Description
''Nepenthes'' species usually consist of a shallow root system and aTaxonomy
About 170 species of ''Nepenthes'' are currently recognised as valid. This number is increasing, with several new species being described each year.Etymology
The genus name ''Nepenthes'' was first published in 1737 inIf this is not Helen's ''Nepenthes'', it certainly will be for all botanists. What botanist would not be filled with admiration if, after a long journey, he should find this wonderful plant. In his astonishment past ills would be forgotten when beholding this admirable work of the Creator! ranslated from Latin by Harry Veitch">Harry_Veitch.html" ;"title="ranslated from Latin by Harry Veitch">ranslated from Latin by Harry Veitch/blockquote> The plant Linnaeus described was ''N. distillatoria'', called'' bāndurā'' (බාඳුරා), a species from Sri Lanka.Phillipps, A. & A. Lamb 1996. ''Pitcher-Plants of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. ''Nepenthes'' was formally published as a genus, generic name in 1753 in Linnaeus's famous ''Species Plantarum'', which establishedbotanical nomenclature Botanical nomenclature is the formal, scientific naming of plants. It is related to, but distinct from Alpha taxonomy, taxonomy. Plant taxonomy is concerned with grouping and classifying plants; botanical nomenclature then provides names for the ...as it exists today. ''Nepenthes distillatoria'' is thetype species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...of the genus. The name "monkey cups" was discussed in the May 1964 issue of ''National Geographic ''National Geographic'' (formerly the ''National Geographic Magazine'', sometimes branded as NAT GEO) is a popular American monthly magazine published by National Geographic Partners. Known for its photojournalism, it is one of the most widely ...'', in which Paul A. Zahl wrote:The carriers called them "monkey cups," a name I had heard elsewhere in reference to ''Nepenthes'', but the implication that monkeys drink the pitcher fluid seemed farfetched. I later proved it true. In Sarawak, I found anThe plants are often called ''kantong semar'' (orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...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.Semar Semar is a character in Javanese mythology who frequently appears in wayang shadow plays. He is one of the punokawan (clowns), but is in fact divine and very wise. He is the dhanyang (guardian spirit) of Java,Geertz, 23. and is regarded by some ...'s pocket) in Indonesia and ''sako ni Hudas'' ( Judas' money bag) in the Philippines.
Evolution and phylogeny
An absence of evidence of intermediate species, fossil or living (i.e. a missing link), does not allow forming aphylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...al timeline for the development of the distinctive traits of modern ''Nepenthes'', which include its relatively rare strictdioecy Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...and carnivorous pitchers. Although ''Nepenthes'' is distantly related to several modern genera, among these, even the carnivorous relatives he sundews (''Drosera''), Venus flytrap (''Dionea muscipula">Drosera.html" ;"title="he sundews (''Drosera">he sundews (''Drosera''), Venus flytrap (''Dionea 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 In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...comparisons of thechloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...'' 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 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 Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ...,Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ..., andSulawesi Sulawesi (), also known as Celebes (), is an island in Indonesia. One of the four Greater Sunda Islands, and the world's eleventh-largest island, it is situated east of Borneo, west of the Maluku Islands, and south of Mindanao and the Sulu Ar ...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 inOceania Oceania (, , ) is a region, geographical region that includes Australasia, Melanesia, Micronesia, and Polynesia. Spanning the Eastern Hemisphere, Eastern and Western Hemisphere, Western hemispheres, Oceania is estimated to have a land area of .... The main complication with this hypothesis is the presence of ''Nepenthes'' on the distant islands ofSeychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, V ...andMadagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa .... The seeds were thought to have been transferred by seabirds andshorebirds 245px, A flock of Dunlins and Red knots">Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflat ..., 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.
Distribution and habitat
The genus ''Nepenthes'' is mostly found within theMalay Archipelago The Malay Archipelago (Indonesian/Malay: , tgl, Kapuluang Malay) is the archipelago between mainland Indochina and Australia. It has also been called the " Malay world," "Nusantara", "East Indies", Indo-Australian Archipelago, Spices Archipe ..., with the greatest biodiversity found on Borneo, Sumatra, and the Philippines, especially in theBorneo montane rain forests The Borneo montane rain forests are an ecoregion, of cloud forest, within the tropical and subtropical moist broadleaf forests biome, of the island of Borneo in Southeast Asia. Location and description This ecoregion consists of tropical moun .... 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 fromIndochina Mainland Southeast Asia, also known as the Indochinese Peninsula or Indochina, is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to the west an ...and throughout the Malay Archipelago to Australia.McPherson, S.R. 2009. ''Pitcher Plants of the Old World ''Pitcher Plants of the Old World'' is a two-volume monograph by Stewart McPherson on the pitcher plants of the genera ''Nepenthes'' and ''Cephalotus''. It was published in May 2009 by Redfern Natural History Productions and covers all species kno ...''. 2 volumes. Redfern Natural History Productions, Poole. Because of the nature of the habitats that ''Nepenthes'' species occupy, they are often graded as either lowland or highland species, depending on theiraltitude Altitude or height (also sometimes known as depth) is a distance measurement, usually in the vertical or "up" direction, between a reference datum and a point or object. The exact definition and reference datum varies according to the context ...abovesea level Mean sea level (MSL, often shortened to sea level) is an average surface level of one or more among Earth's coastal bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical datuma standardised g ..., with 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 ''Nepenthes lamii'' is a tropical pitcher plant endemism, endemic to New Guinea, where it grows at an altitude of up to 3520 m above sea level, higher than any other ''Nepenthes'' species.McPherson, S.R. 2009. ''Pitcher Plants of the Old W ...'' grows at a higher altitude than any other in the genus, up to . Most ''Nepenthes'' species grow in environments that provide highhumidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation, dew, or fog to be present. Humidity depe ...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 inclear-cut Clearcutting, clearfelling or clearcut logging is a forestry/ logging practice in which most or all trees in an area are uniformly cut down. Along with shelterwood and seed tree harvests, it is used by foresters to create certain types of fore ...forest areas, roadsides, and disturbed fields. Other species have adapted to growing insavanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...-like grass communities. The soils in which ''Nepenthes'' species grow are usually acidic and low in nutrients, being composed ofpeat Peat (), also known as turf (), is an accumulation of partially decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, moors, or muskegs. The peatland ecosystem covers and is the most efficien ..., white sand,sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ..., 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 aslithophyte Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are also ...s, while others, such as '' N. inermis'', can grow asepiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...s with no soil contact.
Ecological relationships
The most obvious interaction between ''Nepenthes'' species and their environments, including other organisms, is that ofpredator and prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th .... ''Nepenthes'' species certainly attract and kill their prey, albeit passively, through active production of attractive colours, sugarynectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ..., and even sweet scents. From this relationship, the plants primarily gainnitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...andphosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...to supplement their nutrient requirements for growth, given these soil nutrients are typically lacking. The most frequent prey is an abundant and diverse group ofarthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...s, with ants and otherinsect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...s topping the menu. Other arthropods found frequently includespider Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...s,scorpion Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always end ...s, andcentipede Centipedes (from New Latin , "hundred", and Latin , " foot") are predatory arthropods belonging to the class Chilopoda (Ancient Greek , ''kheilos'', lip, and New Latin suffix , "foot", describing the forcipules) of the subphylum Myriapoda, an ...s, whilesnail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gastro ...s andfrog A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...s 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 trapstermite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...s and produces nearly no nectar. ''Nepenthes albomarginata'' gains its name from the ring of whitetrichome Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a pla ...s 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.
Symbioses
'' N. bicalcarata'' provides space in the hollow tendrils of its upper pitchers for the carpenter ant ''Camponotus schmitzi ''Colobopsis schmitzi'', synonym ''Camponotus schmitzi'', is a species of ant native to Borneo, Schuitemaker, J.P. & A. Stärcke 1933. ''Overdruk uit het Natuurhistorisch Maandblad'' 22(3): 29–31. which is commonly known as the diving ant, swi ...'' to build nests. The ants take larger prey from the pitchers, which may benefit ''N. bicalcarata'' by reducing the amount ofputrefaction Putrefaction is the fifth stage of death, following pallor mortis, algor mortis, rigor mortis, and livor mortis. This process references the breaking down of a body of an animal, such as a human, post-mortem. In broad terms, it can be viewed ...of collected organic matter that could harm the natural community ofinfaunal Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is ''funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. Zool ...species that aid the plant's digestion.Clarke, C.M. 1997. '' Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. '' 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. 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 The mountain treeshrew (''Tupaia montana'') is a treeshrew species within the Tupaiidae. It is endemic to Borneo and inhabits montane forests in Sarawak and Sabah. The first specimen was described by Oldfield Thomas and was part of a zoological ...'').Walker, M. 2010
Giant meat-eating plants prefer to eat tree shrew poo
''BBC Earth News'', March 10, 2010. A similar adaptation 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 toBorneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and eas ..., has a symbiotic partnership withHardwicke's woolly bat Hardwicke's woolly bat (''Kerivoula hardwickii'') is a species of vesper bat in the family Vespertilionidae. Geographic range It is found in Bangladesh, China, India (Assam, Jammu and Kashmir, Karnataka, Meghalaya, Mizoram, Nagaland and West .... During the day, a bat may roost above the digestive fluid inside the pitcher. While a bat is inside, it may defaecate, and the plant can get nitrogen from the droppings.
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 toptrophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...of the infaunal ecosystem, are many species of mosquito larvae. Other infaunal species includefly Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwing ...andmidge A midge is any small fly, including species in several families of non-mosquito Nematoceran Diptera. Midges are found (seasonally or otherwise) on practically every land area outside permanently arid deserts and the frigid zones. Some mid ...larvae, spiders,mite Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...s, ants, and even a species of crab (''Geosesarma malayanum ''Geosesarma malayanum'' is a species of small red crab found in Malaysia. It is famous for its relationship with pitcher plants; as such, it is classified as a nepenthephile. ''G. malayanum'' is known to visit ''Nepenthes ampullaria'' plants a ...''). 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.
Antimicrobial properties
''Nepenthes'' digestive fluids are sterile before pitchers open and contain secondary metabolites and proteins that act asbactericide A bactericide or bacteriocide, sometimes abbreviated Bcidal, is a substance which kills bacteria. Bactericides are disinfectants, antiseptics, or antibiotics. However, material surfaces can also have bactericidal properties based solely on their ...s 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 includingprotease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalyzes (increases reaction rate or "speeds up") proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the ...s, chitinases, pathogenesis-related proteins andthaumatin Thaumatin (also known as talin) is a low-calorie sweetener and flavor modifier. The protein is often used primarily for its flavor-modifying properties and not exclusively as a sweetener. The thaumatins were first found as a mixture of protein ...-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 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 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, the bactericides and fungicides allow plants to maximize nutrient uptake.
Botanical history
The earliest known record of ''Nepenthes'' dates back to the 17th century. In 1658, French colonial governorÉtienne de Flacourt Étienne de Flacourt (1607–1660) was a French governor of Madagascar, born in Orléans in 1607. He was named governor of Madagascar by the French East India Company in 1648. Flacourt restored order among the French soldiers, who had mutinied ...published a description of a pitcher plant in his seminal work ''Histoire de la Grande Isle de Madagascar''. It reads:It is a plant growing about 3 feet high which carries at the end of its leaves, which are 7 inches long, a hollow flower or fruit resembling a small vase, with its own lid, a wonderful sight. There are red ones and yellow ones, the yellow being the biggest. The inhabitants of this country are reluctant to pick the flowers, saying that if somebody does pick them in passing, it 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. ranslated from French in ''Pitcher-Plants of Borneo''">Pitcher-Plants_of_Borneo.html" ;"title="ranslated from French in ''Pitcher-Plants of Borneo">ranslated from French in ''Pitcher-Plants of Borneo''ref name=P&L />Flacourt called the plant ''Amramatico'', after a local name. More than a century later, this species was Species description">formally described as '' N. madagascariensis''.Poiret, J.L.M. 1797. ''Népente''. In: J.B. Lamarck ''Encyclopédie Méthodique Botanique'' Vol. 4. The second species to be described was '' N. distillatoria'', theSri Lanka Sri Lanka (, ; si, ශ්රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...nendemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew .... In 1677, Danish physician Thomas Bartholin made brief mention of it under the name ''Miranda herba'', Latin for "marvellous herb". Three years later, Dutch merchant Jacob Breyne referred to this species as ''Bandura zingalensium'', after a local name for the plant. ''Bandura'' subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined ''Nepenthes'' in 1737. ''Nepenthes distillatoria'' was again described in 1683, this time by Swedish physician and naturalistHerman Niklas Grim Herman may refer to: People * Herman (name), list of people with this name * Saint Herman (disambiguation) * Peter Noone (born 1947), known by the mononym Herman Places in the United States * Herman, Arkansas * Herman, Michigan * Herman, Minneso .... Grim called it ''Planta mirabilis destillatoria'' or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant. Three years later, in 1686, English naturalistJohn Ray John Ray FRS (29 November 1627 – 17 January 1705) was a Christian English naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after ...quoted Grim as saying:The root draws up moisture from the earth which with the help of the sun's rays rises up into the plant itself and then flows down through the stems and nerves of the leaves into the natural utensil to be stored there until used for human needs. ranslated from Latin in ''Pitcher-Plants of Borneo''">Pitcher-Plants_of_Borneo.html" ;"title="ranslated from Latin in ''Pitcher-Plants of Borneo">ranslated from Latin in ''Pitcher-Plants of Borneo''ref name=P&L />One of the earliest illustrations of ''Nepenthes'' appears in Leonard Plukenet's ''Almagestum Botanicum'' of 1696. The plant, called ''Utricaria vegetabilis zeylanensium'', is undoubtedly ''N. distillatoria''. Around the same time, German botanist Georg Eberhard Rumphius discovered two new ''Nepenthes'' species in theMalay Archipelago The Malay Archipelago (Indonesian/Malay: , tgl, Kapuluang Malay) is the archipelago between mainland Indochina and Australia. It has also been called the " Malay world," "Nusantara", "East Indies", Indo-Australian Archipelago, Spices Archipe .... 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 theflora Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''. E ...ofAmbon Island Ambon Island is part of the Maluku Islands of Indonesia. The island has an area of and is mountainous, well watered, and fertile. Ambon Island consists of two territories: the city of Ambon, Maluku, Ambon to the south and various districts ('' .... However, it would not be published until many years after his death. After going blind in 1670, when the manuscript was only partially complete, Rumphius continued work on ''Herbarium Amboinensis'' with the help of clerks and artists. In 1687, with the project nearing completion, at least half of the illustrations were lost in a fire. Persevering, Rumphius and his helpers first completed the book in 1690. However, two years later, the ship carrying the manuscript to the Netherlands was attacked and sunk by the French, forcing them to start over from a copy that had fortunately been retained by Governor-General Johannes Camphuijs. The ''Herbarium Amboinensis'' finally arrived in the Netherlands in 1696. Even then, the first volume did not appear until 1741, 39 years after Rumphius's death. By this time, Linnaeus's name ''Nepenthes'' had become established. ''Nepenthes distillatoria'' was again illustrated inJohannes Burmann Johannes is a Medieval Latin form of the personal name that usually appears as " John" in English language contexts. It is a variant of the Greek and Classical Latin variants (Ιωάννης, '' Ioannes''), itself derived from the Hebrew name '' Y ...'s ''Thesaurus Zeylanicus'' of 1737. The drawing depicts the end of a flowering stem with pitchers. Burmann refers to the plant as ''Bandura zeylanica''. The next mention of tropical pitcher plants was made in 1790, when Portuguese priest João de Loureiro described ''Phyllamphora mirabilis'', or the "marvellous urn-shaped leaf", fromVietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i .... Despite living in the country for around 35 years, it seems unlikely that Loureiro observed living plants of this species, as he stated the lid is a moving part, actively opening and closing. In his most celebrated work, ''Flora Cochinchinensis'', he writes:..(the) leaf-tip ends in a long hanging tendril, twisted spirally in the middle, from which hangs a sort of vase, oblong, pot-bellied, with a smooth lip with a projecting margin and a lid affixed to one side, which of its own nature freely opens and closes in order to receive the dew and store it. A marvellous work of the Lord! ranslated from French in ''Pitcher-Plants of Borneo''">Pitcher-Plants_of_Borneo.html" ;"title="ranslated from French in ''Pitcher-Plants of Borneo">ranslated from French in ''Pitcher-Plants of Borneo''ref name=P&L />''Phyllamphora mirabilis'' was eventually transferred to the genus ''Nepenthes'' by Rafarin in 1869. As such, ''P. mirabilis'' is the basionym of this most cosmopolitan of tropical pitcher plant species. Loureiro's description of a moving lid was repeated by Jean Louis Marie Poiret in 1797. Poiret described two of the four ''Nepenthes'' species known at the time: ''N. madagascariensis'' and ''N. distillatoria''. He gave the former its current name and called the latter ''Nepente de l'Inde'', or simply "''Nepenthes'' of India", although this species is absent from the mainland. InJean-Baptiste Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biologi ...'s ''Encyclopédie Méthodique Botanique'', he included the following account:This urn is hollow, as I have just said, usually full of soft, clear water, and then closed. It opens during the day and more than half the liquid disappears, but this loss is repaired during the night, and the next day the urn is full again and closed by its lid. This is its sustenance, and enough for more than one day because it is always about half-full at the approach of night. ranslated from French in ''Pitcher-Plants of Borneo''">Pitcher-Plants_of_Borneo.html" ;"title="ranslated from French in ''Pitcher-Plants of Borneo">ranslated from French in ''Pitcher-Plants of Borneo''ref name=P&L />With the discovery of new species and Joseph Banks">Sir Joseph Banks' original introduction of specimens to Europe in 1789, interest in ''Nepenthes'' grew throughout the 19th century, culminating in what has been called the "Golden Age of ''Nepenthes''" in the 1880s.Barthlott, W., Porembski, S., Seine, R., and Theisen, I. 2007. ''The Curious World of Carnivorous Plants.'' Portland, Oregon: Timber Press. However, the popularity of the plants dwindled in the early 20th century, before all but disappearing by World War II. This is evidenced by the fact that no new species were described between 1940 and 1966. The revival of global interest in the cultivation and study of ''Nepenthes'' is credited to Japanese botanistShigeo Kurata is a Japanese botanist and '' Nepenthes'' taxonomist whose work in the 1960s and 1970s contributed much to the current popularity of these plants.Clarke, C.M. & C.C. Lee 2004. ''Pitcher Plants of Sarawak''. Natural History Publications (Borneo), ..., whose work in the 1960s and 1970s did much to bring attention to these plants.Clarke, C.M. & C.C. Lee 2004. ''Pitcher Plants of Sarawak This list of ''Nepenthes'' literature is a listing of major published works dealing with the tropical pitcher plants of the genus ''Nepenthes''. It includes specialised standalone publications and taxonomic monographs released as part of larger ...''. Natural History Publications (Borneo), Kota Kinabalu.
Cultivation
''Nepenthes'' may be cultivated in greenhouses. Easier species include '' N. alata'', '' N. ventricosa'', '' N. khasiana'', and '' N. sanguinea''. These four species are highlanders (''N. alata'' has both lowland and highland forms), some easy lowlander species are '' N. rafflesiana'', '' N. bicalcarata'', '' N. mirabilis'', and '' N. hirsuta''. Highland forms are those species that grow in habitats generally higher in elevation, and thus exposed to cooler evening temperatures. Lowland forms are those species growing nearer to sea level. Both forms respond best to rainwater (but some tap water works as long as it is flushed monthly with rainwater or water low in dissolved solid and chemicals), bright light (though some species can grow in full sun), a well-drained medium, good air circulation and relatively high humidity, although easier species such as ''N. alata'' can adapt to lower humidity environments. Highland species must have night-time cooling to thrive in the long term. Chemical fertilisers are best used at low strength. Occasional feeding with frozen (thawed before use)cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by striki ...s may be beneficial. Terrarium culture of smaller plants, such as '' N. bellii'', ''N.'' × ''trichocarpa'' and '' N. ampullaria'', is possible, but most plants will get too large over time. Plants can be propagated by seed, cuttings, andtissue culture Tissue culture is the growth of tissues or cells in an artificial medium separate from the parent organism. This technique is also called micropropagation. This is typically facilitated via use of a liquid, semi-solid, or solid growth medium, su .... Seeds are usually sown on damp chopped '' Sphagnum''moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ..., 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 Tissue culture is the growth of tissues or cells in an artificial medium separate from the parent organism. This technique is also called micropropagation. This is typically facilitated via use of a liquid, semi-solid, or solid growth medium, su ...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 inCITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...Appendix II, with the exception of '' N. rajah'' and '' N. khasiana'' which are listed in CITES Appendix I. The CITES listing means all international trade (including in parts and derivatives) is controlled by the CITES permitting system, with wild sourced specimens of Appendix I species prohibited from commercial international trade.
Hybrids and cultivars
There are manyhybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...''Nepenthes'' and numerous namedcultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...s. Some of the more well-known, artificially produced hybrids and cultivars include: *''N.'' × ''coccinea'' ((''N. rafflesiana'' × ''N. ampullaria'') × ''N. mirabilis'') *''N.'' × ''ventrata'' (''N. ventricosa'' × ''N. alata'') *''N.'' × 'Bloody Mary' (''N. ventricosa'' × ''N. ampullaria'') *''N.'' 'D'amato' (''N. lowii'' × ''N. ventricosa'') *''N.'' × ''mixta'' (''N. northiana'' × ''N. maxima'') *''N.'' 'Syurga' (''N. ventricosa'' × ''N. northiana'') *''N.'' 'Menarik' (''N. rafflesiana'' × ''N. veitchii'') *''N.'' 'Emmarene' (''N. khasiana'' × ''N. ventricosa'') *''N.'' 'Judith Finn' (''N. spathulata'' × ''N. veitchii'')
See also
* ''Nepenthes'' classification * ''Nepenthes'' infauna * List of ''Nepenthes'' endophyte species
References
Danser's Monograph on ''Nepenthes''
(covers species from Malaysia, Indonesia and New Guinea, but not elsewhere)
in: Watson, L., and M. J. Dallwitz (1992 onwards)
The Families of Flowering Plants
Descriptions, Illustrations, Identification, Information Retrieval.
Further reading
* * * * * 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. * * * * Boulay, J. 1997
''Dionée ''Dionée'' is a quarterly French language, French-language Periodical publication, periodical and the official publication of Association Francophone des Amateurs de Plantes Carnivores, a carnivorous plant society based in France.Rice, B. 2010Carn ...'' 38. * * * * * * * * * * * * * * * * * * *
External links
''Nepenthes'' - the Monkey Cups
from theBotanical Society of America The Botanical Society of America (BSA) represents professional and amateur botanists, researchers, educators and students in over 80 countries of the world. It functions as a United States nonprofit 501(c)(3) membership society. History The soci ...
''Nepenthes'': The Interactive Guide
at Tom's Carnivores
How to Grow ''Nepenthes''
at Tom's Carnivores
at the Carnivorous Plant Photo Finder
A video about ''Nepenthes rajah''
from '' The Private Life of Plants''
The Carnivorous Plant FAQ: ''Nepenthes''
by Barry Rice
Evolution -- ''Nepenthes'' Phylogeny
from theInternational Carnivorous Plant Society The International Carnivorous Plant Society (ICPS) is a non-profit organization founded in 1972. It is the International Cultivar Registration Authority for carnivorous plants. As of June 2011, the society had around 1400 members. The ICPS is prob ...
Inner World of ''Nepenthes''
from the John Innes Centre {{Authority control Caryophyllales genera Dioecious plants Taxa named by Carl Linnaeus Articles containing video clips