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

 known only from a very restrictive elevation on 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 ×...

 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...

, Malaysian Borneo. The specific epithet macrophylla is formed from the 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...

 words macro (large) and phylla (leaves).

Nepenthes macrophylla was once thought to be 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 , but differs from that species in several aspects of pitcher and leaf morphology
Plant morphology
Plant morphology or phytomorphology is the study of the physical form and external structure of plants. This is usually considered distinct from plant anatomy, which is the study of the internal structure of plants, especially at the microscopic level...

.

Botanical history

Nepenthes macrophylla was known to grow on 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 ×...

 for a long time prior to its description, although it was not initially considered a distinct species. Shortly after its discovery, it was lumped with , a similar species from 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...

. In his 1976 guide, 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...

, botanist 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....

 wrote:

Mt. Trus Madi, located in the south of Mt. Kinabalu, become known as a locality for this species []. Though this information is not confirmed yet, its existence is expected because this mountain falls within the vertical distribution of this species.


Later, N. macrophylla was treated as a form of , another closely related species. An early collection of N. macrophylla was made by Johannes Marabini in March 1983 and the herbarium
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...

 material, designated as Marabini 2167, deposited at the University of Erlangen-Nuremberg in Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

. Four years later, Marabini formally described
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...

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

 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.

In their 1996 monograph, Pitcher-Plants of Borneo
Pitcher-Plants of Borneo
Pitcher-Plants of Borneo is a monograph by Anthea Phillipps and Anthony Lamb on the tropical pitcher plants of Borneo. It was first published in 1996 by Natural History Publications , in association with the Royal Botanic Gardens, Kew and the Malaysian Nature Society...

, Anthea Phillipps
Anthea Phillipps
Anthea Phillipps B.Sc. is a British botanist. Phillipps was brought up in Sabah, Borneo as a child . She received a Botany degree from the University of Durham, England. She worked at the Sabah Museum before joining the Sabah Parks service from 1980 to 1987 as Park Ecologist, where she studied...

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

 treated N. macrophylla as a species in the process of being described, referring to it as "Nepenthes macrophylla (Marabini) Jebb & Cheek ined.". Botanists 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,...

 had intended to elevate N. macrophylla to species status since at least 1994 and did so upon the publication of their 1997 monograph "A skeletal revision of Nepenthes (Nepenthaceae)
A skeletal revision of Nepenthes (Nepenthaceae)
"A skeletal revision of Nepenthes " is a monograph by Matthew Jebb and Martin Cheek on the tropical pitcher plants of the genus Nepenthes. It was published in the May 1997 issue of the botanical journal Blumea. The work represented the first revision of the entire genus since John Muirhead...

" in the botanical journal Blumea
Blumea (journal)
Blumea - Journal of Plant Taxonomy and Plant Geography is a peer-reviewed journal of botany published by the National Herbarium of the Netherlands.Except for a short period during World War II, Blumea has been published continuously since 1934...

. 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 with each other than do N. edwardsiana and N. macrophylla".

Description

Nepenthes macrophylla is a climbing plant. The stem
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...

 reaches a length of more than 10 m and is up to 10 mm in diameter. Internode
Plant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...

s are up to 35 cm long and circular in cross section.

Leaves are coriaceous and petiolate. The lamina or leaf blade is oblong in shape and reaches exceptionally large dimensions of up to 60 cm by 20 cm. The apex of the lamina is acuminate-obtuse, while the base is abruptly contracted where 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...

 begins. The petiole itself is up to 15 cm long. It is canaliculate (grooved lengthwise) and typically bears wings that form a partially amplexicaul sheath around the stem. One to three longitudinal veins are present on either side of the midrib, but pinnate veins are indistinct. 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 measure up to 35 cm in length.

The pitchers of N. macrophylla are robust and almost woody in texture. Rosette and lower pitchers are shortly cylindrical in shape, becoming narrower around the midpoint. They can be very large: up to 35 cm high by 15 cm wide. A pair of fringed wings (≤10 mm wide) runs down the ventral surface of the pitcher. Terrestrial pitchers usually have a hip in their lower half. The pitcher mouth is ovate and has an oblique insertion. The peristome
Peristome
The word peristome is derived from the Greek peri, meaning 'around' or 'about', and stoma, 'mouth'. It is a term used to describe various anatomical features that surround an opening to an organ or structure. The term is used in plants and invertebrate animals, such as in describing the shells of...

 is cylindrical and expanded, measuring up to 15 mm wide. It bears a series of highly developed ribs, which terminate in prominent downward-pointing teeth. The pitcher lid or operculum
Operculum (botany)
An operculum, in botany, is a term generally used to describe a structure within a plant, moss, or fungus acting as a cap, flap, or lid. In plants, it may also be called a bud cap.Examples of structures identified as opercula include:...

 is ovate, very broad, and has a cordate base. 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...

 (≤9 mm long) is inserted near the base of the lid. Upper pitchers are similar to their lower counterparts in most respects. They differ in having a more cylindrical shape, reduced wings, and the hip always being located in the upper half. Traps may have a range of colours, although the yellow-pitchered form is more common than the red-pitchered form.

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

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

, with both male and female ones having a similar structure. The peduncle
Peduncle (botany)
In botany, a peduncle is a stem supporting an inflorescence, or after fecundation, an infructescence.The peduncle is a stem, usually green and without leaves, though sometimes colored or supporting small leaves...

 is up to 23 cm long and 5 mm wide, 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...

 is up to 55 cm long. 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 one-flowered, 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...

eate, and measure up to 16 mm in length. 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 elliptic in shape and up to 6 mm long.

Most parts of the plants bear an 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 very short hairs, although much of this covering is caducous.

Ecology

Nepenthes macrophylla is known with certainty only from the summit area 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 ×...

 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....

, despite the presence of a number of nearby peaks (such as Mount Pinesowitan) that fall within its altitudinal range. However, there are unconfirmed reports that it also grows "on the summit of a neighbouring peak". Older sources list a number of different values for the altitudinal distribution of N. macrophylla, with some giving a narrow range of 2200–2400 m and others a much wider span of 2000–2600 m. It is now known that this species occurs between 2200 m and the summit at 2642 m.

The typical habitat of N. macrophylla is mossy montane forest along ridge tops, where the vegetation is dominated by 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...

and Rhododendron
Rhododendron
Rhododendron is a genus of over 1 000 species of woody plants in the heath family, most with showy flowers...

. Like most highland Nepenthes species, N. macrophylla experiences marked temperature fluctuations between day and night, from a noon maximum of around 30 °C to a nighttime minimum of 5 °C. 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 always high and in the late evening the summit of Mount Trus Madi is often shrouded in mist and clouds.

Nepenthes macrophylla occurs both terrestrially and as an epiphyte
Epiphyte
An epiphyte is a plant that grows upon another plant non-parasitically or sometimes upon some other object , derives its moisture and nutrients from the air and rain and sometimes from debris accumulating around it, and is found in the temperate zone and in the...

. The species is sympatric with and . A natural hybrid between N. lowii and N. macrophylla, described as , has been recorded.

Due to its restricted natural range
Range (biology)
In biology, the range or distribution of a species is the geographical area within which that species can be found. Within that range, dispersion is variation in local density.The term is often qualified:...

 and the fact that Mount Trus Madi does not receive the same high level of protection as 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...

, N. macrophylla is classified as Critically Endangered
Critically Endangered
Critically Endangered is the highest risk category assigned by the IUCN Red List for wild species. Critically Endangered means that a species' numbers have decreased, or will decrease, by 80% within three generations....

 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...

. This differs slightly from an informal assessment made by botanist Charles Clarke
Charles Clarke (botanist)
Dr. Charles M. Clarke is a botanist and taxonomist specialising in the carnivorous plant genus Nepenthes. Clarke has an honours degree in Botany from Monash University in Melbourne, and a Ph.D. in Ecosystem Management at the University of New England, in Armidale, New South Wales.Clarke first...

 in 1997, who classified the species as Endangered
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

 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. Clarke wrote that the species "requires the highest level of protection if it is to persist in the wild into the next century". N. macrophylla is threatened by over-collection and damage caused to its habitat by visitors. These activities have already depleted populations of this species on the mountain and it is thought to be the most endangered highland Nepenthes of Borneo.

Carnivory

A 2010 study showed that the shape and size of the pitcher orifice of N. macrophylla exactly match the dimensions of a typical tree shrew (Tupaia montana). A similar adaptation was found in N. lowii
Nepenthes lowii
Nepenthes lowii , or Low's Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It is named after Hugh Low, who discovered it on Mount Kinabalu...

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...

, and is also likely to be present in N. ephippiata
Nepenthes ephippiata
Nepenthes ephippiata , or the Saddle-Leaved Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It occurs in the Hose Mountains of central Sarawak, as well as Mount Raya and Bukit Lesung in Kalimantan. Plants from the Hose Mountains appear to lack the decurrent leaf attachment found in...

.

Related species

Nepenthes macrophylla is most closely related to and .

Nepenthes macrophylla differs from both of these species in the structure of its peristome. Although highly developed, the peristome ribs and teeth of N. macrophylla are considerably shorter and more widely-spaced than those of either N. edwardsiana or N. villosa. The pitcher mouth of N. macrophylla is distinctive in that it rises gradually towards the lid, while at the same time not forming a pronounced neck. In addition, the mouth of this species has a much more oblique insertion than its relatives. Nepenthes macrophylla is also distinguished by its broad, ovate lid. The lower pitchers of N. edwardsiana and N. macrophylla are quite similar in shape, although in the latter species the hip is always positioned in the upper portion of the pitcher cup. The upper pitchers of these species are more distinct, with those of N. macrophylla being more ovoid and less elongated. As its name suggests, N. macrophylla produces very large leaves and these may be twice as long as those of either N. edwardsiana or N. villosa.

Whereas N. edwardsiana and N. villosa are restricted to generally ultramafic soils in the 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...

 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 ×...

, which is mostly composed of sandstone
Sandstone
Sandstone is a sedimentary rock composed mainly of sand-sized minerals or rock grains.Most sandstone is composed of quartz and/or feldspar because these are the most common minerals in the Earth's crust. Like sand, sandstone may be any colour, but the most common colours are tan, brown, yellow,...

.

Botanists 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,...

 suggest that N. macrophylla is related to , 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...

.

Natural hybrids

In the wild, N. macrophylla is only known to hybridise with . N. lowii × N. macrophylla was discovered on 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 ×...

 by Johannes Marabini and John Briggs in 1983. Later that year, it was described as by Marabini. Briggs returned to Mount Trus Madi in 1984, but found only one small group of plants.

has petiolate leaves measuring up to 50 cm in length. The pitchers of this hybrid are some of the largest of any Bornean Nepenthes species, reaching 35 cm in height. They are roughly intermediate in form between those of its parent species. The lid is held away from the mouth as in N. lowii and bears short britles on its lower surface. The peristome has prominent ribs and teeth, but is not as developed as that of N. macrophylla. The 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...

 of may be up to 50 cm long and has two-flowered 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....

. Despite the size of the pitchers, this hybrid is not large in stature.

is restricted to the summit ridge of Mount Trus Madi and has been recorded from elevations of 2500 to 2600 m above sea level.

External links

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