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The Nepenthes (from Greek: ne = not, penthos = grief, sorrow; named after the ancient drug Nepenthe), popularly known as Tropical Pitcher Plants or Monkey Cups, are a genus of carnivorous plants in the monotypic family Nepenthaceae that comprises roughly 120 species, numerous natural and many cultivated hybrids. They are vine-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.

Etymology

The name Nepenthes was first published in 1737 in Carolus 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 isn't Helen's Nepenthes, it certainly will be for all botanists. What botanist wouldn't 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! [translatedfrom Latin by H. J. Veitch]
The plant Linnaeus described was Nepenthes distillatoria, a species from Sri Lanka. Nepenthes was formally published as a generic name in 1753 in Linnaeus's famous Species Plantarum, which established botanical nomenclature as it exists today. N. distillatoria is the type species of the genus.

Botanical history

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:
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 won't fail to rain that day. As to that, I and all the other Frenchmen did pick them, but it didn't rain. After rain these flowers are full of water, each one containing a good half-glass. [translatedfrom French in Pitcher-Plants of Borneo]
   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". 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. Grimm called it Planta mirabilis distillatoria or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant.
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. [translatedfrom Latin in Pitcher-Plants of Borneo] The plant, called Utricaria vegetabilis zeylanensium, is undoubtedly N. distillatoria.
   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 didn't appear until 1741, thirty-nine years after Rumphius's death. By this time, Linnaeus's name Nepenthes had become established.
   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:
[...] (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! [translatedfrom French in Pitcher-Plants of Borneo] 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:

Morphology and function

The plants usually consist of a shallow root system and a prostrate or climbing stem, often several metres long, and usually 1 cm or less in diameter, although this may be thicker in a few species (for example N. bicalcarata). From the stems arise leaf-like expanded petioles, similar to certain Citrus spp., ending in a tendril, which in some species aid in climbing, and at the end of which forms the pitcher, considered the true leaf. The pitcher starts as a small bud and gradually expands to form a globe- or tube-shaped trap.
   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.
   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. Nepenthes usually produce two types of pitchers. Appearing near the base of the plant are the large lower traps, which typically sit on the ground, while the upper pitchers may be smaller, colored differently, and have different features than 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 form a loop in the tendril, allowing it to wrap around nearby support. In some species (for example N. rafflesiana) different prey may be attracted by different types of pitchers.
   Prey usually consists of insects, but the largest species (N. rajah, N. rafflesiana, etc.) may occasionally catch small vertebrates, such as rats and lizards. Flowers occur in racemes or more rarely in panicles with male and female flowers on separate plants. Seed is produced in a four-sided capsule which may contain 10-60 or more seeds, consisting of a central ovary and two wings, one on either side. Seeds are wind distributed.

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.
   "Highlander" forms are those species that grow in habitats that are generally higher up in elevation, and thus exposed to cooler evening temperatures. "Lowlander" forms are those species that grow nearer to sealevel.
   Both forms respond best to rainwater (but tap water works as long as you flush it out with rainwater every month or if you've soft water), bright light (not full sun), a well drained medium, good air circulation and a 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 may be sown on damp chopped Sphagnum moss, or on sterile plant tissue culture media once they've been properly disinfected. The seeds generally become inviable soon after harvesting, so seed are not usually the preferred method of propagation. A 50:50 mixture of orchid medium with moss or perlite is suitable 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

More than 100 species of Nepenthes are currently recognised as valid. This number is fast increasing, with one to two new species being described each year.
Species Authority Date Image Distribution Altitudinal distribution IUCN conservation status
Nepenthes adnata Tamin & M.Hotta ex Schlauer 1994 Sumatra 600–1100 m Data Deficient
Nepenthes adrianii Batoro, Wartono, Jebb et al. 2006 Java >950 m
Nepenthes alata Blanco 1837 Philippines 800–2400 m Least Concern
Nepenthes albomarginata Lobb ex Lindl. 1849 Borneo, Peninsular Malaysia, Sumatra 0–1100 m Conservation Dependent
Nepenthes ampullaria Jack 1835 Borneo, Maluku Islands, New Guinea, Peninsular Malaysia, Singapore, Sumatra, Thailand 0–2100 m Least Concern
Nepenthes anamensis Macfarl. 1908 Cambodia, Laos, Thailand, Vietnam ~1500 m Data Deficient
Nepenthes angasanensis Maulder, D.Schub., B.R.Salmon & B.Quinn 1999 Sumatra 2200–2800 m Conservation Dependent
Nepenthes argentii Jebb & Cheek 1997 Philippines ~1400 m Vulnerable
Nepenthes aristolochioides Jebb & Cheek 1997 Sumatra 2000–2500 m Critically Endangered
Nepenthes beccariana Macfarl. 1908 Nias, Sumatra? ?
Nepenthes bellii K.Kondo 1969 Philippines 250–800 m Endangered
Nepenthes benstonei C.Clarke 1999 Peninsular Malaysia 450–600 m
Nepenthes bicalcarata Hook.f. 1873 Borneo 0–?950 m Vulnerable
Nepenthes bongso Korth. 1839 Sumatra 1000–2700 m Vulnerable
Nepenthes boschiana Korth. 1839 Borneo 900–1880 m Endangered
Nepenthes burbidgeae Hook.f. ex Burb. 1882 Borneo 1200–2250 m Endangered
Nepenthes burkei Hort.Veitch ex Mast. 1889 Philippines 1300–1600 m
Nepenthes campanulata Sh.Kurata 1973 Borneo 300–500 m Data Deficient
Nepenthes carunculata Danser 1928 Sumatra 1700–2330 m
Nepenthes chaniana C.Clarke, Chi C.Lee & S.McPherson 2006 Borneo 1500–1800
Nepenthes clipeata Danser 1928 Borneo 600–800 m Critically Endangered
Nepenthes copelandii Merr. ex Macfarl. 1908 Philippines ?
Nepenthes danseri Jebb & Cheek 1997 Halmahera, Waigeo Island 0–300 m Vulnerable
Nepenthes deaniana Macfarl. 1908 Philippines 1300–1500 m Data Deficient
Nepenthes densiflora Danser 1940 Sumatra 1700–3200 m Conservation Dependent
Nepenthes diatas Jebb & Cheek 1997 Sumatra 2400–2900 m Conservation Dependent
Nepenthes distillatoria L. 1753 Sri Lanka 0–700 m Vulnerable
Nepenthes dubia Danser 1928 Sumatra 1800–2700 m Critically Endangered
Nepenthes edwardsiana Low ex Hook.f. 1859 Borneo 1500–2700 m Vulnerable
Nepenthes ephippiata Danser 1928 Borneo 1000–1900 m Vulnerable
Nepenthes eustachya Miq. 1858 Sumatra 0–1600 Least Concern
Nepenthes eymae Sh.Kurata 1984 Sulawesi 1500–1800 m Vulnerable
Nepenthes faizaliana J.H.Adam & Wilcock 1991 Borneo 400–1500 m Vulnerable
Nepenthes fallax Beck 1895 Borneo 900–2600 m Vulnerable
Nepenthes flava Wistuba, Nerz & A.Fleischm. 2007 Sumatra 1800–2200 m
Nepenthes fusca Danser 1928 Borneo 1200–2500 m Vulnerable
Nepenthes glabrata J.R.Turnbull & A.T.Middleton 1984 Sulawesi 1600–2000 m Vulnerable
Nepenthes glandulifera Chi C.Lee 2004 Borneo 1100–1700 m
Nepenthes globosa Sh.Kurata in sched. 2006 Thailand ?
Nepenthes gracilis Korth. 1839 Borneo, Peninsular Malaysia, Singapore, Sulawesi, Sumatra, Thailand 0–1100 m Least Concern
Nepenthes gracillima Ridl. 1908 Peninsular Malaysia 1300–2100 m Endangered
Nepenthes gymnamphora Reinw. ex Nees 1824 Java (at least) 600–2800 m Least Concern
Nepenthes hamata J.R.Turnbull & A.T.Middleton 1984 Sulawesi 1400–2500 m Vulnerable
Nepenthes hirsuta Hook.f. 1873 Borneo 200–1000 m Conservation Dependent
Nepenthes hispida Beck 1895 Borneo 100–800 m Conservation Dependent
Nepenthes hurrelliana Cheek & A.L.Lamb 2003 Borneo >1300 m
Nepenthes inermis Danser 1928 Sumatra 1500–2600 m Vulnerable
Nepenthes insignis Danser 1928 New Guinea 80–800 m Vulnerable
Nepenthes izumiae Troy Davis, C.Clarke & Tamin 2003 Sumatra >1800 m
Nepenthes jacquelineae C.Clarke, Troy Davis & Tamin 2001 Sumatra 1700–2200 m
Nepenthes jamban Chi C.Lee, Hernawati & Akhriadi 2006 Sumatra ?
Nepenthes junghuhnii sensu Macfarl. in sched. nom.nud. 1917 Sumatra 227–648 m
Nepenthes khasiana Hook.f. 1873 India ?
Nepenthes klossii Ridl. 1916 New Guinea 100–2000 m Vulnerable
Nepenthes lamii Jebb & Cheek 1997 New Guinea 1460–3520 m
Nepenthes lavicola Wistuba & Rischer 1996 Sumatra 2000–2600 m Critically Endangered
Nepenthes lingulata Chi C.Lee, Hernawati & Akhriadi 2006 Sumatra ?
Nepenthes longifolia Nerz & Wistuba 1994 Sumatra 300–1100 m
Nepenthes lowii Hook.f. 1859 Borneo 1600–2600 m Vulnerable
Nepenthes macfarlanei Hemsl. 1905 Peninsular Malaysia 1000–2150 m Vulnerable
Nepenthes macrophylla (Marabini) Jebb & Cheek 1997 Borneo 2200–2400 m Critically Endangered
Nepenthes macrovulgaris J.R.Turnbull & A.T.Middleton 1988 Borneo 300–1220 m Vulnerable
Nepenthes madagascariensis Poir. 1797 Madagascar ? Vulnerable
Nepenthes mantalingajanensis Nerz & Wistuba 2007 Philippines ?
Nepenthes mapuluensis J.H.Adam & Wilcock 1990 Borneo 700–800 m Near Threatened
Nepenthes masoalensis Schmid-Höllinger 1977 Madagascar 30–400 m Endangered
Nepenthes maxima Reinw. ex Nees 1824 Maluku Islands, New Guinea, Sulawesi 40–2500 m Least Concern
Nepenthes merrilliana Macfarl. 1911 Philippines 20–1700 m Vulnerable
Nepenthes mikei B.R.Salmon & Maulder 1995 Sumatra 1100–2800 m Vulnerable
Nepenthes mindanaoensis Sh.Kurata 2001 Philippines ?
Nepenthes mira Jebb & Cheek 1998 Philippines 1500–2000 m
Nepenthes mirabilis (Lour.) Druce 1916 Australia, Borneo, Cambodia, China, Hong Kong, Laos, Macau, Maluku Islands, Micronesia, Myanmar, New Guinea, Palau, Peninsular Malaysia, Philippines, Sulawesi, Sumatra, Thailand, Vietnam 0–1500 m Least Concern
Nepenthes mollis Danser 1928 Borneo ~1800 m Data Deficient
Nepenthes muluensis M.Hotta 1966 Borneo 1900–2400 m Vulnerable
Nepenthes murudensis Culham ex Jebb & Cheek 1997 Borneo 2100–2420 m Endangered
Nepenthes naquiyuddinii J.H.Adam & Hafiza 2006 Borneo 1400–1424 m
Nepenthes neoguineensis Macfarl. 1911 D'Entrecasteaux Islands, New Guinea 0–1400 m Least Concern
Nepenthes northiana Hook.f. 1881 Borneo 0–500 m Vulnerable
Nepenthes ovata Nerz & Wistuba 1994 Sumatra 1700–2100 m Vulnerable
Nepenthes paniculata Danser 1928 New Guinea ~1460 m Endangered
Nepenthes papuana Danser 1928 New Guinea 250–1300 m Data Deficient
Nepenthes pectinata Danser 1928 Sumatra 950–2750 m Least Concern
Nepenthes peltata Sh.Kurata 2008 Philippines ?
Nepenthes pervillei Blume 1852 Seychelles 350–500 m Vulnerable
Nepenthes petiolata Danser 1928 Philippines ~1500 m
Nepenthes philippinensis Macfarl. 1908 Philippines 0–500 m
Nepenthes pilosa Danser 1928 Borneo ~1600 m Endangered
Nepenthes platychila Chi C.Lee 2002 Borneo 900–1400 m
Nepenthes rafflesiana Jack 1835 Borneo, Peninsular Malaysia, Singapore, Sumatra 0–1200 m Least Concern
Nepenthes rajah Hook.f. 1859 Borneo 1500–2650 m Endangered
Nepenthes ramispina Ridl. 1908 Peninsular Malaysia 900–2000 m Vulnerable
Nepenthes reinwardtiana Miq. 1852 Borneo, Sumatra 0–2100 m Least Concern
Nepenthes rhombicaulis Sh.Kurata 1973 Sumatra 1700–1900 m Vulnerable
Nepenthes rigidifolia Akhriadi, Hernawati & Tamin 2004 Sumatra 1000–1600 m
Nepenthes rowanae F.M.Bailey 1897 Australia 0–10 m
Nepenthes sanguinea Lindl. 1849 Peninsular Malaysia 300–1800 m Conservation Dependent
Nepenthes saranganiensis Sh.Kurata 2003 Philippines ?
Nepenthes sharifah-hapsahii J.H.Adam & Hafiza 2007 Peninsular Malaysia <1000 m
Nepenthes sibuyanensis Nerz 1998 Philippines 1500–1800 m Vulnerable
Nepenthes singalana Becc. 1886 Sumatra 2000–2900 m Vulnerable
Nepenthes smilesii Hemsl. 1895 Cambodia, Myanmar, Thailand ?
Nepenthes spathulata Danser 1935 Sumatra 1100–2900 m Vulnerable
Nepenthes spectabilis Danser 1928 Sumatra 1400–2200 m Vulnerable
Nepenthes stenophylla Mast. 1890 Borneo ? Least Concern
Nepenthes sumatrana (Miq.) Beck 1895 Sumatra 0–800 m Least Concern
Nepenthes talangensis Nerz & Wistuba 1994 Sumatra 1800–2500 m Endangered
Nepenthes tenax C.Clarke & R.Kruger 2006 Australia ?
Nepenthes tentaculata Hook.f. 1873 Borneo, Sulawesi 700–2550 m Least Concern
Nepenthes tenuis Nerz & Wistuba 1994 Sumatra ~1000 m Data Deficient
Nepenthes thorelii Lecomte 1909 Cambodia, Laos?, Thailand, Vietnam 0–200 m Data Deficient
Nepenthes tobaica Danser 1928 Sumatra 380–2750 m Least Concern
Nepenthes tomoriana Danser 1928 Sulawesi 0–400 m Vulnerable
Nepenthes treubiana Warb. 1891 New Guinea 0–500 m Vulnerable
Nepenthes truncata Macfarl. 1911 Philippines 230–>600 m Endangered
Nepenthes veitchii Hook.f. 1859 Borneo 0–1500 m Near Threatened
Nepenthes ventricosa Blanco 1837 Philippines 1200–1500 m Near Threatened
Nepenthes vieillardii Hook.f. 1873 New Caledonia 30–800 m Least Concern
Nepenthes villosa Hook.f. 1852 Borneo 2340–3240 m Vulnerable
Nepenthes vogelii Schuit. & de Vogel 2002 Borneo 1000–1400 m
Nepenthes xiphioides B.R.Salmon & Maulder 1995 Sumatra 1800–1900 m
Nepenthes zakriana (J.H.Adam & Wilcock)
J.H.Adam & Hafiza
2006 Borneo 1200–1500 m
Hybrids and cultivars There are many hybrid Nepenthes and numerous named cultivars. The following are named natural hybrids:
  • N. × alisaputrana (N. burbidgeae × N. rajah)
  • N. × bauensis (N. gracilis × N. northiana)
  • N. × cantleyi (N. bicalcarata × N. gracilis)
  • N. × cincta (N. albomarginata × N. northiana)
  • N. × ferrugineomarginata (N. albomarginata × N. reinwardtiana)
  • N. × ghazallyana (N. gracilis × N. mirabilis)
  • N. × harryana (N. edwardsiana × N. villosa)
  • N. × hookeriana (N. ampullaria × N. rafflesiana)
  • N. × kinabaluensis (N. rajah × N. villosa)
  • N. × kuchingensis (N. ampullaria × N. mirabilis)
  • N. × merrilliata (N. alata × N. merrilliana)
  • N. × mirabilata (N. alata × N. mirabilis)
  • N. × pangulubauensis (N. mikei × N. pectinata)
  • N. × pyriformis (N. inermis × N. talangensis)
  • N. × sarawakiensis (N. muluensis × N. tentaculata)
  • N. × trichocarpa (N. ampullaria × N. gracilis)
  • N. × truncalata (N. alata × N. truncata)
  • N. × trusmadiensis (N. lowii × N. macrophylla)
  • N. × tsangoya ((N. alata × N. merrilliana) × N. mirabilis)
  • N. × ventrata (N. alata × N. ventricosa) 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)
  • N. 'Ventrata' (N. ventricosa × N. alata)Further Information

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