberg, H., S. Thalhammer, A. Brachmann & G. Heubl 2006. Comparative analysis of a translocated copy of the trnK intron in carnivorous family Nepenthaceae. Molecular Phylogenetics and Evolution 39(2): 478–490. 
 Moran, J.A., W.E. Booth & J.K. Charles 1999.  Annals of Botany 83: 521–528.
  Murniati, Syamswisna & A. Nurdini 2013. Pembuatan flash card dari hasil inventarisasi Nepenthes di hutan adat desa Teluk Bakung . Jurnal Pendidikan dan Pembelajaran 2(1): [unpaginated; 14 pp.] Abstract  
  Nurita, E. 2010. Mikrosporogenesis pada kantung semar (Nepenthes albomarginata T.Lobb ex Lindl). Thesis, Andalas University, Padang. Abstract 
 Osunkoya, O.O., S.D. Daud & F.L. Wimmer 2008. Longevity, lignin content and construction cost of the assimilatory organs of Nepenthes species. Annals of Botany 102(5): 845–853. 
 Renner, T. & C.D. Specht 2011. A sticky situation: assessing adaptations for plant carnivory in the Caryophyllales by means of stochastic character mapping. International Journal of Plant Sciences 172(7): 889–901. 
 Riedel, M., A. Eichner, H. Meimberg & R. Jetter 2007. Chemical composition of epicuticular wax crystals on the slippery zone in pitchers of five Nepenthes species and hybrids. Planta 225(6): 1517–1534. 
 Shivas, R.G. 1984. Pitcher Plants of Peninsular Malaysia & Singapore. Maruzen Asia, Kuala Lumpur.
 Smythies, B.E. 1965. The distribution and ecology of pitcher-plants (Nepenthes) in Sarawak. UNESCO Humid Tropics Symposium, June–July 1963, Kuching, Sarawak.
  Sukamto, L.A. 2005.  In: Laporan Teknik Pusat Penelitian Biologi-LIPI. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 433–438.
  Syamsuardi & R. Tamin 1994. Kajian kekerabatan jenis-jenis Nepenthes di Sumatera Barat. Project report, Andalas University, Padang. Abstract 
  Syamsuardi 1995. Klasifikasi numerik kantong semar (Nepenthes) di Sumatera Barat. [Numerical classification of pitcher plants (Nepenthes) in West Sumatra.] Journal Matematika dan Pengetahuan Alam 4(1): 48–57. Abstract 
 Takeuchi, Y., M.M. Salcher, M. Ushio, R. Shimizu-Inatsugi, M.J. Kobayashi, B. Diway, C. von Mering, J. Pernthaler & K.K. Shimizu 2011. In situ enzyme activity in the dissolved and particulate fraction of the fluid from four pitcher plant species of the genus Nepenthes. PLoS ONE 6(9): e25144. 
 Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York.
 Wan, A.S., R.T. Aexel, R.B. Ramsey & H.J. Nicholas 1972. Nepenthaceae: sterols and triterpenes of the pitcher plant. Phytochemistry 11(1): 456–461. 

 扩展阅读 

 美国种质资源信息网络中白环猪笼草的信息 
 食虫植物照片搜寻引擎中白环猪笼草的照片 
槟榔屿山的白环猪笼草 

Nepenthes albomarginata
albomarginata
Category:婆罗洲植物
Category:苏门答腊植物
Category:馬來西亞植物