Nepenthes bicalcarata
Encyclopedia
Nepenthes bicalcarata also known as the Fanged Pitcher-Plant, is a tropical pitcher plant
endemic to northwestern Borneo
.
Nepenthes bicalcarata was formally described
by Joseph Dalton Hooker
in his 1873 monograph, "Nepenthaceae
", based on specimens collected by Hugh Low
and Odoardo Beccari
near Lawas River in Borneo. The type specimen, Low s.n., is deposited at the Royal Botanic Gardens, Kew
. Seven years later, Spencer Le Marchant Moore
described Nepenthes dyak, based on a specimen collected by Johannes Elias Teijsmann
, Teijsmann 10962, from Kapuas River
near Sintang in western Borneo. This specimen is also held at the Royal Botanic Gardens, Kew and a duplicate is deposited at the National Herbarium of the Netherlands
in Leiden. Nepenthes dyak was later mentioned several more times in the botanical literature, but is now considered conspecific with N. bicalcarta.
Nepenthes bicalcarata was introduced to Europe in 1879 by British explorer Frederick William Burbidge
, who collected plants for the famous Veitch Nursery. These were cultivated to larger size and distributed in 1881.
During this time, interest in Nepenthes had reached its peak. A note in The Gardeners' Chronicle
of 1881 mentions the Veitch Nursery's N. bicalcarata as follows:
Several years after its introduction, N. bicalcarata was still very much a horticultural rarity. In Veitch's catalogue for 1889, N. bicalcarata was priced at £
3.3s
per plant, while the famous giant-pitchered N. northiana
and N. rajah
were selling for £2.2s.
. The cylindrical stem is thicker than that of any other Nepenthes species, measuring up to 3.5 cm in diameter. Internode
s are up to 40 cm long.
The leaves of N. bicalcarata are petiolate and coriaceous in texture. The lamina is obovate-lanceolate in form and also reaches huge dimensions, growing to 80 cm in length and 12 cm in width. It is slightly decurrent on the stem, forming two narrow wings. The lamina has indistinct longitudinal veins and numerous pennate veins. Tendril
s may be up to 60 cm long and 8 mm wide. They are hollow and swollen near the pitcher.
Although most parts of the plant are very large, the pitchers themselves do not rival those of species such as N. rajah
. Nevertheless, they may have a volume of over one litre and grow up to 25 cm high and 16 cm wide. A pair of prominent fringed wings (≤15 mm wide) run down the front of lower pitchers. These are usually reduced to ribs in aerial pitchers. The peristome
(≤20 mm wide) is characteristically flattened and curved inwards. It bears small but distinct teeth. The two sharp spines for which the species is famous are present on the underside of the pitcher lid, and may be 3 cm long. They are derived from the uppermost 10–12 peristome ribs. The pitcher lid or operculum
is reniform to cordate and has no appendages. An unbranched spur
(≤15 mm long) is inserted at the base of the lid.
Nepenthes bicalcarata has a paniculate
inflorescence
. The peduncle
may be up to 40 cm long and the rachis
can reach 100 cm in length. Female inflorescences are usually shorter. Branches on the flower stem are up to 40 mm long and bear up to 15 flowers. Sepal
s are either obovate or lanceolate and up to 4 mm long. A study of 120 pollen
samples taken from a herbarium specimen (Fosberg 43860, altitude not recorded) found the mean pollen diameter to be 28.9 μm
(SE
= 0.4; CV
= 7.5%).
Mature plants are virtually glabrous. Caducous hairs are present on the youngest parts of the plant and on the inflorescences.
Nepenthes bicalcarata varies little across its range. Consequently, no infraspecific taxa have been described.
Nepenthes bicalcarata is endemic to Borneo
. It is most common in the peat swamp forests
of the western coast of the island, which stretch across Sarawak
, Sabah
, Kalimantan
, and Brunei
. There it often grows in the shade of the ubiquitous dipterocarp Shorea albida
. Nepenthes bicalcarata also occurs in kerangas forest
and has even been recorded from white sand heath forest
s in Sarawak
and East Kalimantan
. The species is often sympatric with N. ampullaria
in these habitats.
Specimens growing in undisturbed peat swamp forest, where sunlight is greatly diffused and high humidity prevails, reach the largest dimensions. Nepenthes bicalcarata has a shallow root system that only penetrates the top layer of peat
and leaf litter, to a depth of about 25 cm. Below this, high concentrations of tannin
s and alkaloid
s render the substrate toxic.
Nepenthes bicalcarata is generally found below 300 m in altitude, although Johannes Gottfried Hallier
reported a single collection in 1894 from between 700 and 950 m above sea level.
The conservation status
of N. bicalcarata is listed as Vulnerable
on the 2006 IUCN Red List of Threatened Species
based on an assessment carried out in 2000. In 1997, Charles Clarke
informally classified the species as Near Threatened
based on the IUCN
criteria. This agrees with the conservation status assigned to N. bicalcarata by the World Conservation Monitoring Centre
.
The two thorns that give N. bicalcarata its name are unique to this species and bear some of the largest nectaries in the plant kingdom. The purpose of these structures has long been debated among botanists. Frederick William Burbidge
suggested that they might serve to deter arboreal mammal
s such as tarsier
s, loris
es and monkey
s from stealing the contents of the pitchers. In an article published in 1982, Cliff Dodd speculated on the function of the thorns, but did not believe they play a role in prey capture. Charles Clarke
observed that monkeys and tarsiers rip the pitchers open at the sides in order to feed from them, rather than reaching in through the pitcher mouth. However, he found that the mammals attacked the pitchers of N. bicalcarata less frequently than those of other species, such as N. rafflesiana
. Clarke's observations suggest that the spines likely serve to lure insects into a precarious position over the pitcher mouth, where they may lose their footing and fall into the pitcher fluid, eventually drowning.
A similar trapping method is employed by the Sumatra
n endemic N. lingulata
, which has a single filiform appendage positioned over the pitcher mouth. However, in that species the appendage is a structure of the lid and not the peristome.
Together with N. ampullaria
and N. ventricosa
, N. bicalcarata is unusual in that the glandular region of the pitcher extends almost to the peristome, such that there is little or no conductive waxy zone. The waxy zone functions by causing prey to slip and fall into the digestive fluid. A 2004 study found that the peristome of N. bicalcarata plays a very important role in prey capture. When it is dry, the peristome is very ineffective in catching insect prey, but when wetted, the capture rate increases more than three-fold.
A 1999 study of pitcher morphology and spectral reflectance characteristics showed that N. bicalcarata has no colour contrast maximum between the peristome and pitcher cup in the ultraviolet
waveband, but small maxima of –0.17 at 450 nm (violet
), 0.32 at 548 nm (green
), and –0.16 at 668 nm (red
). This means that the peristome is less reflective than the pitcher cup in the violet and red bands, but more reflective in the green band. The contrast maxima in the green and blue regions seem to correspond to insect visual sensitivity maxima, while the others do not. Based on their findings, the authors made the "tentative prediction" that the upper pitchers of N. bicalcarata are less successful at catching anthophilous (flower-visiting) prey than those of sympatric N. rafflesiana.
that makes its nest in the plant's hollow tendrils. Described as Camponotus schmitzi
in 1933, it is a member of the extremely populous and widespread genus of carpenter ant
s.
This unique animal-plant interaction was noted by Frederick William Burbidge
as early as 1880. In 1904, Odoardo Beccari
suggested that the ants feed on insects found on and around the plant, but may fall prey to it themselves. In 1990, B. Hölldobler and E.O. Wilson proposed that N. bicalcarata and C. schmitzi form a mutually beneficial association. At the time, however, no experimental data existed to support such a hypothesis. A series of observations and experiments carried out in Brunei
by Charles Clarke
in 1992 and 1998, and by Clarke and Roger Kitching in 1993 and 1995, strongly support the mutualism theory. John Thompson has suggested that N. bicalcarata may be the only plant species that obtains nutrients through both insect capture and ant-hosting habits.
Camponotus schmitzi ants feed by descending into the pitcher fluid and retrieving arthropod
s caught by the plant. They seem to ignore smaller insects and only target larger prey items. Hauling food from the pitcher fluid to the peristome—a distance of no more than 5 cm—may take up to 12 hours. In this way the contents of N. bicalcarata pitchers is controlled such that organic matter does not accumulate to the point of putrefaction
, which could lead to the demise of pitcher infauna
(which also appear to benefit the plant) and sometimes the pitcher itself. The ants have also been observed to attack newly caught insects and therefore prevent prey escape. At other times, the ants are very passive, remaining hidden under the inner peristome fold, presumably so as not to dissuade visitation by potential prey species. This behaviour is in stark contrast to other myrmecophytic
ants, which are typically highly territorial. In order to create a favourable environment for its pitcher inhabitants, it appears that N. bicalcarata actively maintains the pH
of the pitcher fluid at a less acidic level than that found in most other Nepenthes species. This might explain the occasional presence of tree frog
eggs, which are sensitive to strongly acidic conditions, in the pitchers of this species.
Research conducted by Dennis and Marlis Merbach has shown that C. schmitzi is also beneficial to N. bicalcarata in that it protects it from pitcher-destroying weevil
s of the genus Alcidodes.
The ants seem to favour upper pitchers and rarely colonise lower pitchers. This is likely due to the fact that terrestrial traps are periodically submerged in water during heavy rains. Flooding of the ants' nest chamber could result in the death of the developing eggs, larva
e, and pupa
e. Camponotus schmitzi nests solely in the tendrils of N. bicalcarata and rarely ventures onto other plants. The species is completely dependent on N. bicalcarata for food and domicile. Nepenthes bicalcarata, on the other hand, is able to survive and reproduce without the presence of the ants; it is a facultative mutualist. This being the case, there appear to be few mature plants over 2 metres in height not colonised by C. schmitzi.
A species of mite
, Naiadacarus nepenthicola, appears to be restricted to the pitchers of N. bicalcarata. It is thought to feed on decomposing leaves and insects that are caught in the pitchers. Deutonymphs of this mite are dispersed through phoresy on C. schmitzi.
None of these hybrids are known to be colonised by C. schmitzi as is the parent species N. bicalcarata.
Since N. ampullaria and N. bicalcarata are often sympatric in the wild, it is not surprising that they occasionally hybridise. Several examples of this natural hybrid are known from Brunei
, where it grows in peat swamp forest
and heath forest
. It was first recorded by Mitsuru Hotta
in 1966.
Pitchers bear the characteristic thorns of N. bicalcarata, although they are greatly reduced in size. The mouth is round and the lid is ovate-cordate in form. This hybrid resembles N. × hookeriana
to a certain extent, but differs in having spines on the underside of the lid and more globose upper pitchers with a smaller lid.
Aerial pitchers of N. ampullaria × N. bicalcarata are usually dominated by characteristics of N. ampullaria. They are often very small and unable to function in a normal manner.
. It produces small speckled pitchers with reduced thorns under the lid. The plant is intermediate in appearance between N. × trichocarpa and N. bicalcarata.
This hybrid has been named N. × cantleyi after Rob Cantley
. The morphology of the pitchers closely resembles N. bicalacarata, although the peristome is more similar to that of N. gracilis. The characteristic spines of N. bicalcarata are greatly reduced and are only present as small bumps. This hybrid has the growth habit of N. gracilis, with the stem scrambling along the ground. It grows in open, sandy areas. The pitcher fluid is notably acid
ic like that of N. gracilis, with pH
values as low as 1.82 being recorded.
, where it involves N. mirabilis var. echinostoma. It closely resembles N. bicalcarata, although it differs from that species in having reduced thorns and a wider peristome. Unlike N. bicalcarata, this hybrid inhabits open areas with sandy substrates.
reported a single plant displaying characters intermediate in form between N. bicalcarata, N. rafflesiana and N. mirabilis var. echinostoma growing among populations of these species. It is likely a complex hybrid, although this cannot be proven without the use of genetic testing
.
Nepenthes bicalcarata may be cultivated in a similar manner to other members of the genus, though it requires relatively hot and humid conditions. It needs large pots to do well, and temporary halts in growth are often due to becoming root-bound. Re-potting to a larger pot will often result in a rapid size increase. Under satisfactory conditions, this plant will grow quite fast for a Nepenthes, reaching a large size relatively rapidly.
Despite naturally occurring in peat swamp forests, N. bicalcarata has been successfully grown in a completely inorganic substrate consisting of one part each of Seramis clay perls, lava gravel, and Lecaton expanded clay perls.
Nepenthes bicalcarata has been artificially crossed with N. hirsuta
. A particularly robust-pitchered cultivar
of this hybrid was named Nepenthes 'Hortulanus Otten', in honour of Karel Otten, former curator of the Botanic Garden in Ghent
, Belgium
.
A cultivar of the artificial cross N. bicalcarata × N. × dyeriana was registered in 1988 under the name Nepenthes 'Nina Dodd'. It is named after a relative of Cliff Dodd, who created the hybrid.
Two further cultivars of manmade hybrids involving N. bicalcarata have been named. These are Nepenthes 'Bella' ((N. ampullaria
× N. bicalcarata) × N. truncata
) and Nepenthes 'Rapa' (N. ampullaria × N. bicalcarata). However, both of these names are not established, as they were published without a description.
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 northwestern 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
Species 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...
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...
in his 1873 monograph, "Nepenthaceae
Nepenthaceae (1873 monograph)
"Nepenthaceae" is a monograph by Joseph Dalton Hooker on the tropical pitcher plants of the genus Nepenthes. It was published in 1873 in the seventeenth and final volume of Augustin Pyramus de Candolle's Prodromus Systematis Naturalis Regni Vegetabilis, which was edited by Augustin's son, Alphonse...
", based on specimens collected 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 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...
near Lawas River in Borneo. The type specimen, Low s.n., 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...
. Seven years later, Spencer Le Marchant Moore
Spencer Le Marchant Moore
Spencer Le Marchant Moore was an English botanist.Moore was born in Hampstead. He worked at the Royal Botanic Gardens, Kew from about 1870 to 1879, wrote a number of botanical papers, and then worked in an unofficial capacity at the Natural History Museum from 1896 until his death.Moore is...
described Nepenthes dyak, based on a specimen collected by Johannes Elias Teijsmann
Johannes Elias Teijsmann
Johannes Elias Teijsmann was a botanist and plant collector. He was born in Arnhem, Netherlands.Teijsmann travelled to Java in 1830 as Gardener of Governor General van den Bosch. He was appointed the Curator of the Buitenzorg Botanic Gardens the following year, a post he held until 1869...
, Teijsmann 10962, from Kapuas River
Kapuas River
The Kapuas River is a river in the Indonesian part of Borneo island, at the geographic center of Maritime Southeast Asia. At in length, it is the longest river of Indonesia and one of the world's longest island rivers...
near Sintang in western Borneo. This specimen is also held at the Royal Botanic Gardens, Kew and a duplicate is deposited at the National Herbarium of the Netherlands
National Herbarium of the Netherlands
The National Herbarium of the Netherlands was established in 1999 through a decentralized merger of the major university herbaria of Leiden , Utrecht and Wageningen...
in Leiden. Nepenthes dyak was later mentioned several more times in the botanical literature, but is now considered conspecific with N. bicalcarta.
Nepenthes bicalcarata was introduced to Europe in 1879 by British explorer 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...
, who collected plants for the famous Veitch Nursery. These were cultivated to larger size and distributed in 1881.
During this time, interest in Nepenthes had reached its peak. A note in 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....
of 1881 mentions the Veitch Nursery's N. bicalcarata as follows:
"Then there is N. bicalcarata, a most robust habited kind with sturdy foliage and bag-like pitchers provided with a vicious-looking rat-trap-like apparatus in its lid which renders it very distinct from its neighbours."
Several years after its introduction, N. bicalcarata was still very much a horticultural rarity. In Veitch's catalogue for 1889, N. bicalcarata was priced at £
Pound sterling
The pound sterling , commonly called the pound, is the official currency of the United Kingdom, its Crown Dependencies and the British Overseas Territories of South Georgia and the South Sandwich Islands, British Antarctic Territory and Tristan da Cunha. It is subdivided into 100 pence...
3.3s
Shilling
The shilling is a unit of currency used in some current and former British Commonwealth countries. The word shilling comes from scilling, an accounting term that dates back to Anglo-Saxon times where it was deemed to be the value of a cow in Kent or a sheep elsewhere. The word is thought to derive...
per plant, while the famous giant-pitchered N. northiana
Nepenthes northiana
Nepenthes northiana , or Miss North's Pitcher-Plant, is a tropical pitcher plant endemic to Borneo, where it grows at elevations ranging from 0 to 500 m above sea level. The specific epithet northiana honours Marianne North, who first illustrated the species...
and 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...
were selling for £2.2s.
Description
Nepenthes bicalcarata plants are the largest in the genus, climbing up to 20 m into the forest canopyCanopy (forest)
In biology, the canopy is the aboveground portion of a plant community or crop, formed by plant crowns.For forests, canopy also refers to the upper layer or habitat zone, formed by mature tree crowns and including other biological organisms .Sometimes the term canopy is used to refer to the extent...
. The cylindrical stem is thicker than that of any other Nepenthes species, measuring up to 3.5 cm 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 up to 40 cm long.
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 be up to 60 cm long and 8 mm wide. They are hollow and swollen near the pitcher.
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...
. Nevertheless, they may have a volume of over one litre and grow up to 25 cm high and 16 cm wide. A pair of prominent fringed wings (≤15 mm wide) run down the front of lower pitchers. These are usually reduced to ribs in aerial pitchers. 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...
(≤20 mm wide) is characteristically flattened and curved inwards. It bears small but distinct teeth. The two sharp spines for which the species is famous are present on the underside of the pitcher lid, and may be 3 cm long. They are derived from the uppermost 10–12 peristome ribs. 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 reniform to cordate and has no appendages. 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...
(≤15 mm long) is inserted at the base of the lid.
Nepenthes bicalcarata has a paniculate
Panicle
A panicle is a compound raceme, a loose, much-branched indeterminate inflorescence with pedicellate flowers attached along the secondary branches; in other words, a branched cluster of flowers in which the branches are racemes....
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 and 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...
can reach 100 cm in length. Female inflorescences are usually shorter. Branches on the flower stem are up to 40 mm long and bear up to 15 flowers. 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 either obovate or lanceolate and up to 4 mm long. A study of 120 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 a herbarium specimen (Fosberg 43860, altitude not recorded) found the mean pollen diameter to be 28.9 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....
= 0.4; 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...
= 7.5%).
Mature plants are virtually glabrous. Caducous hairs are present on the youngest parts of the plant and on the inflorescences.
Nepenthes bicalcarata varies little across its range. Consequently, no infraspecific taxa have been described.
Ecology and conservation status
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 is most common in the peat swamp forests
Borneo peat swamp forests
The Borneo peat swamp forests ecoregion, within the Tropical and subtropical moist broadleaf forests Biome, are on the island of Borneo, which is divided between Brunei, Indonesia and Malaysia.-Location and description:...
of the western coast of the island, which stretch across Sarawak
Sarawak
Sarawak is one of two Malaysian states on the island of Borneo. Known as Bumi Kenyalang , Sarawak is situated on the north-west of the island. It is the largest state in Malaysia followed by Sabah, the second largest state located to the North- East.The administrative capital is Kuching, which...
, 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...
, Kalimantan
Kalimantan
In English, the term Kalimantan refers to the Indonesian portion of the island of Borneo, while in Indonesian, the term "Kalimantan" refers to the whole island of Borneo....
, and Brunei
Brunei
Brunei , officially the State of Brunei Darussalam or the Nation of Brunei, the Abode of Peace , is a sovereign state located on the north coast of the island of Borneo, in Southeast Asia...
. There it often grows in the shade of the ubiquitous dipterocarp Shorea albida
Shorea albida
Shorea albida is a species of plant in the Dipterocarpaceae family. It is found in Brunei, Indonesia, and Malaysia.-Source:* Ashton, P. 1998. . Downloaded on 23 August 2007....
. Nepenthes bicalcarata also occurs in kerangas forest
Kerangas forest
Sundaland heath forest, also known as Kerangas forest, is a type of tropical moist forest found on the island of Borneo, which is divided between Brunei, Indonesia, and Malaysia, as well as on the Indonesian islands of Belitung and Bangka, which lie to the west of Borneo.-Setting:The word Kerangas,...
and has even been recorded from white sand heath forest
Heath forest
Heath forest is a type of tropical moist forest found in areas with acidic, sandy soils that are extremely nutrient-poor. Notable examples are the Rio Negro campinarana of the Amazon Basin in South America, and the Sundaland heath forests of Borneo and neighboring islands.-External links:**...
s in Sarawak
Sarawak
Sarawak is one of two Malaysian states on the island of Borneo. Known as Bumi Kenyalang , Sarawak is situated on the north-west of the island. It is the largest state in Malaysia followed by Sabah, the second largest state located to the North- East.The administrative capital is Kuching, which...
and East Kalimantan
East Kalimantan
East Kalimantan is the second largest Indonesian province, located on the Kalimantan region on the east of Borneo island. The resource-rich province has two major cities, Samarinda and Balikpapan...
. The species is often sympatric with N. ampullaria
Nepenthes ampullaria
Nepenthes ampullaria , the Flask-Shaped Pitcher-Plant, is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea...
in these habitats.
Specimens growing in undisturbed peat swamp forest, where sunlight is greatly diffused and high humidity prevails, reach the largest dimensions. Nepenthes bicalcarata has a shallow root system that only penetrates the top layer of peat
Peat
Peat is an accumulation of partially decayed vegetation matter or histosol. Peat forms in wetland bogs, moors, muskegs, pocosins, mires, and peat swamp forests. Peat is harvested as an important source of fuel in certain parts of the world...
and leaf litter, to a depth of about 25 cm. Below this, high concentrations of tannin
Tannin
A tannin is an astringent, bitter plant polyphenolic compound that binds to and precipitates proteins and various other organic compounds including amino acids and alkaloids.The term tannin refers to the use of...
s and alkaloid
Alkaloid
Alkaloids are a group of naturally occurring chemical compounds that contain mostly basic nitrogen atoms. This group also includes some related compounds with neutral and even weakly acidic properties. Also some synthetic compounds of similar structure are attributed to alkaloids...
s render the substrate toxic.
Nepenthes bicalcarata is generally found below 300 m in altitude, although Johannes Gottfried Hallier
Johannes Gottfried Hallier
Johannes Gottfried Hallier was a German botanist who was a native of Jena.He studied botany and zoology at the University of Jena under Christian Ernst Stahl and Ernst Haeckel , and continued his studies at the University of Munich under Ludwig Radlkofer and Richard Hertwig...
reported a single collection in 1894 from between 700 and 950 m above sea level.
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. bicalcarata 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...
based on an assessment carried out in 2000. 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...
informally classified the species as Near Threatened
Near Threatened
Near Threatened is a conservation status assigned to species or lower taxa that may be considered threatened with extinction in the near future, although it does not currently qualify for the threatened status...
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. This agrees with the conservation status assigned to N. bicalcarata by the World Conservation Monitoring Centre
World Conservation Monitoring Centre
The United Nations Environment Programme's World Conservation Monitoring Centre is an executive agency of the United Nations Environment Programme, based in Cambridge in the United Kingdom. UNEP-WCMC has been part of UNEP since 2000, and has responsibility for biodiversity assessment and support...
.
Carnivory
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...
suggested that they might serve to deter arboreal mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...
s such as tarsier
Tarsier
Tarsiers are haplorrhine primates of the genus Tarsius, a genus in the family Tarsiidae, which is itself the lone extant family within the infraorder Tarsiiformes...
s, loris
Loris
Loris is the common name for the strepsirrhine primates of the subfamily Lorisinae in family Lorisidae. Loris is one genus in this subfamily and includes the slender lorises, while Nycticebus is the genus for the slow lorises....
es and monkey
Monkey
A monkey is a primate, either an Old World monkey or a New World monkey. There are about 260 known living species of monkey. Many are arboreal, although there are species that live primarily on the ground, such as baboons. Monkeys are generally considered to be intelligent. Unlike apes, monkeys...
s from stealing the contents of the pitchers. In an article published in 1982, Cliff Dodd speculated on the function of the thorns, but did not believe they play a role in prey capture. 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...
observed that monkeys and tarsiers rip the pitchers open at the sides in order to feed from them, rather than reaching in through the pitcher mouth. However, he found that the mammals attacked the pitchers of N. bicalcarata less frequently than those of other species, such as N. rafflesiana
Nepenthes rafflesiana
Nepenthes rafflesiana , or Raffles' Pitcher-Plant, is a species of pitcher plant. It has a very wide distribution covering Borneo, Sumatra, Peninsular Malaysia, and Singapore. Nepenthes rafflesiana is extremely variable with numerous forms and varieties described. In Borneo alone, there are at...
. Clarke's observations suggest that the spines likely serve to lure insects into a precarious position over the pitcher mouth, where they may lose their footing and fall into the pitcher fluid, eventually drowning.
A similar trapping method is employed by the Sumatra
Sumatra
Sumatra is an island in western Indonesia, westernmost of the Sunda Islands. It is the largest island entirely in Indonesia , and the sixth largest island in the world at 473,481 km2 with a population of 50,365,538...
n endemic N. lingulata
Nepenthes lingulata
Nepenthes lingulata is a tropical pitcher plant endemic to northern Sumatra. The species is characterised by the highly developed appendage present on the underside of the lid...
, which has a single filiform appendage positioned over the pitcher mouth. However, in that species the appendage is a structure of the lid and not the peristome.
Together with N. ampullaria
Nepenthes ampullaria
Nepenthes ampullaria , the Flask-Shaped Pitcher-Plant, is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea...
and N. ventricosa
Nepenthes ventricosa
Nepenthes ventricosa is a tropical pitcher plant endemic to the Philippines, where it is a highland species, growing at an elevation of 1000–2000 m above sea level. It has been recorded from the islands of Luzon, Panay, and Sibuyan...
, N. bicalcarata is unusual in that the glandular region of the pitcher extends almost to the peristome, such that there is little or no conductive waxy zone. The waxy zone functions by causing prey to slip and fall into the digestive fluid. A 2004 study found that the peristome of N. bicalcarata plays a very important role in prey capture. When it is dry, the peristome is very ineffective in catching insect prey, but when wetted, the capture rate increases more than three-fold.
A 1999 study of pitcher morphology and spectral reflectance characteristics showed that N. bicalcarata has no colour contrast maximum between the peristome and pitcher cup in the ultraviolet
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...
waveband, but small maxima of –0.17 at 450 nm (violet
Violet (color)
As the name of a color, violet is synonymous with a bluish purple, when the word "purple" is used in the common English language sense of any color between blue and red, not including either blue or red...
), 0.32 at 548 nm (green
Green
Green is a color, the perception of which is evoked by light having a spectrum dominated by energy with a wavelength of roughly 520–570 nanometres. In the subtractive color system, it is not a primary color, but is created out of a mixture of yellow and blue, or yellow and cyan; it is considered...
), and –0.16 at 668 nm (red
Red
Red is any of a number of similar colors evoked by light consisting predominantly of the longest wavelengths of light discernible by the human eye, in the wavelength range of roughly 630–740 nm. Longer wavelengths than this are called infrared , and cannot be seen by the naked eye...
). This means that the peristome is less reflective than the pitcher cup in the violet and red bands, but more reflective in the green band. The contrast maxima in the green and blue regions seem to correspond to insect visual sensitivity maxima, while the others do not. Based on their findings, the authors made the "tentative prediction" that the upper pitchers of N. bicalcarata are less successful at catching anthophilous (flower-visiting) prey than those of sympatric N. rafflesiana.
Association with ants and pitcher infauna
Nepenthes bicalcarata plays host to an unusual species of antAnt
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
that makes its nest in the plant's hollow tendrils. Described as Camponotus schmitzi
Camponotus schmitzi
Camponotus schmitzi is a species of carpenter ant native to northwestern Borneo.The ant makes its nest in the hollow tendrils of the pitcher plant Nepenthes bicalcarata....
in 1933, it is a member of the extremely populous and widespread genus of carpenter ant
Carpenter ant
Carpenter ants are large ants indigenous to many parts of the world. They prefer dead, damp wood in which to build nests. They do not eat it, however, unlike termites. Sometimes carpenter ants will hollow out sections of trees. The most likely species to be infesting a house in the United States...
s.
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...
as early as 1880. In 1904, 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...
suggested that the ants feed on insects found on and around the plant, but may fall prey to it themselves. In 1990, B. Hölldobler and E.O. Wilson proposed that N. bicalcarata and C. schmitzi form a mutually beneficial association. At the time, however, no experimental data existed to support such a hypothesis. A series of observations and experiments carried out in Brunei
Brunei
Brunei , officially the State of Brunei Darussalam or the Nation of Brunei, the Abode of Peace , is a sovereign state located on the north coast of the island of Borneo, in Southeast Asia...
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 1992 and 1998, and by Clarke and Roger Kitching in 1993 and 1995, strongly support the mutualism theory. John Thompson has suggested that N. bicalcarata may be the only plant species that obtains nutrients through both insect capture and ant-hosting habits.
Camponotus schmitzi ants feed by descending into the pitcher fluid and retrieving arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...
s caught by the plant. They seem to ignore smaller insects and only target larger prey items. Hauling food from the pitcher fluid to the peristome—a distance of no more than 5 cm—may take up to 12 hours. In this way the contents of N. bicalcarata pitchers is controlled such that organic matter does not accumulate to the point of putrefaction
Putrefaction
Putrefaction is one of seven stages in the decomposition of the body of a dead animal. It can be viewed, in broad terms, as the decomposition of proteins, in a process that results in the eventual breakdown of cohesion between tissues and the liquefaction of most organs.-Description:In terms of...
, which could lead to the demise of pitcher infauna
Nepenthes infauna
Nepenthes infauna are the organisms that inhabit the pitchers of Nepenthes plants. These include fly and midge larvae, spiders, mites, ants, and even a species of crab, Geosesarma malayanum. The most common and conspicuous predators found in pitchers are mosquito larvae, which consume large...
(which also appear to benefit the plant) and sometimes the pitcher itself. The ants have also been observed to attack newly caught insects and therefore prevent prey escape. At other times, the ants are very passive, remaining hidden under the inner peristome fold, presumably so as not to dissuade visitation by potential prey species. This behaviour is in stark contrast to other myrmecophytic
Myrmecophyte
Myrmecophyte is a plant that lives in a mutualistic association with a colony of ants. There are over 100 different genera of myrmecophytes. These plants possess structural adaptations that provide ants with food and/or shelter. These specialized structures include domatia, food bodies, and...
ants, which are typically highly territorial. In order to create a favourable environment for its pitcher inhabitants, it appears that N. bicalcarata actively maintains the 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 the pitcher fluid at a less acidic level than that found in most other Nepenthes species. This might explain the occasional presence of tree frog
Tree frog
Hylidae is a wide-ranging family of frogs commonly referred to as "tree frogs and their allies". However, the hylids include a diversity of frog species, many of which do not live in trees, but are terrestrial or semi-aquatic.-Characteristics:...
eggs, which are sensitive to strongly acidic conditions, in the pitchers of this species.
Research conducted by Dennis and Marlis Merbach has shown that C. schmitzi is also beneficial to N. bicalcarata in that it protects it from pitcher-destroying weevil
Weevil
A weevil is any beetle from the Curculionoidea superfamily. They are usually small, less than , and herbivorous. There are over 60,000 species in several families, mostly in the family Curculionidae...
s of the genus Alcidodes.
The ants seem to favour upper pitchers and rarely colonise lower pitchers. This is likely due to the fact that terrestrial traps are periodically submerged in water during heavy rains. Flooding of the ants' nest chamber could result in the death of the developing eggs, larva
Larva
A larva is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle...
e, and pupa
Pupa
A pupa is the life stage of some insects undergoing transformation. The pupal stage is found only in holometabolous insects, those that undergo a complete metamorphosis, going through four life stages; embryo, larva, pupa and imago...
e. Camponotus schmitzi nests solely in the tendrils of N. bicalcarata and rarely ventures onto other plants. The species is completely dependent on N. bicalcarata for food and domicile. Nepenthes bicalcarata, on the other hand, is able to survive and reproduce without the presence of the ants; it is a facultative mutualist. This being the case, there appear to be few mature plants over 2 metres in height not colonised by C. schmitzi.
A species of mite
Mite
Mites, along with ticks, are small arthropods belonging to the subclass Acari and the class Arachnida. The scientific discipline devoted to the study of ticks and mites is called acarology.-Diversity and systematics:...
, Naiadacarus nepenthicola, appears to be restricted to the pitchers of N. bicalcarata. It is thought to feed on decomposing leaves and insects that are caught in the pitchers. Deutonymphs of this mite are dispersed through phoresy on C. schmitzi.
Natural hybrids
The following natural hybrids involving N. bicalcarata have been recorded.- N. ampullariaNepenthes ampullariaNepenthes ampullaria , the Flask-Shaped Pitcher-Plant, is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea...
× N. bicalcarata - (N. ampullariaNepenthes ampullariaNepenthes ampullaria , the Flask-Shaped Pitcher-Plant, is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea...
× N. gracilisNepenthes gracilisNepenthes gracilis , or the Slender Pitcher-Plant, is a very common lowland pitcher plant that is widespread in the Sunda region. It has been recorded from Borneo, Peninsular Malaysia, Singapore, central Sulawesi, Sumatra, and southernmost Thailand...
) × N. bicalcarata [=N. × trichocarpa × N. bicalcarata] - N. bicalcarata × N. gracilisNepenthes gracilisNepenthes gracilis , or the Slender Pitcher-Plant, is a very common lowland pitcher plant that is widespread in the Sunda region. It has been recorded from Borneo, Peninsular Malaysia, Singapore, central Sulawesi, Sumatra, and southernmost Thailand...
[=N. × cantleyi] - N. bicalcarata × N. mirabilisNepenthes mirabilisNepenthes 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,...
(including N. bicalcarata × N. mirabilis var. echinostoma) - N. bicalcarata × N. rafflesianaNepenthes rafflesianaNepenthes rafflesiana , or Raffles' Pitcher-Plant, is a species of pitcher plant. It has a very wide distribution covering Borneo, Sumatra, Peninsular Malaysia, and Singapore. Nepenthes rafflesiana is extremely variable with numerous forms and varieties described. In Borneo alone, there are at...
- ? (N. bicalcarata × N. rafflesianaNepenthes rafflesianaNepenthes rafflesiana , or Raffles' Pitcher-Plant, is a species of pitcher plant. It has a very wide distribution covering Borneo, Sumatra, Peninsular Malaysia, and Singapore. Nepenthes rafflesiana is extremely variable with numerous forms and varieties described. In Borneo alone, there are at...
) × N. mirabilis var. echinostomaNepenthes mirabilisNepenthes 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,...
None of these hybrids are known to be colonised by C. schmitzi as is the parent species N. bicalcarata.
N. ampullaria × N. bicalcarata
Brunei
Brunei , officially the State of Brunei Darussalam or the Nation of Brunei, the Abode of Peace , is a sovereign state located on the north coast of the island of Borneo, in Southeast Asia...
, where it grows in peat swamp forest
Peat swamp forest
Peat swamp forests are tropical moist forests where waterlogged soils prevent dead leaves and wood from fully decomposing, which over time creates a thick layer of acidic peat...
and heath forest
Heath forest
Heath forest is a type of tropical moist forest found in areas with acidic, sandy soils that are extremely nutrient-poor. Notable examples are the Rio Negro campinarana of the Amazon Basin in South America, and the Sundaland heath forests of Borneo and neighboring islands.-External links:**...
. It was first recorded by Mitsuru Hotta
Mitsuru Hotta
is a Japanese botanist best known for his research on Araceae.Hotta was born in Osaka, Japan in 1937. He graduated from the Agricultural Department of Osaka Prefecture University in 1960. The same year, he took part in the Tonga and Fiji Expedition organised by Kyoto University...
in 1966.
Pitchers bear the characteristic thorns of N. bicalcarata, although they are greatly reduced in size. The mouth is round and the lid is ovate-cordate in form. This hybrid resembles N. × hookeriana
Nepenthes x hookeriana
Nepenthes × hookeriana , or Hooker's Pitcher-Plant, is a common natural hybrid involving N. rafflesiana and N. ampullaria. It was originally described as a species....
to a certain extent, but differs in having spines on the underside of the lid and more globose upper pitchers with a smaller lid.
Aerial pitchers of N. ampullaria × N. bicalcarata are usually dominated by characteristics of N. ampullaria. They are often very small and unable to function in a normal manner.
(N. ampullaria × N. gracilis) × N. bicalcarata
This rare complex hybrid was discovered in the lowland forests of BruneiBrunei
Brunei , officially the State of Brunei Darussalam or the Nation of Brunei, the Abode of Peace , is a sovereign state located on the north coast of the island of Borneo, in Southeast Asia...
. It produces small speckled pitchers with reduced thorns under the lid. The plant is intermediate in appearance between N. × trichocarpa and N. bicalcarata.
N. bicalcarata × N. gracilis
Rob Cantley
Robert Cantley is a conservationist and Managing Director of Borneo Exotics, a Sri Lankan-based plant nursery specialising in tissue-cultured and seed-grown Nepenthes species and hybrids. Cantley has contributed to a number of papers on Nepenthes...
. The morphology of the pitchers closely resembles N. bicalacarata, although the peristome is more similar to that of N. gracilis. The characteristic spines of N. bicalcarata are greatly reduced and are only present as small bumps. This hybrid has the growth habit of N. gracilis, with the stem scrambling along the ground. It grows in open, sandy areas. The pitcher fluid is notably acid
Acid
An acid is a substance which reacts with a base. Commonly, acids can be identified as tasting sour, reacting with metals such as calcium, and bases like sodium carbonate. Aqueous acids have a pH of less than 7, where an acid of lower pH is typically stronger, and turn blue litmus paper red...
ic like that of N. gracilis, with 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...
values as low as 1.82 being recorded.
N. bicalcarata × N. mirabilis var. echinostoma
This natural hybrid is rare and is only known from BruneiBrunei
Brunei , officially the State of Brunei Darussalam or the Nation of Brunei, the Abode of Peace , is a sovereign state located on the north coast of the island of Borneo, in Southeast Asia...
, where it involves N. mirabilis var. echinostoma. It closely resembles N. bicalcarata, although it differs from that species in having reduced thorns and a wider peristome. Unlike N. bicalcarata, this hybrid inhabits open areas with sandy substrates.
N. bicalcarata × N. rafflesiana
This relatively rare natural hybrid is sometimes found growing among populations of N. bicalcarata and N. rafflesiana. It is intermediate in form between its parent species.(N. bicalcarata × N. rafflesiana) × N. mirabilis var. echinostoma
Charles ClarkeCharles 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...
reported a single plant displaying characters intermediate in form between N. bicalcarata, N. rafflesiana and N. mirabilis var. echinostoma growing among populations of these species. It is likely a complex hybrid, although this cannot be proven without the use of genetic testing
Genetic testing
Genetic testing is among the newest and most sophisticated of techniques used to test for genetic disorders which involves direct examination of the DNA molecule itself. Other genetic tests include biochemical tests for such gene products as enzymes and other proteins and for microscopic...
.
Cultivation and cultivars
Despite naturally occurring in peat swamp forests, N. bicalcarata has been successfully grown in a completely inorganic substrate consisting of one part each of Seramis clay perls, lava gravel, and Lecaton expanded clay perls.
Nepenthes bicalcarata has been artificially crossed with N. hirsuta
Nepenthes hirsuta
Nepenthes hirsuta , the Hairy Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It is characterised by an indumentum of thick brown hairs, which is even present on the inflorescence. Pitchers are mostly green throughout with some having red blotches on the inside surfaces.N. hirsuta...
. A particularly robust-pitchered cultivar
Cultivar
A cultivar'Cultivar has two meanings as explained under Formal definition. When used in reference to a taxon, the word does not apply to an individual plant but to all those plants sharing the unique characteristics that define the cultivar. is a plant or group of plants selected for desirable...
of this hybrid was named Nepenthes 'Hortulanus Otten', in honour of Karel Otten, former curator of the Botanic Garden in Ghent
Ghent
Ghent is a city and a municipality located in the Flemish region of Belgium. It is the capital and biggest city of the East Flanders province. The city started as a settlement at the confluence of the Rivers Scheldt and Lys and in the Middle Ages became one of the largest and richest cities of...
, Belgium
Belgium
Belgium , officially the Kingdom of Belgium, is a federal state in Western Europe. It is a founding member of the European Union and hosts the EU's headquarters, and those of several other major international organisations such as NATO.Belgium is also a member of, or affiliated to, many...
.
A cultivar of the artificial cross N. bicalcarata × N. × dyeriana was registered in 1988 under the name Nepenthes 'Nina Dodd'. It is named after a relative of Cliff Dodd, who created the hybrid.
Two further cultivars of manmade hybrids involving N. bicalcarata have been named. These are Nepenthes 'Bella' ((N. ampullaria
Nepenthes ampullaria
Nepenthes ampullaria , the Flask-Shaped Pitcher-Plant, is a very distinctive and widespread species of Nepenthes, present in Borneo, Sumatra, Thailand, Peninsular Malaysia, Singapore, the Maluku Islands, and New Guinea...
× N. bicalcarata) × N. truncata
Nepenthes truncata
Nepenthes truncata is a tropical pitcher plant endemic to the Philippines. It is known from the islands of Dinagat, Leyte, and Mindanao. The species grows at an elevation of 0–1500 m above sea level...
) and Nepenthes 'Rapa' (N. ampullaria × N. bicalcarata). However, both of these names are not established, as they were published without a description.
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.
- Bourke, G. 2011. The Nepenthes of Mulu National Park. Carniflora AustralisCarniflora AustralisCarniflora Australis is a biannual English-language periodical and the official publication of the Australasian Carnivorous Plant Society. Typical articles include matters of horticultural interest, field reports, and scientific studies. The journal was established in March 2003...
8(1): 20–31. - 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. - Fashing, N.J. 2010. In: M.W. Sabelis & J. Bruin (eds.) Trends in Acarology: Proceedings of the 12th International Congress. Springer Science, Dordrecht. pp. 81–84.
- Fretwell, S. 2010. Twelve days in Borneo – a dream expedition: part 4. 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...
98: 6–13. Kubeš, V. 2003. "Panel story" o pěstování láčkovky Nepenthes bicalcarata. TrifidTrifid (journal)Trifid is a quarterly Czech-language periodical and the official publication of Darwiniana, a carnivorous plant society based in the Czech Republic. Typical articles include matters of horticultural interest, field reports, and scientific studies. The journal was established in 1996. It is...
2003(2): 7–10. (page 2, page 3, page 4) Mansur, M. 2001. In: Prosiding Seminar Hari Cinta Puspa dan Satwa Nasional. Lembaga Ilmu Pengetahuan Indonesia, Bogor. pp. 244–253. - Siegara, A. & Yogiara 2009. Bacterial community profiles in the fluid of four pitcher plant species (Nepenthes spp.) grown in a nursery. Microbiology Indonesia 3(3): 109–114.
- Thorogood, C. 2010. The Malaysian Nepenthes: Evolutionary and Taxonomic Perspectives. Nova Science Publishers, New York. Zizka, G. 1990. Pflanzen und Ameisen. Palmengarten Sonderheft 15: 112–116. Zizka, G. 1991. Über die Kannenpflanze Nepenthes bicalcarata J.D. Hooker. Palmengarten 91(3): 39–44.
External links
- Photographs of N. bicalcarata at the Carnivorous Plant Photofinder