Nepenthes bellii
Encyclopedia
Nepenthes bellii 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 the Philippine
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...

 islands of Mindanao
Mindanao
Mindanao is the second largest and easternmost island in the Philippines. It is also the name of one of the three island groups in the country, which consists of the island of Mindanao and smaller surrounding islands. The other two are Luzon and the Visayas. The island of Mindanao is called The...

 and Dinagat
Dinagat Island
' is a province in the Philippines, located on the south side of Leyte Gulf. Leyte is to its west, across Surigao Strait, and Mindanao is to its south...

, where it grows at elevations of 0–800 m above sea level.

The specific epithet bellii honours American botanist Clyde Ritchie Bell.

Botanical history

Nepenthes bellii 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...

 by Katsuhiko Kondo in the November–December 1969 issue of the Bulletin of the Torrey Botanical Club. 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...

, designated as Kondo 11514, was collected on April 14, 1968, from a "[s]wimming pool between Hayangobon & Carrascar" in Surigao, Mindanao
Mindanao
Mindanao is the second largest and easternmost island in the Philippines. It is also the name of one of the three island groups in the country, which consists of the island of Mindanao and smaller surrounding islands. The other two are Luzon and the Visayas. The island of Mindanao is called The...

, at an altitude of 800 m. It was collected by Kondo as part of a 1968 trip to 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...

 alongside J. V. Pancho. The holotype is deposited at 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...

 of the North Carolina Botanical Garden
North Carolina Botanical Garden
The North Carolina Botanical Garden is a botanical garden operated by the University of North Carolina at Chapel Hill in Chapel Hill, North Carolina. The primary goal of the Garden is to research, catalog, and promote the native plant species of North Carolina...

 (NCU); 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 KC and Nagoya herbaria.

Nepenthes globamphora, which is now considered to be a heterotypic synonym of N. bellii, was described by 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....

 and Masami Toyoshima in a 1972 issue of The Gardens' Bulletin Singapore. The holotype of this taxon, designated as Kurata & Toyoshima 1128, was collected on August 22, 1965, from Mount Legaspi in Surigao del Sur
Surigao del Sur
' is a province of the Philippines located in the CARAGA region in Mindanao. Its capital is Tandag City and borders Surigao del Norte to the north, Agusan del Norte and Agusan del Sur to the west, and Davao Oriental to the south...

, Mindanao
Mindanao
Mindanao is the second largest and easternmost island in the Philippines. It is also the name of one of the three island groups in the country, which consists of the island of Mindanao and smaller surrounding islands. The other two are Luzon and the Visayas. The island of Mindanao is called The...

, at an elevation of 270 m. It is deposited at the herbarium of the Nippon Dental College (NDC). A very early mention of N. globamphora (at the time a nomen nudum
Nomen nudum
The phrase nomen nudum is a Latin term, meaning "naked name", used in taxonomy...

as the taxon was undescribed
Undescribed taxon
In taxonomy, an undescribed taxon is a taxon that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. Until such a description has been published, the taxon has no formal or...

) appeared in a 1966 issue (volume 36, page 15) of the Journal of Insectivorous Plant Society
Journal of Insectivorous Plant Society
The is a quarterly Japanese-language periodical and the official publication of the Insectivorous Plant Society of Japan. The journal was established in January 1950. As of 2010, it is published in A4 format and totals around 120 pages annually...

.

The next detailed treatments of N. bellii appeared in 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,...

's 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...

", and their 2001 revision, "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...

". In the latter publication, Cheek and Jebb noted that little ecological data was known about the species and that it was represented in herbaria by only the two type collections (Kondo 11514 and Kurata & Toyoshima 1128) at the time. Stewart McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

's 2009 monograph, Pitcher Plants of the Old World
Pitcher Plants of the Old World
Pitcher Plants of the Old World is a two-volume monograph by Stewart McPherson on the pitcher plants of the genera Nepenthes and Cephalotus. It was published in May 2009 by Redfern Natural History Productions...

, presented an updated description and colour habitat photographs of N. bellii.

Description

Nepenthes bellii is a climbing plant growing to a height of 2.5 m and occasionally even 10 m. The stem, which may be branched, often scrambles through vegetation but may also grow prostrate along the ground. It is terete or slightly angular and up to 5 mm in diameter, with internodes up to 2 cm long.

Leaves are coriaceous and sessile. The lamina (leaf blade) is linear to slightly lanceolate or narrowly elliptic and measures up to 18 cm in length by 3 cm in width. Its apex is acute or obtuse, whereas the base is slightly attenuate and clasps the stem for half to three-quarters of its circumference. It is also slightly auriculate and has an oblique attachment to the stem. The laminar base may be decurrent down the stem to varying degrees or not decurrent at all. Three longitudinal veins are present on either side of the midrib and restricted to the distal third of the lamina. Pinnate veins are abundant and run obliquely to the laminar margin. The 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 proportionately very long, especially those bearing lower pitchers, which may be more than 30 cm long.
Rosette and lower pitchers are usually cylindrical or ellipsoidal throughout, but may also be urceolate or subglobose. They are small, reaching only 9 cm in height by 5 cm in width. The basal half of the pitcher cup may be somewhat swollen, forming a slight hip around the middle. A pair of wings (≤12 mm wide) runs down the ventral surface of the pitcher cup. These wings are densely fringed with filaments up to 9 mm long, which may be grouped in clusters of two or three (the clusters spaced up to 1.3 mm apart), but are often borne singly (particularly in young plants). The pitcher mouth ranges in shape from suborbicular to broadly ovate and is more or less straight, being only slightly oblique. 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 bulbous and cylindrical, subcylindrical, or flattened. It measures up to 10 mm in width, becoming broader and rising slightly towards the top. The peristome bears ribs up to 1 mm high and spaced up to 1.5 mm apart, which terminate in narrow teeth (≤3 mm long) on the inner margin on the peristome. Its outer margin may undulate slightly. The 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:...

 or lid varies in shape from elliptic to ovate or broadly ovate. It has a rounded apex and may have a somewhat cordate base. It measures up to 4 cm in length by 3.5 cm in width. No appendages are present on the lower surface of the lid, although it bears a small number (5 or 6) of sparsely scattered nectar glands. These nectaries are transversely elliptic to circular in shape and measure 0.2–0.4 mm in length. They are unbordered, pit-like, and deep. An unbranched spur up to 9 mm long is inserted near the base of the lid.

Upper pitchers are infundibular throughout, being variably swollen in the upper portion. They are frequently produced, although Cheek and Jebb reported seeing a flowering specimen that lacked upper pitchers. Aerial traps are even smaller than their terrestrial counterparts and some of the smallest pitchers in the genus, reaching only 7.5 cm in height and 3 cm in width. Wings are reduced to a maximum width of only 4 mm with fringe elements up to 3 mm long, but may be absent altogether. The ventral surface between the wings or wing vestiges is usually slightly flattened. The peristome is often cylindrical and bulbous as in lower pitchers, but only up to 5 mm wide. It rises towards the rear, where it is broader. It bears ribs up to 0.5 mm high and spaced up to 1 mm apart, although in some specimens they are inconspicuous. The narrow peristome teeth are up to 1 mm long. The lid, which lacks appendages, is elliptic to ovate and up to 3 cm long by 2 cm wide. The unbranched spur reaches 7 mm in length.

Nepenthes bellii 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...

 up to 15 cm long by 1 cm wide. 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...

 itself reaches up to 9 cm in length, with a basal diameter of 1 mm. Flowers, which number up to 40 per inflorescence, are borne on one-flowered, ebracteate 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....

 up to 4 mm long. Tepal
Tepal
Tepals are elements of the perianth, or outer part of a flower, which include the petals or sepals. The term tepal is more often applied specifically when all segments of the perianth are of similar shape and color, or undifferentiated, which is called perigone...

s are ovate and up to 2.5 mm long by 1.2 mm wide. The androphore is around 1.5 mm long. Fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

s measure up to 20 mm in length.

An inconspicuous 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 reddish or rust-coloured simple (unbranched) hairs measuring 0.1 mm in length may be present on the pitchers and inflorescence. Tepals are minutely tomentose. The stem, laminae and androphores are typically glabrous.

The stem, tendrils and midribs are most commonly yellow to green, but may be tinged orange or red in some specimens; this more intense colouration seems to be associated with drought
Drought
A drought is an extended period of months or years when a region notes a deficiency in its water supply. Generally, this occurs when a region receives consistently below average precipitation. It can have a substantial impact on the ecosystem and agriculture of the affected region...

 stress. The laminae are usually green, but may be orange, red, or even purple when young. These developing leaves gradually turn green with age. Plants often consist of 2 or 3 reddish leaves at the top with many green leaves below. Lower pitchers are mostly green, yellow, orange or sometimes red, with darker blotches of orange to purple. The peristome and lid may be any of the base colours of the pitcher exterior, but without blotches. The inner surface is yellow or green and may have reddish speckles. Upper pitchers are similar but often lack blotches, being a solid green, yellow, orange, or red, although reddish aerial traps occasionally do bear some darker markings. Sometimes upper pitchers may have a contrasting combination of colours in the form of a reddish pitcher body and yellow or green peristome, or vice versa.

No infraspecific taxa of N. bellii have been described.

Ecology

Nepenthes bellii is endemic to 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...

. It is common on Dinagat
Dinagat Island
' is a province in the Philippines, located on the south side of Leyte Gulf. Leyte is to its west, across Surigao Strait, and Mindanao is to its south...

 and northern Mindanao
Mindanao
Mindanao is the second largest and easternmost island in the Philippines. It is also the name of one of the three island groups in the country, which consists of the island of Mindanao and smaller surrounding islands. The other two are Luzon and the Visayas. The island of Mindanao is called The...

; its presence in southern Mindanao is "poorly documented" due to ongoing conflicts. As such, it is native to at least the provinces of Surigao del Norte
Surigao del Norte
Surigao del Norte is a province of the Philippines located in the Caraga region in Mindanao. Its capital is Surigao City. The province consists of two major islands—Siargao Island and Bucas Grande Island—in the Philippine Sea, and a small region at the northernmost tip of the island of...

 and Surigao del Sur
Surigao del Sur
' is a province of the Philippines located in the CARAGA region in Mindanao. Its capital is Tandag City and borders Surigao del Norte to the north, Agusan del Norte and Agusan del Sur to the west, and Davao Oriental to the south...

. It has an altitudinal distribution of 0–800 m above sea level.

The species grows terrestrially in a number of habitats, including lowland heath forest
Heath forest
Heath forest is a type of tropical moist forest found in areas with acidic, sandy soils that are extremely nutrient-poor. Notable examples are the Rio Negro campinarana of the Amazon Basin in South America, and the Sundaland heath forests of Borneo and neighboring islands.-External links:**...

, exposed sites such as cliff faces and landslides, lower montane forest among stunted vegetation, and disturbed or recovering secondary vegetation (such as previously logged dipterocarp forest). It also often colonises the sides of roads running through the forest. The known distribution of this species appears to roughly correlate with ultramafic substrate occurrence. Nepenthes bellii is often sympatric with N. merrilliana
Nepenthes merrilliana
Nepenthes merrilliana is a tropical pitcher plant endemic to the Philippines. It produces some of the largest pitchers in the genus, rivalling those of N. rajah....

and N. mindanaoensis
Nepenthes mindanaoensis
Nepenthes mindanaoensis is a species of pitcher plant endemic to coastal lateritic hills on Mindanao in the Philippines. It is very closely related to N. alata.-Natural hybrids:*N. alata × N. mindanaoensis...

. Although N. bellii is known to form natural hybrids with both of these species, such crosses appear to be rare. Field observations of these three species suggest that they flower at different times of the year.

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. bellii 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 IUCN Red List
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. Stewart McPherson
Stewart McPherson (geographer)
Stewart R. McPherson is a British geographer.He studied at the University of Durham in England, the University of Tübingen in Germany and Yale University in the United States....

 considers the species "not currently threatened" and writes that it is "widespread" across Dinagat and northern Mindanao, where it is represented by "extensive stands". Nevertheless, habitat loss is affecting the species in lowland areas and this problem is particularly severe on Dinagat.

Related species

The combination of subglobular lower pitchers, densely fringed wings (often with clustered filaments) and proportionately long tendrils separate N. bellii from all other species with the possible exception of the miniature N. argentii
Nepenthes argentii
Nepenthes argentii is a highland Nepenthes pitcher plant native to Mount Guiting-Guiting on Sibuyan Island in the Philippines. It is possibly the smallest species in the genus and does not appear to have a climbing stage....

. However, the latter species can hardly be confused with N. bellii as it does not produce a climbing stem, differs markedly in the shape of the lamina, and has a uniquely curved peristome that continues along the lower surface of the lid.

Nepenthes bellii is closely allied to N. merrilliana
Nepenthes merrilliana
Nepenthes merrilliana is a tropical pitcher plant endemic to the Philippines. It produces some of the largest pitchers in the genus, rivalling those of N. rajah....

and N. surigaoensis
Nepenthes surigaoensis
Nepenthes surigaoensis is a tropical pitcher plant endemic to the Philippine island of Mindanao, where it grows at elevations of at least 800–1200 m above sea level.The species is named after Surigao Peninsula, where the type specimen was collected...

and shares with these species a similar morphology of the pitchers and laminae as well as a reddish colouration of the uppermost leaves. It is not easily confused with them, however, because it is much smaller in all respects, particularly in the size of its pitchers and inflorescence. More generally, N. bellii appears to fall under B. H. Danser
B. H. Danser
Benedictus Hubertus Danser , often abbreviated B. H. Danser, was a Dutch taxonomist and botanist...

's classical Insignes group, which also includes N. burkei
Nepenthes burkei
Nepenthes burkei |David Burke]], British plant collector) is a tropical pitcher plant native to the island of Mindoro in the Philippines, where it grows at an elevation of 1100–2000 m. It is very closely related to N. sibuyanensis and N. ventricosa...

, N. insignis
Nepenthes insignis
Nepenthes insignis is a tropical pitcher plant endemic to New Guinea and surrounding islands. The specific epithet insignis is Latin for "distinguished" or "remarkable".-Botanical history:...

, N. merrilliana
Nepenthes merrilliana
Nepenthes merrilliana is a tropical pitcher plant endemic to the Philippines. It produces some of the largest pitchers in the genus, rivalling those of N. rajah....

, and N. ventricosa
Nepenthes ventricosa
Nepenthes ventricosa is a tropical pitcher plant endemic to the Philippines, where it is a highland species, growing at an elevation of 1000–2000 m above sea level. It has been recorded from the islands of Luzon, Panay, and Sibuyan...

, among others, with N. sibuyanensis
Nepenthes sibuyanensis
Nepenthes sibuyanensis is a tropical pitcher plant endemic to Sibuyan Island in the Philippines, after which it is named.-Botanical history:...

being a recent addition.

Nepenthes bellii was also compared to N. micramphora
Nepenthes micramphora
Nepenthes micramphora is a tropical pitcher plant known only from Mount Hamiguitan on the island of Mindanao in the Philippines. It is a highland plant growing at elevations of 1100–1635 m....

in the formal description of the latter, in which the authors noted that the stem, laminae and inflorescence of N. micramphora match those of N. bellii "almost exactly". Indeed, prior to the description of N. micramphora in 2009, the species was misidentified as N. bellii on its native Mount Hamiguitan
Mount Hamiguitan
Mount Hamiguitan is a mountain located in the province of Davao Oriental. It has a height of 1,620 meters . The mountain and its vicinity has one of the most diverse wildlife populations in the Philippines. Among the wildlife found in the area are Philippine eagles and several species of Nepenthes...

. Nepenthes micramphora can be clearly distinguished by its distinct pitchers and even smaller vegetative features.

In their 2001 monograph, 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,...

 and 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....

 also noted a "remarkable resemblance" between the lower pitchers of N. tomoriana
Nepenthes tomoriana
Nepenthes tomoriana is a species of pitcher plant endemic to Sulawesi, where it grows at an elevation of 0–500 m above sea level.-References:...

and those of N. bellii.

Natural hybrids

Two putative natural hybrids involving N. bellii have been recorded: with N. merrilliana
Nepenthes merrilliana
Nepenthes merrilliana is a tropical pitcher plant endemic to the Philippines. It produces some of the largest pitchers in the genus, rivalling those of N. rajah....

and N. mindanaoensis
Nepenthes mindanaoensis
Nepenthes mindanaoensis is a species of pitcher plant endemic to coastal lateritic hills on Mindanao in the Philippines. It is very closely related to N. alata.-Natural hybrids:*N. alata × N. mindanaoensis...

. It is commonly sympatric with these species.

External links

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