Nepenthes smilesii type specimen.jpg

''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and 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 (two species) and the Seychelles (one); southward to Australia (four) and New Caledonia (one); and northward to India (one) and Sri Lanka (one). The greatest diversity occurs on Borneo, Sumatra, and the Philippines, 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 were once thought to drink rainwater from the pitchers.

Description

''Nepenthes'' species usually consist of a shallow root system and a prostrate or climbing stem, often several metres long and up to or more, and usually 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 (the tendril), which in some species aids 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. The shapes can evoke a champagne flute or a misused condom.

The trap contains a fluid of the plant's own production, which may be watery or more viscous, and is used to drown the prey. This fluid contains viscoelastic biopolymers that may be crucial to the retention of insects within the traps of many species. The viscoelastic fluid in pitchers is especially effective in the retention of winged insects. The trapping efficiency of this fluid remains high, even when significantly diluted by water, as inevitably happens in wet conditions.

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. The prey-capture effectiveness of the peristome is further enhanced in moist environments, where condensation may cause a thin water film to form on the surface of the peristome. When wet, the slippery surface of the peristome causes insects to ‘aquaplane’, or slip and fall, into the pitcher. 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.

''Nepenthes'' species 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, coloured differently, 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.

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. Records of cultivated plants trapping small birds have been made. Flowers occur in racemes or more rarely in panicles with male and female flowers on separate plants. Three species have symbiotic relationships with treeshrews, which eat the nectar produced by the plant and defecate into the pitchers, providing valuable nutrients.

''Nepenthes'' are insect-pollinated, the primary agents being flies (including blow flies, midges, and mosquitoes), moths, wasps, and butterflies.Clarke, C.M. 2001. ''Nepenthes of Sumatra and Peninsular Malaysia''. Natural History Publications (Borneo), Kota Kinabalu. Their smells can range from sweet to musty or fungus-like. Seed is typically 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 (though '' N. pervillei'' differs).

The genus is cytologically diploid, with all studied species having a chromosome number of ''2n''=80. This high number is thought to reflect paleopolyploidy (likely 8x or 16x).

Taxonomy

About 170 species of ''Nepenthes'' are currently recognised as valid. This number is increasing, with several new species being described each year.

Etymology

The genus name ''Nepenthes'' was first published in 1737 in Carl Linnaeus's ''Hortus Cliffortianus''. 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. 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! ranslated_from_Latin_by_Harry_Veitch.html" ;"title="Harry_Veitch.html" ;"title="ranslated from Latin by Harry Veitch">ranslated from Latin by Harry Veitch">Harry_Veitch.html" ;"title="ranslated from Latin by Harry Veitch">ranslated from Latin by Harry Veitch/blockquote>

The plant Linnaeus described was ''N. distillatoria'', called'' bāndurā'' (බාඳුරා), a species from Sri Lanka.Phillipps, A. & A. Lamb 1996. ''Pitcher-Plants of Borneo''. Natural History Publications (Borneo), Kota Kinabalu.

''Nepenthes'' was formally published as a genus, 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.

The name "monkey cups" was discussed in the May 1964 issue of '' National Geographic'', in which Paul A. Zahl wrote:

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.

The plants are often called ''kantong semar'' (Semar's pocket) in Indonesia and ''sako ni Hudas'' ( Judas' money bag

Flacourt called the plant ''Amramatico'', after a local name. More than a century later, this species was Species description">formally described as '' N. madagascariensis''.Poiret, J.L.M. 1797. ''Népente''. In: J.B. Lamarck ''Encyclopédie Méthodique Botanique'' Vol. 4.

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

''Nepenthes distillatoria'' was again described in 1683, this time by Swedish physician and naturalist Herman Niklas Grim. Grim called it ''Planta mirabilis destillatoria'' or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant. Three years later, in 1686, English naturalist John Ray quoted Grim as saying:

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. ranslated_from_Latin_in_''Pitcher-Plants_of_Borneo''.html" ;"title="Pitcher-Plants_of_Borneo.html" ;"title="ranslated from Latin in ''Pitcher-Plants of Borneo">ranslated from Latin in ''Pitcher-Plants of Borneo''">Pitcher-Plants_of_Borneo.html" ;"title="ranslated from Latin in ''Pitcher-Plants of Borneo">ranslated from Latin in ''Pitcher-Plants of Borneo''ref name=P&L />

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

Around the same time, 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.

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, 39 years after Rumphius's death. By this time, Linnaeus's name ''Nepenthes'' had become established.

''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''.

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 stated the lid is a moving part, actively opening and closing. In his most celebrated work, ''Flora Cochinchinensis'', he writes:

..(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! ranslated_from_French_in_''Pitcher-Plants_of_Borneo''.html" ;"title="Pitcher-Plants_of_Borneo.html" ;"title="ranslated from French in ''Pitcher-Plants of Borneo">ranslated from French in ''Pitcher-Plants of Borneo''">Pitcher-Plants_of_Borneo.html" ;"title="ranslated from French in ''Pitcher-Plants of Borneo">ranslated from French in ''Pitcher-Plants of Borneo''ref name=P&L />

''Phyllamphora mirabilis'' was eventually transferred to the genus ''Nepenthes'' by Rafarin in 1869. As such, ''P. mirabilis'' is the basionym of this most cosmopolitan of tropical pitcher plant species.

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:

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. ranslated_from_French_in_''Pitcher-Plants_of_Borneo''.html"_;"title="Pitcher-Plants_of_Borneo.html"_;"title="ranslated_from_French_in_''Pitcher-Plants_of_Borneo">ranslated_from_French_in_''Pitcher-Plants_of_Borneo''">Pitcher-Plants_of_Borneo.html"_;"title="ranslated_from_French_in_''Pitcher-Plants_of_Borneo">ranslated_from_French_in_''Pitcher-Plants_of_Borneo''ref_name=P&L_/>

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With the discovery of new species and Joseph Banks">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.Barthlott, W., Porembski, S., Seine, R., and Theisen, I. 2007. ''The Curious World of Carnivorous Plants.'' Portland, Oregon: Timber Press. 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.Clarke, C.M. & C.C. Lee 2004. '' Pitcher Plants of Sarawak''. Natural History Publications (Borneo), Kota Kinabalu.

Cultivation

''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 generally higher in elevation, and thus exposed to cooler evening temperatures. Lowland forms are those species growing nearer to sea level. Both forms respond best to rainwater (but some tap water works as long as it is flushed monthly with rainwater or water low in dissolved solid and chemicals), bright light (though some species can grow in 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, such as '' 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 one to two 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'' species are considered threatened or endangered plants and all of them are listed in CITES Appendix II, with the exception of '' N. rajah'' and '' N. khasiana'' which are listed in CITES Appendix I. The CITES listing means all international trade (including in parts and derivatives) is controlled by the CITES permitting system, with wild
sourced specimens of Appendix I species prohibited from commercial international trade.

Hybrids and cultivars

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

*''N.'' × ''coccinea'' ((''N. rafflesiana'' × ''N. ampullaria'') × ''N. mirabilis'')
*''N.'' × ''ventrata'' (''N. ventricosa'' × ''N. alata'')
*''N.'' × 'Bloody Mary' (''N. ventricosa'' × ''N. ampullaria'')
*''N.'' 'D'amato' (''N. lowii'' × ''N. ventricosa'')
*''N.'' × ''mixta'' (''N. northiana'' × ''N. maxima'')
*''N.'' 'Syurga' (''N. ventricosa'' × ''N. northiana'')
*''N.'' 'Menarik' (''N. rafflesiana'' × ''N. veitchii'')
*''N.'' 'Emmarene' (''N. khasiana'' × ''N. ventricosa'')
*''N.'' 'Judith Finn' (''N. spathulata'' × ''N. veitchii'')

See also

* ''Nepenthes'' classification
* ''Nepenthes'' infauna
* List of ''Nepenthes'' endophyte species

References

Danser's Monograph on ''Nepenthes''
(covers species from Malaysia, Indonesia and New Guinea, but not elsewhere)


in: Watson, L., and M. J. Dallwitz (1992 onwards)
The Families of Flowering Plants
Descriptions, Illustrations, Identification, Information Retrieval.

Further reading

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* Beaver, R.A. 1983. The communities living in ''Nepenthes'' pitcher plants: fauna and food webs. In: J.H. Frank & L.P. Lounibos (eds.) ''Phytotelmata: Plants as Hosts for Aquatic Insect Communities''. Plexus Publishing, New Jersey. pp. 129–159.
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* Boulay, J. 1997

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External links

''Nepenthes'' - the Monkey Cups
from the Botanical Society of America

''Nepenthes'': The Interactive Guide
at Tom's Carnivores

How to Grow ''Nepenthes''
at Tom's Carnivores


at the Carnivorous Plant Photo Finder

A video about ''Nepenthes rajah''
from ''The Private Life of Plants''

The Carnivorous Plant FAQ: ''Nepenthes''
by Barry Rice

Evolution -- ''Nepenthes'' Phylogeny
from the International Carnivorous Plant Society

Inner World of ''Nepenthes''
from the John Innes Centre

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Caryophyllales genera
Dioecious plants
Taxa named by Carl Linnaeus
Articles containing video clips