Nepenthes burbidgeae
Encyclopedia
Nepenthes burbidgeae also known as the painted pitcher plant or Burbidge's Pitcher-Plant, is a tropical pitcher plant
Pitcher plant
Pitcher plants are carnivorous plants whose prey-trapping mechanism features a deep cavity filled with liquid known as a pitfall trap. It has been widely assumed that the various sorts of pitfall trap evolved from rolled leaves, with selection pressure favouring more deeply cupped leaves over...

 with a patchy distribution around Mount Kinabalu
Mount Kinabalu
Mount Kinabalu is a prominent mountain on the island of Borneo in Southeast Asia. It is located in the East Malaysian state of Sabah and is protected as Kinabalu National Park, a World Heritage Site. Kinabalu is the tallest peak in Borneo's Crocker Range and is the tallest mountain in the Malay...

 and neighbouring Mount Tambuyukon
Mount Tambuyukon
Mount Tambuyukon or Tamboyukon is Malaysia's third highest mountain at 2,579 m . It lies close to the famous Mount Kinabalu. The mountain supports a wide range of unique flora and fauna, including a number of pitcher plant species of the genus Nepenthes....

 in Sabah
Sabah
Sabah is one of 13 member states of Malaysia. It is located on the northern portion of the island of Borneo. It is the second largest state in the country after Sarawak, which it borders on its southwest. It also shares a border with the province of East Kalimantan of Indonesia in the south...

, Borneo
Borneo
Borneo is the third largest island in the world and is located north of Java Island, Indonesia, at the geographic centre of Maritime Southeast Asia....

.

Botanical history

Nepenthes burbidgeae was discovered on Mount Kinabalu in 1858 by Hugh Low
Hugh Low
Sir Hugh Low, GCMG was a British colonial administrator and naturalist. After a long residence in various colonial roles in Labuan, he became the first successful British administrator in the Malay Peninsula. His methods became models for future administrators. He made the first documented ascent...

 and Spenser St. John. St. John wrote the following account of finding the species near the Marai Parai
Marai Parai
Marai Parai or Marei Parei is a plateau on the northwestern side of Mount Kinabalu, in Malaysia. The mountain can be climbed from this side, although few parties attempt this route...

 plateau
Plateau
In geology and earth science, a plateau , also called a high plain or tableland, is an area of highland, usually consisting of relatively flat terrain. A highly eroded plateau is called a dissected plateau...

:

Crossing the Hobang, a steep climb led us to the western spur, along which our path lay; here, at about 4000 ft [1200 m], Mr. Low found a beautiful white and spotted pitcher-plant which he considered the prettiest of the twenty-two species of Nepenthes with which he was then acquainted; the pitchers are white and covered in a most beautiful manner with spots of an irregular form, of a rosy pink colour.


Frederick William Burbidge
Frederick William Burbidge
Frederick William Thomas Burbidge was a British explorer who collected many rare tropical plants for the famous Veitch Nurseries. Burbidge's first job was as a gardener at Kew Gardens...

 was one of the first to collect the plant in 1878, although he did not succeed in introducing it into cultivation. The type specimen of N. burbidgeae, Burbidge s.n., was collected on the Marai Parai plateau of Mount Kinabalu and is deposited at the Royal Botanic Gardens, Kew
Royal Botanic Gardens, Kew
The Royal Botanic Gardens, Kew, usually referred to as Kew Gardens, is 121 hectares of gardens and botanical glasshouses between Richmond and Kew in southwest London, England. "The Royal Botanic Gardens, Kew" and the brand name "Kew" are also used as umbrella terms for the institution that runs...

. A duplicate specimen is held at the New York Botanical Garden
New York Botanical Garden
- See also :* Education in New York City* List of botanical gardens in the United States* List of museums and cultural institutions in New York City- External links :* official website** blog*...

.

Nepenthes burbidgeae appeared as an unnamed species in Burbidge's 1880 book The Gardens of the Sun. Joseph Dalton Hooker
Joseph Dalton Hooker
Sir Joseph Dalton Hooker OM, GCSI, CB, MD, FRS was one of the greatest British botanists and explorers of the 19th century. Hooker was a founder of geographical botany, and Charles Darwin's closest friend...

 named N. burbidgeae after Burbidge's wife, though the name only appeared in an unpublished manuscript. The specific epithet is attributed to Burbidge as he used it in a letter to The Gardeners' Chronicle
The Gardeners' Chronicle
The Gardeners' Chronicle was a British horticulture periodical. It lasted as a title in its own right for nearly 150 years and is still extant as part of the magazine Horticulture Week....

 in 1882. It reads:

Nepenthes Burbidgeae, Hook. f. MSS., is a lovely thing, as yet unintroduced : pitchers pure white, semi-translucent like egg-shell, porcelain-white, with crimson or blood-tinted blotches. Lid blotched and dotted with crimson-purple. It is a very distinct plant, with triangular stems, 50 feet long, and the margins of the leaves decurrent.


In 1894, Otto Stapf
Otto Stapf
Otto Stapf FRS was an Austrian born botanist and taxonomist.Stapf trained in Vienna, moving to Royal Botanic Gardens, Kew in 1890. He was keeper of the Herbarium from 1909 to 1920...

 identified specimens belonging to N. burbidgeae as N. phyllamphora, a taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

 that is now considered synonymous with N. mirabilis
Nepenthes mirabilis
Nepenthes mirabilis , or the Common Swamp Pitcher-Plant, is a tropical carnivorous plant species of the pitfall trap variety. It has by far the widest distribution of any Nepenthes species and is known from the following countries and regions: Borneo, Sumatra, Thailand, Peninsular Malaysia,...

.

In two articles authored by Burbidge in 1894 and 1896, the name of this species was written as N. burbidgei. This name is considered a sphalma typographicum (misprint) of N. burbidgeae, although it appeared in a number of other works by authors such as Odoardo Beccari
Odoardo Beccari
Odoardo Beccari was an Italian naturalist perhaps best known for discovering the titan arum, the plant with the largest unbranched inflorescence in the world, in Sumatra in 1878...

 (1886), John Muirhead Macfarlane
John Muirhead Macfarlane
John Muirhead Macfarlane was a Scottish botanist. He was born and educated in Scotland, where he occupied several different academic positions at the University of Edinburgh before emigrating to the United States to assume a professorial chair at the University of Pennsylvania in 1893. He held...

 (1908), and Elmer Drew Merrill
Elmer Drew Merrill
Elmer Drew Merrill was an American botanist, specializing in the flora of the Asia-Pacific region.He was born in East Auburn, Maine, and attended the University of Maine where he received a B.S. in 1898...

 (1921). Herbarium material also bears this spelling of the name.

Seventy years after its discovery, N. burbidgeae remained a poorly known species. This is reflected in the writing of B. H. Danser
B. H. Danser
Benedictus Hubertus Danser , often abbreviated B. H. Danser, was a Dutch taxonomist and botanist...

 in his seminal 1928 monograph, "The Nepenthaceae of the Netherlands Indies
The Nepenthaceae of the Netherlands Indies
"The Nepenthaceae of the Netherlands Indies" is a seminal monograph by B. H. Danser on the tropical pitcher plants of the Dutch East Indies, North Borneo, the Malay Peninsula, and eastern New Guinea...

", where he suggests a close relative in N. pilosa
Nepenthes pilosa
Nepenthes pilosa is a tropical pitcher plant endemic to Borneo. It is characterised by a dense indumentum of long yellow-brown hairs. Pitchers have a distinctive hook-shaped appendage on the underside of the lid...

:

This species has only been found twice on Mt. Kinabalu and is very insufficiently known. I have not ventured to unite it with any other. N. pilosa, though doubtless the most nearly related species, is certainly different.


In 1981, Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

n botanist Allen Lowrie
Allen Lowrie
Allen Lowrie is a West Australian botanist. He is living in Duncraig, a Perth suburb, is married and has two daughters.Lowrie, originally a businessman and inventor, got in contact with the carnivorous flora of western Australia in the late sixties and worked on it as an amateur...

 reported that the fluid in unopened pitchers of N. burbidgeae is effective in stopping external bleeding. Lowrie cited two examples of researchers in the field successfully using this fluid on cuts and wounds.

Description

Nepenthes burbidgeae is a strong climber that quickly enters the vining stage. The stem reaches 15 m in length and is up to 18 mm in diameter. Internode
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...

s are cylindrical to triangular in cross section and up to 12 cm long.

The leaves of this species are coriaceous and petiolate. The lamina or leaf blade is oblong in shape and up to 40 cm long by 10 cm wide. It has an acute apex and its base is typically abruptly attenuate. The petiole
Petiole (botany)
In botany, the petiole is the stalk attaching the leaf blade to the stem. The petiole usually has the same internal structure as the stem. Outgrowths appearing on each side of the petiole are called stipules. Leaves lacking a petiole are called sessile, or clasping when they partly surround the...

 is winged, up to 15 cm long, and clasps the stem. It is often decurrent into two narrow wings that extend down the stem. Three to four longitudinal veins are present on either side of the midrib. Pinnate veins are inconspicuous. Tendril
Tendril
In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape that is used by climbing plants for support, attachment and cellular invasion by parasitic plants, generally by twining around suitable hosts. They do not have a lamina or blade, but they can photosynthesize...

s are up to 30 cm long.

Rosette and lower pitchers are rounded-infundibular or conical in shape. Unlike the pitchers of many other Nepenthes species, those of N. burbidgeae have no obvious constriction in the middle. The lower pitchers are relatively large, being up to 25 cm high by 10 cm wide. A pair of fringed wings, measuring up to 10 mm in width, runs down the front of each pitcher. The gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...

ular region, which bears minute overarched glands, covers the basal half of the pitcher's inner surface. The pitcher mouth is round and elongated into a short neck at the rear. The peristome
Peristome
The word peristome is derived from the Greek peri, meaning 'around' or 'about', and stoma, 'mouth'. It is a term used to describe various anatomical features that surround an opening to an organ or structure. The term is used in plants and invertebrate animals, such as in describing the shells of...

 is flattened and expanded, measuring up to 30 mm in width. Its inner margin is lined with a series of small but distinct teeth. The pitcher lid or operculum
Operculum (botany)
An operculum, in botany, is a term generally used to describe a structure within a plant, moss, or fungus acting as a cap, flap, or lid. In plants, it may also be called a bud cap.Examples of structures identified as opercula include:...

 is ovate and up to 8 cm wide. It bears a distinct keel as well as a characteristic hooked appendage on its lower surface. An unbranched spur
Spur (biology)
A spur in botany is a spike, usually part of a flower.In certain plants, part of a sepal or petal develops into an elongated hollow spike extending behind the flower, containing nectar which is sucked by long-tongued animals . Plants with such structures include Delphinium, Aquilegia, Piperia, and...

 (≤12 mm long) is inserted near the base of the lid.

Upper pitchers are similar to their terrestrial counterparts in most respects, even retaining the same colouration. However, they are smaller, reaching only 13 cm in height and 7 cm in width. They are infundibular in the basal third and globose above. In aerial pitchers, a pair of ribs is present in place of wings.

Nepenthes burbidgeae has a racemose
Raceme
A raceme is a type of inflorescence that is unbranched and indeterminate and bears pedicellate flowers — flowers having short floral stalks called pedicels — along the axis. In botany, axis means a shoot, in this case one bearing the flowers. In a raceme, the oldest flowers are borne...

 inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

. The peduncle
Peduncle (botany)
In botany, a peduncle is a stem supporting an inflorescence, or after fecundation, an infructescence.The peduncle is a stem, usually green and without leaves, though sometimes colored or supporting small leaves...

 is up to 25 cm long, while the rachis
Rachis
Rachis is a biological term for a main axis or "shaft".-In zoology:In vertebrates a rachis can refer to the series of articulated vertebrae, which encase the spinal cord. In this case the rachis usually form the supporting axis of the body and is then called the spine or vertebral column...

 reaches 30 cm in length. Partial peduncles may be one- or two-flowered and are up to 15 mm long. Sepal
Sepal
A sepal is a part of the flower of angiosperms . Collectively the sepals form the calyx, which is the outermost whorl of parts that form a flower. Usually green, sepals have the typical function of protecting the petals when the flower is in bud...

s are ovate and up to 5 mm long.

Most parts of the plant are covered in a sparse indumentum
Indumentum
The indumentum is a covering of fine hairs or bristles on a plant or insect.In plants, the indumentum types are:*pubescent*hirsute*pilose*villous*tomentose*stellate*scabrous*scurfy...

 of short hairs. The margins of the lamina are lined with brown hairs up to 3 mm long.

Nepenthes burbidgeae has a very restricted range and exhibits relatively little variability. As such, no infraspecific taxa have been described.

Habitat and distribution

Nepenthes burbidgeae is endemic to Kinabalu National Park
Kinabalu National Park
Kinabalu National Park or Taman Negara Kinabalu in Malay, established as one of the first national parks of Malaysia in 1964, is Malaysia's first World Heritage Site designated by UNESCO in December 2000 for its "outstanding universal values" and the role as one of the most important biological...

, where it has a patchy distribution around Mount Kinabalu
Mount Kinabalu
Mount Kinabalu is a prominent mountain on the island of Borneo in Southeast Asia. It is located in the East Malaysian state of Sabah and is protected as Kinabalu National Park, a World Heritage Site. Kinabalu is the tallest peak in Borneo's Crocker Range and is the tallest mountain in the Malay...

 and neighbouring Mount Tambuyukon
Mount Tambuyukon
Mount Tambuyukon or Tamboyukon is Malaysia's third highest mountain at 2,579 m . It lies close to the famous Mount Kinabalu. The mountain supports a wide range of unique flora and fauna, including a number of pitcher plant species of the genus Nepenthes....

. Specifically, it has been recorded from the Marai Parai
Marai Parai
Marai Parai or Marei Parei is a plateau on the northwestern side of Mount Kinabalu, in Malaysia. The mountain can be climbed from this side, although few parties attempt this route...

 plateau
Plateau
In geology and earth science, a plateau , also called a high plain or tableland, is an area of highland, usually consisting of relatively flat terrain. A highly eroded plateau is called a dissected plateau...

, Mamut copper mine, and Pig Hill
Pig Hill (Mount Kinabalu)
Pig Hill is an ultramafic area situated at approximately 2000 m above sea level on the southeastern side of Mount Kinabalu in Kinabalu National Park, Sabah, Malaysian Borneo. It is located to the east of Mesilau, some 4 hours' walk from the Mesilau Nature Resort...

. On Pig Hill, it grows at 1900–1950 m and is sympatric with N. rajah
Nepenthes rajah
Nepenthes rajah is an insectivorous pitcher plant species of the Nepenthaceae family. It is endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo. Nepenthes rajah grows exclusively on serpentine substrates, particularly in areas of seeping ground water where the...

, N. tentaculata
Nepenthes tentaculata
Nepenthes tentaculata , or the Fringed Pitcher-Plant, is a tropical pitcher plant with a very wide distribution across Borneo and Sulawesi...

, and the natural hybrid N. × alisaputrana. The altitudinal range of this species is often quoted as 1200–1800 m above sea level, but some sources give a lower limit of 1100 m and upper limit of 2250 m or even 2300 m.

Mount Kinabalu was only formed around 1 million years ago and, during the last ice age (approximately 20,000 to 10,000 years ago), it had an ice cap
Ice cap
An ice cap is an ice mass that covers less than 50 000 km² of land area . Masses of ice covering more than 50 000 km² are termed an ice sheet....

 on its summit. As such, it appears that N. burbidgeae is a relatively recent species in evolutionary terms.

Nepenthes burbidgeae is probably the rarest of the Nepenthes species native to Mount Kinabalu. Its typical habitat consists of mossy forest or montane forest, where it often grows in low scrub
Shrubland
Shrubland, scrubland, scrub or brush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity...

 and exposed areas on the tops of steep ridges. The species is restricted to ultramafic soils. In more exposed areas, N. burbidgeae is often found climbing amongst bushes of Leptospermum javanicum. At some localities it has also been recorded from bamboo
Bamboo
Bamboo is a group of perennial evergreens in the true grass family Poaceae, subfamily Bambusoideae, tribe Bambuseae. Giant bamboos are the largest members of the grass family....

 forest.

Nepenthes burbidgeae can often be found growing amongst populations of N. edwardsiana
Nepenthes edwardsiana
Nepenthes edwardsiana , or the Splendid Pitcher-Plant, is a tropical pitcher plant endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo...

, N. rajah
Nepenthes rajah
Nepenthes rajah is an insectivorous pitcher plant species of the Nepenthaceae family. It is endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo. Nepenthes rajah grows exclusively on serpentine substrates, particularly in areas of seeping ground water where the...

, and N. tentaculata
Nepenthes tentaculata
Nepenthes tentaculata , or the Fringed Pitcher-Plant, is a tropical pitcher plant with a very wide distribution across Borneo and Sulawesi...

, and hybrids with all of these species have been recorded.

Threats and conservation status

The El Niño climatic phenomenon of 1997 to 1998 had a catastrophic effect on the Nepenthes species of Mount Kinabalu. The dry period that followed severely depleted some natural populations. Forest fires
Wildfire
A wildfire is any uncontrolled fire in combustible vegetation that occurs in the countryside or a wilderness area. Other names such as brush fire, bushfire, forest fire, desert fire, grass fire, hill fire, squirrel fire, vegetation fire, veldfire, and wilkjjofire may be used to describe the same...

 broke out in 9 locations in Kinabalu Park, covering a total area of 25 square kilometres and generating large amounts of smog
Smog
Smog is a type of air pollution; the word "smog" is a portmanteau of smoke and fog. Modern smog is a type of air pollution derived from vehicular emission from internal combustion engines and industrial fumes that react in the atmosphere with sunlight to form secondary pollutants that also combine...

. Hugo Steiner
Hugo Steiner
Hugo Steiner is a Swiss citizen who graduated from the University of Zurich as a physician specialising in endocrinology. He worked for several years as the leader of a research group in experimental endocrinology for Hoffmann-La Roche Ltd in Basel, Switzerland. He has also done research work in...

 recalls being struck by the scarcity of N. burbidgeae pitchers observed on Mount Kinabalu during a trip in 1999. At the time of the El Niño, many plants were temporarily transferred to the park nursery. These were later replanted in the "Nepenthes Garden" in Mesilau
Mesilau
Mesilau, named after Mesilau River, is an area situated at approximately 2000 m above sea level on the East Ridge of Mount Kinabalu in Kinabalu National Park, Sabah, Malaysian Borneo. It is the site of the Mesilau Nature Resort, which is owned and operated by Sutera Sanctuary Lodges...

. Since then, Ansow Gunsalam has established a nursery close to the Mesilau Lodge at the base of Kinabalu Park to protect the endangered species of that area, including N. burbidgeae.

The conservation status
Conservation status
The conservation status of a group of organisms indicates whether the group is still extant and how likely the group is to become extinct in the near future...

 of N. burbidgeae is listed as Endangered
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

 on the 2006 IUCN Red List of Threatened Species
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

 based on an assessment carried out in 2000. This does not agree with an informal assessment made by Charles Clarke
Charles Clarke (botanist)
Dr. Charles M. Clarke is a botanist and taxonomist specialising in the carnivorous plant genus Nepenthes. Clarke has an honours degree in Botany from Monash University in Melbourne, and a Ph.D. in Ecosystem Management at the University of New England, in Armidale, New South Wales.Clarke first...

 in 1997, who classified the species as Vulnerable
Vulnerable species
On 30 January 2010, the IUCN Red List of Threatened Species identified 9694 Vulnerable species, subspecies and varieties, stocks and sub-populations.-References:...

 based on the IUCN
World Conservation Union
The International Union for Conservation of Nature and Natural Resources is an international organization dedicated to finding "pragmatic solutions to our most pressing environment and development challenges." The organization publishes the IUCN Red List, compiling information from a network of...

 criteria. However, Clarke noted that since all known populations of N. burbidgeae lie within the boundaries of Kinabalu National Park
Kinabalu National Park
Kinabalu National Park or Taman Negara Kinabalu in Malay, established as one of the first national parks of Malaysia in 1964, is Malaysia's first World Heritage Site designated by UNESCO in December 2000 for its "outstanding universal values" and the role as one of the most important biological...

 and are inaccessible to collectors, they "are unlikely to become threatened in the foreseeable future". Taking this into account, he suggested a revised assessment of Conservation Dependent
Conservation Dependent
Conservation Dependent was an IUCN category assigned to species or lower taxa which were dependent on conservation efforts to prevent the taxon becoming threatened with extinction...

.

Related species

Nepenthes burbidgeae is easily distinguished from other species in the genus on the basis of its pitcher shape and colouration, as well as the hook-shaped appendage on the underside of the lid. The only other Bornean Nepenthes species with a similarly developed appendage are N. chaniana
Nepenthes chaniana
Nepenthes chaniana ]]) is a highland pitcher plant species belonging to the genus Nepenthes. It is characterised by a dense indumentum of long, white hairs. Pitchers are cylindrical and mostly white to yellow in colouration....

 and N. pilosa
Nepenthes pilosa
Nepenthes pilosa is a tropical pitcher plant endemic to Borneo. It is characterised by a dense indumentum of long yellow-brown hairs. Pitchers have a distinctive hook-shaped appendage on the underside of the lid...

.

B. H. Danser
B. H. Danser
Benedictus Hubertus Danser , often abbreviated B. H. Danser, was a Dutch taxonomist and botanist...

 suggested that N. burbidgeae is most closely related to N. pilosa. The latter species is poorly known and was for a long time confused with N. chaniana.

The glandular crest of N. chaniana is very similar to that of N. burbidgeae, particularly in upper pitchers. However, it is difficult to confuse these species as the pitchers are otherwise markedly different in structure; the upper pitchers of N. burbidgeae are short and funnel-shaped, whereas those of N. chaniana are elongated and have a dense indumentum
Indumentum
The indumentum is a covering of fine hairs or bristles on a plant or insect.In plants, the indumentum types are:*pubescent*hirsute*pilose*villous*tomentose*stellate*scabrous*scurfy...

 of white hair.

Natural hybrids

Natural hybrids involving N. burbidgeae appear to be relatively rare and only four have been recorded to date. Three of these (crosses with N. edwardsiana
Nepenthes edwardsiana
Nepenthes edwardsiana , or the Splendid Pitcher-Plant, is a tropical pitcher plant endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo...

, N. fusca
Nepenthes fusca
Nepenthes fusca , or the Dusky Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It is found throughout a wide altitudinal range and is almost always epiphytic in nature, primarily growing in mossy forest....

, and N. tentaculata
Nepenthes tentaculata
Nepenthes tentaculata , or the Fringed Pitcher-Plant, is a tropical pitcher plant with a very wide distribution across Borneo and Sulawesi...

) have received little attention in the scientific literature, but N. burbidgeae × N. rajah
Nepenthes rajah
Nepenthes rajah is an insectivorous pitcher plant species of the Nepenthaceae family. It is endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo. Nepenthes rajah grows exclusively on serpentine substrates, particularly in areas of seeping ground water where the...

 has been described as N. × alisaputrana and is famous for producing huge pitchers rivalling those of N. rajah in size.

N. burbidgeae × N. rajah

Nepenthes × alisaputrana was described in 1992 by J. H. Adam
J. H. Adam
Jumaat Haji Adam is a botanist and taxonomist specialising in the carnivorous pitcher plant genus Nepenthes.Adam has described numerous Nepenthes taxa, mostly with C. C. Wilcock, including the species N. faizaliana and N. mapuluensis, as well as the natural hybrids N. × alisaputrana, N. ×...

 and C. C. Wilcock
C. C. Wilcock
Christopher C. Wilcock is a taxonomist specialising in the carnivorous pitcher plant genus Nepenthes.Together with J. H. Adam, Wilcock has described several Nepenthes taxa, including the species N. faizaliana and N. mapuluensis, as well as the natural hybrids N. × alisaputrana and N. ×...

 and is named in honour of Datuk Lamri Ali
Datuk Lamri Ali
Datuk Lamri Ali is a former Director of Sabah Parks. At the December 1999 World Conservation Union 's regional meeting staged in Pakse, Laos, Ali was awarded the Fred M. Packard WCPA-IUCN award. The award recognises his contribution to nature conservation and protected area movement in Malaysia...

, Director of Sabah Parks
Sabah Parks
Sabah Parks is a conservation-based statutory body established in 1962 with the purpose of conserving the scenic, scientific and historic heritage of the state of Sabah, Malaysia, on the island of Borneo. The organisation is also responsible for the management and promotion of the various protected...

. It is only known from a few remote localities within Kinabalu National Park
Kinabalu National Park
Kinabalu National Park or Taman Negara Kinabalu in Malay, established as one of the first national parks of Malaysia in 1964, is Malaysia's first World Heritage Site designated by UNESCO in December 2000 for its "outstanding universal values" and the role as one of the most important biological...

, where it grows in stunted, open vegetation over serpentine soils at around 2000 m above sea level, often amongst populations of N. burbidgeae.
This plant is notable for combining the best characters of both parent species, not least the size of its pitchers, which rival those of N. rajah in volume (≤35 cm high, ≤20 cm wide). The other hybrids involving N. rajah do not exhibit such impressive proportions. The pitchers of N. × alisaputrana can be distinguished from those of N. burbidgeae by a broader peristome, larger lid and simply by their sheer size. The hybrid differs from its other parent, N. rajah, by its lid structure, indumentum of short, brown hairs, narrower and more cylindrical peristome, and pitcher colour, which is usually yellow-green with red or brown flecking. For this reason, Anthea Phillipps
Anthea Phillipps
Anthea Phillipps B.Sc. is a British botanist. Phillipps was brought up in Sabah, Borneo as a child . She received a Botany degree from the University of Durham, England. She worked at the Sabah Museum before joining the Sabah Parks service from 1980 to 1987 as Park Ecologist, where she studied...

 and Anthony Lamb
Anthony Lamb
Anthony L. Lamb M.A., Dip. Ag., D.T.A. is a British botanist, born in Sri Lanka, and specialising in the flora of Borneo. Lamb was educated at Blundell's School in Tiverton and at St John's College at Cambridge...

 gave it the common name "Leopard Pitcher-Plant". The peristome is green to dark red and striped with purple bands. Leaves are often slightly peltate. The hybrid is a strong climber and frequently produces upper pitchers.

Nepenthes × alisaputrana more closely resembles N. rajah than N. burbidgeae, but it is difficult to confuse this plant with either. However, this mistake has previously been made on at least one occasion; a pitcher illustrated in Adrian Slack
Adrian Slack
Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them ....

's Insect-Eating Plants and How to Grow Them as being N. rajah is in fact N. burbidgeae × N. rajah.
Distribution of phenolic compounds and leucoanthocyanins in
N. burbidgeae, N. rajah, and N. × alisaputrana
Taxon
1
2
3
4
5
6
7
 8 
Specimen
N. burbidgeae
3+
++
3+
3+
-
+
-
-
Jumaat 2484
N. rajah
Nepenthes rajah
Nepenthes rajah is an insectivorous pitcher plant species of the Nepenthaceae family. It is endemic to Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Malaysian Borneo. Nepenthes rajah grows exclusively on serpentine substrates, particularly in areas of seeping ground water where the...

-
-
+
±
++
++
3+
+
Jumaat 2443
N. × alisaputrana
+
++
3+
3+
+
++
3+
+
Jumaat 2442
N. × alisaputrana (in vitro
In vitro
In vitro refers to studies in experimental biology that are conducted using components of an organism that have been isolated from their usual biological context in order to permit a more detailed or more convenient analysis than can be done with whole organisms. Colloquially, these experiments...

)
+
++
3+
3+
+
++
+
+
Key: 1: Phenolic acid
Phenolic acid
Phenolic acids are a type of organic compounds. Included in that class are substances containing a phenolic ring and an organic carboxylic acid function ....

, 2: Ellagic acid
Ellagic acid
Ellagic acid is a natural phenol antioxidant found in numerous fruits and vegetables including blackberries, raspberries, strawberries, cranberries, walnuts, pecans, pomegranates, wolfberry and other plant foods...

, 3: Quercetin
Quercetin
Quercetin , a flavonol, is a plant-derived flavonoid found in fruits, vegetables, leaves and grains. It also may be used as an ingredient in supplements, beverages or foods.-Occurrence:...

, 4: Kaempferol
Kaempferol
Kaempferol is a natural flavonol, a type of flavonoid, that has been isolated from tea, broccoli, Delphinium, Witch-hazel, grapefruit,cabbage, kale, beans, endive, leek, tomato, strawberries, grapes, brussels sprouts, apples and other plant sources. Kaempferol is a yellow crystalline solid with a...

, 5: Luteolin
Luteolin
Luteolin is a yellow crystalline compound. It is a flavonoid; to be specific, it is one of the more common flavones. From preliminary research, it is thought to play a role in the human body possibly as an antioxidant, a free radical scavenger, a promoter of carbohydrate metabolism, or an immune...

, 6: 'Unknown Flavonoid
Flavonoid
Flavonoids , are a class of plant secondary metabolites....

 1', 7: 'Unknown Flavonoid 3', 8: Cyanidin
Cyanidin
Cyanidin is a natural organic compound. It is a particular type of anthocyanidin . It is a pigment found in many red berries including but not limited to grapes, bilberry, blackberry, blueberry, cherry, cranberry, elderberry, hawthorn, loganberry, acai berry and raspberry...


±: very weak spot, +: weak spot, ++: strong spot, 3+: very strong spot, -: absent


In 2002, phytochemical screening
Phytochemical
Phytochemicals are biologically active chemical compounds that occur naturally in plants . Phytochemicals are the molecules responsible for the color and organoleptic properties . For example, the deep purple color of blueberries and the smell of garlic...

 and analytical chromatography were used to study the presence of phenolic compounds and leucoanthocyanins in N. × alisaputrana and its putative parent species. The research was based on leaf material from nine dry herbarium specimens. Eight spots containing phenolic acid
Phenolic acid
Phenolic acids are a type of organic compounds. Included in that class are substances containing a phenolic ring and an organic carboxylic acid function ....

s, flavonols, flavones, leucoanthocyanins and 'unknown flavonoid
Flavonoid
Flavonoids , are a class of plant secondary metabolites....

' 1 and 3 were identified from chromatographic profiles. The distributions of these in the hybrid N. × alisaputrana and its putative parental species N. burbidgeae and N. rajah are shown in the table to the left. A specimen of N. × alisaputrana grown from tissue culture
Tissue culture
Tissue culture is the growth of tissues or cells separate from the organism. This is typically facilitated via use of a liquid, semi-solid, or solid growth medium, such as broth or agar...

 (in vitro
In vitro
In vitro refers to studies in experimental biology that are conducted using components of an organism that have been isolated from their usual biological context in order to permit a more detailed or more convenient analysis than can be done with whole organisms. Colloquially, these experiments...

) was also tested.

Luteolin
Luteolin
Luteolin is a yellow crystalline compound. It is a flavonoid; to be specific, it is one of the more common flavones. From preliminary research, it is thought to play a role in the human body possibly as an antioxidant, a free radical scavenger, a promoter of carbohydrate metabolism, or an immune...

, cyanidin
Cyanidin
Cyanidin is a natural organic compound. It is a particular type of anthocyanidin . It is a pigment found in many red berries including but not limited to grapes, bilberry, blackberry, blueberry, cherry, cranberry, elderberry, hawthorn, loganberry, acai berry and raspberry...

 and 'Unknown Flavonoid 3' were undetected in N. burbidgeae, while concentrations of 'Unknown Flavonoid 1' were found to be weak. Chromatographic patterns of the N. × alisaputrana samples studied showed complementation of its putative parental species.

Myricetin
Myricetin
Myricetin is a naturally occurring flavonol, a flavonoid found in many grapes, berries, fruits, vegetables, herbs, as well as other plants. Walnuts are a rich dietary source. Trace amounts can be found as glycosides. It is one of the phenolic compounds present in red wine.Myricetin has antioxidant...

 was found to be absent from all studied taxa. This agrees with the findings of previous authors and suggests that the absence of a widely distributed compound like myricetin among the Nepenthes examined might provide additional diagnostic information for these taxa
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

.

Cultivation

Little information has been published on the growing requirements of N. burbidgeae. In Insect-Eating Plants and How to Grow Them, Adrian Slack
Adrian Slack
Adrian Slack is an author and authority on carnivorous plants. He is the author of two books: Carnivorous Plants and Insect-Eating Plants and How to Grow Them ....

 wrote that cuttings of N. burbidgeae were more difficult to root than those of other Nepenthes species.

In 2004, professional horticulturist Robert Sacilotto published a summary of measured tolerances of highland Nepenthes species, based on experiments conducted between 1996 and 2001. Nepenthes burbidgeae was found to be tolerant of a fairly wide range of conditions, particularly in terms of temperature and soil composition; it grew in every substrate used in the experiment. However, plants showed stunted growth when grown in a mixture consisting of 50% silica gel
Silica gel
Silica gel is a granular, vitreous, porous form of silica made synthetically from sodium silicate. Despite its name, silica gel is a solid. It is a naturally occurring mineral that is purified and processed into either granular or beaded form...

, 20% Sphagnum
Sphagnum
Sphagnum is a genus of between 151 and 350 species of mosses commonly called peat moss, due to its prevalence in peat bogs and mires. A distinction is made between sphagnum moss, the live moss growing on top of a peat bog on one hand, and sphagnum peat moss or sphagnum peat on the other, the...

 moss, 20% fir
Fir
Firs are a genus of 48–55 species of evergreen conifers in the family Pinaceae. They are found through much of North and Central America, Europe, Asia, and North Africa, occurring in mountains over most of the range...

 bark, and 10% peat moss chunks. The highest growth rates were exhibited by specimens in 50% leached perlite
Perlite
Perlite is an amorphous volcanic glass that has a relatively high water content, typically formed by the hydration of obsidian. It occurs naturally and has the unusual property of greatly expanding when heated sufficiently...

, 30% long fiber Sphagnum moss, 10% peat moss chunks, and 10% fir bark, as well as media without fir bark and with a higher percentage of Sphagnum. Nepenthes burbidgeae was found to tolerate temperatures in the range of 9 to 41°C (48° to 105°F). A nighttime drop in temperature below 18°C (65°F) was necessary for good growth; plants that were not exposed to such a drop grew around 50% slower and produced fewer pitchers. Optimal growth rates were observed with daytime temperatures of 20 to 29°C (68° to 85°F) and nighttime temperatures of 12 to 16°C (54° to 60°F). Soil with a pH
PH
In chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline...

 of 4.8 to 5.5 produced the best results; values below 3.5 corresponded with slower growth. Optimal soil conductivity was between 10 and 24 microsiemens
Siemens (unit)
The siemens is the SI derived unit of electric conductance and electric admittance. Conductance and admittance are the reciprocals of resistance and impedance respectively, hence one siemens is equal to the reciprocal of one ohm, and is sometimes referred to as the mho. In English, the term...

, and prolonged exposure of one week or more to levels of more than 60 microsiemens resulted in foliar burn. The experiments suggested that N. burbidgeae grows best when relative humidity
Relative humidity
Relative humidity is a term used to describe the amount of water vapor in a mixture of air and water vapor. It is defined as the partial pressure of water vapor in the air-water mixture, given as a percentage of the saturated vapor pressure under those conditions...

 is in the range of 68 to 95%. However, constant exposure to high humidity in excess of 90% resulted in disease outbreaks and increased plant death rates. Seedlings of less than one year proved to be particularly vulnerable to this. Optimal light levels varied depending on the light source used: 8100–11000 lx
Lux
The lux is the SI unit of illuminance and luminous emittance, measuring luminous flux per unit area. It is used in photometry as a measure of the intensity, as perceived by the human eye, of light that hits or passes through a surface...

 (750–1000 fc
Foot-candle
A foot-candle is a non-SI unit of illuminance or light intensity widely used in photography, film, television, conservation lighting, and the lighting industry...

) in sunlight, 7000–9700 lx (650–900 fc) under high pressure sodium lamps
Sodium vapor lamp
A sodium vapor lamp is a gas discharge lamp that uses sodium in an excited state to produce light. There are two varieties of such lamps: low pressure and high pressure...

, 6500–9100 lx (600–850 fc) under metal halide lamp
Metal halide lamp
Metal-halide lamps, a member of the high-intensity discharge family of lamps, produce high light output for their size, making them a compact, powerful, and efficient light source. By adding rare earth metal salts to the mercury vapor lamp, improved luminous efficacy and light color is obtained...

s, and 5400–7300 lx (500–680 fc) under fluorescent lamp
Fluorescent lamp
A fluorescent lamp or fluorescent tube is a gas-discharge lamp that uses electricity to excite mercury vapor. The excited mercury atoms produce short-wave ultraviolet light that then causes a phosphor to fluoresce, producing visible light. A fluorescent lamp converts electrical power into useful...

s. Nepenthes burbidgeae could be grown in lower light conditions, but such plants exhibited etiolated growth and reduced colouration. The species was found to respond well to a fertilizer
Fertilizer
Fertilizer is any organic or inorganic material of natural or synthetic origin that is added to a soil to supply one or more plant nutrients essential to the growth of plants. A recent assessment found that about 40 to 60% of crop yields are attributable to commercial fertilizer use...

 that was applied to the pitchers on a monthly basis, but a foliar feed using the same solution produced no visible change in growth rate.

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