Nepenthes benstonei
Encyclopedia
Nepenthes benstonei is a tropical pitcher plant
endemic to Peninsular Malaysia
, where it occurs at elevations of 450–600 m above sea level. The specific epithet benstonei honours botanist Benjamin Clemens Stone
, who was one of the first to collect the species.
", Matthew Jebb
and Martin Cheek
tentatively referred specimens collected from Bukit Bakar, near Macang, Kelantan
, to N. sanguinea
. These were Stone & Chin 15238, deposited at Universiti Kebangsaan Malaysia
near Kuala Lumpur
, and Shah & Shukor 3168, also held at the university as well as the Forest Research Institute of Malaysia in Kepong
. They noted that the plants exhibited some unusual morphological features, such as larger leaves and decurrent, almost petiolate leaf bases, suggesting that they might represent an as-yet undescribed taxon
.
Field studies confirmed that the taxon represented a separate species. It was described as N. benstonei in 1999 by Charles Clarke
.
The holotype
of N. benstonei, Clarke s.n., was collected by Charles Clarke
on July 24, 1998, on Bukit Bakar in Kelantan
at an altitude of between 450 and 550 m. It is deposited at the Forest Research Institute of Malaysia in Kepong
. Isotype
s are held at the Bogor Botanical Gardens
, the Royal Botanic Gardens, Kew
, the National Herbarium of the Netherlands
in Leiden, the Forest Department in Sandakan
, and the Singapore Botanic Gardens
.
on Mount Tahan
in Pahang
. The specimen, Ridley 16097, includes female floral material and is deposited at the herbarium of the Singapore Botanic Gardens
.
In his seminal monograph "The Nepenthaceae of the Netherlands Indies
", B. H. Danser
referred Ridley 16097 to N. alata
. Nepenthes alata was not known from Borneo
and this specimen constituted the sole record of N. alata from Peninsular Malaysia. Danser also treated N. eustachya
from Sumatra
in synonymy with N. alata, making its extreme rarity in Peninsular Malaysia difficult to explain.
In 1990, Ruth Kiew identified Ridley 16097 as belonging to N. gracillima
. Kiew explained the apparent near-absence of N. alata from Peninsular Malaysia as follows:
Jebb and Cheek supported this interpretation in their 1997 monograph, "A skeletal revision of Nepenthes (Nepenthaceae)
".
However, Charles Clarke
disagrees with Kiew's identification. He notes that the inflorescences of both N. gracillima and the closely related N. ramispina
are very short, rarely exceeding 10 and 20 cm, respectively. Both female inflorescences of Ridley 16097 have a long peduncle
and rachis
, each exceeding 20 cm in length. In addition, the flowers of these species are almost always borne on pedicels
, unlike those of Ridley 16097, which mostly have two-flowered partial peduncles. Furthermore, the shape of the lamina is unlike that of N. gracillima or N. ramispina; despite being narrow and lanceolate, it is proportionately considerably longer and has a much narrower, almost sub-petiolate base. Kiew noted that the lamina of Ridley 16097 appears narrower due to the way in which the specimen was preserved, but Clarke states that this cannot fully account for the differences. He also notes that the specimen exhibits a decurrent leaf attachment. Taking all of these morphological features into account, Clarke feels that Ridley 16097 most likely represents N. benstonei. Clarke considers that Kiew grouped both N. benstonei and N. ramispina with N. gracillima.
also belong to N. benstonei. These are Shah et al. 3274, which is deposited at the Forest Research Institute of Malaysia in Kepong, and Shah et al. 3283, held at the Universiti Kebangsaan Malaysia near Kuala Lumpur.
Specimens from peninsula
r Thailand
originally assigned to N. benstonei in Cheek and Jebb's 2001 monograph, "Nepenthaceae
", have since been identified as belonging to a new species, N. thai
.
s are cylindrical and up to 15 cm long.
Leaves are coriaceous and sessile to sub-petiolate. The lamina is usually broadly linear-lanceolate in shape, but may also be slightly spathulate. Its base is a broad, amplexicaul sheath with decurrent margins. The lamina can reach 60 cm in length and 9 cm in width. It has a rounded to acute apex. The margins of the lamina usually meet the tendril unequally on both sides, being up to 3 mm apart. Three to five longitudinal veins are present on either side of the midrib. Pinnate veins are almost indistinct. Tendril
s are up to 60 cm long.
Rosette and lower pitchers reach 20 cm in height and 5 cm in width. They are ovoid in the lower part and cylindrical above, with a pronounced hip in the middle. A pair of fringed wings (≤4 mm wide) runs the whole length of the pitcher cup. The pitcher mouth is round to ovate and oblique throughout. The peristome
is up to 6 mm wide and bears very small but distinct teeth along its inner margin. The pitcher lid is ovate and lacks appendages. It bears a short but distinct keel and often has a very broad insertion. A simple or bifurcate spur
(≤12 mm long) is inserted near the base of the lid.
Upper pitchers are similar in most respects to their lower counterparts. They are up to 20 cm high and 3 cm wide. They are infundibular in the lowermost part, narrowly ovoid in the next part, and cylindrical above. The peristome lacks teeth in upper pitchers. The lid is narrower and has a less obtuse apex. The spur is simple and much smaller, reaching only 5 mm in length.
Nepenthes benstonei has a racemose
inflorescence
. The peduncle
is up to 20 cm long and the rachis
up to 30 cm long. The first flower is generally borne on a pedicel
, sometimes with a simple, lanceolate bract
eole (≤1.5 cm long). Subsequent flowers are produced on pedicels or two-flowered partial peduncles, which lack bracteoles. Sepal
s are ovate and around 4 mm long. Male inflorescences usually bear around twice as many flowers as female ones. N. benstonei is one of the few Nepenthes species known to produce multiple inflorescences concurrently on a single stem. Two to three are usually produced, originating from sequential nodes at the top of the stem. This unusual reproductive habit has also been observed, although much more rarely, in N. alba
, N. ampullaria
, N. attenboroughii
, N. rigidifolia
, N. sanguinea
, and N. thai
. It is seen even more frequently in N. philippinensis
.
The stem and lamina have a sparse indumentum
of simple white hairs. Short, branched reddish-brown hairs line the margins of the lamina. The outer surfaces of the pitchers bear a sparse covering of short, branched red hairs. The same hairs are more densely present on the margins of the lid and upper part of the pitcher directly below the peristome. Immature inflorescences have an indumentum of short white and red hairs throughout.
The stem and leaves of N. benstonei bear a thick, waxy cuticle
that often gives a whitish-blue sheen to the lamina and pitchers. Inflorescences are distinctly waxy throughout.
No infraspecific taxa of N. benstonei have been described.
. It is known with certainty only from the summits of low hills in Kelantan
and northern Terengganu
, although collections and observations from Pahang
and Taman Negara National Park
suggest it may be more widespread in the region.
Nepenthes benstonei grows terrestrially amongst open, secondary vegetation, where it is exposed to direct sunlight. It is very abundant near the summit of Bukit Bakar, where it grows on cuttings beside a paved road leading to a Telekom Malaysia
station at the summit. The exact boundaries of its altitudinal distribution are not yet known, but it has been recorded from elevations of 450–600 m above sea level.
Although the extent of its range is uncertain, N. benstonei appears to have a secure future in the wild as the type locality lies within a protected area and the species's unremarkable appearance means over-collection does not pose a serious threat.
on the leaves. Subsequent field studies have shown that the former is not unique to N. benstonei, but also occasionally occurs in other Nepenthes. Likewise, a number of other species, such as N. hirsuta
from Borneo
, are known to produce a waxy cuticle, although it is less developed than in N. benstonei. Otherwise, N. benstonei lacks remarkable characteristics and is distinguished from related species on the basis of its stem, leaves, peristome, lid, indumentum, and glands of the digestive zone.
Nepenthes benstonei appears to be related to N. sanguinea
, which is also endemic to Peninsular Malaysia. It can be distinguished on the basis of its significantly larger leaves, which are often sub-petiolate and differ in shape. Nepenthes benstonei also has longer tendrils and a denser indumentum. The presence of teeth on the peristome of lower pitchers and of a thick, waxy cuticle on the leaves also serve to distinguish these taxa. In addition, herbarium specimens of N. benstonei tend to dry to a lighter colour than those of N. sanguinea.
The pitchers of N. benstonei also resemble those of N. smilesii
from Indochina
. Clarke suggests that N. benstonei may represent an evolution
ary link between the Nepenthes taxa
of Indochina and Peninsular Malaysia. Nepenthes benstonei also superficially resembles N. macrovulgaris
from Borneo
. It differs in producing multiple inflorescences, which are longer than those of N. macrovulgaris and bear one- or two-flowered partial peduncles, as opposed to exclusively two-flowered in the latter. The waxy coating of its leaves also separates these species. Nepenthes benstonei has also been compared to N. albomarginata
, although the presence of a white band below the peristome, which gives the latter its name, makes identification easy. Upper pitchers of N. benstonei could be confused with those of N. mirabilis
, although all other parts of the plant have little in common.
In 2001, Charles Clarke
performed a cladistic analysis of the Nepenthes species of Sumatra and Peninsular Malaysia
using 70 morphological characteristics of each taxon. The resultant cladogram
placed N. benstonei in an unresolved polytomy
at the base of the Montanae/Nobiles
clade
, together with N. rhombicaulis
.
was discovered by Andrew Hurrell at the foot of Bukit Bakar, where the two species occur sympatrically.
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 Peninsular Malaysia
Peninsular Malaysia
Peninsular Malaysia , also known as West Malaysia , is the part of Malaysia which lies on the Malay Peninsula. Its area is . It shares a land border with Thailand in the north. To the south is the island of Singapore. Across the Strait of Malacca to the west lies the island of Sumatra...
, where it occurs at elevations of 450–600 m above sea level. The specific epithet benstonei honours botanist Benjamin Clemens Stone
Benjamin Clemens Stone
Benjamin Clemens Masterman Stone was a British–American botanist.Stone was born in Shanghai, China to a British father, who worked for the government, and an American mother. He graduated from Pomona College in Claremont, California and, in 1960, received a Ph.D...
, who was one of the first to collect the species.
Botanical history
In 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...
", 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,...
tentatively referred specimens collected from Bukit Bakar, near Macang, Kelantan
Kelantan
Kelantan is a state of Malaysia. The capital and royal seat is Kota Bharu. The Arabic honorific of the state is Darul Naim, ....
, to N. sanguinea
Nepenthes sanguinea
Nepenthes sanguinea is a large and vigorous Nepenthes pitcher plant species, native to the Malay Peninsula, where it grows at 900–1800 m altitude. The pitchers are variable in size, from 10–30 cm tall, and range from green and yellow to orange and red. The insides of the pitchers are usually...
. These were Stone & Chin 15238, deposited at Universiti Kebangsaan Malaysia
Universiti Kebangsaan Malaysia
The National University of Malaysia is a public university located in Bangi, Selangor which is about 35 km south of Kuala Lumpur...
near Kuala Lumpur
Kuala Lumpur
Kuala Lumpur is the capital and the second largest city in Malaysia by population. The city proper, making up an area of , has a population of 1.4 million as of 2010. Greater Kuala Lumpur, also known as the Klang Valley, is an urban agglomeration of 7.2 million...
, and Shah & Shukor 3168, also held at the university as well as the Forest Research Institute of Malaysia in Kepong
Kepong
Kepong is a major township in the borders Kuala Lumpur and Selangor, located about 6 km from the KLCC, to the south of Sungai Buloh, to the east of Bukit Lanjan, to the west of Selayang and to the north of Jalan Duta. The main access roads to Kepong are Damansara-Puchong Expressway, Jalan...
. They noted that the plants exhibited some unusual morphological features, such as larger leaves and decurrent, almost petiolate leaf bases, suggesting that they might represent an as-yet undescribed 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...
.
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...
.
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. benstonei, Clarke s.n., was collected 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...
on July 24, 1998, on Bukit Bakar in Kelantan
Kelantan
Kelantan is a state of Malaysia. The capital and royal seat is Kota Bharu. The Arabic honorific of the state is Darul Naim, ....
at an altitude of between 450 and 550 m. It is deposited at the Forest Research Institute of Malaysia in Kepong
Kepong
Kepong is a major township in the borders Kuala Lumpur and Selangor, located about 6 km from the KLCC, to the south of Sungai Buloh, to the east of Bukit Lanjan, to the west of Selayang and to the north of Jalan Duta. The main access roads to Kepong are Damansara-Puchong Expressway, Jalan...
. Isotype
Isotype
Isotype can refer to:* In crystallography, an "isotype" is a synonym for isomorph* In biology, per the International Code of Botanical Nomenclature, the "isotype" is a duplicate of the holotype....
s are held at the Bogor Botanical Gardens
Bogor Botanical Gardens
The Bogor Botanical Gardens are located 60 km south of the capital of Jakarta in Bogor, Indonesia. The botanical gardens are situated in the city center of Bogor and adjoin the Istana Bogor...
, 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...
, 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, the Forest Department in Sandakan
Sandakan
Sandakan is the second-largest city in Sabah, East Malaysia, on the north-eastern coast of Borneo. It is located on the east coast of the island and it is the administrative centre of Sandakan Division and was the former capital of British North Borneo...
, and the Singapore Botanic Gardens
Singapore Botanic Gardens
The Singapore Botanic Gardens is a 74-hectare botanical garden in Singapore. It is half the size of the Royal Botanic Gardens in Kew or around one-fifth the size of Central Park in New York. It is the only botanic garden in the world that opens from 5 a.m...
.
Ridley 16097
Although only described towards the very end of the twentieth century, N. benstonei was probably first collected in July 1911 by Henry Nicholas RidleyHenry Nicholas Ridley
Henry Nicholas Ridley CMG , MA , FRS, FLS, F.R.H.S. was an English botanist and geologist.Born at West Harling Hall, Norfolk, England...
on Mount Tahan
Mount Tahan
Mount Tahan , at 2,187 m, is the highest point in Peninsular Malaysia. It is located within the Taman Negara national forest, in the state of Pahang. The mountain is part of the Tahan Range and popular to local climbers....
in Pahang
Pahang
Pahang is the third largest state in Malaysia, after Sarawak and Sabah, occupying the huge Pahang River river basin. It is bordered to the north by Kelantan, to the west by Perak, Selangor, Negeri Sembilan, to the south by Johor and to the east by Terengganu and the South China Sea.Its state...
. The specimen, Ridley 16097, includes female floral material and is deposited at the herbarium of the Singapore Botanic Gardens
Singapore Botanic Gardens
The Singapore Botanic Gardens is a 74-hectare botanical garden in Singapore. It is half the size of the Royal Botanic Gardens in Kew or around one-fifth the size of Central Park in New York. It is the only botanic garden in the world that opens from 5 a.m...
.
In his seminal monograph "The Nepenthaceae of the Netherlands Indies
The Nepenthaceae of the Netherlands Indies
"The Nepenthaceae of the Netherlands Indies" is a seminal monograph by B. H. Danser on the tropical pitcher plants of the Dutch East Indies, North Borneo, the Malay Peninsula, and eastern New Guinea...
", B. H. Danser
B. H. Danser
Benedictus Hubertus Danser , often abbreviated B. H. Danser, was a Dutch taxonomist and botanist...
referred Ridley 16097 to N. alata
Nepenthes alata
Nepenthes alata is a tropical pitcher plant endemic to the Philippines. It is found on all the major islands of the archipelago, with the possible exception of Palawan. It is one of the easiest and most popular Nepenthes species in cultivation....
. Nepenthes alata was not known 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 this specimen constituted the sole record of N. alata from Peninsular Malaysia. Danser also treated N. eustachya
Nepenthes eustachya
Nepenthes eustachya is a tropical pitcher plant endemic to Sumatra, where it grows from sea level to an elevation of 1600 m. The specific epithet eustachya, formed from the Greek words eu and stachys , refers to the racemose structure of the inflorescence.-Botanical history:Nepenthes eustachya...
from 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...
in synonymy with N. alata, making its extreme rarity in Peninsular Malaysia difficult to explain.
In 1990, Ruth Kiew identified Ridley 16097 as belonging to N. gracillima
Nepenthes gracillima
Nepenthes gracillima is a highland Nepenthes pitcher plant species, native to Peninsular Malaysia.-Infraspecific taxa:*Nepenthes gracillima f. ramispina Hort.Westphal [=N. ramispina]...
. Kiew explained the apparent near-absence of N. alata from Peninsular Malaysia as follows:
Why has this species not been recollected since 1911 in spite of several botanical expeditions to G. Tahan since then? The answer is quite simple, Ridley's specimen (16097) actually belongs to N. gracillima. Although N. alata Blanco is superficially similar to N. gracillima in its narrow leaf blade which has an attenuate base, the pitchers of these two species are distinct. Those of N. alata are broader (about 4 cm wide) and are distinctly bulbous towards the base compared with the very slender, non-bulbous pitchers of N. gracillima that are 1.5 to 3 cm wide. Nor does the G. Tahan specimen have truly petiolate leaves - its leaf base is narrow and has rolled up during drying. Ridley's specimen from G. Tahan determined by Danser as N. alata bears pitchers typical of N. gracillima and examination of the Tahan population in the field leaves no doubt that it belongs to this species.
Jebb and Cheek supported this interpretation in 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...
".
However, 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...
disagrees with Kiew's identification. He notes that the inflorescences of both N. gracillima and the closely related N. ramispina
Nepenthes ramispina
Nepenthes ramispina is a highland Nepenthes pitcher plant species, native to Peninsular Malaysia. It is very similar to N. gracillma and there is much debate about whether the two should be treated as separate species.-Natural hybrids:...
are very short, rarely exceeding 10 and 20 cm, respectively. Both female inflorescences of Ridley 16097 have a long 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...
and 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...
, each exceeding 20 cm in length. In addition, the flowers of these species are almost always borne on 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....
, unlike those of Ridley 16097, which mostly have two-flowered partial peduncles. Furthermore, the shape of the lamina is unlike that of N. gracillima or N. ramispina; despite being narrow and lanceolate, it is proportionately considerably longer and has a much narrower, almost sub-petiolate base. Kiew noted that the lamina of Ridley 16097 appears narrower due to the way in which the specimen was preserved, but Clarke states that this cannot fully account for the differences. He also notes that the specimen exhibits a decurrent leaf attachment. Taking all of these morphological features into account, Clarke feels that Ridley 16097 most likely represents N. benstonei. Clarke considers that Kiew grouped both N. benstonei and N. ramispina with N. gracillima.
Other specimens
A number of specimens collected from TerengganuTerengganu
Terengganu is a sultanate and constitutive state of federal Malaysia. The state is also known by its Arabic honorific, Darul Iman...
also belong to N. benstonei. These are Shah et al. 3274, which is deposited at the Forest Research Institute of Malaysia in Kepong, and Shah et al. 3283, held at the Universiti Kebangsaan Malaysia near Kuala Lumpur.
Specimens from peninsula
Peninsula
A peninsula is a piece of land that is bordered by water on three sides but connected to mainland. In many Germanic and Celtic languages and also in Baltic, Slavic and Hungarian, peninsulas are called "half-islands"....
r Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...
originally assigned to N. benstonei in Cheek and Jebb's 2001 monograph, "Nepenthaceae
Nepenthaceae (2001 monograph)
"Nepenthaceae" is a monograph by Martin Cheek and Matthew Jebb on the tropical pitcher plants of Malesia, which encompasses Brunei, Indonesia, Malaysia, Papua New Guinea, the Philippines, and Singapore. It was published in 2001 by the National Herbarium of the Netherlands as the fifteenth volume of...
", have since been identified as belonging to a new species, N. thai
Nepenthes thai
Nepenthes thai is a tropical pitcher plant endemic to peninsular Thailand. It grows on limestone hills at elevations of 500–600 m above sea level.Nepenthes thai has no known natural hybrids.-External links:*...
.
Description
Nepenthes benstonei is a climbing plant. The stem can attain a length of 5 m and is up to 0.6 cm in diameter. InternodePlant stem
A stem is one of two main structural axes of a vascular plant. The stem is normally divided into nodes and internodes, the nodes hold buds which grow into one or more leaves, inflorescence , conifer cones, roots, other stems etc. The internodes distance one node from another...
s are cylindrical and up to 15 cm long.
Leaves are coriaceous and sessile to sub-petiolate. The lamina is usually broadly linear-lanceolate in shape, but may also be slightly spathulate. Its base is a broad, amplexicaul sheath with decurrent margins. The lamina can reach 60 cm in length and 9 cm in width. It has a rounded to acute apex. The margins of the lamina usually meet the tendril unequally on both sides, being up to 3 mm apart. Three to five longitudinal veins are present on either side of the midrib. Pinnate veins are almost 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 are up to 60 cm long.
Rosette and lower pitchers reach 20 cm in height and 5 cm in width. They are ovoid in the lower part and cylindrical above, with a pronounced hip in the middle. A pair of fringed wings (≤4 mm wide) runs the whole length of the pitcher cup. The pitcher mouth is round to ovate and oblique throughout. 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 up to 6 mm wide and bears very small but distinct teeth along its inner margin. The pitcher lid is ovate and lacks appendages. It bears a short but distinct keel and often has a very broad insertion. A simple or bifurcate spur
Spur (biology)
A spur in botany is a spike, usually part of a flower.In certain plants, part of a sepal or petal develops into an elongated hollow spike extending behind the flower, containing nectar which is sucked by long-tongued animals . Plants with such structures include Delphinium, Aquilegia, Piperia, and...
(≤12 mm long) is inserted near the base of the lid.
Upper pitchers are similar in most respects to their lower counterparts. They are up to 20 cm high and 3 cm wide. They are infundibular in the lowermost part, narrowly ovoid in the next part, and cylindrical above. The peristome lacks teeth in upper pitchers. The lid is narrower and has a less obtuse apex. The spur is simple and much smaller, reaching only 5 mm in length.
Nepenthes benstonei has a racemose
Raceme
A raceme is a type of inflorescence that is unbranched and indeterminate and bears pedicellate flowers — flowers having short floral stalks called pedicels — along the axis. In botany, axis means a shoot, in this case one bearing the flowers. In a raceme, the oldest flowers are borne...
inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
. The peduncle
Peduncle (botany)
In botany, a peduncle is a stem supporting an inflorescence, or after fecundation, an infructescence.The peduncle is a stem, usually green and without leaves, though sometimes colored or supporting small leaves...
is up to 20 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...
up to 30 cm long. The first flower is generally borne on a pedicel
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....
, sometimes with a simple, lanceolate 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...
eole (≤1.5 cm long). Subsequent flowers are produced on pedicels or two-flowered partial peduncles, which lack bracteoles. Sepal
Sepal
A sepal is a part of the flower of angiosperms . Collectively the sepals form the calyx, which is the outermost whorl of parts that form a flower. Usually green, sepals have the typical function of protecting the petals when the flower is in bud...
s are ovate and around 4 mm long. Male inflorescences usually bear around twice as many flowers as female ones. N. benstonei is one of the few Nepenthes species known to produce multiple inflorescences concurrently on a single stem. Two to three are usually produced, originating from sequential nodes at the top of the stem. This unusual reproductive habit has also been observed, although much more rarely, in N. alba
Nepenthes alba
Nepenthes alba is a tropical pitcher plant endemic to Peninsular Malaysia. The specific epithet alba is derived from the Latin word albus, meaning "white", and refers to the colour of the upper pitchers.-Natural hybrids:...
, 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. attenboroughii
Nepenthes attenboroughii
Nepenthes attenboroughii is a montane species of insectivorous pitcher plant of the genus Nepenthes. It is named after the celebrated broadcaster and naturalist Sir David Attenborough, who is a keen enthusiast of the genus. The species is characterised by its large and distinctive bell-shaped...
, N. rigidifolia
Nepenthes rigidifolia
Nepenthes rigidifolia is a tropical pitcher plant endemic to Sumatra, where it grows at elevations of 1000–1600 m above sea level.The specific epithet rigidifolia is formed from the Latin words rigidus and folia . It refers to the stiff, coriaceous texture of the leaf blade...
, N. sanguinea
Nepenthes sanguinea
Nepenthes sanguinea is a large and vigorous Nepenthes pitcher plant species, native to the Malay Peninsula, where it grows at 900–1800 m altitude. The pitchers are variable in size, from 10–30 cm tall, and range from green and yellow to orange and red. The insides of the pitchers are usually...
, and N. thai
Nepenthes thai
Nepenthes thai is a tropical pitcher plant endemic to peninsular Thailand. It grows on limestone hills at elevations of 500–600 m above sea level.Nepenthes thai has no known natural hybrids.-External links:*...
. It is seen even more frequently in N. philippinensis
Nepenthes philippinensis
Nepenthes philippinensis is a tropical pitcher plant endemic to the Philippines. It is known from Palawan and the Calamian Islands , where it grows at 0–600 m above sea level....
.
The stem and lamina have a sparse indumentum
Indumentum
The indumentum is a covering of fine hairs or bristles on a plant or insect.In plants, the indumentum types are:*pubescent*hirsute*pilose*villous*tomentose*stellate*scabrous*scurfy...
of simple white hairs. Short, branched reddish-brown hairs line the margins of the lamina. The outer surfaces of the pitchers bear a sparse covering of short, branched red hairs. The same hairs are more densely present on the margins of the lid and upper part of the pitcher directly below the peristome. Immature inflorescences have an indumentum of short white and red hairs throughout.
The stem and leaves of N. benstonei bear a thick, waxy cuticle
Plant cuticle
Plant cuticles are a protective waxy covering produced only by the epidermal cells of leaves, young shoots and all other aerial plant organs without periderm...
that often gives a whitish-blue sheen to the lamina and pitchers. Inflorescences are distinctly waxy throughout.
No infraspecific taxa of N. benstonei have been described.
Ecology
Nepenthes benstonei is endemic to Peninsular MalaysiaPeninsular Malaysia
Peninsular Malaysia , also known as West Malaysia , is the part of Malaysia which lies on the Malay Peninsula. Its area is . It shares a land border with Thailand in the north. To the south is the island of Singapore. Across the Strait of Malacca to the west lies the island of Sumatra...
. It is known with certainty only from the summits of low hills in Kelantan
Kelantan
Kelantan is a state of Malaysia. The capital and royal seat is Kota Bharu. The Arabic honorific of the state is Darul Naim, ....
and northern Terengganu
Terengganu
Terengganu is a sultanate and constitutive state of federal Malaysia. The state is also known by its Arabic honorific, Darul Iman...
, although collections and observations from Pahang
Pahang
Pahang is the third largest state in Malaysia, after Sarawak and Sabah, occupying the huge Pahang River river basin. It is bordered to the north by Kelantan, to the west by Perak, Selangor, Negeri Sembilan, to the south by Johor and to the east by Terengganu and the South China Sea.Its state...
and Taman Negara National Park
Taman Negara National Park
Taman Negara was established in Malaysia in 1938/1939 as the King George V National Park. It was renamed to Taman Negara after independence, which literally means "national park" in Malay...
suggest it may be more widespread in the region.
Nepenthes benstonei grows terrestrially amongst open, secondary vegetation, where it is exposed to direct sunlight. It is very abundant near the summit of Bukit Bakar, where it grows on cuttings beside a paved road leading to a Telekom Malaysia
Telekom Malaysia
Telekom Malaysia Berhad , DBA TM, is the largest integrated solutions provider in Malaysia, and one of Asia's leading communications companies, with a market capitalisation of RM13.9 billion and a workforce of 26,629 employees. Established as the Telecommunications Department of Malaya in 1946, it...
station at the summit. The exact boundaries of its altitudinal distribution are not yet known, but it has been recorded from elevations of 450–600 m above sea level.
Although the extent of its range is uncertain, N. benstonei appears to have a secure future in the wild as the type locality lies within a protected area and the species's unremarkable appearance means over-collection does not pose a serious threat.
Related species
In his description of N. benstonei, Charles Clarke noted two characteristics that he considered unique among Nepenthes. These were the production of multiple inflorescences and the presence of a thick, waxy cuticlePlant cuticle
Plant cuticles are a protective waxy covering produced only by the epidermal cells of leaves, young shoots and all other aerial plant organs without periderm...
on the leaves. Subsequent field studies have shown that the former is not unique to N. benstonei, but also occasionally occurs in other Nepenthes. Likewise, a number of other species, such as 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...
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....
, are known to produce a waxy cuticle, although it is less developed than in N. benstonei. Otherwise, N. benstonei lacks remarkable characteristics and is distinguished from related species on the basis of its stem, leaves, peristome, lid, indumentum, and glands of the digestive zone.
Nepenthes benstonei appears to be related to N. sanguinea
Nepenthes sanguinea
Nepenthes sanguinea is a large and vigorous Nepenthes pitcher plant species, native to the Malay Peninsula, where it grows at 900–1800 m altitude. The pitchers are variable in size, from 10–30 cm tall, and range from green and yellow to orange and red. The insides of the pitchers are usually...
, which is also endemic to Peninsular Malaysia. It can be distinguished on the basis of its significantly larger leaves, which are often sub-petiolate and differ in shape. Nepenthes benstonei also has longer tendrils and a denser indumentum. The presence of teeth on the peristome of lower pitchers and of a thick, waxy cuticle on the leaves also serve to distinguish these taxa. In addition, herbarium specimens of N. benstonei tend to dry to a lighter colour than those of N. sanguinea.
The pitchers of N. benstonei also resemble those of N. smilesii
Nepenthes smilesii
Nepenthes smilesii is a tropical pitcher plant native to northeastern Thailand, southern Laos, Cambodia, and western Vietnam. Nepenthes smilesii can tolerate an extended dry season and is most common in open, sandy savannah and grassland....
from Indochina
Indochina
The Indochinese peninsula, is a region in Southeast Asia. It lies roughly southwest of China, and east of India. The name has its origins in the French, Indochine, as a combination of the names of "China" and "India", and was adopted when French colonizers in Vietnam began expanding their territory...
. Clarke suggests that N. benstonei may represent an evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
ary link between the Nepenthes taxa
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...
of Indochina and Peninsular Malaysia. Nepenthes benstonei also superficially resembles N. macrovulgaris
Nepenthes macrovulgaris
Nepenthes macrovulgaris , or the Serpentine Pitcher-Plant, is a tropical pitcher plant endemic to Borneo. It is a lowland plant that typically grows at altitudes ranging from 300 to 1200 m in sub-montane forest clearings and mossy forest...
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....
. It differs in producing multiple inflorescences, which are longer than those of N. macrovulgaris and bear one- or two-flowered partial peduncles, as opposed to exclusively two-flowered in the latter. The waxy coating of its leaves also separates these species. Nepenthes benstonei has also been compared to N. albomarginata
Nepenthes albomarginata
Nepenthes albomarginata , the White-Collared Pitcher-Plant, is a tropical pitcher plant native to Borneo, Peninsular Malaysia, and Sumatra...
, although the presence of a white band below the peristome, which gives the latter its name, makes identification easy. Upper pitchers of N. benstonei could be confused with those of N. mirabilis
Nepenthes mirabilis
Nepenthes mirabilis , or the Common Swamp Pitcher-Plant, is a tropical carnivorous plant species of the pitfall trap variety. It has by far the widest distribution of any Nepenthes species and is known from the following countries and regions: Borneo, Sumatra, Thailand, Peninsular Malaysia,...
, although all other parts of the plant have little in common.
In 2001, 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...
performed a cladistic analysis of the Nepenthes species of Sumatra and Peninsular Malaysia
Peninsular Malaysia
Peninsular Malaysia , also known as West Malaysia , is the part of Malaysia which lies on the Malay Peninsula. Its area is . It shares a land border with Thailand in the north. To the south is the island of Singapore. Across the Strait of Malacca to the west lies the island of Sumatra...
using 70 morphological characteristics of each taxon. The resultant cladogram
Cladogram
A cladogram is a diagram used in cladistics which shows ancestral relations between organisms, to represent the evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of morphological characters, DNA and RNA sequencing data and computational...
placed N. benstonei in an unresolved polytomy
Polytomy
A polytomy , meaning many temporal based branches, is a section of a phylogeny in which the evolutionary relationships can not be fully resolved to dichotomies. In a phylogenetic tree, a polytomy is represented as a node which has more than two immediate descending branches...
at the base of the Montanae/Nobiles
Nepenthes classification
The taxonomy of Nepenthes has been revised several times during the nineteenth and twentieth centuries.-Nineteenth century:The first subgeneric division of the Nepenthes was made by Joseph Dalton Hooker in his 1873 monograph, "Nepenthaceae"...
clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
, together with N. rhombicaulis
Nepenthes rhombicaulis
Nepenthes rhombicaulis is a tropical pitcher plant endemic to Sumatra. The specific epithet rhombicaulis is formed from the Latin words rhombicus, meaning "rhomboid", and caulis, "stem"...
.
Natural hybrids
Only one natural hybrid involving N. benstonei is known. A single example of N. benstonei × N. mirabilisNepenthes mirabilis
Nepenthes mirabilis , or the Common Swamp Pitcher-Plant, is a tropical carnivorous plant species of the pitfall trap variety. It has by far the widest distribution of any Nepenthes species and is known from the following countries and regions: Borneo, Sumatra, Thailand, Peninsular Malaysia,...
was discovered by Andrew Hurrell at the foot of Bukit Bakar, where the two species occur sympatrically.
External links
- Photographs of N. benstonei at the Carnivorous Plant Photofinder