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Nepenthes

February 16, 2024

This article is about the tropical pitcher plant. For other uses, see Nepenthe (disambiguation).

Nepenthes (/nɪˈpɛnθiːz/) 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,[4] 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 edit

 
Nepenthes mirabilis at the Periyar Tiger Reserve, in Southern Western Ghats of India

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 (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.[5]

 
Basic structure of an upper pitcher

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.[6] The trapping efficiency of this fluid remains high, even when significantly diluted by water, as inevitably happens in wet conditions.[7]

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.[8] 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.[5]

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.[5]

Prey usually consists of insects, but the largest species (e.g. N. rajah and N. rafflesiana) may occasionally catch small vertebrates, such as "frogs, birds, and small mammals".[9][10] Records of cultivated plants trapping small birds have been made.[11][12] 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.[13]

Nepenthes are insect-pollinated, the primary agents being flies (including blow flies, midges, and mosquitoes), moths, wasps, and butterflies.[14] Their smells can range from sweet to musty or fungus-like.[15] 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.[16][17] This high number is thought to reflect paleopolyploidy (likely 8x or 16x).[17][18][19][20]

Taxonomy edit

Main articles: List of Nepenthes species and List of Nepenthes species by distribution

About 170 species of Nepenthes are currently recognised as valid. This number is increasing, with several new species being described each year.[21]

Etymology edit

The genus name Nepenthes was first published in 1737 in Carl Linnaeus's Hortus Cliffortianus.[22] It references a passage in Homer's Odyssey, in which the potion "Nepenthes pharmakon" is given to Helen by an Egyptian queen. "Nepenthes" (Ancient Greek: νηπενθές) literally means "without grief" (νη nē = "not", πένθος penthos = "grief") and, in Greek mythology, is a drug that quells all sorrows with forgetfulness.[15][page needed][23] 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][24]

The plant Linnaeus described was N. distillatoria, called bāndurā (බාඳුරා), a species from Sri Lanka.[15][page needed]

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.[25]

 
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:[26]

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) in the Philippines.[citation needed]

Evolution and phylogeny edit

An absence of evidence of intermediate species, fossil or living (i.e. a missing link), does not allow forming a phylogenetic timeline for the development of the distinctive traits of modern Nepenthes, which include its relatively rare strict dioecy and carnivorous pitchers. Although Nepenthes is distantly related to several modern genera, among these, even the carnivorous relatives [the sundews (Drosera), Venus flytrap (Dionaea muscipula), waterwheel plant (Aldrovanda), and dewy pine (Drosophyllum)], all lack those traits. Among known Nepenthes, no protomodern characteristics or large variations are found, which suggests that all extant species radiated from a single close ancestor bearing all the modern traits. Phylogenetic comparisons of the chloroplast matK gene sequences between Nepenthes species and with related species support this conclusion, long genetic distance between Nepenthes and others, and abruptly diverging "pom-pom" grouping of the Nepenthes species .[27]

Fossilized pollen of Nepenthes-like plants living on the northern Tethys Sea from 65 to 35 million years ago indicates that then-warmer Europe may have been where the proto-Nepenthes developed, and then escaped to Asia and India as Africa collided with Europe and the ensuing climate change wiped out the ancestral species in the original habitat. About 20 million years ago, Borneo, Sumatra, and Sulawesi and possibly even the Philippines were connected to mainland Asia, providing a bridge for the colonization of most sites of Nepenthes species radiation. The extensive landbridges in the area 20,000 years ago during the ice age would have provided access to the remaining sites of Nepenthes populations in Oceania. The main complication with this hypothesis is the presence of Nepenthes on the distant islands of Seychelles and Madagascar. The seeds were thought to have been transferred by seabirds and shorebirds, which rest during their migrations in swampy habitats and may have inadvertently picked up the seeds. This hypothesis is possibly reinforced by the success of the lowland swamp-dwelling N. distillatoria in colonizing so many locations.[27]

Distribution and habitat edit

 
Global distribution of Nepenthes
Further information: List of Nepenthes species by distribution

The genus Nepenthes is mostly found within the Malay Archipelago, with the greatest biodiversity found on Borneo, Sumatra, and the Philippines,[28][29] 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, N. parvula, and N. tenax) and New Caledonia (N. vieillardii) in the southeast. Most species are restricted to very small ranges, including some found only on individual mountains. These limited distributions and the inaccessibility of the regions often means some species go decades without being rediscovered in the wild (e.g. N. deaniana, which was rediscovered 100 years after its initial discovery). About 10 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.[5][30][31]

Because of the nature of the habitats that 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 growing 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 grows at a higher altitude than any other in the genus, up to 3,520 m (11,549 ft).[5][31]

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 in which Nepenthes species grow are usually acidic and low in nutrients, being composed of peat, white sand, sandstone, or volcanic soils. Exceptions to these generalities include species that thrive in soils with 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.[5]

Ecological relationships edit

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

The most obvious interaction between Nepenthes species and their environments, 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 these 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 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.[5] While many Nepenthes species 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 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.[32][33]

Prey retention by viscoelastic pitcher fluid of N. rafflesiana[7]
  • The blue bottle fly (Calliphora vomitoria) can escape after landing in water on its ventral surface.
  • The same is true if the fly falls in dorsally (wings-first).
  • But the viscoelastic properties of N. rafflesiana digestive fluid prevent prey escape, whether the fall is ventral..
  • ..or dorsal. (All videos recorded at 500 frames/s)

Symbioses edit

 
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.[34]

N. 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.[35]

N. lowii has also formed a dependent relationship, but 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 between 57 and 100% of the plant's foliar nitrogen uptake comes from the faeces of tree shrews.[36] Another study showed the shape and size of the pitcher orifice of N. lowii exactly match the dimensions of a typical tree shrew (Tupaia montana).[37][38] A similar adaptation was found in N. macrophylla, N. rajah, N. ampullaria, and is also likely to be present in N. ephippiata.[38][39]

Similarly, N. hemsleyana, which is native to Borneo, has a symbiotic partnership with Hardwicke's woolly bat.[40] During the day, a bat may roost above the digestive fluid inside the pitcher. While a bat is inside, it may defaecate, with the plant gaining nitrogen from the droppings. Further research has discovered that the shape and design of the pitcher has evolved to be an acoustic reflector to make it easier for bats to echo-locate, and distinguishes it from other closely related species that don't make good roosts.[41][42]

Infauna edit

Main article: Nepenthes 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, often associated with but 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 plants and infauna is not yet fully understood, but the relationship may be 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.[35][43][44]

Antimicrobial properties edit

Nepenthes digestive fluids are sterile before pitchers open and contain secondary metabolites and proteins that act as bactericides and fungicides after the pitcher opens. While the digestive fluid is being produced, the pitcher is not yet open, so there is no chance of microbial contamination. During pitcher development, at least 29 digestive proteins including proteases, chitinases, pathogenesis-related proteins and thaumatin-like proteins are produced in the pitcher fluid. In addition to breaking down prey, these can act as antimicrobial agents.[45] When the pitchers open, the fluid is exposed to bacteria, fungal spores, insects and rain. Often pitchers have a lid that covers the trap, excepting a few (e.g. N. lowii, N. attenboroughii and N. jamban), preventing rain water from entering. The lid inhibits rainwater from diluting the digestive fluid. Once the bacteria and fungi enter the fluid, secondary metabolites are produced in addition to antimicrobial proteins.[46] Naphthoquinones, a class of secondary metabolite, are commonly produced, and these either kill or inhibit the growth and reproduction of bacteria and fungi.[47] This adaptation could have evolved since Nepenthes plants that could produce secondary metabolites and antimicrobial proteins to kill bacteria and fungi were most likely more fit. Plants that produced antimicrobial compounds could prevent loss of valuable nutrients gained from insects within the pitcher. Since Nepenthes cannot digest certain bacteria and fungi, the bactericides and fungicides allow plants to maximize nutrient uptake.

Botanical history edit

 
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 Étienne de Flacourt published a description of a pitcher plant in his seminal work Histoire de la Grande Isle de Madagascar. It reads:[48]

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][15]

Flacourt called the plant Amramatico, after a local name. More than a century later, this species was formally described as N. madagascariensis.[49]

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".[50] Three years later, Dutch merchant Jacob Breyne referred to this species as Bandura zingalensium, after a local name for the plant.[51] Bandura subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined Nepenthes in 1737.[15]

Nepenthes distillatoria was again described in 1683, this time by Swedish physician and naturalist Herman Niklas Grim.[52] Grim called it Planta mirabilis destillatoria or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant.[15] Three years later, in 1686, English naturalist John Ray quoted Grim as saying:[53]

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][15]

One of the earliest illustrations of Nepenthes appears in Leonard Plukenet's Almagestum Botanicum of 1696.[54] The plant, called Utricaria vegetabilis zeylanensium, is undoubtedly N. distillatoria.[15]

 
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.

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.[55]

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.[15]

 
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.[56]

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:[57]

[...] (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][15]

Phyllamphora mirabilis was eventually transferred to the genus Nepenthes by Rafarin in 1869.[58] As such, P. mirabilis is the basionym of this most cosmopolitan of tropical pitcher plant species.[35]

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:[49]

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][15]

 
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.[5][15] 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.[21]

Cultivation edit

 
Cultivated Nepenthes rajah, Nepenthes aristolochioides and other species
 
Nepenthes cultivated in a garden at Kottayam, Kerala

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.[59]

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.[60][61]

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.[62] 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 edit

 
The complex man-made hybrid N. ventricosa × (N. lowii × N. macrophylla)
See also: List of Nepenthes natural hybrids and List of Nepenthes cultivars

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

  • 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)
  • N. 'Gaya' (N. khasiana × (N. ventricosa × N. maxima))

See also edit

References edit

  1. ^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III" (PDF). Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. Retrieved 2013-07-06.
  2. ^ Cheek, M.; Jebb, M. (2013). "The Nepenthes micramphora (Nepenthaceae) group, with two new species from Mindanao, Philippines". Phytotaxa. 151 (1): 25–34. doi:10.11646/phytotaxa.151.1.2.
  3. ^ "Nepenthes L." Plants of the World Online. Royal Botanical Gardens, Kew. 2023. Retrieved 7 June 2023.
  4. ^ Christenhusz, M. J. M.; Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
  5. ^ a b c d e f g h Barthlott, W., Porembski, S., Seine, R., and Theisen, I. 2007. The Curious World of Carnivorous Plants. Portland, Oregon: Timber Press.
  6. ^ Bonhomme, V. (2011). "Slippery or sticky? Functional diversity in the trapping strategy of Nepenthes carnivorous plants". New Phytologist. 191 (2): 545–554. doi:10.1111/j.1469-8137.2011.03696.x. PMID 21434933.
  7. ^ a b Gaume, L.; Forterre, Y. (2007). "A viscoelastic deadly fluid in carnivorous pitcher plants". PLOS ONE. 2 (11): e1185. Bibcode:2007PLoSO...2.1185G. doi:10.1371/journal.pone.0001185. PMC 2075164. PMID 18030325.
  8. ^ Moran, J.A. (2010). "The carnivorous syndrome in Nepenthes pitcher plants". Plant Signaling & Behavior. 5 (6): 644–648. Bibcode:2010PlSiB...5..644M. doi:10.4161/psb.5.6.11238. PMC 3001552. PMID 21135573.
  9. ^ Phillipps, A (1988). "A second record of rats as prey in Nepenthes rajah" (PDF). Carnivorous Plant Newsletter. 17 (2): 55.
  10. ^ Moran, J.A. 1991. The role and mechanism of Nepenthes rafflesiana pitchers as insect traps in Brunei. Ph.D. thesis, University of Aberdeen, Aberdeen, Scotland.
  11. ^ "Killer plant 'eats' great tit at Somerset nursery". BBC News. 5 August 2011. Retrieved 5 August 2011.
  12. ^ Hewitt-Cooper, N (2012). "A case of bird capture by a cultivated specimen of the hybrid Nepenthes × mixta". Carnivorous Plant Newsletter. 41 (1): 31–33. doi:10.55360/cpn411.nh396. S2CID 247074534.
  13. ^ Clarke, Charles; Moran, Jonathan A.; Chin, Lijin (1 October 2010). "Mutualism between tree shrews and pitcher plants". Plant Signal Behav. 5 (10): 1187–1189. Bibcode:2010PlSiB...5.1187C. doi:10.4161/psb.5.10.12807. PMC 3115346. PMID 20861680.
  14. ^ Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  15. ^ a b c d e f g h i j k l Phillipps, A.; Lamb, A. (1996). Pitcher-Plants of Borneo. Kota Kinabalu, Sabah, Malaysia: Natural History Publications (Borneo).
  16. ^ Lowrey, T.K. 1991. No. 519: Chromosome and isozyme number in the Nepenthaceae. American Journal of Botany 78(6, supplement): 200–201.
  17. ^ a b Heubl, G.R.; Wistuba, A. (1997). "A cytological study of the genus Nepenthes L. (Nepenthaceae)". Sendtnera. 4: 169–174.
  18. ^ Meimberg, H.; Heubl, G. (2006). "Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae". Plant Biology. 8 (6): 831–840. Bibcode:2006PlBio...8..831M. doi:10.1055/s-2006-924676. PMID 17203435. S2CID 8482890.
  19. ^ Cytology of Nepenthes. LMU Department für Biologie.
  20. ^ Brittnacher, J. N.d. Evolution -- Nepenthes Phylogeny 2011-05-14 at the Wayback Machine. International Carnivorous Plant Society.
  21. ^ a b Clarke, C.M. & C.C. Lee 2004. Pitcher Plants of Sarawak. Natural History Publications (Borneo), Kota Kinabalu.
  22. ^ Linnaeus, C. 1737. Nepenthes. Hortus Cliffortianus. Amsterdam.
  23. ^ Gledhill, David (2008). The Names of Plants (4th ed.). Cambridge: Cambridge University Press. p. 271. ISBN 978-0-521-86645-3.
  24. ^ Veitch, H.J. (1897). "Nepenthes". Journal of the Royal Horticultural Society. 21 (2): 226–262.
  25. ^ Linnaeus, C (1753). "Nepenthes". Species Plantarum. 2: 955.
  26. ^ Zahl, P.A. (1964). "Malaysia's Giant Flowers and Insect-trapping Plants". National Geographic. 125 (5): 680–701.
  27. ^ a b Brittnacher, John (2011). . ICPS Webpage. Archived from the original on 2011-05-14. Retrieved 2011-02-23.
  28. ^ Amoroso, Victor B.; Aspiras, Reyno A. (2011-01-01). "Hamiguitan Range: A sanctuary for native flora". Saudi Journal of Biological Sciences. 18 (1): 7–15. doi:10.1016/j.sjbs.2010.07.003. ISSN 1319-562X. PMC 3730927. PMID 23961098.
  29. ^ Amoroso, V. B.; Obsioma, L. D.; Arlalejo, J. B.; Aspiras, R. A.; Capili, D. P.; Polizon, J. J. A.; Sumile, E. B. (2009). "Inventory and conservation of endangered, endemic and economically important flora of Hamiguitan Range, southern Philippines". Blumea - Biodiversity, Evolution and Biogeography of Plants. 54: 71–76. doi:10.3767/000651909X474113. Retrieved 2019-11-03.
  30. ^ McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
  31. ^ a b Jebb, M.; Cheek, M. (1997). "A skeletal revision of Nepenthes (Nepenthaceae)". Blumea. 42: 1–106.
  32. ^ Moran, J.A.; Merbach, M.A.; Livingston, N.J.; Clarke, C.M.; Booth, W.E. (2001). "Termite prey specialization in the pitcher plant Nepenthes albomarginata—evidence from stable isotope analysis". Annals of Botany. 88 (2): 307–311. doi:10.1006/anbo.2001.1460.
  33. ^ Merbach, M.A.; Merbach, D.J.; Maschwitz, U.; Booth, W.E.; Fiala, B.; Zizka, G. (2002). "Mass march of termites into the deadly trap". Nature. 415 (6867): 36–37. doi:10.1038/415036a. PMID 11780106. S2CID 846937.
  34. ^ Robinson, A.S.; Fleischmann, A.S.; McPherson, S.R.; Heinrich, V.B.; Gironella, E.P.; Peña, C.Q. (2009). "A spectacular new species of Nepenthes L. (Nepenthaceae) pitcher plant from central Palawan, Philippines". Botanical Journal of the Linnean Society. 159 (2): 195–202. doi:10.1111/j.1095-8339.2008.00942.x.
  35. ^ a b c Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  36. ^ Clarke, C.M.; Bauer, U.; Lee, C.C.; Tuen, A.A.; Rembold, K.; Moran, J.A. (2009). "Tree shrew lavatories: a novel nitrogen sequestration strategy in a tropical pitcher plant". Biology Letters. 5 (5): 632–635. doi:10.1098/rsbl.2009.0311. PMC 2781956. PMID 19515656.
  37. ^ Chin, L.; Moran, J.A.; Clarke, C. (2010). "Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size". New Phytologist. 186 (2): 461–470. doi:10.1111/j.1469-8137.2009.03166.x. PMID 20100203.
  38. ^ a b Walker, M. 2010. Giant meat-eating plants prefer to eat tree shrew poo. BBC Earth News, March 10, 2010.
  39. ^ Moran, J.A. (2003). "From carnivore to detritivore? Isotopic evidence for leaf litter utilization by the tropical pitcher plant Nepenthes ampullaria". International Journal of Plant Sciences. 164 (4): 635–639. doi:10.1086/375422. hdl:10170/576. S2CID 53573745.
  40. ^ Grafe, T. U.; Schoner, C. R.; Kerth, G.; Junaidi, A. & Schoner, M. G. (2011). "A novel resource-service mutualism between bats and pitcher plants". Biology Letters. 7 (3): 436–439. doi:10.1098/rsbl.2010.1141. PMC 3097880. PMID 21270023.
  41. ^ Schöner, M. G.; Schöner, C. R.; Simon, R.; Grafe, T. Ulmar; Puechmaille, S. J.; Ji, L. L.; Kerth, G. (2015-07-09). "Bats Are Acoustically Attracted to Mutualistic Carnivorous Plants". Current Biology. 25 (14): 1911–1916. doi:10.1016/j.cub.2015.05.054. PMID 26166777.
  42. ^ Thompson, Ken (1 November 2018). Darwin's Most Wonderful Plants: Darwin's Botany Today. Profile Books. p. 146. ISBN 978-1-78283-436-6. Retrieved 19 November 2023.
  43. ^ Mogi, M.; Yong, H.S. (1992). "Aquatic arthropod communities in Nepenthes pitchers: the role of niche differentiation, aggregation, predation and competition in community organization". Oecologia. 90 (2): 172–184. Bibcode:1992Oecol..90..172M. doi:10.1007/BF00317174. PMID 28313712. S2CID 26808467.
  44. ^ Beaver, R.A. (1979). "Fauna and foodwebs of pitcher plants in west Malaysia". Malayan Nature Journal. 33: 1–10.
  45. ^ Rottloff, Sandy; Miguel, Sissi; Biteau, Flore; Nisse, Estelle; Hammann, Philippe; Kuhn, Lauriane; Chicher, Johana; Bazile, Vincent; Gaume, Laurence (2016-03-01). "Proteome analysis of digestive fluids in Nepenthes pitchers". Annals of Botany. 117 (3): 479–495. doi:10.1093/aob/mcw001. ISSN 0305-7364. PMC 4765550. PMID 26912512.
  46. ^ Mithöfer, Axel (2011-09-01). "Carnivorous pitcher plants: Insights in an old topic". Phytochemistry. Plant-Insect Interactions. 72 (13): 1678–1682. Bibcode:2011PChem..72.1678M. doi:10.1016/j.phytochem.2010.11.024. PMID 21185041.
  47. ^ Buch, Franziska; Rott, Matthias; Rottloff, Sandy; Paetz, Christian; Hilke, Ines; Raessler, Michael; Mithöfer, Axel (2012-12-21). "Secreted pitfall-trap fluid of carnivorous Nepenthes plants is unsuitable for microbial growth". Annals of Botany. 111 (3): 375–83. doi:10.1093/aob/mcs287. ISSN 0305-7364. PMC 3579442. PMID 23264234.
  48. ^ de Flacourt, É. 1658. Histoire de la Grande Isle de Madagascar.
  49. ^ a b Poiret, J.L.M. 1797. Népente. In: J.B. Lamarck Encyclopédie Méthodique Botanique Vol. 4.
  50. ^ Bartholinus. "Miranda herba". Acta Medica et Philosophica Hafniensia. 3: 38.
  51. ^ Breyne, J. 1680. Bandura zingalensium etc. Prodromus Fasciculi Rariorum Plantarum 1: 18.
  52. ^ Grimm, H.N. 1683. Planta mirabilis destillatoria. In: Miscellanea curiosa sive Ephemeridum. Med. Phys. Germ. Acad. Nat. Cur. Decuriae 2, ann. prim. p. 363, f. 27.
  53. ^ Ray, J. 1686. Bandura cingalensium etc. Historia Plantarum 1: 721–722.
  54. ^ Plukenet, L. 1696. Utricaria vegetabilis zeylanensium. In: Almagestum Botanicum.
  55. ^ Rumphius, G.E. 1741–1750. Cantharifera. In: Herbarium Amboinense 5, lib. 7, cap. 61, p. 121, t. 59, t. 2.
  56. ^ Burmann, J. 1737. Thesaurus Zeylanicus. Amsterdam.
  57. ^ de Loureiro, J. 1790. Flora Cochinchinensis 2: 606–607.
  58. ^ Schlauer, J. N.d. Nepenthes mirabilis. Carnivorous Plant Database.
  59. ^ James Pietropaolo; Patricia Ann Pietropaolo (1986). Carnivorous Plants of the World. Timber Press. p. 49. ISBN 978-0-88192-066-6.
  60. ^ Peter D'Amato (2 July 2013). The Savage Garden, Revised: Cultivating Carnivorous Plants. Ten Speed Press. p. 89. ISBN 978-1-60774-411-5.
  61. ^ Barry A. Rice (2006). Growing carnivorous plants. Timber Press, Incorporated. ISBN 978-0-88192-807-5.
  62. ^ "Appendices | CITES". cites.org. Retrieved 2022-01-14.
  • 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 edit

  • Amagase, S.; Nakayama, S.; Tsugita, A. (1969). "Acid protease in Nepenthes. II. Study on the specificity of nepenthesin". The Journal of Biochemistry. 66 (4): 431–439. doi:10.1093/oxfordjournals.jbchem.a129166. PMID 5354017.
  • Athauda, S.B.P.; Matsumoto, K.; Rajapakshe, S.; Kuribayashi, M.; Kojima, M.; Kubomura-Yoshida, N.; Iwamatsu, A.; Shibata, C.; Inoue, H.; Takahashi, K. (2004). "Enzymatic and structural characterization of nepenthesin, a unique member of a novel subfamily of aspartic proteinases". Biochemical Journal. 381 (1): 295–306. doi:10.1042/BJ20031575. PMC 1133788. PMID 15035659.
  • Bauer, U.; Bohn, H.F.; Federle, W. (2008). "Harmless nectar source or deadly trap: Nepenthes pitchers are activated by rain, condensation and nectar". Proceedings of the Royal Society B. 275 (1632): 259–265. doi:10.1098/rspb.2007.1402. PMC 2593725. PMID 18048280.
  • Beaver, R.A. (1979). "Biological studies of the fauna of pitcher plants Nepenthes in west Malaysia". Annales de la Société Entomologique de France. 15: 3–17. doi:10.1080/21686351.1979.12278188. S2CID 83749546.
  • 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.
  • Beaver, R.A. (1985). "Geographical variation in food web structure in Nepenthes pitcher plants". Ecological Entomology. 10 (3): 241–248. Bibcode:1985EcoEn..10..241B. doi:10.1111/j.1365-2311.1985.tb00720.x. S2CID 85082186.
  • Beekman, E.M. (2004). "A Note on the Priority of Rumphius' Observation of Decapod Crustacea Living In Nepenthes". Crustaceana. 77 (8): 1019–1021. doi:10.1163/1568540042781748.
  • Bohn, H.F.; Federle, W. (2004). "Insect aquaplaning: Nepenthes pitcher plants capture prey with the peristome, a fully wettable water-lubricated anisotropic surface" (PDF). Proceedings of the National Academy of Sciences. 101 (39): 14138–14143. doi:10.1073/pnas.0405885101. PMC 521129. PMID 15383667.
  • (in French) Boulay, J. 1997. Les Nepenthes. Dionée 38.
  • Carlquist, S (1981). "Wood Anatomy of Nepenthaceae". Bulletin of the Torrey Botanical Club. 108 (3): 324–330. doi:10.2307/2484711. JSTOR 2484711.
  • Chia, T.F.; Aung, H.H.; Osipov, A.N.; Goh, N.K.; Chia, L.S. (2004). "Carnivorous pitcher plant uses free radicals in the digestion of prey". Redox Report. 9 (5): 255–261. doi:10.1179/135100004225006029. PMID 15606978.
  • Edwards, P (2005). "Growing Nepenthes – Part 1" (PDF). Victorian Carnivorous Plant Society Inc. 75: 6–8.
  • Edwards, P (2005). "Growing Nepenthes – Part 2" (PDF). Victorian Carnivorous Plant Society Inc. 76: 6–9.
  • Frazier, C.K. (2000). "The Enduring Controversies Concerning the Process of Protein Digestion in Nepenthes". Carnivorous Plant Newsletter. 29 (2): 56–61. doi:10.55360/cpn292.cf425. S2CID 247153343.
  • Jenkin, A (2005). (PDF). Victorian Carnivorous Plant Society Inc. 75: 12–13. Archived from the original (PDF) on 2011-07-07.
  • Jennings, D.E.; Rohr, JR. (2011). "A review of the conservation threats to carnivorous plants". Biological Conservation. 144 (5): 1356–1363. doi:10.1016/j.biocon.2011.03.013.
  • Karagatzides, J.D.; Ellison, A.M. (2009). "Construction costs, payback times, and the leaf economics of carnivorous plants". American Journal of Botany. 96 (9): 1612–1619. doi:10.3732/ajb.0900054. PMID 21622347. S2CID 42120981.
  • Meimberg, H.; Wistuba, A.; Dittrich, P.; Heubl, G. (2001). "Molecular Phylogeny of Nepenthaceae Based on Cladistic Analysis of Plastid trnK Intron Sequence Data". Plant Biology. 3 (2): 164–175. Bibcode:2001PlBio...3..164M. doi:10.1055/s-2001-12897. S2CID 260252804.
  • Mithöfer, A (2011). "Carnivorous pitcher plants: insights in an old topic". Phytochemistry. 72 (13): 1678–1682. Bibcode:2011PChem..72.1678M. doi:10.1016/j.phytochem.2010.11.024. PMID 21185041.
  • Moran, J.A.; Booth, W.E.; Charles, J.K. (1999). "Aspects of Pitcher Morphology and Spectral Characteristics of Six Bornean Nepenthes Pitcher Plant Species: Implications for Prey Capture". Annals of Botany. 83 (5): 521–528. doi:10.1006/anbo.1999.0857.
  • Nosonovsky, M (2011). "Materials science: slippery when wetted". Nature. 477 (7365): 412–413. Bibcode:2011Natur.477..412N. doi:10.1038/477412a. PMID 21938059. S2CID 205067351.
  • Osunkoya, O.O.; Daud, S.D.; Di-Giusto, B.; Wimmer, F.L.; Holige, T.M. (2007). "Construction Costs and Physico-chemical Properties of the Assimilatory Organs of Nepenthes Species in Northern Borneo". Annals of Botany. 99 (5): 895–906. doi:10.1093/aob/mcm023. PMC 2802909. PMID 17452380.
  • Pavlovič, A.; Masarovičová, E.; Hudák, J. (2007). "Carnivorous Syndrome in Asian Pitcher Plants of the Genus Nepenthes". Annals of Botany. 100 (3): 527–536. doi:10.1093/aob/mcm145. PMC 2533595. PMID 17664255.
  • Poppinga, S.; Koch, K.; Bohn, H.F.; Barthlott, W. (2010). "Comparative and functional morphology of hierarchically structured anti-adhesive surfaces in carnivorous plants and kettle trap flowers". Functional Plant Biology. 37 (10): 952–961. doi:10.1071/FP10061.
  • Riedel, M.; Eichner, A.; Meimberg, H.; Jetter, R. (2007). "Chemical composition of epicuticular wax crystals on the slippery zone in pitchers of five Nepenthes species and hybrids". Planta. 225 (6): 1517–1534. Bibcode:2007Plant.225.1517R. doi:10.1007/s00425-006-0437-3. PMID 17109149. S2CID 23581314.
  • Schulze, W.; Frommer, W.B.; Ward, J.M. (1999). "Transporters for ammonium, amino acids and peptides are expressed in pitchers of the carnivorous plant Nepenthes". The Plant Journal. 17 (6): 637–646. doi:10.1046/j.1365-313x.1999.00414.x. PMID 10230062.
  • Vines, S.H. (1876). "On the Digestive Ferment of Nepenthes". Journal of Anatomy and Physiology. 11 (1): 124–127. PMC 1309768. PMID 17231131.
  • Wong, T.-S.; Kang, S.H.; Tang, S.K.Y.; Smythe, E.J.; Hatton, B.D.; Grinthal, A.; Aizenberg, J. (2011). "Bioinspired self-repairing slippery surfaces with pressure-stable omniphobicity" (PDF). Nature. 477 (7365): 443–447. Bibcode:2011Natur.477..443W. doi:10.1038/nature10447. PMID 21938066. S2CID 4300247.

External links edit

 
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February 16, 2024
nepenthes, this, article, about, tropical, pitcher, plant, other, uses, nepenthe, disambiguation, genus, carnivorous, plants, also, known, tropical, pitcher, plants, monkey, cups, monotypic, family, nepenthaceae, genus, includes, about, species, numerous, natu. This article is about the tropical pitcher plant For other uses see Nepenthe disambiguation Nepenthes n ɪ ˈ p ɛ n 8 iː z 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 4 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 NepenthesA rosette plant of N peltata growing on Mount Hamiguitan Mindanao PhilippinesScientific classificationKingdom PlantaeClade TracheophytesClade AngiospermsClade EudicotsOrder CaryophyllalesFamily NepenthaceaeDumort 1 Genus NepenthesL SpeciesSee below or separate list Diversity 2 150 speciesSynonyms 3 Anurosperma Hallier f Bandura Adans Phyllamphora Lour Contents 1 Description 2 Taxonomy 2 1 Etymology 2 2 Evolution and phylogeny 3 Distribution and habitat 4 Ecological relationships 4 1 Symbioses 4 2 Infauna 4 3 Antimicrobial properties 5 Botanical history 6 Cultivation 7 Hybrids and cultivars 8 See also 9 References 10 Further reading 11 External linksDescription edit nbsp Nepenthes mirabilis at the Periyar Tiger Reserve in Southern Western Ghats of IndiaNepenthes 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 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 5 nbsp Basic structure of an upper pitcherThe 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 6 The trapping efficiency of this fluid remains high even when significantly diluted by water as inevitably happens in wet conditions 7 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 8 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 5 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 5 Prey usually consists of insects but the largest species e g N rajah and N rafflesiana may occasionally catch small vertebrates such as frogs birds and small mammals 9 10 Records of cultivated plants trapping small birds have been made 11 12 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 13 Nepenthes are insect pollinated the primary agents being flies including blow flies midges and mosquitoes moths wasps and butterflies 14 Their smells can range from sweet to musty or fungus like 15 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 16 17 This high number is thought to reflect paleopolyploidy likely 8x or 16x 17 18 19 20 Taxonomy editMain articles List of Nepenthes species and List of Nepenthes species by distribution About 170 species of Nepenthes are currently recognised as valid This number is increasing with several new species being described each year 21 Etymology edit The genus name Nepenthes was first published in 1737 in Carl Linnaeus s Hortus Cliffortianus 22 It references a passage in Homer s Odyssey in which the potion Nepenthes pharmakon is given to Helen by an Egyptian queen Nepenthes Ancient Greek nhpen8es literally means without grief nh ne not pen8os penthos grief and in Greek mythology is a drug that quells all sorrows with forgetfulness 15 page needed 23 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 24 The plant Linnaeus described was N distillatoria calledbandura බ ඳ ර a species from Sri Lanka 15 page needed 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 25 nbsp Nepenthes from Carolus Linnaeus s Species Plantarum of 1753The name monkey cups was discussed in the May 1964 issue of National Geographic in which Paul A Zahl wrote 26 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 in the Philippines citation needed Evolution and phylogeny edit An absence of evidence of intermediate species fossil or living i e a missing link does not allow forming a phylogenetic timeline for the development of the distinctive traits of modern Nepenthes which include its relatively rare strict dioecy and carnivorous pitchers Although Nepenthes is distantly related to several modern genera among these even the carnivorous relatives the sundews Drosera Venus flytrap Dionaea muscipula waterwheel plant Aldrovanda and dewy pine Drosophyllum all lack those traits Among known Nepenthes no protomodern characteristics or large variations are found which suggests that all extant species radiated from a single close ancestor bearing all the modern traits Phylogenetic comparisons of the chloroplast matKgene sequences between Nepenthes species and with related species support this conclusion long genetic distance between Nepenthes and others and abruptly diverging pom pom grouping of the Nepenthes species 27 Fossilized pollen of Nepenthes like plants living on the northern Tethys Sea from 65 to 35 million years ago indicates that then warmer Europe may have been where the proto Nepenthes developed and then escaped to Asia and India as Africa collided with Europe and the ensuing climate change wiped out the ancestral species in the original habitat About 20 million years ago Borneo Sumatra and Sulawesi and possibly even the Philippines were connected to mainland Asia providing a bridge for the colonization of most sites of Nepenthes species radiation The extensive landbridges in the area 20 000 years ago during the ice age would have provided access to the remaining sites of Nepenthes populations in Oceania The main complication with this hypothesis is the presence of Nepenthes on the distant islands of Seychelles and Madagascar The seeds were thought to have been transferred by seabirds and shorebirds which rest during their migrations in swampy habitats and may have inadvertently picked up the seeds This hypothesis is possibly reinforced by the success of the lowland swamp dwelling N distillatoria in colonizing so many locations 27 Distribution and habitat edit nbsp Global distribution of NepenthesFurther information List of Nepenthes species by distribution The genus Nepenthes is mostly found within the Malay Archipelago with the greatest biodiversity found on Borneo Sumatra and the Philippines 28 29 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 N parvula and N tenax and New Caledonia N vieillardii in the southeast Most species are restricted to very small ranges including some found only on individual mountains These limited distributions and the inaccessibility of the regions often means some species go decades without being rediscovered in the wild e g N deaniana which was rediscovered 100 years after its initial discovery About 10 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 5 30 31 Because of the nature of the habitats that 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 growing 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 grows at a higher altitude than any other in the genus up to 3 520 m 11 549 ft 5 31 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 in which Nepenthes species grow are usually acidic and low in nutrients being composed of peat white sand sandstone or volcanic soils Exceptions to these generalities include species that thrive in soils with 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 5 Ecological relationships edit nbsp A drowned lizard found in a freshly opened pitcher of N rajahThe most obvious interaction between Nepenthes species and their environments 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 these 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 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 5 While many Nepenthes species 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 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 32 33 Prey retention by viscoelastic pitcher fluid of N rafflesiana 7 source source source source The blue bottle fly Calliphora vomitoria can escape after landing in water on its ventral surface source source source source The same is true if the fly falls in dorsally wings first source source source source But the viscoelastic properties of N rafflesiana digestive fluid prevent prey escape whether the fall is ventral source source source source or dorsal All videos recorded at 500 frames s Symbioses edit nbsp 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 34 N 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 35 N lowii has also formed a dependent relationship but 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 between 57 and 100 of the plant s foliar nitrogen uptake comes from the faeces of tree shrews 36 Another study showed the shape and size of the pitcher orifice of N lowii exactly match the dimensions of a typical tree shrew Tupaia montana 37 38 A similar adaptation was found in N macrophylla N rajah N ampullaria and is also likely to be present in N ephippiata 38 39 Similarly N hemsleyana which is native to Borneo has a symbiotic partnership with Hardwicke s woolly bat 40 During the day a bat may roost above the digestive fluid inside the pitcher While a bat is inside it may defaecate with the plant gaining nitrogen from the droppings Further research has discovered that the shape and design of the pitcher has evolved to be an acoustic reflector to make it easier for bats to echo locate and distinguishes it from other closely related species that don t make good roosts 41 42 Infauna edit Main article Nepenthes 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 often associated with but 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 plants and infauna is not yet fully understood but the relationship may be 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 35 43 44 Antimicrobial properties edit Nepenthes digestive fluids are sterile before pitchers open and contain secondary metabolites and proteins that act as bactericides and fungicides after the pitcher opens While the digestive fluid is being produced the pitcher is not yet open so there is no chance of microbial contamination During pitcher development at least 29 digestive proteins including proteases chitinases pathogenesis related proteins and thaumatin like proteins are produced in the pitcher fluid In addition to breaking down prey these can act as antimicrobial agents 45 When the pitchers open the fluid is exposed to bacteria fungal spores insects and rain Often pitchers have a lid that covers the trap excepting a few e g N lowii N attenboroughii and N jamban preventing rain water from entering The lid inhibits rainwater from diluting the digestive fluid Once the bacteria and fungi enter the fluid secondary metabolites are produced in addition to antimicrobial proteins 46 Naphthoquinones a class of secondary metabolite are commonly produced and these either kill or inhibit the growth and reproduction of bacteria and fungi 47 This adaptation could have evolved since Nepenthes plants that could produce secondary metabolites and antimicrobial proteins to kill bacteria and fungi were most likely more fit Plants that produced antimicrobial compounds could prevent loss of valuable nutrients gained from insects within the pitcher Since Nepenthes cannot digest certain bacteria and fungi the bactericides and fungicides allow plants to maximize nutrient uptake Botanical history edit nbsp 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 48 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 15 Flacourt called the plant Amramatico after a local name More than a century later this species was formally described as N madagascariensis 49 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 50 Three years later Dutch merchant Jacob Breyne referred to this species as Bandura zingalensium after a local name for the plant 51 Bandura subsequently became the most commonly used name for the tropical pitcher plants until Linnaeus coined Nepenthes in 1737 15 Nepenthes distillatoria was again described in 1683 this time by Swedish physician and naturalist Herman Niklas Grim 52 Grim called it Planta mirabilis destillatoria or the miraculous distilling plant and was the first to clearly illustrate a tropical pitcher plant 15 Three years later in 1686 English naturalist John Ray quoted Grim as saying 53 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 15 One of the earliest illustrations of Nepenthes appears in Leonard Plukenet s Almagestum Botanicum of 1696 54 The plant called Utricaria vegetabilis zeylanensium is undoubtedly N distillatoria 15 nbsp 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 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 55 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 15 nbsp Illustration of Bandura zeylanica N distillatoria from Burmann s Thesaurus Zeylanicus of 1737Nepenthes 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 56 The next mention of tropical pitcher plants was made in 1790 when Portuguese priest Joao 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 57 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 15 Phyllamphora mirabilis was eventually transferred to the genus Nepenthes by Rafarin in 1869 58 As such P mirabilis is the basionym of this most cosmopolitan of tropical pitcher plant species 35 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 Encyclopedie Methodique Botanique he included the following account 49 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 15 nbsp The Nepenthes house of the Veitch Nurseries as illustrated in The Gardeners Chronicle 1872With 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 5 15 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 21 Cultivation edit nbsp Cultivated Nepenthes rajah Nepenthes aristolochioidesand other species nbsp Nepenthes cultivated in a garden at Kottayam KeralaNepenthes 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 59 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 60 61 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 62 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 edit nbsp The complex man made hybrid N ventricosa N lowii N macrophylla See also List of Nepenthes natural hybrids and List of Nepenthes cultivars There are many hybrid Nepenthes and numerous named cultivars Some of the more well known artificially produced hybrids and cultivars include citation needed 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 N Gaya N khasiana N ventricosa N maxima See also editNepenthes classification Nepenthes infauna List of Nepenthes endophyte speciesReferences edit Angiosperm Phylogeny Group 2009 An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants APG III PDF Botanical Journal of the Linnean Society 161 2 105 121 doi 10 1111 j 1095 8339 2009 00996 x Retrieved 2013 07 06 Cheek M Jebb M 2013 The Nepenthes micramphora Nepenthaceae group with two new species from Mindanao Philippines Phytotaxa 151 1 25 34 doi 10 11646 phytotaxa 151 1 2 Nepenthes L Plants of the World Online Royal Botanical Gardens Kew 2023 Retrieved 7 June 2023 Christenhusz M J M Byng J W 2016 The number of known plants species in the world and its annual increase Phytotaxa 261 3 201 217 doi 10 11646 phytotaxa 261 3 1 a b c d e f g h Barthlott W Porembski S Seine R and Theisen I 2007 The Curious World of Carnivorous Plants Portland Oregon Timber Press Bonhomme V 2011 Slippery or sticky Functional diversity in the trapping strategy of Nepenthes carnivorous plants New Phytologist 191 2 545 554 doi 10 1111 j 1469 8137 2011 03696 x PMID 21434933 a b Gaume L Forterre Y 2007 A viscoelastic deadly fluid in carnivorous pitcher plants PLOS ONE 2 11 e1185 Bibcode 2007PLoSO 2 1185G doi 10 1371 journal pone 0001185 PMC 2075164 PMID 18030325 Moran J A 2010 The carnivorous syndrome in Nepenthes pitcher plants Plant Signaling amp Behavior 5 6 644 648 Bibcode 2010PlSiB 5 644M doi 10 4161 psb 5 6 11238 PMC 3001552 PMID 21135573 Phillipps A 1988 A second record of rats as prey inNepenthes rajah PDF Carnivorous Plant Newsletter 17 2 55 Moran J A 1991 The role and mechanism of Nepenthes rafflesiana pitchers as insect traps in Brunei Ph D thesis University of Aberdeen Aberdeen Scotland Killer plant eats great tit at Somerset nursery BBC News 5 August 2011 Retrieved 5 August 2011 Hewitt Cooper N 2012 A case of bird capture by a cultivated specimen of the hybrid Nepenthes mixta Carnivorous Plant Newsletter 41 1 31 33 doi 10 55360 cpn411 nh396 S2CID 247074534 Clarke Charles Moran Jonathan A Chin Lijin 1 October 2010 Mutualism between tree shrews and pitcher plants Plant Signal Behav 5 10 1187 1189 Bibcode 2010PlSiB 5 1187C doi 10 4161 psb 5 10 12807 PMC 3115346 PMID 20861680 Clarke C M 2001 Nepenthes of Sumatra and Peninsular Malaysia Natural History Publications Borneo Kota Kinabalu a b c d e f g h i j k l Phillipps A Lamb A 1996 Pitcher Plants of Borneo Kota Kinabalu Sabah Malaysia Natural History Publications Borneo Lowrey T K 1991 No 519 Chromosome and isozyme number in the Nepenthaceae American Journal of Botany 78 6 supplement 200 201 a b Heubl G R Wistuba A 1997 A cytological study of the genus Nepenthes L Nepenthaceae Sendtnera 4 169 174 Meimberg H Heubl G 2006 Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae Plant Biology 8 6 831 840 Bibcode 2006PlBio 8 831M doi 10 1055 s 2006 924676 PMID 17203435 S2CID 8482890 Cytology of Nepenthes LMU Department fur Biologie Brittnacher J N d Evolution Nepenthes Phylogeny Archived 2011 05 14 at the Wayback Machine International Carnivorous Plant Society a b Clarke C M amp C C Lee 2004 Pitcher Plants of Sarawak Natural History Publications Borneo Kota Kinabalu Linnaeus C 1737 Nepenthes Hortus Cliffortianus Amsterdam Gledhill David 2008 The Names of Plants 4th ed Cambridge Cambridge University Press p 271 ISBN 978 0 521 86645 3 Veitch H J 1897 Nepenthes Journal of the Royal Horticultural Society 21 2 226 262 Linnaeus C 1753 Nepenthes Species Plantarum 2 955 Zahl P A 1964 Malaysia s Giant Flowers and Insect trapping Plants National Geographic 125 5 680 701 a b Brittnacher John 2011 Evolution Nepenthes Phylogeny ICPS Webpage Archived from the original on 2011 05 14 Retrieved 2011 02 23 Amoroso Victor B Aspiras Reyno A 2011 01 01 Hamiguitan Range A sanctuary for native flora Saudi Journal of Biological Sciences 18 1 7 15 doi 10 1016 j sjbs 2010 07 003 ISSN 1319 562X PMC 3730927 PMID 23961098 Amoroso V B Obsioma L D Arlalejo J B Aspiras R A Capili D P Polizon J J A Sumile E B 2009 Inventory and conservation of endangered endemic and economically important flora of Hamiguitan Range southern Philippines Blumea Biodiversity Evolution and Biogeography of Plants 54 71 76 doi 10 3767 000651909X474113 Retrieved 2019 11 03 McPherson S R 2009 Pitcher Plants of the Old World 2 volumes Redfern Natural History Productions Poole a b Jebb M Cheek M 1997 A skeletal revision of Nepenthes Nepenthaceae Blumea 42 1 106 Moran J A Merbach M A Livingston N J Clarke C M Booth W E 2001 Termite prey specialization in the pitcher plant Nepenthes albomarginata evidence from stable isotope analysis Annals of Botany 88 2 307 311 doi 10 1006 anbo 2001 1460 Merbach M A Merbach D J Maschwitz U Booth W E Fiala B Zizka G 2002 Mass march of termites into the deadly trap Nature 415 6867 36 37 doi 10 1038 415036a PMID 11780106 S2CID 846937 Robinson A S Fleischmann A S McPherson S R Heinrich V B Gironella E P Pena C Q 2009 A spectacular new species of Nepenthes L Nepenthaceae pitcher plant from central Palawan Philippines Botanical Journal of the Linnean Society 159 2 195 202 doi 10 1111 j 1095 8339 2008 00942 x a b c Clarke C M 1997 Nepenthes of Borneo Natural History Publications Borneo Kota Kinabalu Clarke C M Bauer U Lee C C Tuen A A Rembold K Moran J A 2009 Tree shrew lavatories a novel nitrogen sequestration strategy in a tropical pitcher plant Biology Letters 5 5 632 635 doi 10 1098 rsbl 2009 0311 PMC 2781956 PMID 19515656 Chin L Moran J A Clarke C 2010 Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size New Phytologist 186 2 461 470 doi 10 1111 j 1469 8137 2009 03166 x PMID 20100203 a b Walker M 2010 Giant meat eating plants prefer to eat tree shrew poo BBC Earth News March 10 2010 Moran J A 2003 From carnivore to detritivore Isotopic evidence for leaf litter utilization by the tropical pitcher plant Nepenthes ampullaria International Journal of Plant Sciences 164 4 635 639 doi 10 1086 375422 hdl 10170 576 S2CID 53573745 Grafe T U Schoner C R Kerth G Junaidi A amp Schoner M G 2011 A novel resource service mutualism between bats and pitcher plants Biology Letters 7 3 436 439 doi 10 1098 rsbl 2010 1141 PMC 3097880 PMID 21270023 Schoner M G Schoner C R Simon R Grafe T Ulmar Puechmaille S J Ji L L Kerth G 2015 07 09 Bats Are Acoustically Attracted to Mutualistic Carnivorous Plants Current Biology 25 14 1911 1916 doi 10 1016 j cub 2015 05 054 PMID 26166777 Thompson Ken 1 November 2018 Darwin s Most Wonderful Plants Darwin s Botany Today Profile Books p 146 ISBN 978 1 78283 436 6 Retrieved 19 November 2023 Mogi M Yong H S 1992 Aquatic arthropod communities in Nepenthes pitchers the role of niche differentiation aggregation predation and competition in community organization Oecologia 90 2 172 184 Bibcode 1992Oecol 90 172M doi 10 1007 BF00317174 PMID 28313712 S2CID 26808467 Beaver R A 1979 Fauna and foodwebs of pitcher plants in west Malaysia Malayan Nature Journal 33 1 10 Rottloff Sandy Miguel Sissi Biteau Flore Nisse Estelle Hammann Philippe Kuhn Lauriane Chicher Johana Bazile Vincent Gaume Laurence 2016 03 01 Proteome analysis of digestive fluids in Nepenthes pitchers Annals of Botany 117 3 479 495 doi 10 1093 aob mcw001 ISSN 0305 7364 PMC 4765550 PMID 26912512 Mithofer Axel 2011 09 01 Carnivorous pitcher plants Insights in an old topic Phytochemistry Plant Insect Interactions 72 13 1678 1682 Bibcode 2011PChem 72 1678M doi 10 1016 j phytochem 2010 11 024 PMID 21185041 Buch Franziska Rott Matthias Rottloff Sandy Paetz Christian Hilke Ines Raessler Michael Mithofer Axel 2012 12 21 Secreted pitfall trap fluid of carnivorous Nepenthes plants is unsuitable for microbial growth Annals of Botany 111 3 375 83 doi 10 1093 aob mcs287 ISSN 0305 7364 PMC 3579442 PMID 23264234 de Flacourt E 1658 Histoire de la Grande Isle de Madagascar a b Poiret J L M 1797 Nepente In J B Lamarck Encyclopedie Methodique Botanique Vol 4 Bartholinus Miranda herba Acta Medica et Philosophica Hafniensia 3 38 Breyne J 1680 Bandura zingalensium etc Prodromus Fasciculi Rariorum Plantarum 1 18 Grimm H N 1683 Planta mirabilis destillatoria In Miscellanea curiosa sive Ephemeridum Med Phys Germ Acad Nat Cur Decuriae 2 ann prim p 363 f 27 Ray J 1686 Bandura cingalensium etc Historia Plantarum 1 721 722 Plukenet L 1696 Utricaria vegetabilis zeylanensium In Almagestum Botanicum Rumphius G E 1741 1750 Cantharifera In Herbarium Amboinense 5 lib 7 cap 61 p 121 t 59 t 2 Burmann J 1737 Thesaurus Zeylanicus Amsterdam de Loureiro J 1790 Flora Cochinchinensis 2 606 607 Schlauer J N d Nepenthes mirabilis Carnivorous Plant Database James Pietropaolo Patricia Ann Pietropaolo 1986 Carnivorous Plants of the World Timber Press p 49 ISBN 978 0 88192 066 6 Peter D Amato 2 July 2013 The Savage Garden Revised Cultivating Carnivorous Plants Ten Speed Press p 89 ISBN 978 1 60774 411 5 Barry A Rice 2006 Growing carnivorous plants Timber Press Incorporated ISBN 978 0 88192 807 5 Appendices CITES cites org Retrieved 2022 01 14 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 editAmagase S Nakayama S Tsugita A 1969 Acid protease in Nepenthes II Study on the specificity of nepenthesin The Journal of Biochemistry 66 4 431 439 doi 10 1093 oxfordjournals jbchem a129166 PMID 5354017 Athauda S B P Matsumoto K Rajapakshe S Kuribayashi M Kojima M Kubomura Yoshida N Iwamatsu A Shibata C Inoue H Takahashi K 2004 Enzymatic and structural characterization of nepenthesin a unique member of a novel subfamily of aspartic proteinases Biochemical Journal 381 1 295 306 doi 10 1042 BJ20031575 PMC 1133788 PMID 15035659 Bauer U Bohn H F Federle W 2008 Harmless nectar source or deadly trap Nepenthes pitchers are activated by rain condensation and nectar Proceedings of the Royal Society B 275 1632 259 265 doi 10 1098 rspb 2007 1402 PMC 2593725 PMID 18048280 Beaver R A 1979 Biological studies of the fauna of pitcher plants Nepenthes in west Malaysia Annales de la Societe Entomologique de France 15 3 17 doi 10 1080 21686351 1979 12278188 S2CID 83749546 Beaver R A 1983 The communities living in Nepenthes pitcher plants fauna and food webs In J H Frank amp L P Lounibos eds Phytotelmata Plants as Hosts for Aquatic Insect Communities Plexus Publishing New Jersey pp 129 159 Beaver R A 1985 Geographical variation in food web structure in Nepenthes pitcher plants Ecological Entomology 10 3 241 248 Bibcode 1985EcoEn 10 241B doi 10 1111 j 1365 2311 1985 tb00720 x S2CID 85082186 Beekman E M 2004 A Note on the Priority of Rumphius Observation of Decapod Crustacea Living In Nepenthes Crustaceana 77 8 1019 1021 doi 10 1163 1568540042781748 Bohn H F Federle W 2004 Insect aquaplaning Nepenthes pitcher plants capture prey with the peristome a fully wettable water lubricated anisotropic surface PDF Proceedings of the National Academy of Sciences 101 39 14138 14143 doi 10 1073 pnas 0405885101 PMC 521129 PMID 15383667 in French Boulay J 1997 Les Nepenthes Dionee 38 Carlquist S 1981 Wood Anatomy of Nepenthaceae Bulletin of the Torrey Botanical Club 108 3 324 330 doi 10 2307 2484711 JSTOR 2484711 Chia T F Aung H H Osipov A N Goh N K Chia L S 2004 Carnivorous pitcher plant uses free radicals in the digestion of prey Redox Report 9 5 255 261 doi 10 1179 135100004225006029 PMID 15606978 Edwards P 2005 Growing Nepenthes Part 1 PDF Victorian Carnivorous Plant Society Inc 75 6 8 Edwards P 2005 Growing Nepenthes Part 2 PDF Victorian Carnivorous Plant Society Inc 76 6 9 Frazier C K 2000 The Enduring Controversies Concerning the Process of Protein Digestion inNepenthes Carnivorous Plant Newsletter 29 2 56 61 doi 10 55360 cpn292 cf425 S2CID 247153343 Jenkin A 2005 Nepenthes pollination PDF Victorian Carnivorous Plant Society Inc 75 12 13 Archived from the original PDF on 2011 07 07 Jennings D E Rohr JR 2011 A review of the conservation threats to carnivorous plants Biological Conservation 144 5 1356 1363 doi 10 1016 j biocon 2011 03 013 Karagatzides J D Ellison A M 2009 Construction costs payback times and the leaf economics of carnivorous plants American Journal of Botany 96 9 1612 1619 doi 10 3732 ajb 0900054 PMID 21622347 S2CID 42120981 Meimberg H Wistuba A Dittrich P Heubl G 2001 Molecular Phylogeny of Nepenthaceae Based on Cladistic Analysis of Plastid trnK Intron Sequence Data Plant Biology 3 2 164 175 Bibcode 2001PlBio 3 164M doi 10 1055 s 2001 12897 S2CID 260252804 Mithofer A 2011 Carnivorous pitcher plants insights in an old topic Phytochemistry 72 13 1678 1682 Bibcode 2011PChem 72 1678M doi 10 1016 j phytochem 2010 11 024 PMID 21185041 Moran J A Booth W E Charles J K 1999 Aspects of Pitcher Morphology and Spectral Characteristics of Six Bornean Nepenthes Pitcher Plant Species Implications for Prey Capture Annals of Botany 83 5 521 528 doi 10 1006 anbo 1999 0857 Nosonovsky M 2011 Materials science slippery when wetted Nature 477 7365 412 413 Bibcode 2011Natur 477 412N doi 10 1038 477412a PMID 21938059 S2CID 205067351 Osunkoya O O Daud S D Di Giusto B Wimmer F L Holige T M 2007 Construction Costs and Physico chemical Properties of the Assimilatory Organs of Nepenthes Species in Northern Borneo Annals of Botany 99 5 895 906 doi 10 1093 aob mcm023 PMC 2802909 PMID 17452380 Pavlovic A Masarovicova E Hudak J 2007 Carnivorous Syndrome in Asian Pitcher Plants of the Genus Nepenthes Annals of Botany 100 3 527 536 doi 10 1093 aob mcm145 PMC 2533595 PMID 17664255 Poppinga S Koch K Bohn H F Barthlott W 2010 Comparative and functional morphology of hierarchically structured anti adhesive surfaces in carnivorous plants and kettle trap flowers Functional Plant Biology 37 10 952 961 doi 10 1071 FP10061 Riedel M Eichner A Meimberg H Jetter R 2007 Chemical composition of epicuticular wax crystals on the slippery zone in pitchers of five Nepenthes species and hybrids Planta 225 6 1517 1534 Bibcode 2007Plant 225 1517R doi 10 1007 s00425 006 0437 3 PMID 17109149 S2CID 23581314 Schulze W Frommer W B Ward J M 1999 Transporters for ammonium amino acids and peptides are expressed in pitchers of the carnivorous plantNepenthes The Plant Journal 17 6 637 646 doi 10 1046 j 1365 313x 1999 00414 x PMID 10230062 Vines S H 1876 On the Digestive Ferment ofNepenthes Journal of Anatomy and Physiology 11 1 124 127 PMC 1309768 PMID 17231131 Wong T S Kang S H Tang S K Y Smythe E J Hatton B D Grinthal A Aizenberg J 2011 Bioinspired self repairing slippery surfaces with pressure stable omniphobicity PDF Nature 477 7365 443 447 Bibcode 2011Natur 477 443W doi 10 1038 nature10447 PMID 21938066 S2CID 4300247 External links edit nbsp Wikispecies has information related to Nepenthes nbsp Wikimedia Commons has media related to Nepenthes 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 Nepenthes photographs 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 Retrieved from https en wikipedia org w index php title Nepenthes amp oldid 1203182409, wikipedia, wiki, book, books, library,

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