The Nepenthes ( /nɨˈpɛnθiːz/), popularly known as tropical pitcher plants or monkey cups, are a genus of carnivorous plants in the monotypic family Nepenthaceae. The genus comprises roughly 140 species, numerous natural and many cultivated hybrids. They are mostly liana-forming plants of the Old World tropics, ranging from South China, Indonesia, Malaysia and the Philippines; westward to Madagascar (2 species) and the Seychelles (1); southward to Australia (3) and New Caledonia (1); and northward to India (1) and Sri Lanka (1). The greatest diversity occurs on Borneo and Sumatra with many endemic species. Many are plants of hot humid lowland areas, but the majority are tropical montane plants, receiving warm days but cool to cold humid nights year round. A few are considered tropical alpine with cool days and nights near freezing. The name monkey cups refers to the fact that monkeys have been observed drinking rainwater from these plants.

Description

Nepenthes species usually consist of a shallow root system and a prostrate or climbing stem, often several metres long and up to 15 m (49 ft) or more, and usually 1 cm (0.4 in) or less in diameter, although this may be thicker in a few species (e.g. N. bicalcarata). From the stems arise alternate sword-shaped leaves with entire leaf margins. An extension of the midrib (tendril), which in some species aid in climbing, protrudes from the tip of the leaf; at the end of the tendril the pitcher forms. The pitcher starts as a small bud and gradually expands to form a globe- or tube-shaped trap.^[2]

Basic structure of an upper pitcher

The trap contains a fluid of the plant's own production, which may be watery or syrupy and is used to drown the prey. Research has shown that this fluid contains viscoelastic biopolymers that may be crucial to the retention of insects within the traps of many species. The trapping efficiency of this fluid remains high, even when significantly diluted by water, as inevitably happens in wet conditions.^[3]

The lower part of the trap contains glands which absorb nutrients from captured prey. Along the upper inside part of the trap is a slick waxy coating which makes the escape of its prey nearly impossible. Surrounding the entrance to the trap is a structure called the peristome (the "lip") which is slippery and often quite colorful, attracting prey but offering an unsure footing. Above the peristome is a lid (the operculum): in many species this keeps rain from diluting the fluid within the pitcher, the underside of which may contain nectar glands which attract prey.^[2]

Nepenthes usually produce two types of pitchers, known as leaf dimorphism. Appearing near the base of the plant are the large lower traps, which typically sit on the ground. The upper or aerial pitchers are usually smaller, differently-coloured, and possess different features from the lower pitchers. These upper pitchers usually form as the plant reaches maturity and the plant grows taller. To keep the plant steady, the upper pitchers often form a loop in the tendril, allowing it to wrap around nearby support. In some species (e.g. N. rafflesiana) different prey may be attracted by the two types of pitchers. This varied morphology also often makes identification of species difficult.^[2]

Prey usually consists of insects, but the largest species (e.g. N. rajah and N. rafflesiana) may occasionally catch small vertebrates, such as rats and lizards.^[4][5] There are even records of cultivated plants trapping small birds.^[6][7] Flowers occur in racemes or more rarely in panicles with male and female flowers on separate plants. They are insect pollinated, the primary agents being flies (including blow flies, midges, and mosquitoes), moths, wasps, and butterflies.^[8] Their smell can range from sweet to musty or fungus-like.^[9] Seed is produced in a four-sided capsule which may contain 50–500 wind-distributed seeds, consisting of a central embryo and two wings, one on either side.

Distribution and habitat

Global distribution of Nepenthes

The genus Nepenthes is mostly found within the Malay Archipelago, with the greatest biodiversity found on Borneo and Sumatra, especially in the Borneo montane rain forests. The full range of the genus includes Madagascar (N. madagascariensis and N. masoalensis), the Seychelles (N. pervillei), Sri Lanka (N. distillatoria), and India (N. khasiana) in the west to Australia (N. mirabilis, N. rowanae, and N. tenax) and New Caledonia (N. vieillardii) in the southeast. Most species are restricted to very small ranges, including some that are only found on individual mountains. These limited distributions and the inaccessibility of the region often means that some species go decades without being rediscovered in the wild (e.g. N. deaniana, which was rediscovered 100 years after its initial discovery). Approximately ten species have population distributions larger than a single island or group of smaller islands. Nepenthes mirabilis has the distinction of being the most widely distributed species in the genus, ranging from Indochina and throughout the Malay Archipelago to Australia.^[2][10][11]

Because of the nature of the habitats which Nepenthes species occupy, they are often graded as either lowland or highland species, depending on their altitude above sea level, with 1,200 m (3,937 ft) the rough delineation between lowland and highland. Species that grow at lower altitudes require continuously warm climates with little difference between day and night temperatures, whereas highland species thrive when they receive warm days and much cooler nights. Nepenthes lamii is the species that holds the record of growing at a higher altitude than any other in the genus, up to 3,520 m (11,549 ft).^[2][11]

Most Nepenthes species grow in environments that provide high humidity and precipitation and moderate to high light levels. A few species, including N. ampullaria, prefer the dense, shaded forests, but most other species thrive on the margins of tree/shrub communities or clearings. Some species (e.g. N. mirabilis) have been found growing in clear-cut forest areas, roadsides, and disturbed fields. Other species have adapted to growing in savanna-like grass communities. The soils that Nepenthes grow in are usually acidic and low in nutrients, being composed of peat, white sand, sandstone, or volcanic soils. There are, however, exceptions to these generalities, including species that thrive in soils that have high heavy metal content (e.g. N. rajah), on sandy beaches in the sea spray zone (e.g. N. albomarginata). Other species grow on inselbergs and as lithophytes, while others, such as N. inermis, can grow as epiphytes with no soil contact.^[2]

Ecological relationships

A drowned lizard found in a freshly opened pitcher of N. rajah

The most obvious interaction between Nepenthes species and its environment, including other organisms, is that of predator and prey. Nepenthes species certainly attract and kill their prey, albeit passively, through active production of attractive colours, sugary nectar, and even sweet scents. From this relationship, the plants primarily gain nitrogen and phosphorus to supplement their nutrient requirements for growth, given that soil nutrients are typically lacking. The most frequent prey is an abundant and diverse group of arthropods, with ants and other insects topping the menu. Other arthropods that are found frequently include spiders, scorpions, and centipedes, while snails and frogs are more unusual but not unheard of. The most uncommon prey for Nepenthes species includes rats found in N. rajah. The composition of prey captured depends on many factors, including location, but can incorporate hundreds of individual insects and many different species.^[2] While many Nepenthes are generalists in what they capture, at least one, N. albomarginata, has specialised and almost exclusively traps termites and produces nearly no nectar. Nepenthes albomarginata gains its name from the ring of white trichomes that are directly beneath the peristome. These trichomes—or "hairs"—are palatable to termites and will attract them to the pitcher. In the course of collecting the edible trichomes, hundreds or thousands of termites will fall into the pitcher.^[12][13]

A lower pitcher of N. attenboroughii supporting a large population of mosquito larvae. The upright lid of this species exposes its pitchers to the elements such that they are often completely filled with fluid.^[14]

Symbioses

Nepenthes bicalcarata provides space in the hollow tendrils of its upper pitchers for the carpenter ant Camponotus schmitzi to build nests. The ants take larger prey from the pitchers, which may benefit N. bicalcarata by reducing the amount of putrefaction of collected organic matter that could harm the natural community of infaunal species that aid the plant's digestion.^[15]

Nepenthes lowii has also formed a dependent relationship, but this time it is with vertebrates instead of insects. The pitchers of N. lowii provide a sugary exudate reward on the reflexed pitcher lid (operculum) and a perch for tree shrew species, which have been found eating the exudate and defecating into the pitcher. A 2009 study, which coined the term "tree shrew lavatories", determined that anywhere between 57 and 100% of the plant's foliar nitrogen uptake comes from the faeces of tree shrews.^[16] Another study published the following year showed that the shape and size of the pitcher orifice of N. lowii exactly match the dimensions of a typical tree shrew (Tupaia montana).^[17][18] A similar adaptation was found in N. macrophylla and N. rajah, and is also likely to be present in N. ephippiata.^[18]

Infauna

Organisms that spend at least part of their lives within the pitchers of Nepenthes species are often called Nepenthes infauna. The most common infaunal species, often representing the top trophic level of the infaunal ecosystem, are many species of mosquito larvae. Other infaunal species include fly and midge larvae, spiders, mites, ants, and even a species of crab (Geosesarma malayanum). Many of these species specialise to one pitcher plant species and are found nowhere else. These specialists are called nepenthebionts. Others that are often associated but are not dependent on Nepenthes species are called nepenthophiles. Nepenthexenes, on the other hand, are rarely found in the pitchers, but will often appear when putrefaction approaches a certain threshold, attracting fly larvae that would normally not be found in the pitcher infaunal community. The complex ecological relationship between pitcher plant and infauna is not yet fully understood, but it has been suggested that the relationship is mutualistic: the infauna is given shelter, food, or protection and the plant that harbours the infauna receives expedited breakdown of captured prey, increasing the rate of digestion and keeping harmful bacterial populations repressed.^[15][19][20]

Botanical history

Plukenet's drawing of N. distillatoria from his Almagestum Botanicum of 1696.

The earliest known record of Nepenthes dates back to the 17th century. In 1658, French colonial governor Etienne de Flacourt published a description of a pitcher plant in his seminal work Histoire de la Grande Isle de Madagascar. It reads:^[21]

It is a plant growing about 3 feet high which carries at the end of its leaves, which are 7 inches long, a hollow flower or fruit resembling a small vase, with its own lid, a wonderful sight. There are red ones and yellow ones, the yellow being the biggest. The inhabitants of this country are reluctant to pick the flowers, saying that if somebody does pick them in passing, it will not fail to rain that day. As to that, I and all the other Frenchmen did pick them, but it did not rain. After rain these flowers are full of water, each one containing a good half-glass. [translated from French in Pitcher-Plants of Borneo]^[9]

Flacourt called the plant Amramatico, after a local name. More than a century later, this species was formally described as N. madagascariensis.^[22]

The second species to be described was N. distillatoria, the Sri Lankan endemic. In 1677, Bartholinus made brief mention of it under the name Miranda herba, Latin for "marvellous herb".^[23] Three years later, Dutch merchant Jacob Breyne referred to this species as Bandura zingalensium, after a local name for the plant.^[24] Bandura subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined Nepenthes in 1737.^[9]

Nepenthes distillatoria was again described in 1683, this time by Swedish physician H. N. Grimm.^[25] Grimm called it Planta mirabilis destillatoria or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant.^[9] Three years later, in 1686, English naturalist John Ray quoted Grimm as saying:^[26]

The root draws up moisture from the earth which with the help of the sun's rays rises up into the plant itself and then flows down through the stems and nerves of the leaves into the natural utensil to be stored there until used for human needs. [translated from Latin in Pitcher-Plants of Borneo]^[9]

One of the earliest illustrations of Nepenthes appears in Leonard Plukenet's Almagestum Botanicum of 1696.^[27] The plant, called Utricaria vegetabilis zeylanensium, is undoubtedly N. distillatoria.^[9]

Cantharifera as illustrated in Rumphius's Herbarium Amboinensis, Volume 5, published in 1747, although probably drawn in the late 17th century. The vine on the right is not a Nepenthes, but a species of Flagellaria.

It was around the same time that German botanist Georg Eberhard Rumphius discovered two new Nepenthes species in the Malay Archipelago. Rumphius illustrated the first one, now considered synonymous with N. mirabilis, and gave it the name Cantharifera, meaning "tankard-bearer". The second, referred to as Cantharifera alba, is thought to have been N. maxima. Rumphius described the plants in his most famous work, the six-volume Herbarium Amboinense, a catalogue of the flora of Ambon Island. However, it would not be published until many years after his death.^[28]

After going blind in 1670, when the manuscript was only partially complete, Rumphius continued work on Herbarium Amboinensis with the help of clerks and artists. In 1687, with the project nearing completion, at least half of the illustrations were lost in a fire. Persevering, Rumphius and his helpers first completed the book in 1690. However, two years later, the ship carrying the manuscript to the Netherlands was attacked and sunk by the French, forcing them to start over from a copy that had fortunately been retained by Governor-General Johannes Camphuijs. The Herbarium Amboinensis finally arrived in the Netherlands in 1696. Even then, the first volume did not appear until 1741, thirty-nine years after Rumphius's death. By this time, Linnaeus's name Nepenthes had become established.^[9]

Illustration of Bandura zeylanica (N. distillatoria) from Burmann's Thesaurus Zeylanicus of 1737.

Nepenthes distillatoria was again illustrated in Johannes Burmann's Thesaurus Zeylanicus of 1737. The drawing depicts the end of a flowering stem with pitchers. Burmann refers to the plant as Bandura zeylanica.^[29]

The next mention of tropical pitcher plants was made in 1790, when Portuguese priest João de Loureiro described Phyllamphora mirabilis, or the "marvellous urn-shaped leaf", from Vietnam. Despite living in the country for around 35 years, it seems unlikely that Loureiro observed living plants of this species, as he states that the lid is a moving part, actively opening and closing. In his most celebrated work, Flora Cochinchinensis, he writes:^[30]

[...] (the) leaf-tip ends in a long hanging tendril, twisted spirally in the middle, from which hangs a sort of vase, oblong, pot-bellied, with a smooth lip with a projecting margin and a lid affixed to one side, which of its own nature freely opens and closes in order to receive the dew and store it. A marvellous work of the Lord! [translated from French in Pitcher-Plants of Borneo]^[9]

Phyllamphora mirabilis was eventually transferred to the genus Nepenthes by George Claridge Druce in 1916.^[31] As such, P. mirabilis is the basionym of this most cosmopolitan of tropical pitcher plant species.^[15]

Loureiro's description of a moving lid was repeated by Jean Louis Marie Poiret in 1797. Poiret described two of the four Nepenthes species known at the time: N. madagascariensis and N. distillatoria. He gave the former its current name and called the latter Nepente de l'Inde, or simply "Nepenthes of India", although this species is absent from the mainland. In Jean-Baptiste Lamarck's Encyclopédie Méthodique Botanique, he included the following account:^[22]

This urn is hollow, as I have just said, usually full of soft, clear water, and then closed. It opens during the day and more than half the liquid disappears, but this loss is repaired during the night, and the next day the urn is full again and closed by its lid. This is its sustenance, and enough for more than one day because it is always about half-full at the approach of night. [translated from French in Pitcher-Plants of Borneo]^[9]

The Nepenthes house of the Veitch Nurseries as illustrated in The Gardeners' Chronicle, 1872

With the discovery of new species and Sir Joseph Banks' original introduction of specimens to Europe in 1789, interest in Nepenthes grew throughout the 19th century, culminating in what has been called the "Golden Age of Nepenthes" in the 1880s.^[2][9] However, the popularity of the plants dwindled in the early 20th century, before all but disappearing by World War II. This is evidenced by the fact that no new species were described between 1940 and 1966. The revival of global interest in the cultivation and study of Nepenthes is credited to Japanese botanist Shigeo Kurata, whose work in the 1960s and 1970s did much to bring attention to these plants.^[32]

Etymology

The name Nepenthes was first published in 1737 in Carolus Linnaeus's Hortus Cliffortianus.^[33] It references a passage in Homer's Odyssey, in which the potion "Nepenthes pharmakon" is given to Helen by an Egyptian queen. "Nepenthe" literally means "without grief" (ne = not, penthos = grief) and, in Greek mythology, is a drug that quells all sorrows with forgetfulness.^[9][34] Linnaeus explained:

If this is not Helen's Nepenthes, it certainly will be for all botanists. What botanist would not be filled with admiration if, after a long journey, he should find this wonderful plant. In his astonishment past ills would be forgotten when beholding this admirable work of the Creator! [translated from Latin by Harry Veitch]^[35]

The plant Linnaeus described was Nepenthes distillatoria, a species from Sri Lanka.^[9]

Nepenthes was formally published as a generic name in 1753 in Linnaeus's famous Species Plantarum, which established botanical nomenclature as it exists today. Nepenthes distillatoria is the type species of the genus.^[36]

Nepenthes from Carolus Linnaeus's Species Plantarum of 1753

The name monkey cups was discussed in the May 1964 issue of National Geographic, in which Paul A. Zahl wrote:^[37]

The carriers called them "monkey cups," a name I had heard elsewhere in reference to Nepenthes, but the implication that monkeys drink the pitcher fluid seemed farfetched. I later proved it true. In Sarawak I found an orangutan that had been raised as a pet and later freed. As I approached it gingerly in the forest, I offered it a half-full pitcher. To my surprise, the ape accepted it and, with the finesse of a lady at tea, executed a delicate bottoms-up.

Cultivation

Cultivated Nepenthes rajah and other species

Nepenthes may be cultivated in greenhouses. Easier species include N. alata, N. ventricosa, N. khasiana, and N. sanguinea. These four species are highlanders (N. alata has both lowland and highland forms), some easy lowlander species are N. rafflesiana, N. bicalcarata, N. mirabilis, and N. hirsuta.

Highland forms are those species that grow in habitats that are generally higher up in elevation, and thus exposed to cooler evening temperatures. Lowland forms are those species that grow nearer to sea level. Both forms respond best to rainwater (but tap water works as long as you flush it out with rainwater every month or if you have soft water), bright light (not full sun), a well drained medium, good air circulation and relatively high humidity, although easier species such as N. alata can adapt to lower humidity environments. Highland species must have night-time cooling to thrive in the long-term. Chemical fertilisers are best used at low strength. Occasional feeding with frozen (thawed before use) crickets may be beneficial. Terrarium culture of smaller plants like N. bellii, N. × trichocarpa and N. ampullaria is possible, but most plants will get too large over time.

Plants can be propagated by seed, cuttings, and tissue culture. Seeds are usually sown on damp chopped Sphagnum moss, or on sterile plant tissue culture media once they have been properly disinfected. The seeds generally become nonviable soon after harvesting, so seed are not usually the preferred method of propagation. A 1:1 mixture of orchid medium with moss or perlite has been used for germination and culture. Seed may take two months to germinate, and two years or more to yield mature plants. Cuttings may be rooted in damp Sphagnum moss in a plastic bag or tank with high humidity and moderate light. They can begin to root in 1–2 months and start to form pitchers in about six months. Tissue culture is now used commercially and helps reduce collection of wild plants, as well as making many rare species available to hobbyists at reasonable prices. Nepenthes are considered threatened or endangered plants and are listed in CITES appendices 1 & 2.

Species

Around 138 species of Nepenthes are currently recognised as valid.^[1] This number is quickly increasing, with several new species being described each year.^[32]

Hybrids and cultivars

The complex man-made hybrid N. ventricosa × (N. lowii × N. macrophylla)

There are many hybrid Nepenthes and numerous named cultivars. Some of the more well known artificially produced hybrids include:

• N. 'Coccinea' ((N. rafflesiana × N. ampullaria) × N. mirabilis)
• N. 'Emmarene' (N. khasiana × N. ventricosa)
• N. 'Gentle' (N. fusca × N. maxima)
• N. 'Judith Finn' (N. veitchii × N. spathulata)
• N. 'Miranda' ((N. maxima × N. northiana) × N. maxima)
• N. 'Mixta' (N. northiana × N. maxima)

See also

• Nepenthes classification
• Nepenthes infauna

References

• Danser's Monograph on Nepenthes (covers species from Malaysia, Indonesia and New Guinea, but not elsewhere)
• Nepenthaceae in: Watson, L., and M. J. Dallwitz (1992 onwards). The Families of Flowering Plants. Descriptions, Illustrations, Identification, Information Retrieval.

Further reading

Nepenthes sp. Anipahan is an undescribed tropical pitcher plant known only from Mount Anipahan in central Palawan, the Philippines, where it grows in upper montane forest at 1200–1400 m above sea level.^[1][2][3] It is very closely allied to N. leonardoi and may be conspecific with it.^[1][2]

References

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Nepenthes × sarawakiensis ( /nɨˈpɛnθiːz səˌrɑːwɑːkiˈɛnsɨs/, after Sarawak, Borneo) is a natural hybrid involving N. muluensis and N. tentaculata. It is quite a rare plant as one of its parent species, N. muluensis, is only known from several isolated mountains.

References

• Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. The ecology and distribution of Bornean Nepenthes.PDF Journal of Tropical Forest Science 5(1): 13–25.
• Adam, J.H. & C.C. Wilcock 1993. One new natural hybrid of Nepenthes from Mt. Mulu. The Sarawak Museum Journal 43: 291–294.
• Adam, J. H., R. Omar & C. C. Wilcock 2002. Phytochemical Screening of Flavonoids in Three Hybrids of Nepenthes (Nepenthaceae) and their Putative Parental Species from Sarawak and Sabah.PDF OnLine Journal of Biological Sciences 2(9): 623–625. doi:10.3923/jbs.2002.623.625
• Clarke, C.M. (1997). Nepenthes of Borneo. Kota Kinabalu: Natural History Publications. p. 109. ISBN 983-812-015-4. 

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Nepenthes × hookeriana ( /nɨˈpɛnθiːz ˌhʊkəriˈɑːnə/; after Joseph Dalton Hooker), or Hooker's Pitcher-Plant,^[4] is a common natural hybrid involving N. rafflesiana and N. ampullaria. It was originally described as a species.

It is a relatively common natural hybrid found in lowland conditions throughout Peninsular Malaysia, Borneo, Singapore and the Indonesian island of Sumatra. Like its parental species, the hybrid is generally found in recently disturbed clearings.

Infraspecific taxa

• Nepenthes hookeri var. elongata Hort.Veitch ex Wilson (1877) sphalm.typogr.
• Nepenthes hookeriana f. elongata (Hort.Veitch ex Wilson) Divers (1879)

References

Further reading

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Nepenthes × alisaputrana ( /nɨˈpɛnθiːz ˈɑːlɨsəpʊˈtrɑːnə/ preferably, or /əˈlɪsəpʊˈtrɑːnə/, after Datuk Lamri Ali), or the Leopard Pitcher-Plant,^[1] is a hybrid of two well-known Nepenthes pitcher plant species: N. burbidgeae and N. rajah. The plant is confined to Mount Kinabalu in Sabah, Borneo.^[2]

N. × alisaputrana (right) is often sympatric with N. rajah (left)

Nepenthes × alisaputrana was described in 1992 by J. H. Adam and C. C. Wilcock and is named in honour of Datuk Lamri Ali, a former Director of Sabah Parks.^[3] It is only known from a few remote localities within Kinabalu National Park, where it grows in stunted, open vegetation over serpentine soils at around 2000 m above sea level, often amongst populations of N. burbidgeae.^[4] It grows alongside both parent species on Pig Hill,^[5] where it is found at 1930–1950 m.^[6]

This plant is notable for combining the best characters of both parent species, not least the size of its pitchers, which rival those of N. rajah in volume (≤35 cm high, ≤20 cm wide).^[4] The other hybrids involving N. rajah do not exhibit such impressive proportions. The pitchers of N. × alisaputrana can be distinguished from those of N. burbidgeae by a broader peristome, larger lid and simply by their sheer size. The hybrid differs from its other parent, N. rajah, by its lid structure, indumentum of short, brown hairs, narrower and more cylindrical peristome, and pitcher colour, which is usually yellow-green with red or brown flecking. For this reason, Anthea Phillipps and Anthony Lamb gave it the common name "Leopard Pitcher-Plant".^[1] The peristome is green to dark red and striped with purple bands. Leaves are often slightly peltate. The hybrid is a strong climber and frequently produces upper pitchers.^[2]

Nepenthes × alisaputrana more closely resembles N. rajah than N. burbidgeae, but it is difficult to confuse this plant with either. However, this mistake has previously been made on at least one occasion; a pitcher illustrated in Adrian Slack's Insect-Eating Plants and How to Grow Them as being N. rajah^[7] is in fact N. burbidgeae × N. rajah.^[2]

In 2002, phytochemical screening and analytical chromatography were used to study the presence of phenolic compounds and leucoanthocyanins in N. × alisaputrana and its putative parent species.^[8] The research was based on leaf material from nine dry herbarium specimens. Eight spots containing phenolic acids, flavonols, flavones, leucoanthocyanins and 'unknown flavonoid' 1 and 3 were identified from chromatographic profiles. The distributions of these in the hybrid N. × alisaputrana and its putative parental species N. burbidgeae and N. rajah are shown in the table to the left. A specimen of N. × alisaputrana grown from tissue culture (in vitro) was also tested.^[8]

Luteolin, cyanidin and 'Unknown Flavonoid 3' were undetected in N. burbidgeae, while concentrations of 'Unknown Flavonoid 1' were found to be weak. Chromatographic patterns of the N. × alisaputrana samples studied showed complementation of its putative parental species.^[8]

Myricetin was found to be absent from all studied taxa. This agrees with the findings of previous authors^[9][10] and suggests that the absence of a widely distributed compound like myricetin among the Nepenthes examined might provide additional diagnostic information for these taxa.^[8]

References

• Adam, J.H. 1997. Prey spectra of Bornean Nepenthes species (Nepenthaceae) in relation to their habitat.PDF Pertanika Journal of Tropical Agricultural Science 20(2–3): 121–134.
• Adam, J.H. & C.C. Wilcock 1999. Palynological study of Bornean Nepenthes (Nepenthaceae).PDF Pertanika Journal of Tropical Agricultural Science 22(1): 1–7.
• Beaman, J.H. & C. Anderson 2004. The Plants of Mount Kinabalu: 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo), Kota Kinabalu.
• Cantley, R., M. Cheek, P.F. Gardner, P. Mann, B. Meyers-Rice, N. Parker & P. Temple 2000. Favorite Anecdotes on Adrian Slack.PDF Carnivorous Plant Newsletter 29(4): 109–116.
• Thong, J. 2006. Travels around North Borneo – Part 1.PDF Victorian Carnivorous Plant Society Journal 81: 12–17.

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Nepenthes sp. Luzon is an undescribed tropical pitcher plant known from a single peak in northern Luzon, the Philippines, where it grows at 1220 m above sea level.^[1] It shows similarities to N. burkei and N. ventricosa.^[1]

References

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Nepenthes × tsangoya ( /nɨˈpɛnθiːz sæŋˈɡɔɪ.ə/; after Peter Tsang) is a tropical pitcher plant. It reportedly represents the complex natural hybrid (N. alata × N. merrilliana) × N. mirabilis.

N. × tsangoya was mentioned as a natural hybrid in Guide to Nepenthes Hybrids (1995).^[1] The known ranges of the parent species only overlap in Mindanao, the Philippines.

References

• CP Database: Nepenthes × tsangoya

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Nepenthes × mirabilata ( /nɨˈpɛnθiːz mɪˌræbɪˈlɑːtə/; a blend of mirabilis and alata) is a natural hybrid involving N. alata and N. mirabilis.^[1]

N. × mirabilata was mentioned as a natural hybrid in Guide to Nepenthes Hybrids (1995). The hybrid is restricted to Mindanao, the Philippines, the only location where the parent species overlap.

References

• CP Database: Nepenthes × mirabilata

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Nepenthes × ventrata ( /nɨˈpɛnθiːz vɛnˈtrɑːtə/; a blend of ventricosa and alata) is a natural hybrid involving N. alata and N. ventricosa. Like its two parent species, it is endemic to the Philippines. The name was originally published in the Carnivorous Plant Newsletter in 1979.^[1]

N. × ventrata is one of the most common tissue cultured Nepenthes plants, although it is often mislabelled as Nepenthes alata. It is relatively easy to grow indoors and is usually the first tropical pitcher plant seen by consumers due to its availability in many garden shops and home centres.

The cultivar N. 'LeeAnn Marie' is a later synonym of N. × ventrata, although the name is not established as it was not validly published.^[2]

References

• CP Database: Nepenthes × ventrata

Further reading

• Gaume, L., P. Perret, E. Gorb, S. Gorb, J.-J. Labat & N. Rowe 2004. How do plant waxes cause flies to slide? Experimental tests of wax-based trapping mechanisms in three pitfall carnivorous plants. Arthropod Structure and Development 33(1): 103–111. doi:10.1016/j.asd.2003.11.005
• Gorb, E., V. Kastner, A. Peressadko, E. Arzt, L. Gaume, N. Rowe & S. Gorb 2004. Structure and properties of the glandular surface in the digestive zone of the pitcher in the carnivorous plant Nepenthes ventrata and its role in insect trapping and retention.PDF The Journal of Experimental Biology 207: 2947–2963. doi:10.1242/jeb.01128