小猪笼草

小猪笼草（学名：Nepenthes gracilis）广布于东南亚低地的热带食虫植物。其种加词来源于拉丁文“gracilis”，意为“细长的”。小猪笼草存在于马来西亚、新加坡、印尼苏门答腊及泰国。经常匍匐生长在没有树荫的开阔草地上。该物种的海拔分布范围也很广，为海拔0至1100米（少数甚至可达1700米），但其大部分都分布于海拔100米以下的地区，极少数植株存在于海拔1000米以上的地区。尽管其分布非常的广泛，但关于小猪笼草的自然杂交种却很少见，最知名的就是毛果猪笼草。
彼得·威廉·科萨尔在1839年发表的专着《关于猪笼草属》中对小猪笼草进行了正式的描述。
产自菲律宾的源小猪笼草（N. abgracilis）由于与小猪笼草非常相似而得名。

**食虫性**
小猪笼草的捕虫笼相对比较普通，唇非常窄小且相对其他猪笼草而言寿命较短。但其捕虫笼笼盖下表面却存在着参差不齐的蜡质层，该特征可能是猪笼草属内唯一的。该蜡质层与捕虫笼内表面蜡质区的蜡质层厚度不一且结构也完全不同。，该蜡质层在下暴雨时会增加昆虫在笼盖下表面的附着难度，使昆虫容易受雨滴撞击而落入捕虫笼内，从而该蜡质层成为小猪笼草捕获猎物机制的一部分。科学家透过X光分析小猪笼草，发现笼盖与植株间有个特别脆弱的点，该点被雨滴击中时会塌陷，连带使笼盖往下弹，让猎物落入捕虫笼后以特定方式复原。这是其近亲莱佛士猪笼草没有的机制。

**分布范围**
位于西马来西亚彭亨的小猪笼草原生地
小猪笼草分布广泛，存在于婆罗洲、西马来西亚、新加坡、苏拉威西中部、苏门答腊及泰国南部。其也存在于许多小型岛屿中，包括邦加岛、巴图群岛、勿里洞岛望加丽岛、兰塔岛、达鲁岛、纳闽岛、浮罗交怡、孟德尔岛、明打威群岛（西比路岛）、美兰地群岛（巴东岛、朗桑岛、直名丁宜岛）、马萨拉岛、尼亚斯岛、槟榔屿、廖内群岛（林加群岛、纳土纳群岛、廖内岛）、鲁帕岛。

**相关物种**
2001年，查尔斯·克拉克对来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”。

**种下类群**
尽管小猪笼草的可变性较小，但其还是存在着大量的种下类群。但大部分命名都不是有效的。
*Nepenthes gracilis f. angustifolia (Mast.) Hort.Westphal (1993)
*Nepenthes gracilis var. angustifolia (Mast.) Hort.Weiner in sched. (1985)
*Nepenthes gracilis var. arenaria Ridl. ex Macf. (1908)
*Nepenthes gracilis var. elongata Bl. (1852)
*Nepenthes gracilis var. longinodis Beck (1895)
*Nepenthes gracilis var. major Hort.Lindsay ex Dixon (1889)
*Nepenthes gracilis var. teysmanniana (Miq.) Beck (1895)

**自然杂交种**
以下是已发现的关于小猪笼草的自然杂交种。
*N. albomarginata × N. gracilis
*N. ampullaria × N. gracilis =N. × trichocarpa
*(N. ampullaria × N. gracilis) × N. bicalcarata =N. × trichocarpa × N. bicalcarata
*N. bicalcarata × N. gracilis =N. × cantleyi
*? N. eustachya × N. gracilis
*N. gracilis × N. mirabilis =N. × sharifah-hapsahii, N. × ghazallyana, N. × grabilis, N. neglecta?
*N. gracilis × N. northiana =N. × bauensis
*N. gracilis × N. rafflesiana
*N. gracilis × N. reinwardtiana
*N. gracilis × N. sumatrana

**参考文献**
 Adam, J.H. 1997.  Prey spectra of Bornean Nepenthes species (Nepenthaceae) in relation to their habitat. Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.- Adam, J.H. & C.C. Wilcock 1999.  Palynological study of Bornean Nepenthes (Nepenthaceae). Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.- Adam, J. H., R. Omar & C. C. Wilcock 2002. 1= Phytochemical Screening of Flavonoids in Three Hybrids of Nepenthes (Nepenthaceae) and their Putative Parental Species from Sarawak and Sabah. OnLine Journal of Biological Sciences 2(9): 623–625. 10.3923/jbs.2002.623.625- Adam, J.H., E.M. Nurulhuda, H. Abdul-Halim, O. Abdul-Rahim, A.H. Hafiza, G.K. Gopir, L.M. Pilik, R. Omar, M.B. Qasim, J. Salimon, S. Abdul-Rahim & M.M. Hanafiah 2005.  Pitcher plants recorded from BRIS forest in Jambu Bongkok, Kuala Trengganu, Malaysia date=20110707022420 . Wetland Science 3(3): 183–189.-  Adam, J.H., J.N. Maisarah, A.T.S. Norhafizah, A.H. Hafiza, M.Y. Harun & O.A. Rahim et al. 2009. Ciri Tanih Pada Habitat Nepenthes (Nepenthaceae) di Padang Tujuh, Taman Negeri Endau-Rompin Pahang. Soil Properties in Nepenthes (Nepenthaceae) Habitat at Padang Tujuh, Endau-Rompin State Park, Pahang. In: J.H. Adam, G.M. Barzani & S. Zaini (eds.) Bio-Kejuruteraan and Kelestarian Ekosistem. Bio-Engineering and Sustainable Ecosystem. Kumpulan Penyelidikan Kesihatan Persekitaran, Pusat Penyelidikan Bukit Fraser and Universiti Kebangsaan, Malaysia. pp. 147–157.- Adam, J.H., H.A. Hamid, M.A.A. Juhari, S.N.A. Tarmizi & W.M.R. Idris 2011. Species composition and dispersion pattern of pitcher plants recorded from Rantau Abang in Marang District, Terengganu State of Malaysia. International Journal of Botany 7(2): 162–169. 10.3923/ijb.2011.162.169-  Akhriadi, P. 2007.  Kajian taksonomi hibrid alami Nepenthes (Nepenthaceae) di Kerinci. Working paper, Andalas University, Padang.  Abstract - Athauda, S.B.P., K. Matsumoto, S. Rajapakshe, M. Kuribayashi, M. Kojima, N. Kubomura-Yoshida, A. Iwamatsu, C. Shibata, H. Inoue & K. Takahashi 2004. PDFlink| Enzymatic and structural characterization of nepenthesin, a unique member of a novel subfamily of aspartic proteinases.|1.32 MiB (manuscript BJ20031575) Biochemical Journal 381(1): 295–306. 10.1042/BJ20031575- Aung, H.H., L.S. Chia, N.K. Goh, T.F. Chia, A.A. Ahmed, P.W. Pare & T.J. Mabry 2002. Phenolic constituents from the leaves of the carnivorous plant Nepenthes gracilis. Fitoterapia 73(5): 445–447. 12165348 10.1016/S0367-326X(02)00113-2-  Baloari, G., R. Linda & Mukarlina 2013.  Keanekaragaman jenis dan pola distribusi Nepenthes spp di Gunung Semahung Kecamatan Sengah Temila Kabupaten Landak date=20131229042208 . Protobiont 2(1): 1–6.  Abstract date=20131229042145  - 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- Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.- 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 191(2): 545–554. 10.1111/j.1469-8137.2011.03696.x- Bourke, G. 2010.  The climbing pitcher plants of the Kelabit highlands. Captive Exotics Newsletter 1(1): 4–7. - Brearley, F.Q. & M. Mansur 2012. Nutrient stoichiometry of Nepenthes species from a Bornean peat swamp forest. Carnivorous Plant Newsletter 41(3): 105–108.- 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 111(3): 375–383. 10.1093/aob/mcs287- Chia, T.F., H.H. Aung, A. Osipov, N.K. Goh & L.S. Chia 2004. Carnivorous pitcher plant uses free radicals in the digestion of prey. Redox Report 9(5): 255–261. 10.1179/135100004225006029- Chua, T.J.L. & M.L.M. Lim 2012. Cross-habitat predation in Nepenthes gracilis: the red crab spider Misumenops nepenthicola influences abundance of pitcher dipteran larvae. Journal of Tropical Ecology 28(1): 97–104. 10.1017/S0266467411000629- downloaded=10 May 2006- Damayanti, F., M. Mansur & I. Roostika 2011.  Diversity of Nepenthes spp. in West Kalimantan. date=20201203231914  International Journal of Biodiversity and Conservation 3(13): 705–708. -  Enjelina, W. 2012.  Analisis hibrid alam kantung semar (Nepenthes) di Bukit Taratak Kabupaten Pesisir Selatan Sumatera Barat dengan teknik RAPD. M.Sc. thesis, Andalas University, Padang. - Fan, D.-H., H. Wang, D. Zhi & Y.-M. Shen 2010. CE analysis of endogenous flavonoid gallate esters from Nepenthes gracilis (Nepenthaceae). Chromatographia 72(9–10): 1013–1016. 10.1365/s10337-010-1729-0- Fashing, N.J. 2010.  Two novel adaptations for dispersal in the mite family Histiostomatidae (Astigmata). In: M.W. Sabelis & J. Bruin (eds.)  Trends in Acarology: Proceedings of the 12th International Congress date=20131228222929 . Springer Science, Dordrecht. pp. 81–84. 10.1007/978-90-481-9837-5- Frazier, C.K. 2000.  Reproductive isolating mechanisms and fitness among tropical pitcher plants (Nepenthes) and their hybrids date=20140522104435 . video The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.-  Handayani, T. 1999.  Konservasi Nepenthes di kebun raya Indonesia. Conservation of Nepenthes in Indonesian botanic gardens. In: A. Mardiastuti, I. Sudirman, K.G. Wiryawan, L.I. Sudirman, M.P. Tampubolon, R. Megia & Y. Lestari (eds.) Prosiding II: Seminar Hasil-Hasil Penelitian Bidang Ilmu Hayat. Pusat Antar Universitas Ilmu Hayat IPB, Bogor. pp. 365–372.-  Harahap, A.S. 2010.  Mikropropogasi tunas kantong semar (Nepenthes gracillis Korth.) dengan pemberian NAA dan BAP secara in vitro date=20110406170055 . Student paper, University of North Sumatra, Medan.- 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 date=20130703194740 . The Transactions of the Linnean Society of London 22(4): 415–424. 10.1111/j.1096-3642.1856.tb00113.x- Kato, M. 1993. Floral biology of Nepenthes gracilis (Nepenthaceae) in Sumatra. American Journal of Botany 80(8): 924–927. 10.2307/2445512- 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 date=20121113173602 . Cambridge University Press, Cambridge.- 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. 10.1111/j.1438-8677.2012.00709.x- Lam, S.Y. 1982. Tripteroides aranoides (Theobald) in two pitcher plants, Nepenthes ampullaria Jack and N. gracilis Korth., at Kent Ridge (Diptera: Culicidae). B.Sc (Hons.) Thesis, National University of Singapore.- Lee, C.C. 2000.  Recent Nepenthes Discoveries date=20140605173342 . video The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.- Lim, A.L. & N. Prakash 1973. Life history of Nepenthes gracilis. Malaysian Journal of Science 2(A): 45–53.- Lim, S.H., D.C.Y. Phua & H.T.W. Tan 2000. Primer design and optimization for RAPD analysis of Nepenthes. Biologia Plantarum 43(1): 153–155. 10.1023/A:1026535920714- Ma Hnin Hnin Aung 2004.  Phenolic compounds from Nepenthes gracilis Korth: isolation, identification and antioxidant studies. Ph.D. thesis, Nanyang Technological University, Singapore.- Macfarlane, J.M. 1914.  Family XCVI. Nepenthaceæ. date=20131228130100  pp. 279–288 In: J.S. Gamble.  Materials for a flora of the Malayan Peninsula, No. 24. date=20131228130100  Journal & Proceedings of the Asiatic Society of Bengal 75(3): 279–391.- 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.-  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. 2008.  Penelitian ekologi Nepenthes di Laboratorium Alam Hutan Gambut Sabangau Kereng Bangkirai Kalimantan Tengah date=20160304025237 . Ecological studies on Nepenthes at Peat Swamps Forest Natural Laboratory, Kereng Bangkirai Sabangau, Central Kalimantan. Jurnal Teknologi Lingkungan 9(1): 67–73.  Abstract date=20130912085709  - Mansur, M. & F.Q. Brearley 2008.  Ecological studies on Nepenthes at Barito Ulu, Central Kalimantan, Indonesia. Jurnal Teknologi Lingkungan 9(3): 271–276.- Moore, D. 1872.  On the culture of Nepenthes at Glasnevin. date=20131016193222  The Gardeners' Chronicle and Agricultural Gazette 1872(11): 359–360. - Masters, M.T. 1872.  The cultivated species of Nepenthes. date=20130923043952  The Gardeners' Chronicle and Agricultural Gazette 1872(16): 540–542.- 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- 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. 10.1016/j.ympev.2005.11.023-  Meriko, L. 2010.  Perkembangan androesium beberapa jenis Nepenthes (N. ampullaria Jack., N. gracilis Korth. dan N. reinwardtiana Miq.). M.Sc. thesis, Andalas University, Padang.  Abstract - Mithöfer, A. 2011. Carnivorous pitcher plants: insights in an old topic. Phytochemistry 72(13): 1678–1682. 10.1016/j.phytochem.2010.11.024- Miyagi, I. & T. Toma 2007. A new mosquito of the genus Topomyia (Diptera, Culicidae) from a Nepenthes pitcher plant in a Bario highland of Sarawak, Malaysia. Medical Entomology and Zoology 58(3): 167–174.  Abstract date=20110520193350 - Mogi, M. & K.L. Chan 1997. Variation in communities of dipterans in Nepenthes pitchers in Singapore: Predators increase prey community diversity. Annals of the Entomological Society of America 90(2): 177–183.- Mokkamul, P., A. Chaveerach, R. Sudmoon & T. Tanee 2007.  Species Identification and Sex Determination of the Genus Nepenthes (Nepenthaceae). date=20110724055841  Pakistan Journal of Biological Sciences 10(4): 561–567. 10.3923/pjbs.2007.561.567- Moran, J.A., W.E. Booth & J.K. Charles 1999.  Aspects of Pitcher Morphology and Spectral Characteristics of Six Bornean Nepenthes Pitcher Plant Species: Implications for Prey Capture. Annals of Botany 83: 521–528.- Mullins, J. & M. Jebb 2009.  Phylogeny and biogeography of the genus Nepenthes. National Botanic Gardens, Glasnevin. -  Murniati, Syamswisna & A. Nurdini 2013.  Pembuatan flash card dari hasil inventarisasi Nepenthes di hutan adat desa Teluk Bakung date=20131229042230 . Jurnal Pendidikan dan Pembelajaran 2(1): unpaginated; 14 pp.  Abstract date=20130629055105  - Normawati, Y. 2002. The effect of stem length on pitcher and inflorescence production in Nepenthes gracilis and Nepenthes mirabilis at Serendah Selangor. B.Sc. Thesis. Universiti Kebangsaan Malaysia.-  Oikawa, T. 1992. Nepenthes gracilis Korth.. In: Nihongo
Muyū kusa – Nepenthes. The Grief Vanishing. Parco Co., Japan. pp. 38–39.- 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. 10.1093/aob/mcn162- Pollard, S.D. 2012. Fishing spiders in the hanging stomachs of Borneo. Carnivorous Plant Newsletter 41(1): 20–23.-  Puspitaningtyas, D.M. & H. Wawangningrum 2007.  Keanekaragaman Nepenthes di Suaka Alam Sulasih Talang - Sumatera Barat. Nepenthes diversity in Sulasih Talang Nature Reserve - West Sumatra. -  Rahim, S.A., T. Lihan, M. Baba, A. Laming, Z.A. Rahman, W.M.R. Idris, M.B. Gasim, A. Hashim, S.M. Yusof & L.H. Yin 2008. Pengambilan logam berat oleh Nepenthes gracilis dan N. Hookeriana dalam tanih bekas lombong besi dan timah, Pelepah Kanan, Kota Tinggi, Johor. Heavy metal uptake by Nepenthes gracilis and N. Hookeriana in ex-iron and tin mine soil, Pelepah Kanan, Kota Tinggi, Johor. Sains Malaysiana 37(1): 39–49.  Abstract date=20120503014240 - Robinson, A. 1995 '1994/95'. Plant findings in Malaysia. The Carnivorous Plant Society Journal 18: 44–47.- 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- Russell, C. & E. Ossian 1990. Opportunistic feeding involving the pitcher plants Nepenthes hirsuta, Nepenthes gracilis and the epiphytic orchid Schomburgkia tibicinis, or natural ant eradication, the rube goldberg method. The Orchid Digest 54(4): 182–184.- Shivas, R.G. 1984. Pitcher Plants of Peninsular Malaysia & Singapore. Maruzen Asia, Kuala Lumpur.- Siegara, A. & Yogiara 2009.  Bacterial community profiles in the fluid of four pitcher plant species (Nepenthes spp.) grown in a nurseryfix-attempted=yes . Microbiology Indonesia 3(3): 109–114.- Slack, A. 1979. Nepenthes gracilis. In: Carnivorous Plants. Ebury Press, London. p. 86.- Sun, W.F. 2008.  In vitro induction of polyploidy in Nepenthes gracilis date=2013-02-21 . M.Sc. thesis, University of Malaya, Kuala Lumpur.-  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 - Takahashi, K., S.B.P. Athauda, K. Matsumoto, S. Rajapakshe, M. Kuribayashi, M. Kojima, N. Kubomura-Yoshida, A. Iwamatsu, C. Shibata & H. Inoue 2005. Nepenthesin, a unique member of a novel subfamily of aspartic proteinases: enzymatic and structural characteristics. Current Protein and Peptide Science 6(6): 513–525. 10.2174/138920305774933259- 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. 10.1371/journal.pone.0025144- Tan, S. 2006.  Development of capillary electrophoresis techniques for plant natural products analyses. M.Sc. thesis, Nanyang Technological University, Singapore.- Teo, L.L. 2001.  Study of natural hybridisation in some tropical plants using amplified fragment length polymorphism analysis. M.Sc. thesis, Nanyang Technological University, Singapore.- Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York.-  Toufar, P. 1996.  Ještě jednou k Nepenthes gracilis date=20120315080708 . Trifid 1996(2): 6–7. ( page 2 date=20120315080728 )-  Uji, T. 2003.  Keanekaragaman dan potensi flora di Cagar Alam Muara Kendawangan, Kalimantan Barat. date=20131228034639  Flora diversity and its potential in Muara Kendawangan Nature Reserve, West Kalimantan. Biodiversitas 4(1): 112–117. -  James Wong and the Malaysian Garden date=20121103180025 . video BBC Two.

**扩展阅读**
 食虫植物照片搜寻引擎中 小猪笼草的照片 date=20110519123112 。