Nepenthes lingulata
Encyclopedia
Nepenthes lingulata 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 northern 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...

. The species is characterised by the highly developed appendage present on the underside of the lid. The specific epithet lingulata is derived 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...

 word lingula, meaning "small tongue", and refers to this unique morphological feature.

Botanical history

Nepenthes lingulata was discovered during a series a field trips to 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...

 between 2004 and 2005. The first collection of the species was made on April 22, 2005, south of Padang Sidempuan in the Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...

n province of North Sumatra
North Sumatra
North Sumatra is a province of Indonesia on the Sumatra island. Its capital is Medan. It is the most populous Indonesian province outside of Java. It is slightly larger than Sri Lanka in area.- Geography and population :...

.

The formal description of N. lingulata was published in 2006 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...

. The herbarium specimen Lee, Hernawati, Akhriadi NP 432 was designated as 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...

. It is deposited at the Herbarium of Andalas University
Andalas University
Andalas University is the oldest university in Indonesia outside of Java. The university consists of eleven faculties, with most located at the main campus Limau Manis, 12 km from the center of Padang, West Sumatra. This campus occupies 5 km², at an elevation of about 100 m. The Faculty...

 (ANDA).

Description

Nepenthes lingulata is a climbing plant. The stem, which is usually less than 4 mm thick, grows to 5 m in length and is terete in cross section. 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 9 cm long.

Leaves are sessile and coriaceous in texture. The lamina of rosettes is lanceolate and up to 9.6 cm long and 3 cm wide. Climbing stems produce smaller leaves, with the elliptical-ovate lamina reaching 4.8 cm in length and 2.1 cm in width. Two longitudinal veins are present on either side of the midrib. Tendril
Tendril
In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape that is used by climbing plants for support, attachment and cellular invasion by parasitic plants, generally by twining around suitable hosts. They do not have a lamina or blade, but they can photosynthesize...

s are up to 25 cm long in rosettes and up to 14 cm long in climbing stems.

Rosette and lower pitchers are relatively large, reaching 28 cm in height and 4.5 cm in width. They are infundibular in the lower half and cylindrical in the upper half. The inner surface of the pitcher is only glandular in the lower half, with the waxy zone continuing to the top of the pitcher. A pair of wings (≤2.5 mm wide) with fringe elements up to 1 cm long runs down the front of the pitcher. The pitcher mouth is ovate and bears a flattened 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...

 up to 3 cm wide. The inner margin of the peristome is lined with a series of teeth up to 2 mm long. 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 broadly triangular and may be up to 7.5 cm long and 5.5 cm wide. Curiously, no nectar gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...

s are present on the underside of the lid. A single filiform appendage (≤4 cm long) hangs over the pitcher mouth, the apical end of which is covered with scattered raised nectar glands. A branched 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...

 (≤1.5 cm long) is inserted near the base of the lid.
Upper pitchers are smaller than lower pitchers, growing to 12.3 cm in height and 2 cm in width. Their wings are reduced to ribs. The pitcher mouth is ovate to rounded and bears a rounded or slightly flattened peristome (≤4 mm wide). The lid measures up to 2 cm in length and 2 cm in width. As in lower pitchers, a single highly developed appendage is located on the underside of the lid.

Nepenthes lingulata 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...

. In male plants, 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 2.3 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 4.5 cm long, whereas in female plants the peduncle is up to 5.5 cm long and the rachis up to 3.5 cm long. 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....

s are around 3–4 mm long and bear a single flower. 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 2 mm long in male inflorescences and 5 mm long in female inflorescences.

A very dense indumentum
Indumentum
The indumentum is a covering of fine hairs or bristles on a plant or insect.In plants, the indumentum types are:*pubescent*hirsute*pilose*villous*tomentose*stellate*scabrous*scurfy...

 of long woolly grey-brown hairs is present on immature tendrils and the outer surface of pitchers. The hairs on the margins of the leaves are caducous. Scattered grey-brown stellate hairs cover the upper surface and margins of the lower surface of the lid.

Leaves are dark green on their upper surface and pale green on their lower surface. The midrib and edges of the leaves are purple in some plants. Rosette plants have light green stems, which gradually become dark purple as the plant begins to climb. Tendrils are dark purple. As in the closely related N. izumiae
Nepenthes izumiae
Nepenthes izumiae is a tropical pitcher plant endemic to Sumatra, where it grows in montane forest at 1700–1900 m above sea level. It appears to be most closely related to N. lingulata and N...

, the pitchers are dark purple to black on their outer surface, while the inside surface is pale bluish-green with purple spots. Inflorescences are pale green. Sepals range in colour from light green to reddish-purple.

Ecology

Nepenthes lingulata is endemic to the Barisan Mountains
Barisan Mountains
The Bukit Barisan or the Barisan Mountains are a mountain range on the western side of Sumatra, Indonesia, covering nearly 1,700 km from the north to the south of the island. The Bukit Barisan range consists primarily of volcanoes shrouded in dense jungle cover, including Sumatran tropical pine...

 that line the western side of 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...

. It is known only from the type locality, south of Padang Sidempuan in North Sumatra
North Sumatra
North Sumatra is a province of Indonesia on the Sumatra island. Its capital is Medan. It is the most populous Indonesian province outside of Java. It is slightly larger than Sri Lanka in area.- Geography and population :...

. The species has an altitudinal distribution of 1700–2100 m above sea level.

Nepenthes lingulata grows in upper montane mossy forest. It 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...

.

In the wild, N. lingulata occurs sympatrically with N. bongso
Nepenthes bongso
Nepenthes bongso is a tropical pitcher plant endemic to Sumatra, where it has an altitudinal distribution of 1000–2700 m above sea level. The specific epithet bongso refers to the Indonesian legend of Putri Bungsu , the spirit guardian of Mount Marapi.The species was formally described by Pieter...

, N. dubia
Nepenthes dubia
Nepenthes dubia is a tropical pitcher plant endemic to Sumatra. The specific epithet dubia is the Latin word for "doubtful".-Botanical history:...

, N. gymnamphora
Nepenthes gymnamphora
Nepenthes gymnamphora is a tropical pitcher plant native to the Indonesian islands of Java and Sumatra. There is much debate surrounding the taxonomic status of this species and the taxa N. pectinata and N. xiphioides....

, and N. jamban
Nepenthes jamban
Nepenthes jamban is a tropical pitcher plant endemic to northern Sumatra. The specific epithet jamban is the Indonesian word for "toilet" and refers to the shape of the pitchers.-Botanical history:...

. A natural hybrid with N. jamban has been recorded.

Carnivory

Many Nepenthes species, including the closely related N. izumiae
Nepenthes izumiae
Nepenthes izumiae is a tropical pitcher plant endemic to Sumatra, where it grows in montane forest at 1700–1900 m above sea level. It appears to be most closely related to N. lingulata and N...

, have a glandular crest on the underside of the lid. However, the highly developed filiform appendage of N. lingulata is unique in its size.

Nepenthes lingulata is also unusual in that the undersurface of the lid is completely devoid of glands, except for a small group concentrated at the end of the tongue-like appendage. This structure likely serves to lure insects into a precarious position over the pitcher mouth, where they may lose their footing and fall into the pitcher fluid, eventually drowning.

A similar trapping method is employed by the Bornean
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....

 endemic N. bicalcarata
Nepenthes bicalcarata
Nepenthes bicalcarata , also known as the Fanged Pitcher-Plant, is a tropical pitcher plant endemic to northwestern Borneo.-Botanical history:...

, which has a pair of spines positioned over the pitcher mouth. However, in that species the spines are structures of the peristome and not the lid.

Related species

Nepenthes lingulata is thought to be most closely related to N. izumiae
Nepenthes izumiae
Nepenthes izumiae is a tropical pitcher plant endemic to Sumatra, where it grows in montane forest at 1700–1900 m above sea level. It appears to be most closely related to N. lingulata and N...

, with which it shares the general morphology and colouration of its pitchers. However, it can be easily distinguished from that species on the basis of the highly developed filiform appendage that hangs over the pitcher mouth. It also differs in having a triangular lid, as opposed to the orbicular lid of N. izumiae. Furthermore, N. lingulata completely lacks nectar glands on the underside of the lid and has a very dense woolly indumentum.

Nepenthes lingulata is more distantly related to the Sumatran species N. densiflora
Nepenthes densiflora
Nepenthes densiflora is a tropical pitcher plant endemic to Sumatra, where it grows at an altitude of between 1700 and 3200 m above sea level.No forms or varieties of N. densiflora have been described.-Taxonomy:...

, N. diatas
Nepenthes diatas
Nepenthes diatas is a tropical pitcher plant endemic to Sumatra, where it grows at an altitude of between 2400 and 2900 m above sea level....

, N. singalana
Nepenthes singalana
Nepenthes singalana is a highland Nepenthes pitcher plant species native to the island of Sumatra. It is most closely related to N. diatas and N. spathulata.-Taxonomy:...

, and N. spathulata
Nepenthes spathulata
Nepenthes spathulata is a tropical pitcher plant native to Java and Sumatra, where it grows at elevations of between 1100 and 2900 m above sea level. The specific epithet spathulata is derived from the Latin word spathulatus, meaning "spatula shaped", and refers to the shape of the...

.

Natural hybrids

A natural hybrid involving N. jamban
Nepenthes jamban
Nepenthes jamban is a tropical pitcher plant endemic to northern Sumatra. The specific epithet jamban is the Indonesian word for "toilet" and refers to the shape of the pitchers.-Botanical history:...

and N. lingulata has been recorded.
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