Nepenthes ampullaria

Nepenthes ampullaria
Rosette pitcher of Nepenthes ampullaria from Bako National Park, Borneo
Conservation status
Least Concern (IUCN 2.3)
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Eudicots
(unranked):	Core eudicots
Order:	Caryophyllales
Family:	Nepenthaceae
Genus:	Nepenthes
Species:	N. ampullaria
Binomial name
Nepenthes ampullaria Jack (1835)
Synonyms
Nepenthes ampullacea Low (1848) sphalm.typogr. Nepenthes ampullaria auct. non Jack: Jeanneney (1894) [=N. vieillardii]

Nepenthes ampullaria (pronounced /nɨˈpɛnθiːz ˌæmpəˈlɛəriə/, Latin: ampulla "flask"), the Flask-Shaped Pitcher-Plant,^[1] is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea. It is not generally considered to be closely related to any other species in the genus and is thus often used as the 'outlier' species for cladistic analyses.

Habitat

Nepenthes ampullaria generally grows in damp, shady forest from sea-level to 2100 m altitude.^[2] In Borneo, it occurs usually on relatively flat terrain in kerangas forest, peat swamp forest, and degraded swamp forest, at elevations of 0 to 1000 m.^[3]

In Sumatra and Peninsular Malaysia, it grows from sea-level to 1100 m altitude on flat terrain in heath forest, padang (meaning "field" in Malay), belukar, peat swamp forest, degraded swamp forest, and in padi fields.^[4]

In New Guinea, it is predominantly present in Araucaria forests. The species has also been recorded from secondary forests, open microphyllous vegetation, and swamp grassland.^[2]

Description

N. ampullaria with climbing stems and rosette pitchers.

Due to its unique pitcher morphology and unusual growth habit, it is difficult to confuse N. ampullaria with any other species in the genus. F. E. Lloyd translated Troll's 1932 account of this species as follows:^[5]

"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."

The stem of N. ampullaria is light brown in colour and may climb to 15 m in height. Leaves are light green, up to 25 cm long, and 6 cm wide. Pitchers are produced at the ends of short tendrils no more than 15 cm long.^[3]

The urceolate pitchers are generally small, rarely exceeding 10 cm in height and 7 cm in width. Upper pitchers are very rarely produced and are considerably smaller than those formed on rosettes or offshoots. Pitchers range in colouration from light green throughout to completely dark red, with many intermediate forms recorded. The pitchers of N. ampullaria from Sumatra and Peninsular Malaysia are almost exclusively green throughout or green with red speckles; the red forms are mostly confined to Borneo. A large-pitchered form has been recorded from New Guinea.^[3][4]

The inflorescence of N. ampullaria is a dense panicle. It is the only Nepenthes species recorded from Sumatra or Peninsular Malaysia that produces paniculate inflorescences.^[4]

All parts of the plant are densely covered with short, brown hairs when young. The indumentum of mature plants is more sparse, except on the inflorescenes.^[3]

Carnivory

See also: Protocarnivorous plant

N. ampullaria is well-adapted to capturing leaf litter.

Nepenthes ampullaria has largely moved away from carnivory and acquires a substantial portion of its nutrients from digesting leaf matter that falls to the forest floor. It is thus partially detritivorous.

The species has developed several unique traits as a consequence of its adaptation to trapping leaf litter:

• It is one of the few species in the genus to lack "lunate" cells in its pitchers.^[6][7] These are modified stomatal guard cells which, it is thought, deny prey a foothold in the pitcher.^[5]
• The pitcher lid is atypical, being very small and reflexed, such that leaf litter is allowed to fall directly into the pitcher.^[4]
• Nectar glands, which play an important role in prey capture, are very rare and in some cases completely absent from the pitcher lid.^[4]
• The marginal glands of the peristome are greatly reduced compared to those of other species.^[4]
• In terrestrial pitchers, the glandular region extends almost to the peristome, such that there is little or no conductive waxy zone.^{[3][7][8][9][10]} The waxy zone functions by causing prey to slip and fall into the digestive fluid.^[4]
• The plant's architecture, consisting of subsurface runners and offshoots, is unusual for the genus. The species often forms a "carpet" of pitchers covering the soil. This serves to maximise the area over which falling debris may be intercepted.^[4]
• The pitchers of N. ampullaria are relatively long-lived, as the species relies on a slow accumulation of nutrients over time.^[4]
• It is thought that infaunal organisms, such as mosquito larvae, facilitate breakdown of leaf litter and aid in the transfer of nitrogen from it to the plant by means of the excretion of ammonium ions. Bacterial breakdown of leaf matter is also known to produce ammonium ions.^[4]

It has been shown that foliar stable nitrogen isotope (¹⁵N) abundance in N. ampullaria plants growing under forest canopy (litterfall present) is significantly lower than in plants without access to litterfall. Conversely, total nitrogen concentrations are higher in these plants compared to those growing in open sites with no litterfall. It has been estimated that N. ampullaria plants growing under forest canopy derive 35.7% (±0.1%) of their foliar nitrogen from leaf litter.^[11]

Ecology

Fifty-nine infaunal species have been recorded from the pitchers of N. ampullaria.^[12] Among these are the smallest known species of Old World frog, Microhyla nepenthicola, and the crab spider, Misumenops nepenthicola.^[13][14]

Infraspecific taxa

A red form of N. ampullaria

The most recently described variety, N. ampullaria var. racemosa, occurs in Sarawak and has a racemose inflorescence. B. H. Danser considered the other varieties to be unimportant.^[15]

• N. ampullaria var. geelvinkiana Becc. (1886)
• N. ampullaria var. guttata D.Moore (1872)
• N. ampullaria var. longicarpa Becc. (1886)
• N. ampullaria var. microsepala Macfarl. (1911)
• N. ampullaria var. papuana Becc. in sched. nom.nud.
• N. ampullaria var. picta Hort. ex Nichols. (1885)
• N. ampullaria var. racemosa J.H.Adam & Wilcock (1990)
• N. ampullaria var. vittata Hort. ex G.Beck (1895)
• N. ampullaria var. vittata-major Mast. (1872)

Natural hybrids

N. ampullaria flowers once or twice annually for several weeks at a time. Its flowering period often coincides with those of other Nepenthes species; consequently, it readily forms natural hybrids. The following natural hybrids involving N. ampullaria have been recorded.

• N. albomarginata × N. ampullaria^[3]
• N. ampullaria × N. bicalcarata^[3]
• N. ampullaria × N. eustachya^[16]
• N. ampullaria × N. gracilis [=N. × trichocarpa]^[3]
• (N. ampullaria × N. gracilis) × N. bicalcarata [=N. × trichocarpa × N. bicalcarata]
• N. ampullaria × N. hirsuta^[3]
• N. ampullaria × N. mirabilis [=N. × kuchingensis, Nepenthes cutinensis]^[3]
• N. ampullaria × N. neoguineensis^[16]
• N. ampullaria × N. rafflesiana [=N. × hookeriana]^[3]
• ? (N. ampullaria × N. rafflesiana) × N. mirabilis [=N. × hookeriana × N. mirabilis]^[17]
• N. ampullaria × N. reinwardtiana^[4]
• N. ampullaria × N. tobaica^[4]

• 

  ? N. ampullaria × N. eustachya

• 

  Nepenthes × trichocarpa

• 

  N. ampullaria × N. neoguineensis

• 

  Nepenthes × hookeriana

References

1. ^ Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
2. ^ ^{a b} Jebb, M.H.P. & M.R. Cheek 1997. A skeletal revision of Nepenthes (Nepenthaceae). Blumea 42(1): 1–106.
3. ^ ^{a b c d e f g h i j k} Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
4. ^ ^{a b c d e f g h i j k l} Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
5. ^ ^{a b} Lloyd, F.E. 1942. The Carnivorous Plants. Chronica Botanica 9. Ronald Press Company, New York, U.S.A. xvi + 352 pp.
6. ^ Pant, D.D. & S. Bhatnagar 1977. Morphological studies in Nepenthes (Nepenthaceae). Phytomorphology 27: 13–34.
7. ^ ^{a b} 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 Botany 61(5): 1365–1374. doi:10.1093/jxb/erq004
8. ^ Macfarlane, J.M. 1893. Observations on pitchered insectivorous plants. (Part II.). Annals of Botany 7(4): 403–458.
9. ^ Jebb, M.H.P. 1991. An account of Nepenthes in New Guinea. Science in New Guinea 17(1): 7–54.
10. ^ Di Giusto, B., M. Guéroult, N. Rowe & L. Gaume 2009. Chapter 7: The Waxy Surface in Nepenthes Pitcher Plants: Variability, Adaptive Significance and Developmental Evolution. In: S.N. Gorb (ed.) Functional Surfaces in Biology: Adhesion Related Phenomena. Volume 2. Springer. pp. 183–204.
11. ^ Moran, J.A., C.M. Clarke & B.J. Hawkins 2003. From carnivore to detritivore? Isotopic evidence for leaf litter utilization by the tropical pitcher plant Nepenthes ampullaria. International Journal of Plant Sciences 164(4): 635–639. doi:10.1086/375422
12. ^ Adlassnig, W., M. Peroutka & T. Lendl 2011. Traps of carnivorous pitcher plants as a habitat: composition of the fluid, biodiversity and mutualistic activities. Annals of Botany 107(2): 181–194. doi:10.1093/aob/mcq238
13. ^ Das, I. & A. Haas. 2010. New species of Microhyla from Sarawak: Old World’s smallest frogs crawl out of miniature pitcher plants on Borneo (Amphibia: Anura: Microhylidae).PDF Zootaxa 2571: 37–52.
14. ^ Gururaja, K.V. 2010. Old World’s smallest frog discovered residing in killer plants.PDF Current Science 99(8): 1000.
15. ^ Danser, B.H. 1928. The Nepenthaceae of the Netherlands Indies. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.
16. ^ ^{a b} McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
17. ^ Lowrie, A. 1983. Sabah Nepenthes Expeditions 1982 & 1983.PDF (1.25 MiB) Carnivorous Plant Newsletter 12(4): 88–95.

Further reading

• Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes.PDF Journal of Tropical Forest Science 5(1): 13–25.
• Adam, J.H. 1997. Prey spectra of Bornean Nepenthes species (Nepenthaceae) in relation to their habitat.PDF Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.
• Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae).PDF Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.
• Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.
• (French) Besnard, J. 1991. Nepenthes ampullaria "Cantley's red". Dionée 23.
• 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. doi:10.1111/j.1469-8137.2011.03696.x
• Bourke, G. 2011. The Nepenthes of Mulu National Park. Carniflora Australis 8(1): 20–31.
• Clarke, C.M., R. Cantley, J. Nerz, H. Rischer & A. Witsuba (2000). Nepenthes ampullaria. 2006. IUCN Red List of Threatened Species. IUCN 2006. www.iucnredlist.org. Retrieved on 12 May 2006.
• Clarke, C.M. & J.A. Moran 1994. A further record of aerial pitchers in Nepenthes ampullaria Jack. Malayan Nature Journal 47: 321–323.
• Cresswell, J.E. 1998. Morphological correlates of necromass accumulation in the traps of an Eastern tropical pitcher plant, Nepenthes ampullaria Jack, and observations on the pitcher infauna and its reconstitution following experimental removal. Oecologia 113(3): 383–390. doi:10.1007/s004420050390
• Fashing, N.J. 2010. Two novel adaptations for dispersal in the mite family Histiostomatidae (Astigmata).PDF In: M.W. Sabelis & J. Bruin (eds.) Trends in Acarology: Proceedings of the 12th International Congress. Springer Science, Dordrecht. pp. 81–84. doi:10.1007/978-90-481-9837-5
• Green, T.L. & S. Green 1964. Stem pitchers on Nepenthes ampullaria. Malayan Nature Journal 18: 209–211.
• (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.
• (Indonesian) Handayani, T. & R. Hendrian 1999. Strategi konservasi Nepenthes ampullaria Jack.PDF [Conservation strategy of Nepenthes ampullaria Jack.] In: Workshop & Promosi Flora Kawasan Timur Indonesia. Kebun Raya Ekakarya Bali, Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 1–6.
• Handayani, T., D. Latifah & Dodo 2005. Diversity and growth behaviour of Nepenthes (pitcher plants) in Tanjung Puting National Park, Central Kalimantan Province.PDF Biodiversitas 6(4): 251–255.
• Hansen, E. 2001. Where rocks sing, ants swim, and plants eat animals: finding members of the Nepenthes carnivorous plant family in Borneo. Discover 22(10): 60–68.
• Hernawati & P. Akhriadi 2006. A Field Guide to the Nepenthes of Sumatra. PILI-NGO Movement, Bogor.
• Hirst, S. 1928. A new tyroglyphid mite (Zwickia nepenthesiana sp. n.) from the pitchers of Nepenthes ampullaria. Journal of the Malayan Branch of the British Royal Asiatic Society 6: 19–22.
• (Czech) Koudela, I. 1998. Klíčí nebo neklíčí. Trifid 1998(2): 36–37. (page 2)
• 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.
• (Indonesian) Mansur, M. 2001. Koleksi Nepenthes di Herbarium Bogoriense: prospeknya sebagai tanaman hias.PDF In: Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253.
• 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.
• 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.PDF Annals of Botany 83: 521–528.
• Rice, B. 2007. Carnivorous plants with hybrid trapping strategies. Carnivorous Plant Newsletter 36(1): 23–27.
• Shivas, R.G. 1984. Pitcher Plants of Peninsular Malaysia & Singapore. Maruzen Asia, Kuala Lumpur.
• Steffan, W.A. & N.L. Evenhuis 1982. A new species of Toxorhynchites from Papua New Guinea (Diptera: Culicidae).PDF Mosquito Systematics 14(1): 1–13.
• Tan, W.K. & C.L. Wong 1996. Aerial pitchers of Nepenthes ampullaria. Nature Malaysiana 21(1): 12–14.
• 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.
• Wong, D. 1996. Thoughts, reflections, and upper Nepenthes ampullaria pitcher.PDF Carnivorous Plant Newsletter 25(1): 10–14.
• (Indonesian) Yogiara 2004. Analisis komunitas bakteri cairan kantung semar (Nepenthes spp.) menggunakan teknik terminal restriction fragment length polymorphism (T-RFLP) dan amplified ribosomul DNA restriction analysis (ARDRA).PDF M.Sc. thesis, Bogor Agricultural University, Bogor.
• Yogiara, A. Suwanto & M.T. Suhartono 2006. A complex bacterial community living in pitcher plant fluid. Jurnal Mikrobiologi Indonesia 11(1): 9–14.

External links

• Photographs of N. ampullaria at the Carnivorous Plant Photofinder

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Nepenthes (list)

Species

N. adnata N. alata N. alba N. albomarginata N. ampullaria N. andamana N. angasanensis N. argentii N. aristolochioides N. attenboroughii N. beccariana N. bellii N. benstonei N. bicalcarata N. bokorensis N. bongso N. boschiana N. burbidgeae N. burkei N. campanulata N. ceciliae N. chang

N. chaniana N. clipeata N. copelandii N. danseri N. deaniana N. densiflora N. diatas N. distillatoria N. dubia N. edwardsiana N. ephippiata N. eustachya N. eymae N. faizaliana N. flava N. fusca N. gantungensis N. glabrata N. glandulifera N. gracilis N. gracillima N. gymnamphora

N. hamata N. hamiguitanensis N. hirsuta N. hispida N. holdenii N. hurrelliana N. inermis N. insignis N. izumiae N. jacquelineae N. jamban N. junghuhnii N. kampotiana N. kerrii N. khasiana N. klossii N. kongkandana N. lamii N. lavicola N. leonardoi N. lingulata N. longifolia

N. lowii N. macfarlanei N. macrophylla N. macrovulgaris N. madagascariensis N. mantalingajanensis N. mapuluensis N. masoalensis N. maxima N. merrilliana N. micramphora N. mikei N. mindanaoensis N. mira N. mirabilis N. mollis N. muluensis N. murudensis N. naga N. neoguineensis N. northiana N. ovata

N. palawanensis N. paniculata N. papuana N. peltata N. pervillei N. petiolata N. philippinensis N. pilosa N. pitopangii N. platychila N. pulchra N. rafflesiana N. rajah N. ramispina N. reinwardtiana N. rhombicaulis N. rigidifolia N. rowanae N. sanguinea N. saranganiensis N. sibuyanensis N. singalana

N. smilesii N. spathulata N. spectabilis N. stenophylla N. sumatrana N. suratensis N. surigaoensis N. talangensis N. tenax N. tentaculata N. tenuis N. thai N. thorelii N. tobaica N. tomoriana N. treubiana N. truncata N. veitchii N. ventricosa N. vieillardii N. villosa N. vogelii

Incompletely diagnosed taxa: N. sp. Misool • N. sp. Papua • N. sp. Sulawesi
Possible extinct species: N. echinatus • N. echinosporus • N. major

Natural hybrids

N. × alisaputrana N. × bauensis N. × cantleyi N. × cincta

N. × ferrugineomarginata N. × harryana N. × hookeriana N. × kinabaluensis

N. × kuchingensis N. × merrilliata N. × mirabilata N. × pangulubauensis

N. × pyriformis N. × sarawakiensis N. × sharifah-hapsahii N. × trichocarpa

N. × truncalata N. × trusmadiensis N. × tsangoya N. × ventrata

Related topics

Nepenthes taxonomy • Nepenthes infauna • Nepenthes species by distribution • Nepenthes cultivars • Nepenthes literature
Nepenthesin • Nepenthales