翼状猪笼草

翼状猪笼草（学名：Nepenthes alata），又名红猪笼草，是菲律宾特有的热带食虫植物。其种加词“alata”来源于拉丁文“alatus”，意为“翼状的”。翼状猪笼草形态多样性极高，其下类群分类地位不断更新。
在园艺上，翼状猪笼草是最容易栽培且栽培得最广泛的物种之一，在台湾曾广泛栽培。但因其与葫芦猪笼草（N. ventricosa）的杂交种——红瓶猪笼草（N. × ventrata）更易种植且外观更优越。园艺上，翼状猪笼草逐渐被红瓶猪笼草代替，已少有人栽培。然而，近期有观点表明，园艺上曾被广泛栽培的“翼状猪笼草”已经不再属于翼状猪笼草这一物种，很有可能随着翼状猪笼草分类地位的更新被分配到了小花猪笼草名下。

**形态特征**
翼状猪笼草的花序
翼状猪笼草个体的颜色和形态差异巨大。雄花花程式为✶ K4 A4+4+1* G0（雄花顶端/*/可能不存在），雌花为✶ K4 A0 G(4)。

**分类关系**
翼状猪笼草长期以来被视为是一种形态多样性极高且分布范围涵盖整个菲律宾群岛的物种（巴拉望岛除外）。在这个宽广的界定下其分布海拔0至1900米，范围包括保和岛、卡米金岛、宿雾岛、库利昂岛、莱特岛、吕宋岛、棉兰老岛、民都洛岛、内格罗斯岛、班乃岛、萨马岛及锡布延岛。
2013年，马修·杰布与马丁·奇克重新分类了仅分布于吕宋岛北部及中部的物种，这些物种的捕虫笼均具有显着的毛被（这些类群在园艺上被称之为“多毛翼状猪笼草”）。根据这种分类，分布于更南部的类群则被指向一个被恢复的物种——小花猪笼草（N. graciliflora，园艺上的“典型翼状猪笼草”，其分布于保和岛、莱特岛、吕宋岛、棉兰老岛、民都洛岛、班乃岛、萨马岛及锡布延岛）及新描述的内格罗斯岛猪笼草small|（N. negros，比利兰岛及内格罗斯岛）与拉莫斯猪笼草（N. negros，棉兰老岛）。分布于迪纳加特岛与萨马岛的绿猪笼草（N. viridis）也为其近亲，此外还有塞西尔猪笼草（N. ceciliae，棉兰老岛）、科普兰猪笼草（N. copelandii，迪纳加特岛与棉兰老岛）、绝灭猪笼草（N. extincta，棉兰老岛）、汉密吉伊坦山猪笼草（N. hamiguitanensi，棉兰老岛）、奇坦兰山猪笼草（N. kitanglad，棉兰老岛）、仓田猪笼草（N. kurata，棉兰老岛）、莱特岛猪笼草（N. leyte，莱特岛）、萨兰加尼猪笼草（N. saranganiensis，棉兰老岛）及超基猪笼草（N. ultra，吕宋岛）。这些物种属于“翼状猪笼草组”分类群，之间存在着许多共同特征，比如叶柄具翼，捕虫笼笼盖下表面基部存在附属物，及上位笼基部通常较宽。
翼状猪笼草与许多其他的猪笼草物种间存在着密切的近缘关系，包括科普兰猪笼草、棉兰老岛猪笼草 和萨兰加尼猪笼草。产自苏门答腊的真穗猪笼草（N. eustachya）曾一度因模式标本的误释而被认为是翼状猪笼草的一个变种；实际上这两个物种间并没有密切的近缘关系。

**种下类群**
下位笼
*N. alata f. variegata Hort. ex P.Mann (1996) nom.nud.
*N. alata var. biflora Macfarl. (1908)=N. negros
*N. alata var. ecristata Macfarl. (1908)=N. mindanaoensis

**自然杂交种**
N. alata × N. burkei
*N. alata × N. merrilliana =N. × merrilliata
*? (N. alata × N. merrilliana) × N. mirabilis =N. × tsangoya
*N. alata × N. mindanaoensis
*N. alata × N. mirabilis =N. × mirabilata
*?N. alata × N. petiolata
*N. alata × N. pulchra
*N. alata × N. truncata =N. × truncalata
*N. alata × N. ventricosa =N. × ventrata

**参考文献**
 Alejandro, G.J.D., J.P.C. Baysa, B.O.C. Lemana, G.M. Madulara, R.S. Madulid & D.A. Madulid 2007. Conspecificity of Nepenthes alata Blco. population found in Mt. Guisguis, Zambales inferred from Internal Transcribed Spacer (nrDNA) sequence data. Acta Manilana 55: 15–21. 10.3860/acta.v55i0.1482- Alejandro, G.J.D., R.S. Madulid & D.A. Madulid 2008. The utility of Internal Transcribed Spacer (nrDNA) sequence data for phylogenetic reconstruction in endemic Philippine Nepenthes L. (Nepenthaceae). The Philippine Scientist 45: 99–110. 10.3860/psci.v45i0.994- Amoroso, V.B. & R.A. Aspiras 2011. Hamiguitan Range: a sanctuary for native flora. Saudi Journal of Biological Sciences 18(1): 7–15. 10.1016/j.sjbs.2010.07.003- Amoroso, V.B., L.D. Obsioma, J.B. Arlalejo, R.A. Aspiras, D.P. Capili, J.J.A. Polizon & E.B. Sumile 2009. Inventory and conservation of endangered, endemic and economically important flora of Hamiguitan Range, southern Philippines. Blumea 54(1–3): 71–76. 10.3767/000651909X474113- An, C.-I., E.-I. Fukusaki & A. Kobayashi 2001. Plasma-membrane H+-ATPases are expressed in pitchers of the carnivorous plant Nepenthes alata Blanco. Planta 212(4): 547–555. 10.1007/s004250000455- An, C.-I., E.-I. Fukusaki & A. Kobayashi 2002. Aspartic proteinases are expressed in pitchers of the carnivorous plant Nepenthes alata Blanco. Planta 214(5): 661–667. 10.1007/s004250100665- An, C.-I., S. Takekawa, A. Okazawa, E.-I. Fukusaki & A. Kobayashi 2002. Degradation of a peptide in pitcher fluid of the carnivorous plant Nepenthes alata Blanco. Planta 215(3): 472–477. 10.1007/s00425-002-0768-7- 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. 10.1111/j.1420-9101.2011.02406.x- Bauer, U. & W. Federle 2009. The insect-trapping rim of Nepenthes pitchers: surface structure and function. Plant Signaling & Behavior 4(11): 1019–1023. 10.4161/psb.4.11.9664- Benz, M.J., E.V. Gorb & S.N. Gorb 2012. Diversity of the slippery zone microstructure in pitchers of nine carnivorous Nepenthes taxa. Arthropod-Plant Interactions 6(1): 147–158. 10.1007/s11829-011-9171-2-  Blanco, F.M. 1845.  Nepenthes. date=20200524141834  In: Flora de Filipinas, segun el sistema sexual de Linneo. Segunda impresion, corregida y aumentada por el mismo autor. D. Miguel Sanchez, Manila. pp. 555–557. - Buch, F., M. Rott, S. Rottloff, C. Paetz, I. Hilke, M. Raessler & A. Mithöfer 2012. Secreted pitfall-trap fluid of carnivorous Nepenthes plants is unsuitable for microbial growth. Annals of Botany, published online on December 21, 2012. 10.1093/aob/mcs287- Cantley, R. 2000.  Nepenthes of the Philippines date=20140523213541 . video The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.- Cheers, G. 1992. Nepenthes alata. In: Letts Guide to Carnivorous Plants of the World. Letts of London House, London. p. 92.- Clarke, C. & C.C. Lee 2012.  A revision of Nepenthes (Nepenthaceae) from Gunung Tahan, Peninsular Malaysia. Gardens' Bulletin Singapore 64(1): 33–49.- Clarke, C.M. 2006. Introduction. In: Danser, B.H. The Nepenthaceae of the Netherlands Indies. Natural History Publications (Borneo), Kota Kinabalu. pp. 1–15.- Co, L. & W. Suarez 2012.  Nepenthaceae date=20130728150437 . Co's Digital Flora of the Philippines.-  Eilingsfeld, A. 2008. Endoskop-Aufnahmen aus der Verdauungsflüssigkeit einer Nepenthes alata. Das Taublatt 60(1): 4–7.- Fontanilla, J.R. & R. Madulid 2002. Morphometric analysis of Nepenthes alata Blco. in Mt. Guisguis, Sta. Cruz, Zambales. Acta Manilana 50: 25–31.- Gaume, L., S. Gorb & N. Rowe 2002. Function of epidermal surfaces in the trapping efficiency of Nepenthes alata pitchers. New Phytologist 156(3): 479–489. 10.1046/j.1469-8137.2002.00530.x- Cheek, M.R. & M.H.P. Jebb 1999. Nepenthes (Nepenthaceae) in Palawan, Philippines. Kew Bulletin 54(4): 887–895. 10.2307/4111166- Gorb, E.V., K. Haas, A. Henrich, S. Enders, N. Barbakadze & S. Gorb 2005. Composite structure of the crystalline epicuticular wax layer of the slippery zone in the pitchers of the carnivorous plant Nepenthes alata and its effect on insect attachment. Journal of Experimental Biology 208: 4651–4662. 10.1242/jeb.01939- Gorb, E.V. & S.N. Gorb 2006. Physicochemical properties of functional surfaces in pitchers of the carnivorous plant Nepenthes alata Blanco (Nepenthaceae). Plant Biology (Stuttgart, Germany) 8(6): 841–848. 17203436- Gorb, E.V. & S.N. Gorb 2009.  Chapter 8: Functional Surfaces in the Pitcher of the Carnivorous Plant Nepenthes alata: A Cryo-Sem Approach date=20121111045140 . In: S.N. Gorb (ed.) Functional Surfaces in Biology: Adhesion Related Phenomena. Volume 2. Springer. pp. 205–238.-  Gronemeyer, T. 2008. Nepenthes auf den Philippinen – Ein Reisebericht. Das Taublatt 60(1): 15–27.-  Gronemeyer, T. & V. Heinrich 2008. Wiederentdeckung von Nepenthes surigaoensis am Naturstandort auf den Philippinen. Das Taublatt 60(1): 28–33.- Hatano, N. & T. Hamada 2008. Proteome analysis of pitcher fluid of the carnivorous plant Nepenthes alata. Journal of Proteome Research 7(2): 809–816. 10.1021/pr700566d- Hatano, N. & T. Hamada 2012. Proteomic analysis of secreted protein induced by a component of prey in pitcher fluid of the carnivorous plant Nepenthes alata. Journal of Proteomics 75(15): 4844–4852. 10.1016/j.jprot.2012.05.048- Ishisaki, K., Y. Honda, H. Taniguchi, N. Hatano & T. Hamada 2012. Heterogonous expression and characterization of a plant class IV chitinase from the pitcher of the carnivorous plant Nepenthes alata. Glycobiology 22(3): 345–351. 10.1093/glycob/cwr142 - Ishisaki, K., S. Arai, T. Hamada & Y. Honda 2012. Biochemical characterization of a recombinant plant class III chitinase from the pitcher of the carnivorous plant Nepenthes alata. Carbohydrate Research 361: 170–174. 10.1016/j.carres.2012.09.001- Kajii, E., T. Kamesaki, S. Ikemoto & Y. Miura 1988. Decomposing enzymes against human blood-group antigens in the extract of Nepenthes alata. Die Naturwissenschaften 75(5): 258–259. 3405311- Kajii, E., T. Kamesaki & S. Ikemoto 1991. The effect of the Nepenthes alata extract on the cold agglutinin-associated antigens. Nihon Hōigaku Zasshi 45(1): 30–32. 2046171- Kamesaki, T., E. Kajii & S. Ikemoto 1989. Purification of the decomposing enzyme from Nepenthes alata against glycophorin B of human red blood cells by high-performance liquid chromatography. Journal of Chromatography 489(2): 384–389. 2666423- 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 - Kiew, R.G. 1990. Pitcher plants of Gunung TahaN. Journal of Wildlife and National Parks (Malaysia) 10: 34–37.- Lecoufle, M. 1990. Nepenthes alata. In: Carnivorous Plants: Care and Cultivation. Blandford, London. pp. 118–121.- Macfarlane, J.M. 1927. The Philippine species of Nepenthes. The Philippine Journal of Science 33(2): 127–140.-  Mansur, M. 2001.  Koleksi Nepenthes di Herbarium Bogoriense: prospeknya sebagai tanaman hias. In: Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253.-  McPherson, S. & T. Gronemeyer 2008. Die Nepenthesarten der Philippinen Eine Fotodokumentation. Das Taublatt 60(1): 34–78.- Meimberg, H., P. Dittrich, G. Bringmann, J. Schlauer & G. Heubl 2000. Molecular phylogeny of Caryophyllidae s.l. based on matK sequences with special emphasis on carnivorous taxa. Plant Biology 2(2): 218–228. 10.1055/s-2000-9460- 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. 10.1055/s-2001-12897-  Meimberg, H. 2002.  Molekular-systematische Untersuchungen an den Familien Nepenthaceae und Ancistrocladaceae sowie verwandter Taxa aus der Unterklasse Caryophyllidae s. l.. 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. 10.1055/s-2006-924676- Merrill, E.D. 1905.  Nepenthaceæ. date=20200524141858  In: A review of the identifications of the species described in Blanco's Flora de Filipinas. Bureau of Public Printing, Manila. p. 72.- Merrill, E.D. 1918.  Nepenthaceae date=20200524141901 . In: Species Blancoanae: a critical revision of the Philippine species of plants described by Blanco and by Llanos. Bureau of Printing, Manila. p. 160.- Mey, F.S. 2013.  Neotypification of Nepenthes blancoi and description of N. abalata a new species from the Philippines date=20160304074706 . Strange Fruits: A Garden's Chronicle, February 11, 2013.- Mey, F.S. 2013.  Nepenthes alzapan and an army of new species from the Philippines date=20160304130928 . Strange Fruits: A Garden's Chronicle, July 15, 2013.- Mithöfer, A. 2011. Carnivorous pitcher plants: insights in an old topic. Phytochemistry 72(13): 1678–1682. 10.1016/j.phytochem.2010.11.024- Owen, T.P. & K.A. Lennon 1999.  Structure and development of the pitchers from the carnivorous plant Nepenthes alata (Nepenthaceae). American Journal of Botany 86(10): 1382–1390.- Owen, T.P., K.A. Lennon, M.J. Santo & A.N. Anderson 1999.  Pathways for nutrient transport in the pitchers of the carnivorous plant Nepenthes alata. Annals of Botany 84: 459–466. 10.1006/anbo.1998.0944- Pavlovič, A., E. Masarovičová & J. Hudák 2007. Carnivorous syndrome in Asian pitcher plants of the genus Nepenthes. Annals of Botany 100(3): 527–536. 10.1093/aob/mcm145- 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. 10.1086/660882- Riedel, M., A. Eichner & R. Jetter 2003. Slippery surfaces of carnivorous plants: composition of epicuticular wax crystals in Nepenthes alata Blanco pitchers. Planta 218(1): 87–97. 10.1007/s00425-003-1075-7- Rottloff, S., R. Stieber, H. Maischak, F.G. Turini, G. Heubl & A. Mithöfer 2011. Functional characterization of a class III acid endochitinase from the traps of the carnivorous pitcher plant genus, Nepenthes. Journal of Experimental Botany 62(13): 4639–4647. 10.1093/jxb/err173- Santo, M.J., J.S. Massa, & T.P. Owen 1998. Glandular secretion and absorption in the carnivorous pitcher plant Nepenthes alata. American Journal of Botany 85(supplement): 92.- Schlauer, J. 2000.  Literature Reviews. Carnivorous Plant Newsletter 29(2): 53.- Scholz, I., M. Bückins, L. Dolge, T. Erlinghagen, A. Weth, F. Hischen, J. Mayer, S. Hoffmann, M. Riederer, M. Riedel & W. Baumgartner 2010. Slippery surfaces of pitcher plants: Nepenthes wax crystals minimize insect attachment via microscopic surface roughness. Journal of Experimental Biology 213: 1115–1125. 10.1242/jeb.035618- Slack, A. 1979. Nepenthes alata. In: Carnivorous Plants. Ebury Press, London. pp. 85–86.- Sota, T., M. Mogi & K. Kato 1998. Local and regional-scale food web structure in Nepenthes alata pitchers. Biotropica 30(1): 82–91. 10.1111/j.1744-7429.1998.tb00371.x-  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  - Thornhill, A.H., I.S. Harper & N.D. Hallam 2008. The development of the digestive glands and enzymes in the pitchers of three Nepenthes species: N. alata, N. tobaica, and N. ventricosa (Nepenthaceae). International Journal of Plant Sciences 169(5): 615–624. 10.1086/533599- Uy, B.D. 1990.  The granchina Nepenthes of the Phillipines in retrospect. Carnivorous Plant Newsletter 19(1–2): 18–19.- Wang, L.-X. & Q. Zhou 2010.  Numerical characterization of surface structures of slippery zone in Nepenthes alata pitchers and its mechanism of reducing locust's attachment date=20160304001127 . Advances in Natural Science 3(2): 152–160.-  Wu, X., J. Li, Q. Zhong & X. Wu 2005.  猪笼草组培快繁技术 date=20110707022756 . Tissue culture and fast propagation in Nepenthes alata. Nonwood Forest Research 23(4): 48–50.

**扩展阅读**
 食虫植物照片搜寻引擎中 翼状猪笼草的照片 date=20210405143850