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''Nepenthes'' ( ) is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
carnivorous plants Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods, and occasionally small mammals and birds. They have adapted to grow in waterlo ...
, also known as tropical pitcher plants, or monkey cups, in the
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
family Nepenthaceae. The genus includes about 170 species, and numerous
natural Nature is an inherent character or constitution, particularly of the ecosphere or the universe as a whole. In this general sense nature refers to the laws, elements and phenomena of the physical world, including life. Although humans are part ...
and many cultivated hybrids. They are mostly
liana A liana is a long-Plant stem, stemmed Woody plant, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the Canopy (biology), canopy in search of direct sunlight. T ...
-forming plants of the
Old World The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
tropics The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
, ranging from South
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
,
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...
,
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
, and the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
; westward to
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
(two species) and the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (; Seychellois Creole: ), is an island country and archipelagic state consisting of 155 islands (as per the Constitution) in the Indian Ocean. Its capital and largest city, Victoria, ...
(one); southward to
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
(four) and
New Caledonia New Caledonia ( ; ) is a group of islands in the southwest Pacific Ocean, southwest of Vanuatu and east of Australia. Located from Metropolitan France, it forms a Overseas France#Sui generis collectivity, ''sui generis'' collectivity of t ...
(one); and northward to
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
(one) and
Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...
(one). The greatest diversity occurs on
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
,
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 list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...
, and the Philippines, with many
endemic Endemism is the state of a species being found only 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 foun ...
species. Many are plants of hot, humid, lowland areas, but most are tropical,
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
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
monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...
s were once thought to drink rainwater from the pitchers.


Description

''Nepenthes'' species usually consist of a shallow root system and a
prostrate Prostrate may refer to:- *Prostration, a position of submission in religion etc. *Prone position, a face-down orientation of the body *Prostrate shrub A prostrate shrub is a woody plant, most of the branches of which lie upon or just above the ...
or climbing stem, often several metres long and up to or more, and usually or less in diameter, although this may be thicker in a few species (e.g. '' N. bicalcarata''). From the stems arise alternate, sword-shaped leaves with entire
leaf margin A leaf (: leaves) is a principal appendage of the plant stem, stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leav ...
s. An extension of the
midrib A primary vein, also known as the midrib, is the main vascular structure running through the center of a leaf. The primary vein is crucial for the leaf’s efficiency in photosynthesis and overall health, as it ensures the proper flow of material ...
(the
tendril In botany, a tendril is a specialized Plant stem, stem, leaf or Petiole (botany), petiole with a thread-like shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as ''Cuscuta''. There ar ...
), 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 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 In materials science and continuum mechanics, viscoelasticity is the property of materials that exhibit both Viscosity, viscous and Elasticity (physics), elastic characteristics when undergoing deformation (engineering), deformation. Viscous mate ...
biopolymers Biopolymers are natural polymers produced by the cells of living organisms. Like other polymers, biopolymers consist of monomeric units that are covalently bonded in chains to form larger molecules. There are three main classes of 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 the trap contains glands that 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 Peristome (from the Greek language, Greek ''peri'', meaning 'around' or 'about', and ''stoma'', 'mouth') is an anatomical feature that surrounds an opening to an organ or structure. Some plants, fungi, and shelled gastropods have peristomes. In mo ...
(the "lip"), which is slippery and often quite colorful, attracting prey, but offering an unsure footing. The prey-capture effectiveness of the peristome is further enhanced in moist environments, where condensation may cause a thin water film to form on the surface of the peristome. When wet, the slippery surface of the peristome causes insects to "aquaplane", or slip and fall, into the pitcher. Above the peristome is a lid (the operculum); in many species, this keeps rain from diluting the fluid within the pitcher, the underside of which may contain
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
glands that attract prey. ''Nepenthes'' species usually produce two types of pitchers, known as leaf dimorphism. Appearing near the base of the plant are the large, lower traps, which typically sit on the ground. The upper or aerial pitchers are usually smaller, coloured differently, and possess different features from the lower pitchers. These upper pitchers usually form as the plant reaches maturity and the plant grows taller. To keep the plant steady, the upper pitchers often form a loop in the tendril, allowing it to wrap around nearby support. In some species (e.g. '' N. rafflesiana''), different prey may be attracted by the two types of pitchers. This varied morphology also often makes identification of species difficult. Prey usually consists of
insects Insects (from Latin ') are 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 of jointed ...
, but the largest species (e.g. '' N. rajah'' and ''N. rafflesiana'') may occasionally catch small
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s, such as "frogs, birds, and small mammals". Records of cultivated plants trapping small birds have been made.
Flower Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
s occur in
raceme A raceme () or racemoid is an unbranched, indeterminate growth, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are ...
s or more rarely in
panicle In botany, a panicle is a much-branched inflorescence. (softcover ). Some authors distinguish it from a compound spike inflorescence, by requiring that the flowers (and fruit) be pedicellate (having a single stem per flower). The branches of a p ...
s with
male Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
and
female An organism's sex is female ( symbol: ♀) if it produces the ovum (egg cell), the type of gamete (sex cell) that fuses with the male gamete (sperm cell) during sexual reproduction. A female has larger gametes than a male. Females and ...
flowers on separate plants. Three species have symbiotic relationships with
treeshrew The treeshrews (also called tree shrews or banxrings) are small mammals native to the tropical forests of South and Southeast Asia. They make up the entire order Scandentia (from Latin ''scandere'', "to climb"), which split into two families: the ...
s, which eat the nectar produced by the plant and defecate into the pitchers, providing valuable nutrients. ''Nepenthes'' are
insect-pollinated Entomophily or insect pollination is a form of pollination whereby pollen of plants, especially but not only of flowering plants, is distributed by insects. Flowers pollinated by insects typically advertise themselves with bright colours, somet ...
, the primary agents being flies (including
blow flies The Calliphoridae (commonly known as blowflies, blow flies, blow-flies, carrion flies, bluebottles, or greenbottles) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing bait, ...
,
midge A midge is any small fly, including species in several family (biology), families of non-mosquito nematoceran Diptera. Midges are found (seasonally or otherwise) on practically every land area outside permanently arid deserts and the frigid ...
s, and
mosquito Mosquitoes, the Culicidae, are a Family (biology), family of small Diptera, flies consisting of 3,600 species. The word ''mosquito'' (formed by ''Musca (fly), mosca'' and diminutive ''-ito'') is Spanish and Portuguese for ''little fly''. Mos ...
es), moths, wasps, and butterflies.Clarke, C.M. 2001. ''
Nepenthes of Sumatra and Peninsular Malaysia ''Nepenthes of Sumatra and Peninsular Malaysia'' is a monograph by Charles Clarke on the tropical pitcher plants of Sumatra, Peninsular Malaysia, and their minor surrounding islands. It was published in 2001 by Natural History Publications (Born ...
''. Natural History Publications (Borneo), Kota Kinabalu. Their smells can range from sweet to musty or
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
-like.
Seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
is typically produced in a four-sided capsule which may contain 50–500 wind-distributed seeds, consisting of a central
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
and two wings, one on either side (though '' N. pervillei'' differs). The genus is cytologically
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
, with all studied species having a
chromosome number Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
of ''2n''=80. This high number is thought to reflect
paleopolyploidy Paleopolyploidy is the result of genome duplications which occurred at least several million years ago (MYA). Such an event could either double the genome of a single species (autopolyploidy) or combine those of two species (allopolyploidy). Bec ...
(likely 8x or 16x).


Taxonomy

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


Etymology

The genus name ''Nepenthes'' was first published in 1737 in
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
's ''
Hortus Cliffortianus The ''Hortus Cliffortianus'' is a work of early botanical literature published in 1737. The work was a collaboration between Carl Linnaeus and the illustrator Georg Dionysius Ehret, financed by George Clifford in 1735–1736. Clifford was a we ...
''. It references a passage in
Homer Homer (; , ; possibly born ) was an Ancient Greece, Ancient Greek poet who is credited as the author of the ''Iliad'' and the ''Odyssey'', two epic poems that are foundational works of ancient Greek literature. Despite doubts about his autho ...
's ''
Odyssey The ''Odyssey'' (; ) is one of two major epics of ancient Greek literature attributed to Homer. It is one of the oldest surviving works of literature and remains popular with modern audiences. Like the ''Iliad'', the ''Odyssey'' is divi ...
'', in which the
potion A potion is a liquid "that contains medicine, poison, or something that is supposed to have magic powers." It derives from the Latin word ''potio'' which refers to a drink or the act of drinking. The term philtre is also used, often specifica ...
"Nepenthes pharmakon" is given to Helen by an
Egypt Egypt ( , ), officially the Arab Republic of Egypt, is a country spanning the Northeast Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via the Sinai Peninsula. It is bordered by the Mediterranean Sea to northe ...
ian queen. "''
Nepenthe Nepenthe (, ) is a possibly fictional medicine for sorrow – a "drug of forgetfulness" mentioned in ancient Greek literature and Greek mythology, depicted as originating in Egypt.. The carnivorous plant genus '' Nepenthes'' is named after th ...
s''" () literally means "without grief" ( ''nē'' = "not", ''penthos'' = "grief") and, in
Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
, is a drug that quells all sorrows with forgetfulness. Linnaeus explained:
If this is not Helen's ''Nepenthes'', it certainly will be for all botanists. What botanist would not be filled with admiration if, after a long journey, he should find this wonderful plant. In his astonishment, past ills would be forgotten when beholding this admirable work of the Creator! ranslated from Latin by Harry Veitch">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. ''Nepenthes'' was formally published as a genus">generic Generic or generics may refer to: In business * Generic term, a common name used for a range or class of similar things not protected by trademark * Generic brand, a brand for a product that does not have an associated brand or trademark, other ...
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 ''National Geographic'' (formerly ''The National Geographic Magazine'', sometimes branded as ''Nat Geo'') is an American monthly magazine published by National Geographic Partners. The magazine was founded in 1888 as a scholarly journal, nine ...
'', in which Paul A. Zahl wrote:
The carriers called them "monkey cups", a name I had heard elsewhere in reference to ''Nepenthes'', but the implication that monkeys drink the pitcher fluid seemed farfetched. I later proved it true. In Sarawak, I found an
orangutan 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.
The plants are often called ''kantong semar'' (
Semar Semar (Javanese script: ꦱꦼꦩꦂ) is a character in Javanese mythology who frequently appears in wayang shadow plays. He is one of the punokawan (clowns) but is divine and very wise. He is the Hyang, dhanyang (guardian spirit) of Java,Geertz ...
's pocket) in Indonesia and ''sako ni Hudas'' ( Judas'
money bag A money bag (or money sack) is a bag normally used to hold and transport coins and banknotes, often closed with a drawstring.
) in the Philippines.


Evolution and phylogeny

An absence of evidence of intermediate species, fossil or living (i.e. a
missing link Missing link may refer to: Biology * Missing link (human evolution), a non-scientific term typically referring to transitional fossils * Piltdown Man, a hoax in which bone fragments were presented as the "missing link" between ape and man Geog ...
), does not allow forming a
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
timeline for the development of the distinctive traits of modern ''Nepenthes'', which include its relatively rare strict
dioecy Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
and carnivorous pitchers. Although ''Nepenthes'' is distantly related to several modern genera, among these, even the carnivorous relatives he sundews (''Drosera''), Venus flytrap (''Dionaea muscipula">Drosera.html" ;"title="he sundews (''Drosera">he 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 In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
comparisons of the
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
'' matK ''gene sequences between ''Nepenthes'' species and with related species support this conclusion, long genetic distance between ''Nepenthes'' and others, and abruptly diverging "pom-pon" grouping of the ''Nepenthes'' species . Fossilized pollen of ''Nepenthes''-like plants living on the northern
Tethys Sea The Tethys Ocean ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
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 Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
wiped out the ancestral species in the original habitat. About 20 million years ago,
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
,
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 list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...
, and
Sulawesi Sulawesi ( ), also known as Celebes ( ), is an island in Indonesia. One of the four Greater Sunda Islands, and the List of islands by area, world's 11th-largest island, it is situated east of Borneo, west of the Maluku Islands, and south of Min ...
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 land bridges in the area 20,000 years ago during the ice age would have provided access to the remaining sites of ''Nepenthes'' populations in
Oceania Oceania ( , ) is a region, geographical region including Australasia, Melanesia, Micronesia, and Polynesia. Outside of the English-speaking world, Oceania is generally considered a continent, while Mainland Australia is regarded as its co ...
. The main complication with this hypothesis is the presence of ''Nepenthes'' on the distant islands of
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (; Seychellois Creole: ), is an island country and archipelagic state consisting of 155 islands (as per the Constitution) in the Indian Ocean. Its capital and largest city, Victoria, ...
and
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
. The seeds were thought to have been transferred by
seabirds Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same envi ...
and
shorebirds FIle:Vadare - Ystad-2021.jpg, 245px, A flock of Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflats in order to foraging, forage for food c ...
, 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 the
Malay Archipelago The Malay Archipelago is the archipelago between Mainland Southeast Asia and Australia, and is also called Insulindia or the Indo-Australian Archipelago. The name was taken from the 19th-century European concept of a Malay race, later based ...
, with the greatest biodiversity found on Borneo, Sumatra, and the Philippines, especially in the
Borneo montane rain forests The Borneo montane rain forests is an ecoregion on the island of Borneo in Southeast Asia. It includes montane tropical and subtropical moist broadleaf forests, also known as a cloud forests. The ecoregion is partly in East Malaysia (Sabah and S ...
. 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 Mainland Southeast Asia (historically known as Indochina and the Indochinese Peninsula) 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 th ...
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 ...
''. 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 their
altitude Altitude is a distance measurement, usually in the vertical or "up" direction, between a reference datum (geodesy), datum and a point or object. The exact definition and reference datum varies according to the context (e.g., aviation, geometr ...
above
sea level Mean sea level (MSL, often shortened to sea level) is an mean, average surface level of one or more among Earth's coastal Body of water, bodies of water from which heights such as elevation may be measured. The global MSL is a type of vertical ...
, 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 endemic to New Guinea, where it grows at an altitude of up to 3,520 m above sea level, higher than any other ''Nepenthes'' species.McPherson, S.R. 2009. ''Pitcher Plants of the Old World''. 2 ...
'' grows at a higher altitude than any other in the genus, up to . Most ''Nepenthes'' species grow in environments that provide high
humidity 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 (meteorology), precipitation, dew, or fog t ...
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 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 for ...
forest areas, roadsides, and disturbed fields. Other species have adapted to growing in
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
-like grass communities. The soils in which ''Nepenthes'' species grow are usually acidic and low in nutrients, being composed of
peat Peat is an accumulation of partially Decomposition, decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, Moorland, moors, or muskegs. ''Sphagnum'' moss, also called peat moss, is one of the most ...
, white sand,
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
, 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 Sea spray consists of aerosol particles formed from the ocean, primarily by ejection into Earth's atmosphere through bursting bubbles at the air-sea interface Sea spray contains both organic matter and inorganic salts that form sea salt aeroso ...
zone (e.g. '' N. albomarginata''). Other species grow on
inselberg An inselberg or monadnock ( ) is an isolated rock hill, knob, ridge, or small mountain that rises abruptly from a gently sloping or virtually level surrounding plain. In Southern Africa, a similar formation of granite is known as a koppie, an ...
s and as
lithophyte 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 als ...
s, while others, such as '' N. inermis'', can grow as
epiphyte An epiphyte is a plant or plant-like organism that grows on the surface of another 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 epiphyt ...
s with no soil contact.


Ecological relationships

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 Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
, and sweet scents. From this relationship, the plants primarily gain
nitrogen Nitrogen is a chemical element; it has Symbol (chemistry), symbol N and atomic number 7. Nitrogen is a Nonmetal (chemistry), nonmetal and the lightest member of pnictogen, group 15 of the periodic table, often called the Pnictogen, pnictogens. ...
and
phosphorus Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
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
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s, with
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s and other
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s topping the menu. Other arthropods found frequently include
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s,
scorpion Scorpions are predatory arachnids of the Order (biology), order Scorpiones. They have eight legs and are easily recognized by a pair of Chela (organ), grasping pincers and a narrow, segmented tail, often carried in a characteristic forward cur ...
s, and
centipede Centipedes (from Neo-Latin , "hundred", and Latin , "foot") are predatory arthropods belonging to the class Chilopoda (Ancient Greek , ''kheilos'', "lip", and Neo-Latin suffix , "foot", describing the forcipules) of the subphylum Myriapoda, ...
s, while
snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...
s and
frog A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order (biology), order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough ski ...
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 traps
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
s and produces nearly no nectar. ''Nepenthes albomarginata'' gains its name from the ring of white
trichome 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 plant ...
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 of
putrefaction Putrefaction is the fifth stage of death, following pallor mortis, livor mortis, algor mortis, and rigor mortis. This process references the breaking down of a body of an animal Post-mortem interval, post-mortem. In broad terms, it can be view ...
of collected organic matter that could harm the natural community of
infaunal Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and ''funga'', respectively. Flora, fauna, funga and other forms of life are collectively r ...
species that aid the plant's digestion.Clarke, C.M. 1997. ''
Nepenthes of Borneo ''Nepenthes of Borneo'' is a monograph by Charles Clarke on the tropical pitcher plants of Borneo.Clarke, C.M. 1997. ''Nepenthes of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. It was first published in 1997 by Natural History ...
''. 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 The treeshrews (also called tree shrews or banxrings) are small mammals native to the tropical forests of South and Southeast Asia. They make up the entire Order (biology), order Scandentia (from Latin ''scandere'', "to climb"), which split into ...
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 Feces (also known as faeces American and British English spelling differences#ae and oe, or fæces; : faex) are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the ...
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, Malaysia, and Kalimantan, Indonesia. The first specimen was described by Oldfield ...
'').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 to
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
, 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 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.


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 The trophic level of an organism is the position it occupies in a food web. Within 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 ...
of the infaunal ecosystem, are many species of
mosquito Mosquitoes, the Culicidae, are a Family (biology), family of small Diptera, flies consisting of 3,600 species. The word ''mosquito'' (formed by ''Musca (fly), mosca'' and diminutive ''-ito'') is Spanish and Portuguese for ''little fly''. Mos ...
larvae. Other infaunal species include
fly Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced ...
and
midge A midge is any small fly, including species in several family (biology), families of non-mosquito nematoceran Diptera. Midges are found (seasonally or otherwise) on practically every land area outside permanently arid deserts and the frigid ...
larvae, spiders,
mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
s, 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 ''Nepenthes'' infauna are the organisms that inhabit the pitchers of ''Nepenthes'' plants. These include fly and midge larvae, spiders, mites, ants, and even a species of crab, '' Geosesarma malayanum''. The most common and conspicuous pr ...
. 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 as
bactericide 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 p ...
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 including
protease A protease (also called a peptidase, proteinase, or proteolytic enzyme) is an enzyme that catalysis, catalyzes proteolysis, breaking down proteins into smaller polypeptides or single amino acids, and spurring the formation of new protein products ...
s,
chitinase Chitinases (, chitodextrinase, 1,4-β-poly-N-acetylglucosaminidase, poly-β-glucosaminidase, β-1,4-poly-N-acetyl glucosamidinase, poly ,4-(N-acetyl-β-D-glucosaminide)glycanohydrolase, (1→4)-2-acetamido-2-deoxy-β-D-glucan glycanohydrola ...
s,
pathogenesis-related protein Pathogenesis-related (PR) proteins are proteins produced in plants in the event of a pathogen attack. They are induced as part of systemic acquired resistance. Infections activate genes that produce PR proteins. Some of these proteins are antimicro ...
s and
thaumatin Thaumatin (also known as talin) is a low-calorie sweetener and taste 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.
Naphthoquinone Naphthoquinones constitute a class of organic compounds structurally related to naphthalene. Two isomers are common for the parent naphthoquinones: * 1,2-Naphthoquinone * 1,4-Naphthoquinone Natural products * Alkannin * Hexahydroxy-1,4-nap ...
s, 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 governor
Étienne 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 A species description is a formal scientific description of a newly encountered species, typically articulated through a scientific publication. Its purpose is to provide a clear description of a new species of organism and explain how it differ ...
 as '' N. madagascariensis''.Poiret, J.L.M. 1797. ''Népente''. In: J.B. Lamarck ''Encyclopédie Méthodique Botanique'' Vol. 4. The second species to be described was '' N. distillatoria'', the
Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...
n
endemic Endemism is the state of a species being found only 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 foun ...
. In 1677, Danish physician Thomas Bartholin made brief mention of it under the name ''Miranda herba'', Latin for "marvellous herb". Three years later, Dutch merchant Jacob Breyne referred to this species as ''Bandura zingalensium'', after a local name for the plant. ''Bandura'' subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined ''Nepenthes'' in 1737. ''Nepenthes distillatoria'' was again described in 1683, this time by Swedish physician and naturalist Herman Niklas Grim. Grim called it ''Planta mirabilis destillatoria'' or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant. Three years later, in 1686, English naturalist
John Ray John Ray Fellow of the Royal Society, FRS (November 29, 1627 – January 17, 1705) was a Christian England, English Natural history, naturalist widely regarded as one of the earliest of the English parson-naturalists. Until 1670, he wrote his ...
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 the
Malay Archipelago The Malay Archipelago is the archipelago between Mainland Southeast Asia and Australia, and is also called Insulindia or the Indo-Australian Archipelago. The name was taken from the 19th-century European concept of a Malay race, later based ...
. 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 Flora (: floras or florae) is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. The corresponding term for animals is ''fauna'', and for f ...
of
Ambon 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 three districts (''k ...
. However, it would not be published until many years after his death. After going blind in 1670, when the manuscript was only partially complete, Rumphius continued work on ''Herbarium Amboinensis'' with the help of clerks and artists. In 1687, with the project nearing completion, at least half of the illustrations were lost in a fire. Persevering, Rumphius and his helpers first completed the book in 1690. However, two years later, the ship carrying the manuscript to the Netherlands was attacked and sunk by the French, forcing them to start over from a copy that had fortunately been retained by Governor-General Johannes Camphuijs. The ''Herbarium Amboinensis'' finally arrived in the Netherlands in 1696. Even then, the first volume did not appear until 1741, 39 years after Rumphius's death. By this time, Linnaeus's name ''Nepenthes'' had become established. ''Nepenthes distillatoria'' was again illustrated in Johannes Burmann's ''Thesaurus Zeylanicus'' of 1737. The drawing depicts the end of a flowering stem with pitchers. Burmann refers to the plant as ''Bandura zeylanica''. The next mention of tropical pitcher plants was made in 1790, when Portuguese priest
João de Loureiro João de Loureiro (1717, Lisbon – 18 October 1791) was a Portuguese people, Portuguese Jesuit missionary and botanist. Biography After receiving admission to the Jesuit Order, João de Loureiro served as a missionary in Goa, capital of Port ...
described ''Phyllamphora mirabilis'', or the "marvellous urn-shaped leaf", from
Vietnam Vietnam, officially the Socialist Republic of Vietnam (SRV), is a country at the eastern edge of mainland Southeast Asia, with an area of about and a population of over 100 million, making it the world's List of countries and depende ...
. 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. In
Jean-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 botanist
Shigeo 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), K ...
, 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 games, bat-and-ball game played between two Sports team, teams of eleven players on a cricket field, field, at the centre of which is a cricket pitch, pitch with a wicket at each end, each comprising two Bail (cr ...
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, and
tissue culture Tissue culture is the growth of tissue (biology), tissues or cell (biology), 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-s ...
. Seeds are usually sown on damp chopped ''
Sphagnum ''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
''
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
, 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 Perlite is an amorphous volcanic glass that has a relatively high water content, typically formed by the Hydrate, hydration of obsidian. It occurs naturally and has the unusual property of greatly expanding when heated sufficiently. It is an indu ...
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 tissue (biology), tissues or cell (biology), 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-s ...
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 CITES (shorter acronym 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 inte ...
Appendix II, with the exception of '' N. rajah'' and '' N. khasiana'' which are listed in CITES Appendix I. The CITES listing means all international trade (including in parts and derivatives) is controlled by the CITES permitting system, with wild sourced specimens of Appendix I species prohibited from commercial international trade.


Hybrids and cultivars

There are many
hybrid 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 diff ...
''Nepenthes'' and numerous named
cultivar A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...
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'') * ''N.'' 'Gaya' (''N. khasiana'' × (''N. ventricosa'' × ''N. maxima''))


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 periodical and the official publication of Association Francophone des Amateurs de Plantes Carnivores, a carnivorous plant society based in France France, officially the French Republic, is a country ...
'' 38. * * * * * * * * * * * * * * * * * * *


External links


''Nepenthes'' – the Monkey Cups
from the
Botanical 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 soc ...

''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 Private Life of Plants'' is a BBC nature documentary series written and presented by David Attenborough, first shown in the United Kingdom from 11 January 1995. A study of the growth, movement, reproduction and survival of plants, it wa ...
''
The Carnivorous Plant FAQ: ''Nepenthes''
by Barry Rice
Evolution – ''Nepenthes'' Phylogeny
from the
International 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 publis ...

Inner World of ''Nepenthes''
from the John Innes Centre {{Authority control Caryophyllales genera Dioecious plants Taxa named by Carl Linnaeus Articles containing video clips