Nepenthes villosa
Encyclopedia
Nepenthes villosa or the Villose Pitcher-Plant, is a tropical pitcher plant
endemic to Mount Kinabalu
and neighbouring Mount Tambuyukon
in northeastern Borneo
. It grows at higher altitudes than any other Bornean Nepenthes
species, occurring at elevations of over 3200 m. Nepenthes villosa is characterised by its highly-developed and intricate peristome
, which distinguishes it from the closely related N. edwardsiana
and N. macrophylla
.
The specific epithet villosa is Latin
for "hairy" and refers to the dense indumentum
of this species.
in 1852 by Joseph Dalton Hooker
. The description was published in Icones plantarum and accompanied by an illustration.
The species was first collected in 1858 by Hugh Low
when he made his second ascent of Mount Kinabalu
together with Spenser St. John.
In an issue of Curtis's Botanical Magazine
published in 1858, an illustration of an upper pitcher of N. veitchii
was incorrectly identified as N. villosa by J. D. Hooker's father, William Jackson Hooker
.
That year, N. villosa was also covered in Flore des Serres et des Jardins de l'Europe
by Louis van Houtte
.
In 1859, N. villosa was again described and illustrated in J. D. Hooker's treatment of the genus published in The Transactions of the Linnean Society of London. The illustration and description were reproduced in Spenser St. John's Life in the Forests of the Far East, published in 1862.
B. H. Danser
treated N. edwardsiana in synonymy with N. villosa in his seminal monograph "The Nepenthaceae of the Netherlands Indies
", published in 1928. The work included a revised Latin
diagnosis and botanical description of N. villosa.
Danser listed four herbarium specimens that he identified as belonging to N. villosa. These include two collected by George Darby Haviland
from Mount Kinabalu
in 1892. One of these, Haviland 1656/1232, was collected at an altitude of 2400 m. It includes male floral material and is deposited at the Herbarium of the Sarawak Museum. The second specimen, Haviland 1813/1353, was collected from the Marai Parai
plateau at an altitude of 1650 m; it likely represents N. edwardsiana
. It is also deposited at the Herbarium of the Sarawak Museum. It does not include floral material. G. D. Haviland explored the Mount Kinabalu area with his brother H. A. Haviland between March and April, 1892, and must have collected these specimens during this time.
Additionally, Danser lists two specimens collected by Joseph Clemens
in 1915. The first, Clemens 10627, was collected on November 13 from Paka Cave to Low's Peak. It includes female floral material. The second, Clemens 10871, was collected at Marai Parai between November 22 and November 23, and does not include floral material. Both specimens are deposited at the Bogor Botanical Gardens
(formerly the Herbarium of the Buitenzorg Botanic Gardens) in Java
.
In Letts Guide to Carnivorous Plants of the World, published in 1992, a specimen of the natural hybrid N. × kinabaluensis (N. rajah
× N. villosa) is labeled as N. villosa.
s are cylindrical and up to 10 cm long.
Leaves are coriaceous and petiolate. The lamina is spathulate to oblong and may be up to 25 cm long and 6 cm wide. The apex of the lamina is emarginate. The petiole
is canaliculate, up to 10 cm long, and bears an amplexicaul sheath. One to three longitudinal veins are present on either side of the midrib. Tendril
s may reach 50 cm in length.
Lower and upper pitchers are very similar. They are urceolate to ovate in shape. The pitchers grow up to 25 cm high and 9 cm wide. A pair of fringed wings (≤15 mm wide) runs down the front of the pitcher, although it may be reduced to ribs in aerial traps. The pitcher mouth is oblique and elongated into a neck at the rear. The glands on the inner surface are overarched and occur at a density of 200 to 1300 per square centimetre. The peristome
is cylindrical in cross section and up to 20 mm wide. It bears well developed teeth and ribs. The lid or operculum
is cordate and has a pointed apex. An unbranched spur
(≤20 mm long) is inserted at the base of the lid.
Nepenthes villosa has a racemose
inflorescence
. The peduncle
may be up to 40 cm long, while the rachis
grows to 20 cm in length. Pedicels
are filiform-bract
eolate and up to 15 mm long. Sepal
s are round to elliptic and up to 4 mm long. A study of 490 pollen
samples taken from two herbarium specimens (J.H.Adam 1124 and J.H.Adam 1190, collected at an altitude of 1800–3400 m) found the mean pollen diameter to be 37.2 μm
(SE
= 0.2; CV
= 6.7%).
The species has a dense indumentum
of long, brown hairs that covers all parts of the plant.
and neighbouring Mount Tambuyukon
in Sabah
, Borneo
. It generally grows at 2300–3240 m above sea level, the highest altitude of all Bornean Nepenthes species; only N. lamii
and N. sp. Papua
from New Guinea
are found at greater elevations. On Mount Kinabalu, N. villosa is common along the Mesilau Trail (between Pondok Magnolia and the meeting point with the old summit trail) and almost all the way up to the Laban Rata rest house; a particularly large population has been reported at around 3047 m. On Mount Tambuyukon, an altitudinal inversion has been noted, whereby N. villosa is more common at much lower elevations of 1600–1900 m, being replaced by N. rajah
towards the summit. The exposed, uppermost slopes of Mount Tambuyukon can become very hot during the day and this might explain the inability of N. villosa to colonise them.
Nepenthes villosa often grows in mossy forest and sub-alpine forest dominated by species of the genera Dacrydium
and Leptospermum
, particularly Leptospermum recurvum
. It has also been recorded growing among shrubs, grass, and boulders in open areas. Here the soil may become relatively dry, although relative humidity
is usually close to 100% as the slopes are often enveloped in clouds. Like many Nepenthes from the Mount Kinabalu area, it is endemic to ultramafic soils.
The conservation status
of N. villosa is listed as Vulnerable
on the 2006 IUCN Red List of Threatened Species
. Although many plants grow along Mount Kinabalu's summit trail and are easily accessible to climbers, all known populations of the species grow within Kinabalu National Park
and so their collection is illegal. In 1997, Charles Clarke
suggested a revised assessment of Conservation Dependent
based on this. Clarke writes that N. villosa "has a secure future", although he adds that climbers have had a significant impact on populations of the species growing along the summit trail, with the number of plants having declined in recent years. A 2002 study found 1180 individual N. villosa growing in 11 plots, each measuring 0.01 hectare
s, at elevations of between 2610 m and 2970 m on Mount Kinabalu. This number constituted 94% of the pitcher plants recorded from the plots, the rest being N. × kinabaluensis.
and N. macrophylla
. There has been much taxonomic confusion surrounding the status of these three taxa
.
Joseph Dalton Hooker
, who described both N. edwardsiana and N. villosa, noted the similarity between the two species as follows:
Günther Beck von Mannagetta und Lerchenau was the first to treat N. edwardsiana in synonymy with N. villosa when he published his monograph on the genus in 1895.
In his 1908 monograph, John Muirhead Macfarlane
wrote the following with regards to the two species: "Examinatione microscopica probatur, illas species distinctas esse". This is probably "based on the old belief that plants, which differ anatomically, can not be forms of the same species".
B. H. Danser
united the species "[w]ith some hesitation" in his 1928 monograph "The Nepenthaceae of the Netherlands Indies
". He suggested that N. villosa is a stunted form of N. edwardsiana from higher altitudes, which flowers at a "juvenile stage of development". Danser acknowledged that the indumentum
of N. villosa is more dense than that of N. edwardsiana, but noted that it "is a difference only of degree".
The two taxa differ considerably in their altitudinal distributions. Nepenthes villosa usually occurs at ultrahighland elevations (2300–3240 m), whereas N. edwardsiana is found between 1500 and 2700 m. Where their altitudinal distributions overlap, they are still identifiable as distinct species.
Nepenthes macrophylla
was originally described in 1987 as a subspecies
of N. edwardsiana by Johannes Marabini. It was later elevated to species status by Matthew Jebb
and Martin Cheek
. This interpretation was supported by Charles Clarke
, who noted that N. edwardsiana and N. villosa "have more in common" than N. edwardsiana and N. macrophylla. Whereas N. edwardsiana and N. villosa are restricted to the Kinabalu area, N. macrophylla is only found near the summit of Mount Trus Madi
.
Matthew Jebb and Martin Cheek suggest that N. villosa is related to N. mira
, a species endemic to Palawan
in the Philippines
. N. villosa also shows affinities to N. peltata
of Mindanao
.
Nepenthes × harryana is the natural hybrid between N. edwardsiana
and N. villosa. Its two parent species are very closely related and so N. × harryana, which is intermediate in form, may be difficult to distinguish from either of them.
It was originally described as a species in 1882 by Frederick William Burbidge
. John Muirhead Macfarlane
was the first to realise its hybrid origin and described it as such in his monograph of 1908. Danser wrote that N. × harryana could be a hybrid as Macfarlane suggested, or a form of N. villosa together with N. edwardsiana.
Nepenthes × harryana can be distinguished from N. villosa on the basis of its pitcher morphology. The pitchers of the hybrid are more cylindrical than those of N. villosa, whereas the indumentum
is more dense than that of N. edwardsiana. The hip of the pitcher cup, which is found just below the peristome in N. villosa and in the lower quarter of N. edwardsiana pitchers, is located around the middle of N. × harryana pitchers. However, N. villosa plants from Mount Tambuyukon
are easier to confuse with this hybrid, as they produce pitchers that may be elongated slightly above the hip.
Nepenthes × harryana is known from a ridge above the Upper Kolopis River
and from two locations along the Kinabalu summit trail. Since N edwardsiana does not grow along the summit trail, it cannot be confused with this hybrid there. Burbidge wrote that N. edwardsiana, N. × harryana, and N. villosa "are quite distinct in zone of the mountain".
and N. villosa. It was first collected near Kambarangoh
on Mount Kinabalu by Lilian Gibbs
in 1910 and later mentioned by John Muirhead Macfarlane
as "Nepenthes sp." in 1914. Although Macfarlane did not formally name the plant, he noted that "[a]ll available morphological details suggest that this is a hybrid between N. villosa and N. rajah". It was finally described in 1976 by Shigeo Kurata
as N. × kinabaluensis. The name was published in Nepenthes of Mount Kinabalu
, but it is a nomen nudum
, as it had an inadequate description and lacked information on the type specimen
. The name was subsequently republished by Kurata in 1984 and by J. H. Adam
and C. C. Wilcock
in 1998.
The pitchers of N. × kinabaluensis may be quite large, but do not compare to those of N. rajah or N. × alisaputrana (N. burbidgeae
× N. rajah). N. × kinabaluensis can only be found on Mount Kinabalu (hence the name) and nearby Mount Tambuyukon, where the two parent species occur sympatrically
. More specifically, plants are known from a footpath near Paka Cave and several places along an unestablished route on a south-east ridge, which lies on the west side of the Upper Kolopis River
. The only accessible location from which this hybrid is known is the Kinabalu summit trail, between Layang-Layang and the helipad
, where it grows at about 2900 m in a clearing dominated by Dacrydium gibbsiae
and Leptospermum recurvum
trees. N. × kinabaluensis has an altitudinal distribution of 2420 to 3030 m. It grows in open areas in cloud forest.
The hybrid is generally intermediate in appearance between its parent species. Raised ribs line the inner edge of the peristome and end with elongated teeth. These are more prominent than those found in N. rajah and smaller than those of N. villosa. The peristome is coarse and expanded at the margin (but not scalloped like that of N. rajah), the lid orbiculate or reniform and almost flat. In general, pitchers are larger than those of N. villosa and the tendril joins the apex about 1–2 cm below the leaf tip, a feature which is characteristic of N. rajah. In older plants, the tendril can be almost woody. N. × kinabaluensis is an indumentum of villous hairs covering the pitchers and leaf margins, which is approximately intermediate between the parents. Lower pitchers have two fringed wings, whereas the upper pitchers usually lack these. The colour of the pitcher varies from yellow to scarlet. N. × kinabaluensis seems to produce upper pitchers more readily than either of its parents. In all respects N. × kinabaluensis is intermediate between the two parent species and it is easy to distinguish from all other Nepenthes of Borneo. However, it has been confused once before, when the hybrid was identified as N. rajah in Letts Guide to Carnivorous Plants of the World.
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...
endemic to 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 northeastern 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....
. It grows at higher altitudes than any other Bornean Nepenthes
Nepenthes
The Nepenthes , popularly known as tropical pitcher plants or monkey cups, are a genus of carnivorous plants in the monotypic family Nepenthaceae. The genus comprises roughly 130 species, numerous natural and many cultivated hybrids...
species, occurring at elevations of over 3200 m. Nepenthes villosa is characterised by its highly-developed and intricate 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...
, which distinguishes it from the closely related 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...
and N. macrophylla
Nepenthes macrophylla
Nepenthes macrophylla , the Large-Leaved Pitcher-Plant, is a tropical pitcher plant known only from a very restrictive elevation on Mount Trus Madi in Sabah, Malaysian Borneo...
.
The specific epithet villosa is Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
for "hairy" and refers to the 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 this species.
Botanical history
Nepenthes villosa was formally describedSpecies description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...
in 1852 by 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...
. The description was published in Icones plantarum and accompanied by an illustration.
The species was first collected 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...
when he made his second ascent of 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...
together with Spenser St. John.
In an issue of Curtis's Botanical Magazine
Curtis's Botanical Magazine
The Botanical Magazine; or Flower-Garden Displayed, is an illustrated publication which began in 1787. The longest running botanical magazine, it is widely referred to by the subsequent name Curtis's Botanical Magazine....
published in 1858, an illustration of an upper pitcher of N. veitchii
Nepenthes veitchii
Nepenthes veitchii |James Veitch]], nurseryman of the Veitch Nurseries), or Veitch's Pitcher-Plant, is a Nepenthes species from the island of Borneo. The plant is widespread in north-western Borneo and can also be found in parts of Kalimantan. N...
was incorrectly identified as N. villosa by J. D. Hooker's father, William Jackson Hooker
William Jackson Hooker
Sir William Jackson Hooker, FRS was an English systematic botanist and organiser. He held the post of Regius Professor of Botany at Glasgow University, and was the first Director of the Royal Botanic Gardens, Kew. He enjoyed the friendship and support of Sir Joseph Banks for his exploring,...
.
That year, N. villosa was also covered in Flore des Serres et des Jardins de l'Europe
Flore des Serres et des Jardins de l'Europe
Flore des Serres et des Jardins de l'Europe was one of the finest horticulture journals produced in Europe during the 19th century, spanning 23 volumes and over 2000 coloured plates with French, German and English text...
by Louis van Houtte
Louis van Houtte
Louis Benoit van Houtte was a Belgian horticulturist who was with the Jardin Botanique de Brussels between 1836 and 1838 and is best-known for the journal Flore des Serres et des Jardins de l'Europe, produced with Charles Lemaire and M...
.
In 1859, N. villosa was again described and illustrated in J. D. Hooker's treatment of the genus published in The Transactions of the Linnean Society of London. The illustration and description were reproduced in Spenser St. John's Life in the Forests of the Far East, published in 1862.
B. H. Danser
B. H. Danser
Benedictus Hubertus Danser , often abbreviated B. H. Danser, was a Dutch taxonomist and botanist...
treated N. edwardsiana in synonymy with N. villosa in his seminal 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...
", published in 1928. The work included a revised Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
diagnosis and botanical description of N. villosa.
Danser listed four herbarium specimens that he identified as belonging to N. villosa. These include two collected by George Darby Haviland
George Darby Haviland
Dr George Darby Haviland was a British surgeon and naturalist. He was born at Warbleton, Sussex in England. He served as Director of the Raffles Museum in Singapore as well as being a medical officer in Sarawak and Curator of the Sarawak Museum in Kuching from 1891 to 1893...
from 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...
in 1892. One of these, Haviland 1656/1232, was collected at an altitude of 2400 m. It includes male floral material and is deposited at the Herbarium of the Sarawak Museum. The second specimen, Haviland 1813/1353, was collected 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 at an altitude of 1650 m; it likely represents 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...
. It is also deposited at the Herbarium of the Sarawak Museum. It does not include floral material. G. D. Haviland explored the Mount Kinabalu area with his brother H. A. Haviland between March and April, 1892, and must have collected these specimens during this time.
Additionally, Danser lists two specimens collected by Joseph Clemens
Joseph Clemens
Joseph Clemens was an American Methodist Episcopalian chaplain, missionary and plant collector who served and worked in South East Asia and elsewhere. He was born in the rugged western English county of Cornwall. Later, his family migrated to Williamsport, Pennsylvania, and then moved to...
in 1915. The first, Clemens 10627, was collected on November 13 from Paka Cave to Low's Peak. It includes female floral material. The second, Clemens 10871, was collected at Marai Parai between November 22 and November 23, and does not include floral material. Both specimens are deposited at the Bogor Botanical Gardens
Bogor Botanical Gardens
The Bogor Botanical Gardens are located 60 km south of the capital of Jakarta in Bogor, Indonesia. The botanical gardens are situated in the city center of Bogor and adjoin the Istana Bogor...
(formerly the Herbarium of the Buitenzorg Botanic Gardens) in Java
Java
Java is an island of Indonesia. With a population of 135 million , it is the world's most populous island, and one of the most densely populated regions in the world. It is home to 60% of Indonesia's population. The Indonesian capital city, Jakarta, is in west Java...
.
In Letts Guide to Carnivorous Plants of the World, published in 1992, a specimen of the natural hybrid N. × kinabaluensis (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. villosa) is labeled as N. villosa.
Description
Nepenthes villosa is a weak climber, rarely exceeding 60 cm in height, although the stem may grow to 8 m in length and 10 mm in diameter. InternodePlant 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 and up to 10 cm long.
Leaves are coriaceous and petiolate. The lamina is spathulate to oblong and may be up to 25 cm long and 6 cm wide. The apex of the lamina is emarginate. 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 canaliculate, up to 10 cm long, and bears an amplexicaul sheath. One to three longitudinal veins are present on either side of the midrib. 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 may reach 50 cm in length.
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 cylindrical in cross section and up to 20 mm wide. It bears well developed teeth and ribs. The 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 cordate and has a pointed apex. 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...
(≤20 mm long) is inserted at the base of the lid.
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...
may be up to 40 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...
grows to 20 cm in length. Pedicels
Pedicel (botany)
A pedicel is a stem that attaches single flowers to the main stem of the inflorescence. It is the branches or stalks that hold each flower in an inflorescence that contains more than one flower....
are filiform-bract
Bract
In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis, or cone scale. Bracts are often different from foliage leaves. They may be smaller, larger, or of a different color, shape, or texture...
eolate and 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 round to elliptic and up to 4 mm long. A study of 490 pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...
samples taken from two herbarium specimens (J.H.Adam 1124 and J.H.Adam 1190, collected at an altitude of 1800–3400 m) found the mean pollen diameter to be 37.2 μm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...
(SE
Standard error (statistics)
The standard error is the standard deviation of the sampling distribution of a statistic. The term may also be used to refer to an estimate of that standard deviation, derived from a particular sample used to compute the estimate....
= 0.2; CV
Coefficient of variation
In probability theory and statistics, the coefficient of variation is a normalized measure of dispersion of a probability distribution. It is also known as unitized risk or the variation coefficient. The absolute value of the CV is sometimes known as relative standard deviation , which is...
= 6.7%).
The species has 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 long, brown hairs that covers all parts of the plant.
Ecology
Nepenthes villosa is endemic to the upper slopes of Mount KinabaluMount 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....
. It generally grows at 2300–3240 m above sea level, the highest altitude of all Bornean Nepenthes species; only N. lamii
Nepenthes lamii
Nepenthes lamii is a tropical pitcher plant endemic to New Guinea, where it grows at an altitude of up to 3520 m above sea level, higher than any other Nepenthes species...
and N. sp. Papua
Nepenthes sp. Papua
Nepenthes sp. Papua, also known as Nepenthes sp. West-Papua and Nepenthes sp. Doormans Top 1, is an undescribed tropical pitcher plant known from a single mountain in central West Papua. The species is an extreme highlander, growing at elevations of 3100–3500 m above sea level, and has on occasion...
from New Guinea
New Guinea
New Guinea is the world's second largest island, after Greenland, covering a land area of 786,000 km2. Located in the southwest Pacific Ocean, it lies geographically to the east of the Malay Archipelago, with which it is sometimes included as part of a greater Indo-Australian Archipelago...
are found at greater elevations. On Mount Kinabalu, N. villosa is common along the Mesilau Trail (between Pondok Magnolia and the meeting point with the old summit trail) and almost all the way up to the Laban Rata rest house; a particularly large population has been reported at around 3047 m. On Mount Tambuyukon, an altitudinal inversion has been noted, whereby N. villosa is more common at much lower elevations of 1600–1900 m, being replaced by 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...
towards the summit. The exposed, uppermost slopes of Mount Tambuyukon can become very hot during the day and this might explain the inability of N. villosa to colonise them.
Nepenthes villosa often grows in mossy forest and sub-alpine forest dominated by species of the genera Dacrydium
Dacrydium
Dacrydium is a genus of conifers belonging to the podocarp family Podocarpaceae. Sixteen species of evergreen dioecious trees and shrubs are presently recognized. The genus was first described by Solander in 1786, and formerly included many more species, which were divided into sections A, B, and C...
and Leptospermum
Leptospermum
Leptospermum is a genus of about 80-86 species of plants in the myrtle family Myrtaceae. Most species are endemic to Australia, with the greatest diversity in the south of the continent; but one species extends to New Zealand, another to Malaysia, and L. recurvum is endemic to Malaysia.They...
, particularly Leptospermum recurvum
Leptospermum recurvum
Leptospermum recurvum is a species of Leptospermum native to Malaysia. It grows on Mount Kinabalu on ultramafic soil and is notable for being able to tolerate the high levels of toxic metals present in these soils....
. It has also been recorded growing among shrubs, grass, and boulders in open areas. Here the soil may become relatively dry, although 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 usually close to 100% as the slopes are often enveloped in clouds. Like many Nepenthes from the Mount Kinabalu area, it is endemic to ultramafic soils.
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. villosa is listed 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:...
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...
. Although many plants grow along Mount Kinabalu's summit trail and are easily accessible to climbers, all known populations of the species grow 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...
and so their collection is illegal. In 1997, 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...
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...
based on this. Clarke writes that N. villosa "has a secure future", although he adds that climbers have had a significant impact on populations of the species growing along the summit trail, with the number of plants having declined in recent years. A 2002 study found 1180 individual N. villosa growing in 11 plots, each measuring 0.01 hectare
Hectare
The hectare is a metric unit of area defined as 10,000 square metres , and primarily used in the measurement of land. In 1795, when the metric system was introduced, the are was defined as being 100 square metres and the hectare was thus 100 ares or 1/100 km2...
s, at elevations of between 2610 m and 2970 m on Mount Kinabalu. This number constituted 94% of the pitcher plants recorded from the plots, the rest being N. × kinabaluensis.
Related species
Nepenthes villosa is most closely related to N. edwardsianaNepenthes 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...
and N. macrophylla
Nepenthes macrophylla
Nepenthes macrophylla , the Large-Leaved Pitcher-Plant, is a tropical pitcher plant known only from a very restrictive elevation on Mount Trus Madi in Sabah, Malaysian Borneo...
. There has been much taxonomic confusion surrounding the status of these three 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...
.
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...
, who described both N. edwardsiana and N. villosa, noted the similarity between the two species as follows:
This most remarkable plant [N. villosa] resembles that of edwardsiana in so many respects, especially in the size, form and disposition of the distant lamellae of the mouth, that I am inclined to suspect that it may be produced by young plants of that species, before it arrives at a stage when the pitchers have elongated necks.
Günther Beck von Mannagetta und Lerchenau was the first to treat N. edwardsiana in synonymy with N. villosa when he published his monograph on the genus in 1895.
In his 1908 monograph, 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...
wrote the following with regards to the two species: "Examinatione microscopica probatur, illas species distinctas esse". This is probably "based on the old belief that plants, which differ anatomically, can not be forms of the same species".
B. H. Danser
B. H. Danser
Benedictus Hubertus Danser , often abbreviated B. H. Danser, was a Dutch taxonomist and botanist...
united the species "[w]ith some hesitation" in his 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...
". He suggested that N. villosa is a stunted form of N. edwardsiana from higher altitudes, which flowers at a "juvenile stage of development". Danser acknowledged that the 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 N. villosa is more dense than that of N. edwardsiana, but noted that it "is a difference only of degree".
The two taxa differ considerably in their altitudinal distributions. Nepenthes villosa usually occurs at ultrahighland elevations (2300–3240 m), whereas N. edwardsiana is found between 1500 and 2700 m. Where their altitudinal distributions overlap, they are still identifiable as distinct species.
Nepenthes macrophylla
Nepenthes macrophylla
Nepenthes macrophylla , the Large-Leaved Pitcher-Plant, is a tropical pitcher plant known only from a very restrictive elevation on Mount Trus Madi in Sabah, Malaysian Borneo...
was originally described in 1987 as a subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...
of N. edwardsiana by Johannes Marabini. It was later elevated to species status by Matthew Jebb
Matthew Jebb
Dr. Matthew H. P. Jebb is an Irish taxonomist and botanist specialising in the ant plant genera Squamellaria, Myrmecodia, Hydnophytum, Myrmephytum and Anthorrhiza, as well as the carnivorous plant genus Nepenthes....
and Martin Cheek
Martin Cheek
Dr. Martin Roy Cheek is a taxonomist and botanist specialising in the carnivorous plant genus Nepenthes.-Research:Cheek has described several new Nepenthes species, mostly with Matthew Jebb, including: N. argentii, N. aristolochioides, N. danseri, N. diatas,...
. This interpretation was supported 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...
, who noted that N. edwardsiana and N. villosa "have more in common" than N. edwardsiana and N. macrophylla. Whereas N. edwardsiana and N. villosa are restricted to the Kinabalu area, N. macrophylla is only found near the summit of Mount Trus Madi
Mount Trus Madi
Mount Trus Madi or Trusmadi is Malaysia's second highest mountain at . It lies in the state of Sabah, close to Mount Kinabalu. The mountain supports a wide range of unique flora and fauna, perhaps most notably Nepenthes macrophylla, a species of pitcher plant.The natural hybrid Nepenthes ×...
.
Matthew Jebb and Martin Cheek suggest that N. villosa is related to N. mira
Nepenthes mira
Nepenthes mira is a highland pitcher plant endemic to Palawan in the Philippines. It grows at elevations of 1550–1605 m above sea level....
, a species endemic to Palawan
Palawan
Palawan is an island province of the Philippines located in the MIMAROPA region or Region 4. Its capital is Puerto Princesa City, and it is the largest province in the country in terms of total area of jurisdiction. The islands of Palawan stretch from Mindoro in the northeast to Borneo in the...
in the Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...
. N. villosa also shows affinities to N. peltata
Nepenthes peltata
Nepenthes peltata is a tropical pitcher plant known only from the upper slopes of Mount Hamiguitan on the island of Mindanao in the Philippines. It is characterised by a peltate tendril attachment and conspicuous indumentum...
of Mindanao
Mindanao
Mindanao is the second largest and easternmost island in the Philippines. It is also the name of one of the three island groups in the country, which consists of the island of Mindanao and smaller surrounding islands. The other two are Luzon and the Visayas. The island of Mindanao is called The...
.
Natural hybrids
Two natural hybrids involving N. villosa have been recorded.N. edwardsiana × N. villosa
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...
and N. villosa. Its two parent species are very closely related and so N. × harryana, which is intermediate in form, may be difficult to distinguish from either of them.
It was originally described as a species in 1882 by 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...
. 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...
was the first to realise its hybrid origin and described it as such in his monograph of 1908. Danser wrote that N. × harryana could be a hybrid as Macfarlane suggested, or a form of N. villosa together with N. edwardsiana.
Nepenthes × harryana can be distinguished from N. villosa on the basis of its pitcher morphology. The pitchers of the hybrid are more cylindrical than those of N. villosa, whereas the 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...
is more dense than that of N. edwardsiana. The hip of the pitcher cup, which is found just below the peristome in N. villosa and in the lower quarter of N. edwardsiana pitchers, is located around the middle of N. × harryana pitchers. However, N. villosa plants from 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....
are easier to confuse with this hybrid, as they produce pitchers that may be elongated slightly above the hip.
Nepenthes × harryana is known from a ridge above the Upper Kolopis River
Kolopis River
The Kolopis River is one of the major rivers that flows through Kinabalu National Park. An area adjacent to the upper Kolopis River is home to a number of pitcher plants of the genus Nepenthes, including N. edwardsiana, N. rajah, and N. villosa, as well as two natural hybrids...
and from two locations along the Kinabalu summit trail. Since N edwardsiana does not grow along the summit trail, it cannot be confused with this hybrid there. Burbidge wrote that N. edwardsiana, N. × harryana, and N. villosa "are quite distinct in zone of the mountain".
N. rajah × N. villosa
Nepenthes × kinabaluensis is the natural hybrid between N. rajahNepenthes 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. villosa. It was first collected near Kambarangoh
Kambarangoh
Kambarangoh is an area along the summit trail to Low's Peak on Mount Kinabalu in Sabah, Borneo. It lies between the Power Station and Layang-Layang. It is named after the only telecommunications station on the mountain, which is owned by Kambarangoh Telecoms.The road leading up to the Power...
on Mount Kinabalu by Lilian Gibbs
Lilian Gibbs
Lilian Suzette Gibbs was a British botanist who worked for the British Museum in London. She was the first woman and the first botanist to ascend Mount Kinabalu in February 1910. Gibbs collected many plants new to science, several of which are named in her honour .-References:* Vickery, R. ....
in 1910 and later mentioned by 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...
as "Nepenthes sp." in 1914. Although Macfarlane did not formally name the plant, he noted that "[a]ll available morphological details suggest that this is a hybrid between N. villosa and N. rajah". It was finally described in 1976 by Shigeo Kurata
Shigeo Kurata
is a Japanese botanist and Nepenthes taxonomist whose work in the 1960s and 1970s contributed much to the current popularity of these plants. Of particular note is his 1976 guide, Nepenthes of Mount Kinabalu....
as N. × kinabaluensis. The name was published in Nepenthes of Mount Kinabalu
Nepenthes of Mount Kinabalu
Nepenthes of Mount Kinabalu is a monograph by Shigeo Kurata on the tropical pitcher plants of Mount Kinabalu and the surrounding area of Kinabalu National Park in Sabah, Borneo. It was published in 1976 by Sabah National Parks Trustees as the second booklet of the Sabah National Parks series...
, but it is a nomen nudum
Nomen nudum
The phrase nomen nudum is a Latin term, meaning "naked name", used in taxonomy...
, as it had an inadequate description and lacked information on the type specimen
Holotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
. The name was subsequently republished by Kurata in 1984 and 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. ×...
in 1998.
The pitchers of N. × kinabaluensis may be quite large, but do not compare to those of N. rajah or N. × alisaputrana (N. burbidgeae
Nepenthes burbidgeae
Nepenthes burbidgeae , also known as the painted pitcher plant or Burbidge's Pitcher-Plant, is a tropical pitcher plant with a patchy distribution around Mount Kinabalu and neighbouring Mount Tambuyukon in Sabah, Borneo.-Botanical history:...
× N. rajah). N. × kinabaluensis can only be found on Mount Kinabalu (hence the name) and nearby Mount Tambuyukon, where the two parent species occur sympatrically
Sympatric speciation
Sympatric speciation is the process through which new species evolve from a single ancestral species while inhabiting the same geographic region. In evolutionary biology and biogeography, sympatric and sympatry are terms referring to organisms whose ranges overlap or are even identical, so that...
. More specifically, plants are known from a footpath near Paka Cave and several places along an unestablished route on a south-east ridge, which lies on the west side of the Upper Kolopis River
Kolopis River
The Kolopis River is one of the major rivers that flows through Kinabalu National Park. An area adjacent to the upper Kolopis River is home to a number of pitcher plants of the genus Nepenthes, including N. edwardsiana, N. rajah, and N. villosa, as well as two natural hybrids...
. The only accessible location from which this hybrid is known is the Kinabalu summit trail, between Layang-Layang and the helipad
Helipad
Helipad is a common abbreviation for helicopter landing pad, a landing area for helicopters. While helicopters are able to operate on a variety of relatively flat surfaces, a fabricated helipad provides a clearly marked hard surface away from obstacles where a helicopter can safely...
, where it grows at about 2900 m in a clearing dominated by Dacrydium gibbsiae
Dacrydium gibbsiae
Dacrydium gibbsiae is a conifer species native to Malaysia. It grows on Mount Kinabalu on ultramafic soil and is notable for being able to tolerate the high levels of toxic metal compounds present in these soils....
and Leptospermum recurvum
Leptospermum recurvum
Leptospermum recurvum is a species of Leptospermum native to Malaysia. It grows on Mount Kinabalu on ultramafic soil and is notable for being able to tolerate the high levels of toxic metals present in these soils....
trees. N. × kinabaluensis has an altitudinal distribution of 2420 to 3030 m. It grows in open areas in cloud forest.
The hybrid is generally intermediate in appearance between its parent species. Raised ribs line the inner edge of the peristome and end with elongated teeth. These are more prominent than those found in N. rajah and smaller than those of N. villosa. The peristome is coarse and expanded at the margin (but not scalloped like that of N. rajah), the lid orbiculate or reniform and almost flat. In general, pitchers are larger than those of N. villosa and the tendril joins the apex about 1–2 cm below the leaf tip, a feature which is characteristic of N. rajah. In older plants, the tendril can be almost woody. N. × kinabaluensis is an indumentum of villous hairs covering the pitchers and leaf margins, which is approximately intermediate between the parents. Lower pitchers have two fringed wings, whereas the upper pitchers usually lack these. The colour of the pitcher varies from yellow to scarlet. N. × kinabaluensis seems to produce upper pitchers more readily than either of its parents. In all respects N. × kinabaluensis is intermediate between the two parent species and it is easy to distinguish from all other Nepenthes of Borneo. However, it has been confused once before, when the hybrid was identified as N. rajah in Letts Guide to Carnivorous Plants of the World.
Further reading
- Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. Journal of Tropical Forest Science 5(1): 13–25.
- Adam, J.H. 1997. Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.
- Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.
- Clarke, C.M. 2006. Introduction. In: Danser, B.H. The Nepenthaceae of the Netherlands IndiesThe 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...
. Natural History Publications (Borneo), Kota Kinabalu. pp. 1–15. - Denton, B. 2001. Mt. Kinabalu - much more than Nepenthes utopia. Bulletin of the Australian Carnivorous Plant SocietyBulletin of the Australian Carnivorous Plant SocietyThe Bulletin of the Australian Carnivorous Plant Society was a quarterly periodical and the official publication of the Australian Carnivorous Plant Society. Established in April 1982 as Bulletin / South Australian Carnivorous Plant Society, it continued publication until 2003...
20(3): 1–4. Mansur, M. 2001. In: Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253. - Sacilotto, R. 2004. Experiments with highland Nepenthes seedlings: a summary of measured tolerances. Carnivorous Plant NewsletterCarnivorous Plant NewsletterThe Carnivorous Plant Newsletter is the official publication of the International Carnivorous Plant Society , the largest such organization in the world.-History and editorship:...
33(1): 26–31. - Shafer, J. 2003. Carnivorous Plant NewsletterCarnivorous Plant NewsletterThe Carnivorous Plant Newsletter is the official publication of the International Carnivorous Plant Society , the largest such organization in the world.-History and editorship:...
32(1): 20–23. - Thong, J. 2006. Victorian Carnivorous Plant Society JournalVictorian Carnivorous Plant Society JournalThe Victorian Carnivorous Plant Society Journal is a quarterly periodical and the official publication of the Victorian Carnivorous Plant Society of Australia. Typical articles include matters of horticultural interest, field reports, literature reviews, and plant descriptions. The journal was...
81: 12–17. - Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York.
External links
- Danser, B.H. 1928. 51. Nepenthes villosa HOOK. F. In: The Nepenthaceae of the Netherlands IndiesThe 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...
. Bulletin du Jardin Botanique de Buitenzorg, Série III, 9(3–4): 249–438.