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Nepenthes albomarginata
	A pair of lower pitchers of N. albomarginata from Bako National Park, Borneo, where it often grows in heath forest and scrub vegetation
Conservation status
	; Least Concern (IUCN 3.1)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Order:	Caryophyllales
Family:	Nepenthaceae
Genus:	Nepenthes
Species:	N. albomarginata
Binomial name
	Nepenthes albomarginata; T.Lobb ex Lindl. (1849)
Synonyms
	Synonyms Nepenthes albocincta; Hort. ex Macfarl. (1908) nom.nud. ; Nepenthes albocincta var. rubra; Hort. ex Macfarl. (1908) ; Nepenthes laevis; auct. non Lindl.: C.Morren (1852) ; Nepenthes teysmanniana; Miq. (1858); [=N. albomarginata/N. gracilis] ; Nepenthes tomentella; Miq. (1855) ; Nepenthes tupmanniana; Bonst. (1931) sphalm.typogr. ;

Nepenthes albomarginata /nɪˈpɛnθiːz ˌælboʊmɑːrdʒɪˈnɑːtə/ is a tropical pitcher plant native to Borneo, Peninsular Malaysia, and Sumatra.^[4]^[5]

The specific epithet albomarginata, formed from the Latin words albus (white) and marginatus (margin), refers to the white band of trichomes that is characteristic of this species.^[4]

Botanical history

Nepenthes albomarginata was first collected by Thomas Lobb in 1848. It was formally described a year later by John Lindley in The Gardeners' Chronicle.^[3]^[6]

The species was introduced into cultivation in the United Kingdom in 1856.^[6]

In the 1996 book Pitcher-Plants of Borneo, N. albomarginata is given the vernacular name white-collared pitcher-plant.^[6] This name, along with all others, was dropped from the much-expanded second edition, published in 2008.^[7]

Description

Nepenthes albomarginata is a climbing plant. The stem may reach lengths of up to 4 metres (13 ft) and is up to 5 millimetres (0.20 in) in diameter. Internodes are cylindrical in cross section and up to 15 centimetres (5.9 in) long.^[8]

Upper pitchers of two colour forms from Bako National Park, Sarawak

Leaves are coriaceous in texture. The lamina or leaf blade is lanceolate in shape and up to 25 centimetres (9.8 in) long by 2 centimetres (0.79 in) wide. It has an acute apex and its base is gradually attenuate and amplexicaul. The leaves of this species are characteristic in that they completely lack a petiole. Longitudinal veins are indistinct. Tendrils are up to 20 centimetres (7.9 in) long.^[8]

Rosette and lower pitchers are bulbous in the basal third and cylindrical above. They are relatively small, reaching only 15 centimetres (5.9 in) in height by 4 centimetres (1.6 in) in width. A pair of fringed wings up to 5 millimetres (0.20 in) wide runs down the front of each pitcher. The pitcher mouth is round and rises to form a short neck at the rear. The peristome is cylindrical in cross section, up to 2 millimetres (0.079 in) wide, and bears indistinct teeth.^[8] The inner portion of the peristome accounts for around 34% of its total cross-sectional surface length.^[9] A dense band of short white trichomes is present directly below the peristome, although these may be missing from pitchers that have caught termites. The glandular region covers the bulbous portion of the pitcher's inner surface. The lid or operculum is suborbicular and lacks appendages. An unbranched spur (≤3 millimetres (0.12 in) long) is inserted near the base of the lid.^[8]

Upper pitchers are similar to their lower counterparts in most respects. They are cylindrical-infundibular throughout and have a pair of ribs in place of wings.^[8]

Nepenthes albomarginata has a racemose inflorescence that is usually longer in male plants. The peduncle is up to 25 centimetres (9.8 in) long, while the rachis reaches lengths of up to 40 centimetres (16 in). Partial peduncles are one- or two-flowered, up to 30 millimetres (1.2 in) long, and lack a bract. Sepals are obovate to oblong in shape and up to 4 millimetres (0.16 in) long.^[8] A study of 120 pollen samples taken from a herbarium specimen (J.H.Adam 2417, collected in Borneo at an altitude of 0–30 metres (0–98 ft)) found the mean pollen diameter to be 31.8 μm (0.00125 in) (SE = 0.4; CV = 6.2%).^[10]

Most parts of the plant are covered in a dense indumentum of very short, stellate white hairs. However, the underside of the lamina bears a dense covering of long hairs.^[8]

Ecology

Nepenthes albomarginata is a widespread species, occurring in Borneo, Peninsular Malaysia, and Sumatra. It is also found on smaller islands such as Nias and Penang.^[11]^[12] It has an altitudinal distribution of 0–1200 m above sea level.^[13]

Its typical habitat consists of kerangas forest, but it has also been recorded from the summit vegetation of lowland peaks.^[8] It is known from peat and limestone substrates.^[8]^[14]

Carnivory

A lower pitcher with an intact band of trichomes (left) and one lacking them (right)

Nepenthes albomarginata is notable for specializing in termites; most of the species in the genus Nepenthes are unselective about their prey. According to botanist Marlis A. Merbach and coworkers, this specialization to a single prey taxon is unique amongst carnivorous plants.^[15]^[16]^[17]^[18]^[19]

Nepenthes albomarginata has a unique morphological feature: a rim of living white trichomes directly below the peristome. The rim's hairs tend to be missing from pitchers that have caught termites. Merbach said "For several days, nothing would happen, then — after a single night — pitchers would fill with termites and their rim hairs would disappear."

Merbach investigated this phenomenon by placing fresh intact pitchers, together with pitchers with their white rims removed, near to the head of foraging columns of the termite Hospitalitermes bicolor.^[15] When the column found the pitcher, termites grazed on the rim.

While grazing, many termites (both workers and soldiers) fell into the pitchers. Once in the pitcher, they were unable to climb out. Merbach counted up to 22 individuals per minute falling into the pitchers and noted that the capture rate could easily exceed this for denser columns. After about an hour, the hairs were all gone and the pitcher was evidently no longer attractive to termites (and was filled with termites trying to escape).

It is not known how the trichomes lure termites to the plant. Merbach detected no long-range olfactory attraction during his experiments and noted that "all contacts seemed to happen by chance, with termites often missing pitchers less than 1 cm away from them."

Merbach also points out that N. albomarginata is the only plant species to offer its tissue as a bait.

Related species

In 2001, Clarke performed a cladistic analysis of the Nepenthes species of Sumatra and Peninsular Malaysia using 70 morphological characteristics of each taxon. The following is a portion of the resultant cladogram, showing "Clade 6", which is only weakly supported at 50%. The sister pair of N. angasanensis and N. mikei has 79% support.^[4]

Lower pitchers of N. adnata (left) and a purple form of N. albomarginata (right)

50%

	N. albomarginata

N. adnata

	N. gracilis

	N. reinwardtiana

N. tobaica

79%

	N. angasanensis

	N. mikei

Infraspecific taxa

N. albomarginata f. sanguinea Toyoda ex Hinode-Kadan (1985) nom.nud.
N. albomarginata var. rubra (Hort. ex Macfarl.) Macfarl. (1908)^[20]
N. albomarginata var. tomentella (Miq.) Beck (1895)^[21]
N. albomarginata var. typica Beck (1895)^[21] nom.illeg.
N. albomarginata var. villosa Hook.f. (1873)^[22]

Natural hybrids

N. albomarginata × N. ampullaria^[8]
? N. albomarginata × N. chaniana^[23]
N. albomarginata × N. clipeata^[8]
N. albomarginata × N. eustachya^[4]
N. albomarginata × N. gracilis
N. albomarginata × N. hirsuta^[8]
N. albomarginata × N. macrovulgaris^[23]
N. albomarginata × N. northiana [=N. × cincta]^[8]^[24]
N. albomarginata × N. rafflesiana^[23]^[25]
N. albomarginata × N. reinwardtiana [=N. × ferrugineomarginata]^[8]
? N. albomarginata × N. sanguinea^[23]^[26]
N. albomarginata × N. veitchii^[8]

N. albomarginata × N. northiana

Nepenthes × cincta is a rare plant and, due to the localised distribution of N. northiana, only grows at a few sites in Bau, Sarawak, usually on a substrate of limestone.

The traits of N. albomarginata are very dominant in this hybrid; the wide flared peristome of its larger parent species (N. northiana) is almost completely lost. Pitchers are narrowly infundibulate (funnel-shaped) throughout and range in colour from cream to dusky purple with red or black spots.^[27]

N. albomarginata × N. reinwardtiana

Its natural range covers the islands Borneo and Sumatra. The type specimen was collected by Shigeo Kurata in Kenukat, West Kalimantan, in 1981. Kurata described the hybrid the following year.

References

^ Ashton, P.S. 1971. The plants and vegetation of Bako National Park. Malayan Nature Journal 24: 151–162.
^ Clarke, C.M. (2018). "Nepenthes albomarginata". IUCN Red List of Threatened Species. 2018: e.T39639A143958253. doi:10.2305/IUCN.UK.2018-1.RLTS.T39639A143958253.en. Retrieved 19 November 2021.
^ ^a ^b Lindley, J. 1849. Familiar botany. — The pitcher plant. The Gardeners' Chronicle and Agricultural Gazette 1849(37): 580–581.
^ ^a ^b ^c ^d Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
^ Adam, J.H., C.C. Wilcock & M.D. Swaine 1989. Ecology and taxonomy of Bornean Nepenthes. University of Aberdeen Tropical Biology Newsletter 56: 2–4.
^ ^a ^b ^c Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
^ Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
^ 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. doi:10.1111/j.1420-9101.2011.02406.x
^ Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae). Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.
^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Peninsular Malaysia and Indochina. Redfern Natural History Productions, Poole.
^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.
^ Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes. Archived 2011-07-22 at the Wayback Machine Journal of Tropical Forest Science 5(1): 13–25.
^ Anderson, J.A.R. 1965. Limestone habitat in Sarawak. Proceedings of the Symposium on Ecological Research in Humid Tropics Vegetation, July 1963, Kuching, Sarawak. pp. 49–57.
^ ^a ^b Merbach, M.A., D.J. Merbach, U. Maschwitz, W.E. Booth, B. Fiala & G. Zizka 2002. Mass march of termites into the deadly trap. Nature 415: 36–37. doi:10.1038/415036a
^ Clarke, T. 2002. Plant has taste for termites. Nature News, January 3, 2002. doi:10.1038/news020101-4
^ Moran, J.A., M.A. Merbach, N.J. Livingston, C.M. Clarke & W.E. Booth 2001. Termite prey specialization in the pitcher plant Nepenthes albomarginata—evidence from stable isotope analysis. Annals of Botany 88: 307–311. doi:10.1006/anbo.2001.1460
^ Merbach, M.A., D.J. Merbach, W.E. Booth, U. Maschwitz, G. Zizka & B. Fiala 2000. A unique niche in plant carnivory: Nepenthes albomarginata feeds on epigaeically mass foraging termites. Tagungsband gtö 2000 13. Jahrestagung der Deutschen Gesellschaft für Tropenökologie 1–3. March 2000 in Würzburg Lehrstuhl für Tierökologie und Tropenbiologie Universität Würzburg. p. 105.
^ (in German) Merbach, D. & M. Merbach 2002. Auf der Suche nach Nahrung in die Todesfalle. Über die merkwürdigen Ernährungsgewohnheiten der fleischfressenden Kannenpflanze Nepenthes albomarginata. Forschung Frankfurt 2002(3): 74–77.
^ Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler. Das Pflanzenreich IV, III, Heft 36: 1–91.
^ ^a ^b (in German) Beck, G. 1895. Die Gattung Nepenthes. Wiener Illustrirte Garten-Zeitung 20(3–6): 96–107, 141–150, 182–192, 217–229.
^ (in Latin) Hooker, J.D. 1873. Ordo CLXXV bis. Nepenthaceæ. In: A. de Candolle Prodromus Systematis Naturalis Regni Vegetabilis 17: 90–105.
^ ^a ^b ^c ^d McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
^ Masters, M.T. 1884. New garden plants. Nepenthes cincta (Mast.), n. sp.. The Gardeners' Chronicle, new series, 21(540): 576–577.
^ Lowrie, A. 1983. Sabah Nepenthes Expeditions 1982 & 1983. Carnivorous Plant Newsletter 12(4): 88–95.
^ Shivas, R.G. 1985. Variation in Nepenthes albo-marginata. Carnivorous Plant Newsletter 14(1): 13–14.
^ Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.