Species of pitcher plant from Southeast Asia From Wikipedia, the free encyclopedia
Nepenthes rafflesiana (/nɪˈpɛnθiːzræˌfliːziˈɑːnə/; after Stamford Raffles), or Raffles' pitcher-plant,[3] is a species of tropical pitcher plant. It has a very wide distribution covering Borneo, Sumatra, Peninsular Malaysia, and Singapore. Nepenthes rafflesiana is extremely variable, with numerous forms and varieties described. In Borneo alone, there are at least three distinct varieties. The giant form of this species produces enormous pitchers rivalling those of N. rajah in size.
Nepenthes rafflesiana generally occurs in open, sandy, wet areas. It has been recorded from kerangas forest, secondary formations, margins of peat swamp forest, heath forest, and seaside cliffs. It grows at elevations ranging from sea-level to 1200m[4] or even 1500m.[8]
Nepenthes rafflesiana is a scrambling vine. The stem may climb to a height of 15 m and is up to 10mm thick. Internodes are up to 20cm long.[4]Tendrils may be over 110cm long.
The lower pitchers of N. rafflesiana are bulbous and possess well-developed fringed wings. These terrestrial traps rarely exceed 20cm in height, although the giant form of N. rafflesiana is known produce pitchers up to 35cm long and 15cm wide. Upper pitchers are funnel-shaped and often bear a distinctive raised section at the front of the peristome. Both types of pitchers have a characteristically elongated peristome neck that may be 3cm or more in length.
Pitcher colouration varies greatly from dark purple to almost completely white. The typical form of N. rafflesiana is light green throughout with heavy purple blotches on the lower pitchers and cream-coloured aerial pitchers.
The inflorescence is a raceme and grows between 16 and 70cm tall. The red or purple flowers usually occur singly, or sometimes in pairs, on each flower-stalk.
Young plants are wholly covered with long, caducous, brown or white hairs. Mature plants often have a sparse indumentum of short, brown hairs, though they may be completely glabrous.
Nepenthes rafflesiana is found in tropical lowlands. It produces two distinct types of pitchers (heavily modified leaves), which are used to capture and kill insect prey for nutrients. The lower pitchers are generally round, squat and 'winged', while the upper pitchers are more narrow at their base. The species is widely variable and comes in a variety of shapes and colors– most contain varying amounts of green, white, and maroon streaks.
All Nepenthes are passive carnivores with no moving parts, unlike their distant cousins the Venus flytrap. Nepenthes rafflesiana kills by luring its prey into its pitchers, whose peristomes secrete a sweet-tasting nectar. Once the insect is inside, it quickly finds the walls of the pitcher too slippery to scale and drowns. Digestive enzymes released by the plant into the liquid break down the prey and release soluble nutrients, which are absorbed by the plant through the walls of the pitcher. The carnivorous nature of Nepenthes is supposedly a consequence of living in nutrient-poor soils; since the main method of nutrient absorption in most plants (the root) is insufficient in these soils, the plants have evolved other ways to gain nutrients. As a result, the roots of Nepenthes and most other carnivorous plants are slight and fragile; hence care must be taken when repotting. All Nepenthes are dioecious, meaning that each individual plant has only male or female characteristics.
Nepenthes rafflesiana was discovered by Dr. William Jack in 1819. In a letter from Singapore published in Curtis's Botanical Magazine, Jack wrote the following account:[3]
It is impossible to conceive anything more beautiful than the approach to Singapore, through the Archipelago of islands that lie at the extremity of the Straits of Malacca. Seas of glass wind among innumerable islets, clothed in all the luxuriance of tropical vegetation and basking in the full brilliance of a tropical sky... I have just arrived in time to explore the woods before they yield to the axe, and have made many interesting discoveries, particularly of two new and splendid species of pitcher-plant [Nepenthes rafflesiana and Nepenthes ampullaria], far surpassing any yet known in Europe. I have completed two perfect drawings of them with ample descriptions. Sir S. Raffles is anxious that we should give publicity to our researches in one way or other and has planned bringing out something at Bencoolen. He proposes sending home these pitcher-plants that such splendid things may appear under all the advantages of elegant execution, by way of attracting attention to the subject of Sumatran botany.
At the time the largest known species in the genus, N. rafflesiana was described in the Gardener's and Farmer's Journal for 1850 as follows:[3]
Whoever has seen this plant in a living state must undoubted be constrained to consider it as one of the most astonishing productions of the whole vegetable kingdom. The resemblance that a portion of it bears to our more familiar domestic utensils leaves a lasting impression on the minds of spectators that is not easily eradicated; it is the largest and most magnificent of the genus, far surpassing any hitherto known in Europe.
Nepenthes rafflesiana is very popular in cultivation; it is a lowland Nepenthes (enjoying hot, humid conditions most of the time, as found in tropical jungle lowlands) but can be grown as an intermediate, with cooler nights and less humidity. It is a comparatively hardy Nepenthes that is commonly recommended as a "first plant" to new Nepenthes growers. The plant should be grown in shaded conditions, diffuse sunlight, or in a large grow chamber under artificial lights. Watering and misting should be performed frequently, and preferably with distilled water, to avoid mineral build-up that is not only unsightly but that may damage the delicate roots of Nepenthes (and most other carnivorous plants). Standing water is inadvisable. A wet, well-draining potting medium is a necessity. Methods of feeding are varied– some growers feed freeze-dried bloodworms or Koi pellets (both available in the fish section of most pet stores); others prefer orchid mixes. No carnivorous plant should ever be fed mammalian meat– this will result not only in an unpleasant smell but also the probable rotting of the pitcher and potential death of the plant. The digestive enzymes present have not evolved to handle large prey items, and the rotting material gives opportunistic bacteria and fungi a chance to take hold.
Left:Nepenthes rafflesiana var. insignis, an apparently "large-pitchered variety with unusually dark purple markings",[3] as illustrated in The Gardeners' Chronicle, 1882 Right: A lower pitcher of N.hemsleyana, previously known under a variety of names including N.rafflesiana var. elongata and N.baramensis
Across its expansive range, N. rafflesiana exhibits great variability in both pitcher morphology and colour. The following infraspecific taxa of N. rafflesiana have appeared in the literature. Most of these are not considered valid today, and a number represent different taxa altogether. The elongate plant often referred to informally as N.rafflesiana var. elongata, and described as N.baramensis,[9] is now known under the name N.hemsleyana.[10]
Nepenthes rafflesiana f. alba Hort.Westphal (2000) nom.nud.
N. rafflesiana lower pitcher with a winged tendril
Giant form of N. rafflesiana with wavy laminar margins
Lower traps of a light pitchered variant
Giant form
Vegetative parts of the giant form (from left to right): a lower pitcher measuring over 30cm; a tendril measuring over 110cm; and an upper pitcher measuring around 45cm.
Giant plants of N.rafflesiana have been recorded from a number of isolated localities on the northwestern coast of Borneo and one population has been found near the seaside town of Sematan, around 110km west of Kuching.[16] The typical habitat of this form is dense heath forest, especially around vegetation boundaries.[16]
The giant form is a much larger plant than the typical form in all respects. The stem may climb to a height of 15m. Leaf blades are around two and a half times as long as usual. Lower pitchers reach 35cm in height by 15cm in width and sometimes exceed 1 litre in volume, making them some of the largest in the genus.[16] They vary widely in pigmentation, from white with red blotches to dark purple. Upper pitchers may be spotted or green throughout. The inflorescence is also massive, reaching over 1m in length. The individual flowers measure up to 1.5cm in diameter and have dark red tepals.[16]
In addition to its size, the giant form is distinguished by the colour of its developing leaves, which have a bronze sheen. Both this characteristic and the plant's exceptional size are exhibited by cultivated specimens and thus they cannot be due to unusual environmental factors.[16]
The following natural hybrids involving N. rafflesiana have been recorded.
Most wild populations of Nepenthes, including N. rafflesiana, are endangered due to habitat destruction and (to a lesser extent) poaching. N. rafflesiana is currently listed as a CITES Appendix II plant, so it does have some international trade restrictions (though not an outright ban). Today, most N. rafflesiana plants on the market are propagated by plant tissue culture or other forms of vegetative propagation. When purchasing any plant, especially those protected by CITES, it is important to ask the vendor about the plant's provenance.
(in Indonesian) Mansur, M. 2012. Keanekaragaman jenis tumbuhan pemakan serangga dan laju fotosintesisnya di Pulau Natuna. [Diversity on insectivorous plants and its photosynthetic rate in Natuna Island.] Berita Biologi11(1): 33–42. AbstractArchived 2013-09-21 at the Wayback Machine
Clarke, C., J.A. Moran & C.C. Lee 2011. Nepenthes baramensis (Nepenthaceae)– a new species from north-western Borneo. Blumea56(3): 229–233. doi:10.3767/000651911X607121
Scharmann, M. & T.U. Grafe 2013. Reinstatement of Nepenthes hemsleyana (Nepenthaceae), an endemic pitcher plant from Borneo, with a discussion of associated Nepenthes taxa. Blumea58(1): 8–12. doi:10.3767/000651913X668465
(in French) Teysmann, M.J.E. 1859. Énumération des plantes envoyées de Java au jardin botanique de l'Université de Leide.Annales d'horticulture et de botanique, ou Flore des jardins du royaume des Pays-Bas, et histoire des plantes cultivées les plus intéressantes des possessions néerlandaises aux Indes orientales, de l'Amérique et du Japon2: 133–142.
Yulita, K.S. & M. Mansur 2012. The occurrence of hybrid in Nepenthes hookeriana Lindl. from Central Kalimantan can be detected by RAPD and ISSR markers. HAYATI Journal of Biosciences19(1): 18–24. doi:10.4308/hjb.19.1.18
(in Malay) 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.
Adams, R.M. & G.W. Smith 1977. An S.E.M. survey of the five carnivorous pitcher plant genera. American Journal of Botany64(3): 265–272. doi:10.2307/2441969
Bauer, U., C. Willmes & W. Federle 2009. Effect of pitcher age on trapping efficiency and natural prey capture in carnivorous Nepenthes rafflesiana plants. Annals of Botany103(8): 1219–1226. doi:10.1093/aob/mcp065
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 Biology25(1): 90–102. doi:10.1111/j.1420-9101.2011.02406.x
Bauer, U., B. Di Giusto, J. Skepper, T.U. Grafe & W. Federle 2012. With a flick of the lid: a novel trapping mechanism in Nepenthes gracilis pitcher plants. PLoS ONE7(6): e38951. doi:10.1371/journal.pone.0038951
Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.
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 Interactions6(1): 147–158. doi:10.1007/s11829-011-9171-2
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 Phytologist191(2): 545–554. doi:10.1111/j.1469-8137.2011.03696.x
Brearley, F.Q. & M. Mansur 2012. Nutrient stoichiometry of Nepenthes species from a Bornean peat swamp forest. Carnivorous Plant Newsletter41(3): 105–108.
Cannon, J., V. Lojanapiwatna, C. Raston, W. Sinchai & A. White 1980. The Quinones of Nepenthes rafflesiana. The Crystal Structure of 2,5-Dihydroxy-3,8-dimethoxy-7-methylnaphtho-1,4-quinone (Nepenthone-E) and a Synthesis of 2,5-Dihydroxy-3-Methoxy-7-methylnaphtho-1,4-quinone (Nepenthone-C). Australian Journal of Chemistry33(5): 1073–1093. doi:10.1071/CH9801073
Chung, A.Y.C. 2006. Biodiversity and Conservation of The Meliau Range: A Rain Forest in Sabah's Ultramafic Belt. Natural History Publications (Borneo), Kota Kinabalu. ISBN9838121169.
Di Giusto, B., V. Grosbois, E. Fargeas, D.J. Marshall & L. Gaume 2008. Contribution of pitcher fragrance and fluid viscosity to high prey diversity in a Nepenthes carnivorous plant from Borneo. Journal of Biosciences33(1): 121–136. doi:10.1007/s12038-008-0028-5
Di Giusto, B., J.-M. Bessière, M. Guéroult, L.B.L. Lim, D.J. Marshall, M. Hossaert-McKey & L. Gaume 2010. Flower-scent mimicry masks a deadly trap in the carnivorous plant Nepenthes rafflesiana. Journal of Ecology98(4): 845–856. doi:10.1111/j.1365-2745.2010.01665.x
Dixon, W.E. 1889. Nepenthes.The Gardeners' Chronicle, series 3, 6(144): 354.
Gaume, L. & Y. Forterre 2007. A viscoelastic deadly fluid in carnivorous pitcher plants. PLoS ONE2(11): e1185. doi:10.1371/journal.pone.0001185
(in French) Gaume, L. & Y. Forterre 2008. Un piège viscoélastique chez Nepenthes rafflesiana. Dionée71: 28–32.
Gaume, L. & B. Di Giusto 2009. Adaptive significance and ontogenetic variability of the waxy zone in Nepenthes rafflesiana. Annals of Botany104(7): 1281–1291. doi:10.1093/aob/mcp238
(in Indonesian) Handayani, T. & Syamsuddin 1998. Nepenthes rafflesiana Jack. dan keturunannya. Warta Kebun Raya2(3): 1–8.
(in Indonesian) Handayani, T. 1999. "Konservasi Nepenthes di kebun raya Indonesia"(PDF). [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.
Korthals, P.W. 1839. Over het geslacht Nepenthes. In: C.J. Temminck 1839–1842. Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche overzeesche bezittingen; Kruidkunde. Leiden. pp.1–44, t. 1–4, 13–15, 20–22.
Kurup, R., A.J. Johnson, S. Sankar, A.A. Hussain, C.S. Kumar & S. Baby 2013. Fluorescent prey traps in carnivorous plants. Plant Biology15(3): 611–615. doi:10.1111/j.1438-8677.2012.00709.x
Lecoufle, M. 1990. Nepenthes rafflesiana. In: Carnivorous Plants: Care and Cultivation. Blandford, London. pp.136–137.
Lee, C.C. 2000. Recent Nepenthes Discoveries. [video] The 3rd Conference of the International Carnivorous Plant Society, San Francisco, USA.
Lim, S.H., D.C.Y. Phua & H.T.W. Tan 2000. Primer design and optimization for RAPD analysis of Nepenthes. Biologia Plantarum43(1): 153–155. doi:10.1023/A:1026535920714
(in Indonesian) Mansur, M. 2007. Keanekaragaman jenis Nepenthes (kantong semar) dataran rendah di Kalimantan Tengah. [Diversity of lowland Nepenthes (kantong semar) in Central Kalimantan.] Berita Biologi8(5): 335–341. Abstract
Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. Plant Biology3(2): 164–175. doi:10.1055/s-2001-12897
Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. Plant Biology8(6): 831–840. doi: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 Evolution39(2): 478–490. doi:10.1016/j.ympev.2005.11.023
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 America90(2): 177–183.
Moran, J.A. 1991. The role and mechanism of Nepenthes rafflesiana pitchers as insect traps in Brunei. Ph.D. thesis, University of Aberdeen, Aberdeen, Scotland.
Moran, J.A. 1993. Visitors to the flowers of the pitcher plant Nepenthes rafflesiana. Brunei Museum Journal8: 73–75.
Moran, J.A. 1993. Pitcher allocation strategy of the pitcher plant Nepenthes rafflesiana. Brunei Museum Journal8: 77–80.
Moran, J.A. 1993. The effect of pitcher wing removal on prey capture by the pitcher plant Nepenthes rafflesiana. Brunei Museum Journal8: 81–82.
Moran, J.A. 1993. Misumenops nepenthicola: the top aquatic predator of the Nepenthes food web? Brunei Museum Journal8: 83–84.
Moran, J.A. 1996. Pitcher dimorphism, prey composition and the mechanisms of prey attraction in the pitcher plant Nepenthes rafflesiana in Borneo. Journal of Ecology84(4): 515–525. doi:10.2307/2261474
Moran, J.A. & A.J. Moran 1998. Foliar Reflectance and Vector Analysis Reveal Nutrient Stress in Prey-Deprived Pitcher Plants (Nepenthes rafflesiana). International Journal of Plant Sciences159(6): 996–1001. doi:10.1086/314086
Moran, J.A., B.J. Hawkins, B.E. Gowen & S.L. Robbins 2010. Ion fluxes across the pitcher walls of three Bornean Nepenthes pitcher plant species: flux rates and gland distribution patterns reflect nitrogen sequestration strategies. Journal of Experimental Botany61(5): 1365–1374. doi:10.1093/jxb/erq004
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 Botany102(5): 845–853. doi:10.1093/aob/mcn162
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 Sciences172(7): 889–901. doi:10.1086/660882
Rizzacasa, M.A. & M.V. Sargent 1987. The structure and synthesis of nepenthone-A, a naphthoquinone from Nepenthes rafflesiana. Journal of the Chemical Society, Perkin Transactions 1: 2017–2022. doi:10.1039/P19870002017
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 Botany62(13): 4639–4647. doi:10.1093/jxb/err173
Ruxton, G.D. & H.M. Schaefer 2011. Alternative explanations for apparent mimicry. Journal of Ecology99(4): 899–904. doi:10.1111/j.1365-2745.2011.01806.x
Smythies, B.E. 1965. The distribution and ecology of pitcher-plants (Nepenthes) in Sarawak. UNESCO Humid Tropics Symposium, June–July 1963, Kuching, Sarawak.