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Nepenthes ampullaria

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Nepenthes ampullaria
N. ampullaria from Sarawak.
Scientific classification
Kingdom:
Plantae
Division:
Magnoliophyta
Class:
Magnoliopsida
Order:
Caryophyllales
Family:
Nepenthaceae
Genus:
Nepenthes
Species:
N. ampullaria
Binomial name
Nepenthes ampullaria
Jack (1835)
Synonyms

Nepenthes ampullaria (Latin: ampulla = a flask-like bladder) is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, 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

N. ampullaria generally grows in damp, shady forest from sea-level to 2100 m altitude.[1] 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.[2]

In Sumatra and Peninsular Malaysia, it grows from sea-level to 1100 m altitude on flat terrain in heath forest, padang, belukar, peat swamp forest, degraded swamp forest, and in padi fields.[3]

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.[1]

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:[4]

"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.[2]

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.[2][3]

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.[3]

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.[2]

Carnivory

See also: Protocarnivorous plant
N. ampullaria is well-adapted to capture leaf litter.

N. ampullaria has largely moved away from carnivory and acquires most of its nutrients from digesting leaf matter that falls to the forest floor. It could thus be called a detritivore.

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

  • It is the only species in the genus to lack "lunate" cells in its pitchers.[5] These are modified stomatal guard cells which, it is thought, deny prey a foothold in the pitcher.[4]
  • The pitcher lid is atypical, being very small and reflexed, such that leaf litter is allowed to fall directly into the pitcher.[3]
  • Nectar glands, which play an important role in prey capture, are very rare and in some cases completely absent from the pitcher lid.[3]
  • The marginal glands of the peristome are greatly reduced compared to those of other species.[3]
  • In terrestrial pitchers, the glandular region extends almost to the peristome, such that there is little or no conductive waxy zone.[6][7][2] The waxy zone functions by causing prey to slip and fall into the digestive fluid.[3]
  • 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.[3]
  • The pitchers of N. ampullaria are relatively long-lived, as the species relies on a slow accumulation of nutrients over time.[3]
  • 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.[3]

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.[8]

Forms and varieties

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.[9]

  • N. ampullaria var. geelvinkiana Becc. (1886)
  • N. ampullaria var. guttata D.Moore (1872)
  • N. ampullaria var. longicarpa Becc. (1886)
  • N. ampullaria var. microsepala Macf. (1911)
  • N. ampullaria var. papuana Becc. in sched. nom.nud.
  • N. ampullaria var. picta Hort. ex Nichols. (1885)
  • N. ampullaria var. racemosa 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.[3][2]

  • Nepenthes × trichocarpa
    Nepenthes × trichocarpa
  • Nepenthes × hookeriana
    Nepenthes × hookeriana

References

Wikimedia Commons has media related to Nepenthes ampullaria.
  1. ^ a b Jebb, M. & M. Cheek 1997. A Skeletal Revision of Nepenthes (Nepenthaceae). Blumea 42: 1-106.
  2. ^ a b c d e f Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications, Kota Kinabalu, pp. 65-68.
  3. ^ a b c d e f g h i j k Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications, Kota Kinabalu, pp. 59-60.
  4. ^ a b Lloyd, F.E. 1942. The Carnivorous Plants. Chronica Botanica 9. Ronald Press Company, New York, U.S.A. xvi + 352 pp.
  5. ^ Pant, D.D. & S. Bhatnagar 1977. Morphological studies in Nepenthes (Nepenthaceae). Phytomorphology 27: 13-34.
  6. ^ Macfarlane, J.M. 1893. Observations on pitchered insectivorous plants. Part II. Annals of Botany 7: 403-458.
  7. ^ Jebb, M.H.P. 1991. An account of Nepenthes in New Guinea. Science in New Guinea 17(1): 7-54.
  8. ^ 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: 635–639.
  9. ^ Danser, B.H. 1928. The Nepenthaceae of the Netherlands Indies. Bulletin de Jardin de Botanique, Buitenzorg, Série III, 9(3-4): 249-438.
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