Nepenthes albomarginata

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Nepenthes albomarginata Template:IPAc-en is a tropical pitcher plant native to Borneo, Peninsular Malaysia, and Sumatra.<ref name=Clarke01 /><ref>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.</ref>

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.<ref name=Clarke01 />

Botanical history

File:Nepenthes albomarginata (Bogor botanical park).jpg
Nepenthes albomarginata in the Bogor Botanical Gardens

Nepenthes albomarginata was first collected by Thomas Lobb in 1848. It was formally described a year later by John Lindley in The Gardeners' Chronicle.<ref name=Lindley1849 /><ref name=P&L />

The species was introduced into cultivation in the United Kingdom in 1856.<ref name=P&L />

In the 1996 book Pitcher-Plants of Borneo, N. albomarginata is given the vernacular name white-collared pitcher-plant.<ref name=P&L>Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.</ref> This name, along with all others, was dropped from the much-expanded second edition, published in 2008.<ref name=P&L2008>Phillipps, A., A. Lamb & C.C. Lee 2008. Pitcher Plants of Borneo. Second Edition. Natural History Publications (Borneo), Kota Kinabalu.</ref>

Description

Nepenthes albomarginata is a climbing plant. The stem may reach lengths of up to Template:Convert and is up to Template:Convert in diameter. Internodes are cylindrical in cross section and up to Template:Convert long.<ref name=Clarke97 />

Template:Multiple image Leaves are coriaceous in texture. The lamina or leaf blade is lanceolate in shape and up to Template:Convert long by Template:Convert 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 Template:Convert long.<ref name=Clarke97 />

Rosette and lower pitchers are bulbous in the basal third and cylindrical above. They are relatively small, reaching only Template:Convert in height by Template:Convert in width. A pair of fringed wings up to Template:Convert 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 Template:Convert wide, and bears indistinct teeth.<ref name=Clarke97 /> The inner portion of the peristome accounts for around 34% of its total cross-sectional surface length.<ref name=Baueretal2012>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. {{#invoke:CS1 identifiers|main|_template=doi}}</ref> 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 (≤Template:Convert long) is inserted near the base of the lid.<ref name=Clarke97 />

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.<ref name=Clarke97 />

Nepenthes albomarginata has a racemose inflorescence that is usually longer in male plants. The peduncle is up to Template:Convert long, while the rachis reaches lengths of up to Template:Convert. Partial peduncles are one- or two-flowered, up to Template:Convert long, and lack a bract. Sepals are obovate to oblong in shape and up to Template:Convert long.<ref name=Clarke97 /> A study of 120 pollen samples taken from a herbarium specimen (J.H.Adam 2417, collected in Borneo at an altitude of Template:Convert) found the mean pollen diameter to be Template:Convert (SE = 0.4; CV = 6.2%).<ref>Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae). Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.</ref>

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.<ref name=Clarke97 />

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.<ref name=MalaysiaIndochina>McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Peninsular Malaysia and Indochina. Redfern Natural History Productions, Poole.</ref><ref>McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.</ref> It has an altitudinal distribution of 0–1200 m above sea level.<ref>Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes. Template:Webarchive Journal of Tropical Forest Science 5(1): 13–25.</ref>

File:Nepenthes albomarginata purple.jpg
N. albomarginata growing in Sumatran heath forest

Its typical habitat consists of kerangas forest, but it has also been recorded from the summit vegetation of lowland peaks.<ref name=Clarke97 /> It is known from peat and limestone substrates.<ref name=Clarke97 /><ref>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.</ref>

Carnivory

Template:Multiple image 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.<ref name=Merbach>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. {{#invoke:CS1 identifiers|main|_template=doi}} </ref><ref>Clarke, T. 2002. Plant has taste for termites. Nature News, January 3, 2002. {{#invoke:CS1 identifiers|main|_template=doi}}</ref><ref>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. {{#invoke:CS1 identifiers|main|_template=doi}}</ref><ref>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.</ref><ref>Template:In lang 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. </ref>

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.<ref name=Merbach/> 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.

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.<ref name=Clarke01 />

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Infraspecific taxa

Natural hybrids

File:N. albomarginata x N. gracilis.jpg
N. albomarginata × N. gracilis

N. albomarginata × N. northiana

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File:Nepenthes cincta.jpg
An upper pitcher of N. × cincta

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.<ref name=Sarawak>Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.</ref>

N. albomarginata × N. reinwardtiana

{{#invoke:Labelled list hatnote|labelledList|Main article|Main articles|Main page|Main pages}} 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

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Further reading

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