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77-610: The term "pitcher plant" generally refers to members of the Nepenthaceae and Sarraceniaceae families, but similar pitfall traps are employed by the monotypic Cephalotaceae and some members of the Bromeliaceae . The families Nepenthaceae and Sarraceniaceae are the most species-rich families of pitcher plants. The Nepenthaceae contains a single genus , Nepenthes , containing over 100 species and numerous hybrids and cultivars. In this genus of Old World pitcher plants,

154-400: A cavity formed by the cupped leaf, often by visual lures such as anthocyanin pigments, and nectar . Many pitcher plants exhibit patterns of ultraviolet coloration which may play a role in attracting insects. Some species, such as Cephalotus follicularis , likely use camouflage to trap insects, as their coloration matches that of the surrounding environment and the plants are often embedded in

231-464: A central embryo and two wings, one on either side (though N. pervillei differs). The genus is cytologically diploid , with all studied species having a chromosome number of 2n =80. This high number is thought to reflect paleopolyploidy (likely 8x or 16x). About 170 species of Nepenthes are currently recognised as valid. This number is increasing, with several new species being described each year. The genus name Nepenthes

308-560: A copy that had fortunately been retained by Governor-General Johannes Camphuijs. The Herbarium Amboinensis finally arrived in the Netherlands in 1696. Even then, the first volume did not appear until 1741, 39 years after Rumphius's death. By this time, Linnaeus's name Nepenthes had become established. Nepenthes distillatoria was again illustrated in Johannes Burmann 's Thesaurus Zeylanicus of 1737. The drawing depicts

385-586: 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. 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

462-407: A few species (e.g. N. bicalcarata ). From the stems arise alternate, sword-shaped leaves with entire leaf margins . An extension of the midrib (the tendril ), which in some species aids in climbing, protrudes from the tip of the leaf; at the end of the tendril the pitcher forms. The pitcher starts as a small bud and gradually expands to form a globe- or tube-shaped trap. The trap contains

539-495: 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 the pitchers is especially effective in the retention of winged insects. The trapping efficiency of this fluid remains high, even when significantly diluted by water, as inevitably happens in wet conditions. The lower part of

616-528: A good half-glass. [translated from French in Pitcher-Plants of Borneo ] Flacourt called the plant Amramatico , after a local name. More than a century later, this species was formally described as N. madagascariensis . 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

693-887: A higher altitude than any other in the genus, up to 3,520 m (11,549 ft). 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

770-447: 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

847-432: 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. Naphthoquinones , a class of secondary metabolite, are commonly produced, and these either kill or inhibit

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924-401: 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

1001-458: 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. 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 ). A similar adaptation

1078-509: 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

1155-637: A sharpened straight edge and a blunt crescent shaped back. The word originates from the Latin word lunatus which means to bend like a crescent, and from luna meaning moon in Latin. A lunate object can be typically used as a decorative piece or as a stone tool. In the earlier findings of Epipaleolithic lunate in the Natufian , Harifian , and Negev Kebaran periods in Israel, they were roughly 10–40 mm long and were formed on small blades or bladelets. While

1232-555: A simple rolled-leaf pitcher, at the tip of which is a spoon-like structure that secretes nectar. They are restricted to areas of high rainfall in South America . The North American genus Sarracenia are the trumpet pitchers, which have a more complex trap than Heliamphora , with an operculum , which prevents excess accumulation of rainwater in most of the species. The single species in the California genus Darlingtonia

1309-440: A solution of amino acids , peptides , phosphates , ammonium and urea , from which the plant obtains its mineral nutrition (particularly nitrogen and phosphorus ). Like all carnivorous plants, pitcher plants all grow in locations where the soil is too poor in minerals and/or too acidic for most plants to survive. Pitcher plants supplement available nutrients and minerals (which plants normally obtain through their roots) with

1386-530: 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. Nepenthes distillatoria is the type species of the genus. The name "monkey cups" was discussed in the May 1964 issue of National Geographic , in which Paul A. Zahl wrote: The carriers called them "monkey cups,"

1463-400: Is a crescent or moon-shaped microlith . In the specialized terminology of lithic reduction , a lunate flake is a small, crescent-shaped flake removed from a stone tool during the process of pressure flaking . In the Natufian period , a lunate was a small crescent-shaped stone tool that was sometimes used to harvest grasses. In archaeology a lunate is a small stone artifact, that has

1540-731: Is a monotypic family with but one genus and species, Cephalotus follicularis . This species has a small (2–5 cm) pitcher similar in form to those of Nepenthes . Unlike in Nepenthes , in Cephalotus follicularis the petiole is attached to the rear of the upper trap rim rather than to the base of the pitcher. The species occurs in only one location in southwestern Australia . A few species of bromeliads ( Bromeliaceae ), such as Brocchinia reducta and Catopsis berteroniana , are known or suspected to be carnivorous. Foraging, flying, or crawling insects such as flies are attracted to

1617-410: Is associated with increased insect retention to help capture flying insects such as flies, whereas increased fluid acidity can decrease insect killing-time, which can help capture crawling insects such as ants. Some pitcher plants contain mutualistic insect larvae , which feed on trapped prey, and whose excreta the plant absorbs. Whatever the mechanism of digestion, the prey items are converted into

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1694-427: 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

1771-635: 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 ,

1848-464: 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 ] The plant Linnaeus described was N. distillatoria , called bāndurā (බාඳුරා),

1925-445: 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. When the pitchers open, the fluid is exposed to bacteria, fungal spores, insects and rain. Often pitchers have

2002-460: Is popularly known as the cobra plant, due to its possession of an inflated "lid" with elegant false-exits, and a forked "tongue", which serves to ferry ants and other prey to the entrance of the pitcher. The species in the genus Sarracenia readily hybridize, making their classification a complex matter. The purple pitcher plant, Sarracenia purpurea , is the floral emblem of the province of Newfoundland and Labrador , Canada. The Cephalotaceae

2079-624: Is possible, but most plants will get too large over time. Plants can be propagated by seed, cuttings, and tissue culture . Seeds are usually sown on damp chopped Sphagnum moss , or on sterile plant tissue culture media once they have been properly disinfected. The seeds generally become nonviable soon after harvesting, so seed are not usually the preferred method of propagation. A 1:1 mixture of orchid medium with moss or perlite has been used for germination and culture. Seed may take two months to germinate, and two years or more to yield mature plants. Cuttings may be rooted in damp Sphagnum moss in

2156-611: 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

2233-490: 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. The most obvious interaction between Nepenthes species and their environments, including other organisms, is that of predator and prey . Nepenthes species attract their prey through active production of attractive colours, sugary nectar , and sweet scents. From this relationship,

2310-496: The "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant. Three years later, in 1686, English naturalist John Ray quoted Grim as saying: The root draws up moisture from the earth which with the help of the sun's rays rises up into the plant itself and then flows down through the stems and nerves of the leaves into the natural utensil to be stored there until used for human needs. [translated from Latin in Pitcher-Plants of Borneo ] One of

2387-725: The 1960s and 1970s did much to bring attention to these plants. Nepenthes may be cultivated in greenhouses. Easier species include N. alata , N. ventricosa , N. khasiana , and N. sanguinea . These four species are highlanders ( N. alata has both lowland and highland forms), some easy lowlander species are N. rafflesiana , N. bicalcarata , N. mirabilis , and N. hirsuta . Highland forms are those species that grow in habitats generally higher in elevation, and thus exposed to cooler evening temperatures. Lowland forms are those species growing nearer to sea level. Both forms respond best to rainwater (but some tap water works as long as it

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2464-451: The 19th century, culminating in what has been called the "Golden Age of Nepenthes " in the 1880s. 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

2541-626: The Azor tombs in Israel and was far south as south Sinai in this particular region. Lunate artefacts have been discovered among early Māori stone carving in New Zealand . The original lunate pendant found in New Zealand appears to be of clear transparent pounamu (greenstone), from Ruapuke Island , in Foveaux Strait . Its characteristics include a notched edge and the stone itself is thought to originate from Tangiwai, New Zealand. There

2618-617: The Malay Archipelago to Australia. 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

2695-479: 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

2772-491: 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

2849-430: The bactericides and fungicides allow plants to maximize nutrient uptake. 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: It is a plant growing about 3 feet high which carries at the end of its leaves, which are 7 inches long,

2926-509: 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

3003-563: The bats. It is widely assumed pitfall traps evolved by epiascidiation (infolding of the leaf with the adaxial or upper surface becoming the inside of the pitcher), with selection pressure favouring more deeply cupped leaves over evolutionary time. The pitcher trap evolved independently in three eudicot lineages and one monocot lineage, representing a case of convergent evolution . Some pitcher plant families (such as Nepenthaceae) are placed within clades consisting mostly of flypaper traps , indicating that some pitchers may have evolved from

3080-478: The common ancestors of today's flypaper traps by loss of mucilage. Nepenthaceae See below or separate list . Nepenthes ( / n ɪ ˈ p ɛ n θ iː z / nih- PEN -theez ) is a genus of carnivorous plants , also known as tropical pitcher plants , or monkey cups , in the monotypic family Nepenthaceae . The genus includes about 170 species , and numerous natural and many cultivated hybrids. They are mostly liana -forming plants of

3157-449: The constituents of their insect prey. Mature plants of Nepenthes lowii attract tree shrews ( Tupaia montana ), which feed on nectar that the plant produces but also defecate into the pitcher, providing nitrates and other nutrients. The plant and tree shrew have a symbiotic relationship . The rim of N. lowii is not slippery so that tree shrews can easily get in and out; it provides more nectar than other pitcher plants. The shape of

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3234-486: 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 ] With the discovery of new species and Sir Joseph Banks' original introduction of specimens to Europe in 1789, interest in Nepenthes grew throughout

3311-490: The dew and store it. A marvellous work of the Lord! [translated from French in Pitcher-Plants of Borneo ] Phyllamphora mirabilis was eventually transferred to the genus Nepenthes by Rafarin in 1869. As such, P. mirabilis is the basionym of this most cosmopolitan of tropical pitcher plant species. Loureiro's description of a moving lid was repeated by Jean Louis Marie Poiret in 1797. Poiret described two of

3388-443: 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. 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

3465-571: The earliest illustrations of Nepenthes appears in Leonard Plukenet's Almagestum Botanicum of 1696. The plant, called Utricaria vegetabilis zeylanensium , is undoubtedly N. distillatoria . Around the same time, German botanist Georg Eberhard Rumphius discovered two new Nepenthes species in the Malay Archipelago . Rumphius illustrated the first one, now considered synonymous with N. mirabilis , and gave it

3542-431: The end of a flowering stem with pitchers. Burmann refers to the plant as Bandura zeylanica . The next mention of tropical pitcher plants was made in 1790, when Portuguese priest João de Loureiro described Phyllamphora mirabilis , or the "marvellous urn-shaped leaf", from Vietnam . Despite living in the country for around 35 years, it seems unlikely that Loureiro observed living plants of this species, as he stated

3619-511: The end of the 4th millennium B.C. These Lunate tools were most likely used as barbs in arrow shafts, or as transverse arrowheads coated with poison. The Lunate are also a very rare artifact from the Early Bronze Age because there was not as much emphasis on hunting during that period. The reappearance of Lunate after several millennia could shed some light on the hunting emphasis in the society. Lunate have been found as far north as

3696-613: The exception of N. rajah and N. khasiana which are listed in CITES Appendix I. The CITES listing means all international trade (including in parts and derivatives) is controlled by the CITES permitting system, with wild sourced specimens of Appendix I species prohibited from commercial international trade. There are many hybrid Nepenthes and numerous named cultivars . Some of the more well-known, artificially produced hybrids and cultivars include: lunate Lunate

3773-481: The four Nepenthes species known at the time: N. madagascariensis and N. distillatoria . He gave the former its current name and called the latter Nepente de l'Inde , or simply " Nepenthes of India", although this species is absent from the mainland. In Jean-Baptiste Lamarck 's Encyclopédie Méthodique Botanique , he included the following account: This urn is hollow, as I have just said, usually full of soft, clear water, and then closed. It opens during

3850-481: The ground in forest clearings, or as epiphytes on trees. The New World pitcher plants (Sarraceniaceae), which comprise three genera, are ground-dwelling herbs whose pitchers arise from a horizontal rhizome . In this family, the entire leaf forms the pitcher, as opposed to Nepenthaceae, where the pitcher arises from the terminal portion of the leaf. The species of the genus Heliamphora , which are popularly known as marsh pitchers (or erroneously as sun pitchers), have

3927-419: The growth and reproduction of bacteria and fungi. 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,

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4004-446: 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

4081-452: The later findings Natufian and Harifian range of lengths varied then between 9 mm and 17 mm. In the later period, the lunate resulted in three specific types: The differences among these three types are also associated with the length of the lunate objects, with Helwan lunate normally being the longest and bipolar being the shortest. For unknown reasons, the epipaleolithic lunate tool type disappeared and did not reappear until around

4158-406: 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

4235-555: 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. 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

4312-463: 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 . 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

4389-406: The name Cantharifera , meaning "tankard-bearer". The second, referred to as Cantharifera alba , is thought to have been N. maxima . Rumphius described the plants in his most famous work, the six-volume Herbarium Amboinense , a catalogue of the flora of Ambon Island . However, it would not be published until many years after his death. After going blind in 1670, when the manuscript

4466-503: The name Miranda herba , Latin for "marvellous herb". Three years later, Dutch merchant Jacob Breyne referred to this species as Bandura zingalensium , after a local name for the plant. Bandura subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined Nepenthes in 1737. Nepenthes distillatoria was again described in 1683, this time by Swedish physician and naturalist Herman Niklas Grim . Grim called it Planta mirabilis destillatoria or

4543-435: The nectar produced by the plant and defecate into the pitchers, providing valuable nutrients. Nepenthes are insect-pollinated , the primary agents being flies (including blow flies , midges , and mosquitoes ), moths, wasps, and butterflies. Their smells can range from sweet to musty or fungus -like. Seed is typically produced in a four-sided capsule which may contain 50–500 wind-distributed seeds, consisting of

4620-409: 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

4697-561: The peristome is further enhanced in moist environments, where condensation may cause a thin water film to form on the surface of the peristome. When wet, the slippery surface of the peristome causes insects to ‘aquaplane’, or slip and fall, into the pitcher. Above the peristome is a lid (the operculum ); in many species, this keeps rain from diluting the fluid within the pitcher, the underside of which may contain nectar glands which attract prey. Nepenthes species usually produce two types of pitchers, known as leaf dimorphism. Appearing near

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4774-741: The pitcher rim and the position of the nectar ensure that the animal's hindquarters are over the rim while it feeds. Nepenthes rafflesiana var. elongata has a similar relationship with Hardwicke's woolly bats ( Kerivoula hardwickii ). The bats roost inside the pitchers and the plants derive much of their foliar nitrogen from the feces of the bats. Compared to other varieties of Nepenthes rafflesiana that do not exhibit this form of mutualism, N. rafflesiana var. elongata has elongated pitchers that can accommodate both single bats and mother-juvenile pairs. As well as its elongated shape, N. rafflesiana var. elongata has reduced volumes of pitcher fluid compared to other species, leaving more space to accommodate

4851-417: The pitcher traps are called phytotelmata . They drown the insect, whose body is gradually dissolved. This may occur by bacterial action (the bacteria being washed into the pitcher by rainfall), or by digestive enzymes secreted by the plant itself. Pitcher trap fluids largely vary in their viscoelasticity and acidity, which then dictates which type of prey they can target. For example, increased viscoelasticity

4928-430: The pitchers are borne at the end of tendrils that extend from the midrib of an otherwise unexceptional leaf. Old World pitcher plants are typically characterized as having reduced and symmetrical pitchers with a comprehensive waxy coating on the surface of the inner pitcher wall. The plants themselves are often climbers, accessing the canopy of their habitats using the aforementioned tendrils, although others are found on

5005-426: 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. 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

5082-406: The plant that harbours the infauna receives expedited breakdown of captured prey, increasing the rate of digestion and keeping harmful bacterial populations repressed. 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

5159-917: 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. 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

5236-436: 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

5313-457: 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. 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

5390-517: The seeds. This hypothesis is possibly reinforced by the success of the lowland swamp-dwelling N. distillatoria in colonizing so many locations. The genus Nepenthes is mostly found within the Malay Archipelago , with the greatest biodiversity found on Borneo, Sumatra, and the Philippines, especially in the Borneo montane rain forests . The full range of the genus includes Madagascar ( N. madagascariensis and N. masoalensis ),

5467-586: The substrate such that the traps are flush with the ground. Olfactory cues can also play a role in attraction. For example, Nepenthes rafflesian a uses flower-scent mimicry to attract insects to its pitchers. The rim of the pitcher (peristome) is slippery when moistened by condensation or nectar, causing insects to fall into the trap. The walls of the pitfall may be covered with waxy scales, protruding aldehyde crystals, cuticular folds, downward-pointing hairs, or guard-cell-originating lunate cells, to help prevent escape. The small bodies of liquid contained within

5544-479: 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

5621-402: 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

5698-559: The two types of pitchers. This varied morphology also often makes identification of species difficult. Prey usually consists of insects , but the largest species (e.g. N. rajah and N. rafflesiana ) may occasionally catch small vertebrates , such as "frogs, birds, and small mammals". Records of cultivated plants trapping small birds have been made. Flowers occur in racemes or more rarely in panicles with male and female flowers on separate plants. Three species have symbiotic relationships with treeshrews , which eat

5775-621: Was first published in 1737 in Carl Linnaeus 's Hortus Cliffortianus . It references a passage in Homer 's Odyssey , in which the potion "Nepenthes pharmakon" is given to Helen by an Egyptian queen. " 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. Linnaeus explained: If this

5852-424: Was found in N. macrophylla , N. rajah , N. ampullaria , and is also likely to be present in N. ephippiata . Similarly, N. hemsleyana , which is native to Borneo , has a symbiotic partnership with Hardwicke's woolly bat . 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

5929-490: 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

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