Nepenthes murudensis
Encyclopedia
Nepenthes murudensis or the Murud 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...

 endemic to Mount Murud
Mount Murud
Mount Murud or Muru is a sandstone mountain located in the Malaysian part of Borneo. At 2,423 m , it is the highest mountain in the state of Sarawak....

 in 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....

, after which it is named. It is of putative hybrid origin: its two original parent species are thought to be N. reinwardtiana
Nepenthes reinwardtiana
Nepenthes reinwardtiana , Reinwardt's Pitcher-Plant, is a Nepenthes species found on the islands of Borneo and Sumatra. Although some sources have included Peninsular Malaysia and Singapore within the range of this species, these records appear to be erroneous.Nepenthes reinwardtiana has an...

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

.

Botanical history

Nepenthes murudensis was known for some time prior to its description, although authors differed as to its identity. In 1988, 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...

 suggested that it might represent a natural hybrid between N. reinwardtiana
Nepenthes reinwardtiana
Nepenthes reinwardtiana , Reinwardt's Pitcher-Plant, is a Nepenthes species found on the islands of Borneo and Sumatra. Although some sources have included Peninsular Malaysia and Singapore within the range of this species, these records appear to be erroneous.Nepenthes reinwardtiana has an...

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

. However, 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...

the authors treated it as a species in the process of being described, referring to it as "Nepenthes murudensis Culham ined.", a name given to it informally by Alistair Culham that was already in use in 1994.

Nepenthes murudensis was 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...

 in 1997 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,...

 in their 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...

", published 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...

. At the time of its description, botanists were divided on whether N. murudensis should be considered a species, although the consensus view held that it was of hybridogenic origin.

The holotype
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...

 of N. murudensis, Yii Puan Ching S 44623, was collected on September 13, 1982, between the first and second summits of Mount Murud
Mount Murud
Mount Murud or Muru is a sandstone mountain located in the Malaysian part of Borneo. At 2,423 m , it is the highest mountain in the state of Sarawak....

. It 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...

. Another specimen, Beaman 11461, was collected by John H. Beaman between April 10 and April 17, 1995, from the summit ridge of Mount Murud at an elevation of between 2300 and 2400 m above sea level. This latter specimen was collected as part of the eighth botanical expedition to Mount Murud since Eric Mjöberg
Eric Mjöberg
Dr Eric Georg Mjöberg was a Swedish zoologist and ethnographer who led the first Swedish scientific expeditions to Australia in the early 1900s, and worked in Indonesia. The plant Vaccinium mjoebergii J.J.Sm...

's first ascent in 1922.

The close relationship between N. murudensis and its putative parent species has led to some confusion in the literature. One example of this appears in the article "Nepenthes of Gunung Murud", authored by John De Witte and published in a 1996 issue of the Carnivorous Plant Newsletter
Carnivorous Plant Newsletter
The Carnivorous Plant Newsletter is the official publication of the International Carnivorous Plant Society , the largest such organization in the world.-History and editorship:...

, where an upper pitcher of N. murudensis is identified as N. tentaculata
Nepenthes tentaculata
Nepenthes tentaculata , or the Fringed Pitcher-Plant, is a tropical pitcher plant with a very wide distribution across Borneo and Sulawesi...

.

Description

Nepenthes murudensis is a climbing plant. The stem can attain a length of 5 m and is up to 5 mm wide. 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 triangular in cross section and up to 10 cm long.

Leaves are coriaceous and adnate. The lamina is oblong-elliptic in shape and reaches 12 cm in length by 4 cm in width. The apex of the lamina is rounded-obtuse and the base is decurrent for up to 2 cm. Up to 5 longitudinal veins are present on either side of the midrib. Pinnate veins are indistinct.

Rosette and lower pitchers are ovoid in the lower part, becoming sub-cylindrical above. They reach 20 cm in height and 5 cm in width. A pair of fringed wings (≤6 mm wide) runs down the front of the pitcher. 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 a flattened cylinder in cross section and is up to 5 mm wide. 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 ovate-obovate in shape and lacks appendages. It has a rounded apex, a truncate base, and measures up to 6 cm by 5 cm. Large nectar glands are concentrated along the midrib of the lid. 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, but differ in being more cylindrical and elongate. They are also larger, growing to 30 cm in height. Wings are reduced to a pair of prominent ribs in upper pitchers. Both lower and upper pitchers have an unusually long waxy zone, which in the latter extends for as much as three-quarters or more of the pitcher length.

Nepenthes murudensis 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 male and female inflorescences differing little in structure. It is very compact: 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...

 reaches 3 cm in length and 7 mm in diameter, 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...

 may be up to 6 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...

eolate, and up to 7 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 elliptic and up to 5 mm long.

Most parts of the plant are virtually glabrous, although a short, 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 velvety brown hairs is present on the stem, inflorescences, and lamina midribs.

Ecology

Nepenthes murudensis is endemic to the summit area of Mount Murud
Mount Murud
Mount Murud or Muru is a sandstone mountain located in the Malaysian part of Borneo. At 2,423 m , it is the highest mountain in the state of Sarawak....

 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...

, 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 has an altitudinal distribution of 2000–2423 m above sea level.

The typical habitat of this species consists of stunted montane scrub
Shrubland
Shrubland, scrubland, scrub or brush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity...

 and ridge vegetation, which rarely exceeds 1.5 m in height. The species also occurs in mossy forest. Nepenthes murudensis has no known natural hybrids, although N. hurrelliana
Nepenthes hurrelliana
Nepenthes hurrelliana is a tropical pitcher plant endemic to Borneo, where it has been recorded from northern Sarawak, southwestern Sabah, and Brunei. It is of putative hybrid origin; its two original parent species are thought to be N. fusca and N. veitchii...

, 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...

, N. muluensis
Nepenthes muluensis
Nepenthes muluensis , or the Mulu Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It grows in highland habitats at elevations of 1700 to 2400 m above sea level.-Botanical history:...

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

also occur on the mountain. In 1996, John De Witte reported observing N. reinwardtiana on Mount Murud, but other authors have failed to find it on the mountain.

Nepenthes murudensis often scrambles over low vegetation, but also grows as rosettes in open areas. Botanist Andrew Hurrell has described plants growing on the summit as small rosettes rarely exceeding 30 cm in height, with proportionately huge pitchers sometimes measuring over 30 cm themselves.

Two true toad
True toad
Bufonidae is the family of the true toads, members of the order Anura . They are the only family of anurans all members of which are known as "toads." The bufonids now comprise more than 35 genera, Bufo being the most widespread and well known.-Characteristics:True toads are widespread and occur...

 species from Mount Murud, Pelophryne linanitensis
Pelophryne linanitensis
Pelophryne linanitensis is a species of toad in the Bufonidae family. It is endemic to Borneo....

and Pelophryne murudensis
Pelophryne murudensis
Pelophryne murudensis is a species of toad in the Bufonidae family. It is endemic to Borneo....

, are sympatric with N. murudensis and may breed in its pitchers.

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

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

 based on an assessment carried out in 2000. This agrees with an informal assessment made by Charles Clarke
Charles Clarke (botanist)
Dr. Charles M. Clarke is a botanist and taxonomist specialising in the carnivorous plant genus Nepenthes. Clarke has an honours degree in Botany from Monash University in Melbourne, and a Ph.D. in Ecosystem Management at the University of New England, in Armidale, New South Wales.Clarke first...

 in 1997, who also classified the species as Endangered 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.

Hybrid origin

The pitchers of N. murudensis are roughly intermediate in appearance between those of N. reinwardtiana and N. tentaculata, although significantly larger than either. This has led to speculation regarding the lineage of this species, with a number of authors suggesting a possible hybridogenic origin.

Botanist Clive A. Stace
Clive A. Stace
Clive Anthony Stace is a British botanist and botanical author. His academic career was based at the University of Leicester, where he held the post of Professor of Plant taxonomy...

 writes that one may speak of "stabilised hybrids when they have developed a distributional, morphological or genetic set of characters which is no longer strictly related to that of its parents, [...] if the hybrid has become an independent, recognisable, self-producing unit, it is de facto a separate species". This would tend to support the status of N. murudensis as a species, since populations of this 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...

 appear to be stabilised as well as highly homogeneous, and it is one of the most abundant Nepenthes on the summit ridge of Mount Murud. Examples of other Nepenthes species with a putative hybrid origin include N. hamiguitanensis
Nepenthes hamiguitanensis
Nepenthes hamiguitanensis is a tropical pitcher plant endemic to a single peak on the Philippine island of Mindanao, where it grows at elevations of 1200–1600 m above sea level. Once thought to be a natural hybrid between N. micramphora and N. peltata, this plant is now considered a...

, N. hurrelliana
Nepenthes hurrelliana
Nepenthes hurrelliana is a tropical pitcher plant endemic to Borneo, where it has been recorded from northern Sarawak, southwestern Sabah, and Brunei. It is of putative hybrid origin; its two original parent species are thought to be N. fusca and N. veitchii...

, and N. petiolata
Nepenthes petiolata
Nepenthes petiolata is a highland Nepenthes pitcher plant species endemic to Mindanao island in the Philippines, where it grows at an elevation of 1450–1900 m above sea level.-Natural hybrids:*? N. alata × N. petiolata...

.

Alternatively, N. murudensis may have evolved from populations of N. tentaculata that underwent speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...

, possibly in isolation.

Related species

Nepenthes murudensis belongs to what has been called the "Hamata group", which also includes four other closely related species from 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....

 and Sulawesi
Sulawesi
Sulawesi is one of the four larger Sunda Islands of Indonesia and is situated between Borneo and the Maluku Islands. In Indonesia, only Sumatra, Borneo, and Papua are larger in territory, and only Java and Sumatra have larger Indonesian populations.- Etymology :The Portuguese were the first to...

: N. glabrata
Nepenthes glabrata
Nepenthes glabrata is a tropical pitcher plant endemic to Sulawesi. The species grows in open, high forest at elevations of 1600 to 2100 m. It produces dainty, colourful pitchers reaching only a few centimetres in height...

, N. hamata
Nepenthes hamata
Nepenthes hamata is a tropical pitcher plant endemic to Sulawesi, where it grows at elevations of 1400–2500 m above sea level.The specific epithet hamata is derived from the Latin word hamatus, meaning "hooked"...

, N. muluensis
Nepenthes muluensis
Nepenthes muluensis , or the Mulu Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It grows in highland habitats at elevations of 1700 to 2400 m above sea level.-Botanical history:...

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

.

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

and it is undoubtedly closely related to this species. Nepenthes murudensis differs in lacking filiform hairs on the upper surface of the lid, being more robust in all respects, and having 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...

 on 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...

s and some vegetative parts. However, a number of populations of N. tentaculata from northern 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...

 produce pitchers exceeding 20 cm in height and these may be very similar in appearance to N. murudensis.

Nepenthes murudensis also differs in that its aerial pitchers lack wings. Although N. tentaculata is variable in this respect, plants from Mount Murud usually produce upper pitchers with wings.

Nepenthes murudensis can be distinguished from its other suspected parent species, N. reinwardtiana
Nepenthes reinwardtiana
Nepenthes reinwardtiana , Reinwardt's Pitcher-Plant, is a Nepenthes species found on the islands of Borneo and Sumatra. Although some sources have included Peninsular Malaysia and Singapore within the range of this species, these records appear to be erroneous.Nepenthes reinwardtiana has an...

, on the basis of lacking "eye spots" on the inside surface of its pitchers and having distinctive leaf bases that completely clasp the stem. In addition, the lower pitchers of N. murudensis have a pair of fringed wings, whereas those of N. reinwardtiana typically have ribs or, rarely, short wings without fringe elements.
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