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| − | {{About|the tropical [[pitcher plant]]|other uses|Nepenthe (disambiguation)}}
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| − | '''''Nepenthes''''' ({{IPAc-en|n|ᵻ|ˈ|p|ɛ|n|θ|iː|z}}<!--The OED and Random House have [nɪ]; Merriam-Webster and the author of these pages have [nə]; other Webster's dictionaries have [ə̇] (written ''schwa-dot'' etc.)-->), also known as '''tropical pitcher plants''', is a [[genus]] of [[carnivorous plants]] in the [[monotypic]] family '''Nepenthaceae'''. The genus comprises roughly [[List of Nepenthes species|150 species]],<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press }}</ref> and numerous [[List of Nepenthes natural hybrids|natural]] and many cultivated hybrids. They are mostly [[liana]]-forming plants of the [[Old World]] [[tropics]], ranging from South [[China]], [[Indonesia]], [[Malaysia]] and the [[Philippines]]; westward to [[Madagascar]] (two species) and the [[Seychelles]] (one); southward to [[Australia]] (three) and [[New Caledonia]] (one); and northward to [[India]] (one) and [[Sri Lanka]] (one). The [[List of Nepenthes species by distribution|greatest diversity]] occurs on [[Borneo]], [[Sumatra]], and the [[Philippines]], with many [[endemism|endemic]] species. Many are plants of hot, humid, lowland areas, but the majority are tropical [[montane]] plants, receiving warm days but cool to cold, humid nights year round. A few are considered tropical alpine, with cool days and nights near freezing. The name "monkey cups" refers to the fact that [[monkey]]s have been observed drinking rainwater from these plants.
| + | The '''pitcher plants''' ('''''Nepenthes''''') are carnivorous plants. In Malaysian Borneo they are stuffed with coconut rice and eaten.<ref>{{cite web|url=http://www.bbc.com/travel/story/20170502-where-people-eat-carnivorous-plants|title=Where people eat carnivorous plants}}</ref> |
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| − | ==Description==
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| − | [[File:Nepenthes in the Southern Western Ghats.jpg|thumb|''Nepenthes'' at the [[Periyar Tiger Reserve]], in Southern [[Western ghats]] of [[India]].]]
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| − | ''Nepenthes'' species usually consist of a shallow root system and a [[prostrate shrub|prostrate]] or climbing stem, often several metres long and up to {{convert|15|m|ft|0|abbr=on}} or more, and usually {{convert|1|cm|in|1|abbr=on}} or less in diameter, although this may be thicker in a few species (e.g. ''[[Nepenthes bicalcarata|N. bicalcarata]]''). From the stems arise alternate, sword-shaped leaves with entire [[leaf margin]]s. An extension of the [[midrib]] (the [[tendril]]), which in some species aids in climbing, protrudes from the tip of the leaf; at the end of the tendril the pitcher forms. The pitcher starts as a small bud and gradually expands to form a globe- or tube-shaped trap.<ref name="Barthlott 2007" /> | |
| − | [[File:Nepenthes pitcher morphology upper.svg|thumb|250px|left|Basic structure of an upper pitcher]]
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| − | The trap contains a fluid of the plant's own production, which may be watery or [[syrup]]y, and is used to drown the prey. Research has shown this fluid contains [[viscoelastic]] [[biopolymers]] that may be crucial to the retention of insects within the traps of many species. The viscoelastic fluid in pitchers is especially effective in the retention of winged insects.<ref>{{cite journal|last1=Bonhomme|first1=V.|title=Slippery or sticky? Functional diversity in the trapping strategy of Nepenthes carnivorous plants|journal=New Phytologist|date=2011|volume=191|issue=2|pages=545–554|doi=10.1111/j.1469-8137.2011.03696.x|pmid=21434933}}</ref> The trapping efficiency of this fluid remains high, even when significantly diluted by water, as inevitably happens in wet conditions.<ref name=viscoelastic_pitcher_fluid>{{cite journal | last1 = Gaume | first1 = L. | last2 = Forterre | first2 = Y. | year = 2007 | title = A viscoelastic deadly fluid in carnivorous pitcher plants | url = | journal = PLoS ONE | volume = 2 | issue = 11| page = e1185 | doi = 10.1371/journal.pone.0001185 | pmid=18030325 | pmc=2075164}}</ref>
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| − | The lower part of the trap contains glands which absorb nutrients from captured prey. Along the upper inside part of the trap is a slick, waxy coating which makes the escape of its prey nearly impossible. Surrounding the entrance to the trap is a structure called the [[peristome]] (the "lip") which is slippery and often quite colorful, attracting prey, but offering an unsure footing. The prey-capture effectiveness of the peristome is further enhanced in moist environments, where condensation may cause a thin water film to form on the surface of the peristome. When wet, the slippery surface of the peristome causes insects to ‘aquaplane’, or slip and fall, into the pitcher.<ref name="Moran 2010">{{cite journal|last1=Moran|first1=J.A.|title=The carnivorous syndrome in Nepenthes pitcher plants|journal=Plant signaling & behavior|date=2010|volume=5|issue=6|pages=644–648|doi=10.4161/psb.5.6.11238|pmid=21135573|pmc=3001552}}</ref> Above the peristome is a lid (the [[operculum (botany)|operculum]]); in many species, this keeps [[rain]] from diluting the fluid within the pitcher, the underside of which may contain [[nectar]] glands which attract prey.<ref name="Barthlott 2007" />
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| − | ''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.<ref name="Barthlott 2007" />
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| − | Prey usually consists of [[insects]], but the largest species (e.g. ''[[Nepenthes rajah|N. rajah]]'' and ''N. rafflesiana'') may occasionally catch small [[vertebrate]]s, such as rats and lizards.<ref>{{cite journal | last1 = Phillipps | first1 = A | year = 1988 | title = ''A second record of rats as prey in ''Nepenthes rajah | url = http://www.carnivorousplants.org/cpn/articles/CPNv17n2p55.pdf | format = PDF | journal = [[Carnivorous Plant Newsletter]] | volume = 17 | issue = 2| page = 55 }}</ref><ref>Moran, J.A. 1991. The role and mechanism of ''Nepenthes rafflesiana'' pitchers as insect traps in Brunei. Ph.D. thesis, University of Aberdeen, Aberdeen, Scotland.</ref> There are even records of cultivated plants trapping small birds.<ref>{{Cite web
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| − | | title = Killer plant 'eats' great tit at Somerset nursery
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| − | | publisher = BBC News
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| − | | date = 5 August 2011
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| − | | url = http://www.bbc.co.uk/news/uk-england-somerset-14416809
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| − | | accessdate = 5 August 2011}}</ref><ref>{{cite journal | last1 = Hewitt-Cooper | first1 = N | year = 2012 | title = A case of bird capture by a cultivated specimen of the hybrid ''Nepenthes'' × ''mixta'' | url = | journal = [[Carnivorous Plant Newsletter]] | volume = 41 | issue = 1| pages = 31–33 }}</ref> [[Flower]]s occur in [[raceme]]s or more rarely in [[panicle]]s with [[male]] and [[female]] flowers on separate plants. They are [[Entomophily|insect-pollinated]], the primary agents being flies (including [[Calliphoridae|blow flies]], [[midge]]s, and [[mosquito]]es), moths, wasps, and butterflies.<ref name=Clarke2001>Clarke, C.M. 2001. ''[[Nepenthes of Sumatra and Peninsular Malaysia]]''. Natural History Publications (Borneo), Kota Kinabalu.</ref> Their smells can range from sweet to musty or [[fungus]]-like.<ref name=P&L /> [[Seed]] is typically produced in a four-sided capsule which may contain 50–500 wind-distributed seeds, consisting of a central [[embryo]] and two wings, one on either side (though see ''[[Nepenthes pervillei|N. pervillei]]'').
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| − | The genus is [[Cell biology|cytologically]] [[diploid]], with all studied species having a [[chromosome number]] of ''2n''=80.<ref>Lowrey, T.K. 1991. No. 519: Chromosome and isozyme number in the Nepenthaceae. ''American Journal of Botany'' '''78'''(6, supplement): 200–201.</ref><ref name=cyto>{{cite journal | last1 = Heubl | first1 = G.R. | last2 = Wistuba | first2 = A. | year = 1997 | title = A cytological study of the genus ''Nepenthes'' L. (Nepenthaceae) | url = | journal = Sendtnera | volume = 4 | issue = | pages = 169–174 }}</ref> This high number is thought to reflect [[paleopolyploidy]] (likely 8x or 16x).<ref name=cyto /><ref>{{cite journal | last1 = Meimberg | first1 = H. | last2 = Heubl | first2 = G. | year = 2006 | title = Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae | url = | journal = Plant Biology | volume = 8 | issue = 6| pages = 831–840 | doi = 10.1055/s-2006-924676 | pmid=17203435}}</ref><ref>[http://www.sysbot.biologie.uni-muenchen.de/botsyst/karyol.html Cytology of ''Nepenthes'']. LMU Department für Biologie.</ref><ref>Brittnacher, J. {{Tooltip| N.d. | No date}} [http://www.carnivorousplants.org/cp/EvolutionNepenthes.php Evolution -- ''Nepenthes'' Phylogeny]. [[International Carnivorous Plant Society]].</ref>
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| − | ==Taxonomy ==
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| − | {{Main|List of Nepenthes species|List of Nepenthes species by distribution}}
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| − | Around [[List of Nepenthes species|160 species]] of ''Nepenthes'' are currently recognised as valid. This number is quickly increasing, with several new species being described each year.<ref name=Sarawak />
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| − | ===Etymology===
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| − | The genus name ''Nepenthes'' was first published in 1737 in [[Carl Linnaeus]]'s ''[[Hortus Cliffortianus]]''.<ref>Linnaeus, C. 1737. ''Nepenthes''. ''Hortus Cliffortianus''. Amsterdam.</ref> It references a passage in [[Homer]]'s ''[[Odyssey]]'', in which the potion "Nepenthes pharmakon" is given to [[Helen of Troy|Helen]] by an [[Egypt]]ian queen. "[[Nepenthe]]" literally means "without grief" (''ne'' = not, ''penthos'' = grief) and, in [[Greek mythology]], is a drug that quells all sorrows with forgetfulness.<ref name=P&L /><ref>Gledhill, D. 2008. [https://books.google.com/books?id=NJ6PyhVuecwC&pg=PA271&lpg=PA271 ''The Names of Plants'']. Fourth Edition. Cambridge University Press, Cambridge.</ref> Linnaeus explained:
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| − | <blockquote>If this is not Helen's ''Nepenthes'', it certainly will be for all botanists. What botanist would not be filled with admiration if, after a long journey, he should find this wonderful plant. In his astonishment past ills would be forgotten when beholding this admirable work of the Creator! [translated from Latin by [[Harry Veitch]]]<ref>{{cite journal | last1 = Veitch | first1 = H.J. | year = 1897 | title = Nepenthes | url = | journal = Journal of the Royal Horticultural Society | volume = 21 | issue = 2| pages = 226–262 }}</ref></blockquote>
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| − | The plant Linnaeus described was ''N. distillatoria'', called bāndurā (බාඳුරා), a species from [[Sri Lanka]].<ref name=P&L>Phillipps, A. & A. Lamb 1996. ''[[Pitcher-Plants of Borneo]]''. Natural History Publications (Borneo), Kota Kinabalu.</ref>
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| − | ''Nepenthes'' was formally published as a [[genus|generic]] name in 1753 in Linnaeus's famous ''[[Species Plantarum]]'', which established [[botanical nomenclature]] as it exists today. ''Nepenthes distillatoria'' is the [[type species]] of the genus.<ref>{{cite journal | last1 = Linnaeus | first1 = C | year = 1753 | title = Nepenthes | url = | journal = Species Plantarum | volume = 2 | issue = | page = 955 }}</ref>
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| − | [[File:Nepenthes Species Plantarum.jpg|thumb|400px|''Nepenthes'' from Carolus Linnaeus's ''Species Plantarum'' of 1753]]
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| − | The name "monkey cups" was discussed in the May 1964 issue of ''[[National Geographic (magazine)|National Geographic]]'', in which Paul A. Zahl wrote:<ref>{{cite journal | last1 = Zahl | first1 = P.A. | year = 1964 | title = Malaysia's Giant Flowers and Insect-trapping Plants | url = | journal = National Geographic | volume = 125 | issue = 5| pages = 680–701 }}</ref>
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| − | <blockquote>The carriers called them "monkey cups," a name I had heard elsewhere in reference to ''Nepenthes'', but the implication that monkeys drink the pitcher fluid seemed farfetched. I later proved it true. In Sarawak I found an [[orangutan]] that had been raised as a pet and later freed. As I approached it gingerly in the forest, I offered it a half-full pitcher. To my surprise, the ape accepted it and, with the finesse of a lady at tea, executed a delicate bottoms-up.</blockquote>
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| − | The plants are often called ''kantong semar'' ([[Semar]]'s pocket) in [[Indonesia]] and ''sako ni Hudas'' ([[Judas Iscariot|Judas']] [[Money bag#History|money bag]]) in the [[Philippines]].
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| − | ===Evolution and phylogeny===
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| − | There is an absence of evidence of intermediate species, fossil or living (i.e. a [[Transitional fossil|missing link]]), to inform a [[phylogenetic]]al timeline for the development of the distinctive traits of modern ''Nepenthes'', which include its relatively rare strict [[dioecy]] and carnivorous pitchers. Although ''Nepenthes'' is distantly related to several modern genera, among these, even the carnivorous relatives (the Sundews ([[Drosera]]), Venus Flytrap ([[Dionea muscipula]]), Waterwheel plant ([[Aldrovanda]]), and Dewy Pine ([[Drosophyllum]])), all lack those traits. Among known ''Nepenthes'', no proto-modern characteristics or large variations are found, which suggests that all extant species radiated from a single close ancestor bearing all the modern traits. [[Phylogenetic]] comparisons of the [[chloroplast]] [[Maturase K|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 .<ref name=ICPS_phylogeny>{{cite webpage | last1 = Brittnacher | first1 = John | year = 2011 | title = Evolution -- Nepenthes Phylogeny | url = http://www.carnivorousplants.org/cp/EvolutionNepenthes.php | journal = ICPS Webpage}}</ref>
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| − | Fossilized pollen of ''Nepenthes''-like plants living on the northern [[Tethys Sea]] from 65 to 35 million years ago indicate 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. 20 million years ago, [[Borneo]], [[Sumatra]] and [[Sulawesi]] and possibly even the [[Philippines]] were connected to mainland Asia, providing a bridge for the colonization of most sites of ''Nepenthes'' species radiation. The extensive landbridges in the area 20,000 years ago during the ice age would have provided access to the remaining sites of ''Nepenthes'' populations in [[Oceania]]. The main complication with this hypothesis is the presence of ''Nepenthes'' on the distant islands of [[Seychelles]] and [[Madagascar]]. It has been proposed that the seeds were transferred by [[seabirds]] and [[shorebirds]], which rest during their migrations in swampy habitats and may have inadvertently picked up the seeds. This hypothesis is possibly reinforced by the success of the lowland swamp-dwelling ''[[Nepenthes distillatoria|N. distillatoria]]'' in colonizing so many locations.<ref name=ICPS_phylogeny />
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| − | ==Distribution and habitat==
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| − | [[File:Nepenthes distribution.svg|thumb|280px|right|Global distribution of ''Nepenthes'']]
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| − | {{further|List of Nepenthes species by distribution|l1=List of ''Nepenthes'' species by distribution}}
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| − | The genus ''Nepenthes'' is mostly found within the [[Malay Archipelago]], with the [[List of Nepenthes species by distribution|greatest biodiversity]] found on Borneo, Sumatra, and the Philippines, especially in the [[Borneo montane rain forests]]. The full range of the genus includes Madagascar (''[[Nepenthes madagascariensis|N. madagascariensis]]'' and ''[[Nepenthes masoalensis|N. masoalensis]]''), the Seychelles (''[[Nepenthes pervillei|N. pervillei]]''), Sri Lanka (''[[Nepenthes distillatoria|N. distillatoria]]''), and India (''[[Nepenthes khasiana|N. khasiana]]'') in the west to Australia (''[[Nepenthes mirabilis|N. mirabilis]]'', ''[[Nepenthes rowanae|N. rowanae]]'', and ''[[Nepenthes tenax|N. tenax]]'') and New Caledonia (''[[Nepenthes vieillardii|N. vieillardii]]'') in the southeast. Most species are restricted to very small ranges, including some only found 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. ''[[Nepenthes deaniana|N. deaniana]]'', which was rediscovered 100 years after its initial discovery). About 10 species have population distributions larger than a single island or group of smaller islands. ''Nepenthes mirabilis'' has the distinction of being the most widely distributed species in the genus, ranging from [[Indochina]] and throughout the Malay Archipelago to Australia.<ref name="Barthlott 2007" /><ref name=McPherson>McPherson, S.R. 2009. ''[[Pitcher Plants of the Old World]]''. 2 volumes. Redfern Natural History Productions, Poole.</ref><ref name="Jebb and Cheek">{{cite journal | last1 = Jebb | first1 = M. | last2 = Cheek | first2 = M. | year = 1997 | title = [[A skeletal revision of Nepenthes (Nepenthaceae)|A skeletal revision of ''Nepenthes'' (Nepenthaceae)]]. | url = | journal = Blumea | volume = 42 | issue = | pages = 1–106 }}</ref>
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| − | Because of the nature of the habitats which ''Nepenthes'' species occupy, they are often graded as either lowland or highland species, depending on their [[altitude]] above [[sea level]], with {{convert|1200|m|ft|0|abbr=on}} the rough delineation between lowland and highland. Species growing at lower altitudes require continuously warm climates with little difference between day and night temperatures, whereas highland species thrive when they receive warm days and much cooler nights. ''[[Nepenthes lamii]]'' grows at a higher altitude than any other in the genus, up to {{convert|3520|m|ft|0|abbr=on}}.<ref name="Barthlott 2007" /><ref name="Jebb and Cheek" />
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| − | Most ''Nepenthes'' species grow in environments that provide high [[humidity]] and precipitation and moderate to high light levels. A few species, including ''[[Nepenthes ampullaria|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 [[clearcutting|clear-cut]] forest areas, roadsides, and disturbed fields. Other species have adapted to growing in [[savanna]]-like grass communities. The soils in which ''Nepenthes'' species grow are usually acidic and low in nutrients, being composed of [[peat]], white sand, [[sandstone]], or volcanic soils. Exceptions to these generalities include species that thrive in soils with high [[heavy metal (chemistry)|heavy metal]] content (e.g. ''[[Nepenthes rajah|N. rajah]]''), on sandy beaches in the [[sea spray]] zone (e.g. ''[[Nepenthes albomarginata|N. albomarginata]]''). Other species grow on [[inselberg]]s and as [[lithophyte]]s, while others, such as ''[[Nepenthes inermis|N. inermis]]'', can grow as [[epiphyte]]s with no soil contact.<ref name="Barthlott 2007" />
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| − | ==Ecological relationships==
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| − | [[File:Rajahlizard.jpg|thumb|A drowned lizard found in a freshly opened pitcher of ''[[Nepenthes rajah|N. rajah]]'']]
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| − | The most obvious interaction between ''Nepenthes'' species and their environments, including other organisms, is that of [[predation|predator and prey]]. ''Nepenthes'' species certainly attract and kill their prey, albeit passively, through active production of attractive colours, sugary [[nectar]], and even sweet scents. From this relationship, the plants primarily gain [[nitrogen]] and [[phosphorus]] to supplement their nutrient requirements for growth, given these soil nutrients are typically lacking. The most frequent prey is an abundant and diverse group of [[arthropod]]s, with [[ant]]s and other [[insect]]s topping the menu. Other arthropods found frequently include [[spider]]s, [[scorpion]]s, and [[centipede]]s, while [[snail]]s and [[frog]]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.<ref name="Barthlott 2007" /> While many ''Nepenthes'' species are generalists in what they capture, at least one, ''[[Nepenthes albomarginata|N. albomarginata]]'', has specialised and almost exclusively traps [[termite]]s and produces nearly no nectar. ''Nepenthes albomarginata'' gains its name from the ring of white [[trichome]]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.<ref name="Moran 2001">{{cite journal | last1 = Moran | first1 = J.A. | last2 = Merbach | first2 = M.A. | last3 = Livingston | first3 = N.J. | last4 = Clarke | first4 = C.M. | last5 = Booth | first5 = W.E. | year = 2001 | title = Termite prey specialization in the pitcher plant ''Nepenthes albomarginata''—evidence from stable isotope analysis | url = http://aob.oxfordjournals.org/cgi/reprint/88/2/307.pdf | format = PDF | journal = Annals of Botany | volume = 88 | issue = 2| pages = 307–311 | doi = 10.1006/anbo.2001.1460 }}</ref><ref name="Merbach et al 2002">{{cite journal | last1 = Merbach | first1 = M.A. | last2 = Merbach | first2 = D.J. | last3 = Maschwitz | first3 = U. | last4 = Booth | first4 = W.E. | last5 = Fiala | first5 = B. | last6 = Zizka | first6 = G. | year = 2002 | title = Mass march of termites into the deadly trap | url = https://www.researchgate.net/publication/232780033_Carnivorous_plants_Mass_march_of_termites_into_the_deadly_trap | journal = Nature | volume = 415 | issue = 6867| pages = 36–37 | doi = 10.1038/415036a | pmid=11780106}}</ref>
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| − | {{Gallery
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| − | |title=Prey retention by viscoelastic pitcher fluid of ''[[Nepenthes rafflesiana|N. rafflesiana]]''<ref name=viscoelastic_pitcher_fluid />
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| − | |File:Calliphora vomitoria water escape (ventral).ogv|The [[blue bottle fly]] (''Calliphora vomitoria'') can escape after landing in water on its [[ventral]] surface.
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| − | |File:Calliphora vomitoria water escape (dorsal).ogv|The same is true if the fly falls in [[Dorsum (anatomy)|dorsal]]ly (wings-first).
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| − | |File:Calliphora vomitoria in digestive fluid (ventral).ogv|But the [[viscoelasticity|viscoelastic]] properties of ''N. rafflesiana'' digestive fluid prevent prey escape, whether the fall is ventral..
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| − | |File:Calliphora vomitoria in digestive fluid (dorsal).ogv|..or dorsal. (All videos recorded at 500 frames/s)
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| − | ===Symbioses===
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| − | [[File:Attlwmosq.jpg|thumb|right|A lower pitcher of ''[[Nepenthes attenboroughii|N. attenboroughii]]'' supporting a large population of mosquito larvae. The upright lid of this species exposes its pitchers to the elements such that they are often completely filled with fluid.<ref>{{cite journal | last1 = Robinson | first1 = A.S. | last2 = Fleischmann | first2 = A.S. | last3 = McPherson | first3 = S.R. | last4 = Heinrich | first4 = V.B. | last5 = Gironella | first5 = E.P. | last6 = Peña | first6 = C.Q. | year = 2009 | title = A spectacular new species of ''Nepenthes'' L. (Nepenthaceae) pitcher plant from central Palawan, Philippines | url = | journal = Botanical Journal of the Linnean Society | volume = 159 | issue = 2| pages = 195–202 | doi = 10.1111/j.1095-8339.2008.00942.x }}</ref>]]
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| − | ''[[Nepenthes bicalcarata]]'' provides space in the hollow tendrils of its upper pitchers for the carpenter ant ''[[Camponotus schmitzi]]'' to build nests. The ants take larger prey from the pitchers, which may benefit ''N. bicalcarata'' by reducing the amount of [[putrefaction]] of collected organic matter that could harm the natural community of [[Nepenthes infauna|infaunal]] species that aid the plant's digestion.<ref name="Clarke 1997">Clarke, C.M. 1997. ''[[Nepenthes of Borneo]]''. Natural History Publications (Borneo), Kota Kinabalu.</ref>
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| − | ''[[Nepenthes 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.<ref name="Clarke 2009">{{cite journal | last1 = Clarke | first1 = C.M. | last2 = Bauer | first2 = U. | last3 = Lee | first3 = C.C. | last4 = Tuen | first4 = A.A. | last5 = Rembold | first5 = K. | last6 = Moran | first6 = J.A. | year = 2009 | title = Tree shrew lavatories: a novel nitrogen sequestration strategy in a tropical pitcher plant | url = http://rsbl.royalsocietypublishing.org/content/5/5/632.full.pdf | format = PDF | journal = Biology Letters | volume = 5 | issue = 5| pages = 632–635 | doi = 10.1098/rsbl.2009.0311 | pmid=19515656 | pmc=2781956}}</ref> 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]]'').<ref name=CMC2010>{{cite journal | last1 = Chin | first1 = L. | last2 = Moran | first2 = J.A. | last3 = Clarke | first3 = C. | year = 2010 | title = Trap geometry in three giant montane pitcher plant species from Borneo is a function of tree shrew body size | url = | journal = New Phytologist | volume = 186 | issue = 2| pages = 461–470 | doi = 10.1111/j.1469-8137.2009.03166.x | pmid=20100203}}</ref><ref name=Walker>Walker, M. 2010. [http://news.bbc.co.uk/earth/hi/earth_news/newsid_8552000/8552157.stm Giant meat-eating plants prefer to eat tree shrew poo]. ''BBC Earth News'', March 10, 2010.</ref> A similar adaptation was found in ''[[Nepenthes macrophylla|N. macrophylla]]'', ''[[Nepenthes rajah|N. rajah]]'', ''[[Nepenthes ampullaria|N. ampullaria]]'', and is also likely to be present in ''[[Nepenthes ephippiata|N. ephippiata]]''.<ref name=Walker /><ref name="Moran 2003">{{cite journal|last1=Moran|first1=J.A.|title=From carnivore to detritivore? Isotopic evidence for leaf litter utilization by the tropical pitcher plant Nepenthes ampullaria|journal=International Journal of Plant Sciences|date=2003|volume=164|issue=4|pages=635–639|doi=10.1086/375422}}</ref>
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| − | ===Infauna===
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| − | {{Main|Nepenthes infauna}}
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| − | Organisms that spend at least part of their lives within the pitchers of ''Nepenthes'' species are often called ''Nepenthes'' infauna. The most common infaunal species, often representing the top [[trophic level]] of the infaunal ecosystem, are many species of [[mosquito]] larvae. Other infaunal species include [[fly]] and [[midge]] larvae, spiders, [[mite]]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]]. 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 [[Mutualism (biology)|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.<ref name="Clarke 1997" /><ref name="Mogi 1992">{{cite journal | last1 = Mogi | first1 = M. | last2 = Yong | first2 = H.S. | year = 1992 | title = Aquatic arthropod communities in ''Nepenthes'' pitchers: the role of niche differentiation, aggregation, predation and competition in community organization | url = | journal = Oecologia | volume = 90 | issue = 2| pages = 172–184 | doi = 10.1007/BF00317174 }}</ref><ref name="Beaver 1979">{{cite journal | last1 = Beaver | first1 = R.A. | year = 1979 | title = Fauna and foodwebs of pitcher plants in west Malaysia | url = | journal = Malayan Nature Journal | volume = 33 | issue = | pages = 1–10 }}</ref>
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| − | === Antimicrobial properties ===
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| − | ''Nepenthes'' digestive fluids are sterile before pitchers open and contain secondary metabolites and proteins that act as [[bactericide]]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]]s, [[chitinase]]s, [[pathogenesis-related protein]]s and [[thaumatin]]-like proteins are produced in the pitcher fluid. In addition to breaking down prey, these can act as antimicrobial agents.<ref>{{Cite journal|last=Rottloff|first=Sandy|last2=Miguel|first2=Sissi|last3=Biteau|first3=Flore|last4=Nisse|first4=Estelle|last5=Hammann|first5=Philippe|last6=Kuhn|first6=Lauriane|last7=Chicher|first7=Johana|last8=Bazile|first8=Vincent|last9=Gaume|first9=Laurence|date=2016-03-01|title=Proteome analysis of digestive fluids in Nepenthes pitchers|url=http://aob.oxfordjournals.org/content/117/3/479|journal=Annals of Botany|language=en|volume=117|issue=3|pages=479–495|doi=10.1093/aob/mcw001|issn=0305-7364|pmc=4765550|pmid=26912512}}</ref> 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. ''[[Nepenthes lowii|N. lowii]]'', ''[[Nepenthes attenboroughii|N. attenboroughii]]'' and ''[[Nepenthes jamban|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.<ref>{{Cite journal|last=Mithöfer|first=Axel|date=2011-09-01|title=Carnivorous pitcher plants: Insights in an old topic|url=http://www.sciencedirect.com/science/article/pii/S0031942210004450|journal=Phytochemistry|series=Plant-Insect Interactions|volume=72|issue=13|pages=1678–1682|doi=10.1016/j.phytochem.2010.11.024|pmid=21185041}}</ref> [[Naphthoquinone]]s, a class of secondary metabolite, are commonly produced, and these either kill or inhibit the growth and reproduction of bacteria and fungi.<ref>{{Cite journal|last=Buch|first=Franziska|last2=Rott|first2=Matthias|last3=Rottloff|first3=Sandy|last4=Paetz|first4=Christian|last5=Hilke|first5=Ines|last6=Raessler|first6=Michael|last7=Mithöfer|first7=Axel|date=2012-12-21|title=Secreted pitfall-trap fluid of carnivorous Nepenthes plants is unsuitable for microbial growth|url=http://aob.oxfordjournals.org/content/early/2012/12/20/aob.mcs287|journal=Annals of Botany|language=en|pages=mcs287|doi=10.1093/aob/mcs287|issn=0305-7364|pmc=3579442|pmid=23264234|volume=111}}</ref> 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.
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| − | ==Botanical history==
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| − | [[File:Utricaria vegetabilis zeylanensium.jpg|thumb|right|Plukenet's drawing of ''N. distillatoria'' from his ''Almagestum Botanicum'' of 1696.]]
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| − | The earliest known record of ''Nepenthes'' dates back to the 17th century. In 1658, French colonial governor [[Étienne de Flacourt]] published a description of a pitcher plant in his seminal work ''Histoire de la Grande Isle de Madagascar''. It reads:<ref>de Flacourt, É. 1658. ''Histoire de la Grande Isle de Madagascar''.</ref>
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| − | <blockquote>It is a plant growing about 3 feet high which carries at the end of its leaves, which are 7 inches long, a hollow flower or fruit resembling a small vase, with its own lid, a wonderful sight. There are red ones and yellow ones, the yellow being the biggest. The inhabitants of this country are reluctant to pick the flowers, saying that if somebody does pick them in passing, it will not fail to rain that day. As to that, I and all the other Frenchmen did pick them, but it did not rain. After rain these flowers are full of water, each one containing a good half-glass. [translated from French in ''[[Pitcher-Plants of Borneo]]'']<ref name=P&L /></blockquote>
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| − | Flacourt called the plant ''Amramatico'', after a local name. More than a century later, this species was [[Species description|formally described]] as ''[[Nepenthes madagascariensis|N. madagascariensis]]''.<ref name=Poiret>Poiret, J.L.M. 1797. ''Népente''. In: J.B. Lamarck ''Encyclopédie Méthodique Botanique'' Vol. 4.</ref>
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| − | The second species to be described was ''[[Nepenthes distillatoria|N. distillatoria]]'', the [[Sri Lanka]]n [[endemism|endemic]]. In 1677, Danish physician [[Thomas Bartholin]]<!--Bartolinus--> made brief mention of it under the name ''Miranda herba'', Latin for "marvellous herb".<ref>{{cite journal | last1 = Bartholinus | year = | title = Miranda herba | url = | journal = Acta Medica et Philosophica Hafniensia | volume = 3 | issue = | page = 38 }}</ref> Three years later, Dutch merchant [[Jacob Breyne]] referred to this species as ''Bandura zingalensium'', after a local name for the plant.<ref>Breyne, J. 1680. ''Bandura zingalensium'' etc. ''Prodromus Fasciculi Rariorum Plantarum'' 1: 18.</ref> ''Bandura'' subsequently became the most commonly used name for the tropical pitcher plants, until Linnaeus coined ''Nepenthes'' in 1737.<ref name=P&L />
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| − | ''Nepenthes distillatoria'' was again described in 1683, this time by Swedish physician and naturalist [[Herman Niklas Grim]]<!--also known as Hermann Nicolaus Grimm, Hermannus Nicolaus Grimmius-->.<ref>Grimm, H.N. 1683. ''Planta mirabilis destillatoria''. In: Miscellanea curiosa sive Ephemeridum. ''Med. Phys. Germ. Acad. Nat. Cur. Decuriae'' 2, ann. prim. p. 363, f. 27.</ref> Grim called it ''Planta mirabilis destillatoria'' or the "miraculous distilling plant", and was the first to clearly illustrate a tropical pitcher plant.<ref name=P&L /> Three years later, in 1686, English naturalist [[John Ray]] quoted Grim as saying:<ref>Ray, J. 1686. ''Bandura cingalensium'' etc. ''Historia Plantarum'' 1: 721–722.</ref>
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| − | <blockquote>The root draws up moisture from the earth which with the help of the sun's rays rises up into the plant itself and then flows down through the stems and nerves of the leaves into the natural utensil to be stored there until used for human needs. [translated from Latin in ''[[Pitcher-Plants of Borneo]]'']<ref name=P&L /></blockquote>
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| − | One of the earliest illustrations of ''Nepenthes'' appears in Leonard Plukenet's ''Almagestum Botanicum'' of 1696.<ref>Plukenet, L. 1696. ''Utricaria vegetabilis zeylanensium''. In: ''Almagestum Botanicum''.</ref> The plant, called ''Utricaria vegetabilis zeylanensium'', is undoubtedly ''N. distillatoria''.<ref name =P&L />
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| − | [[File:Cantharifera - Herbarium amboinense (1747).jpg|thumb|left|''Cantharifera'' as illustrated in Rumphius's ''Herbarium Amboinensis'', Volume 5, published in 1747, although probably drawn in the late 17th century. The vine on the right is not a ''Nepenthes'', but a species of ''[[Flagellaria]]''.]]
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| − | Around the same time, German botanist [[Georg Eberhard Rumphius]] discovered two new ''Nepenthes'' species in the [[Malay Archipelago]]. Rumphius illustrated the first one, now considered synonymous with ''[[Nepenthes mirabilis|N. mirabilis]]'', and gave it the name ''Cantharifera'', meaning "tankard-bearer". The second, referred to as ''Cantharifera alba'', is thought to have been ''[[Nepenthes maxima|N. maxima]]''. Rumphius described the plants in his most famous work, the six-volume ''Herbarium Amboinense'', a catalogue of the [[flora]] of [[Ambon Island]]. However, it would not be published until many years after his death.<ref>Rumphius, G.E. 1741–1750. ''Cantharifera''. In: ''Herbarium Amboinense'' 5, lib. 7, cap. 61, p. 121, t. 59, t. 2.</ref>
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| − | 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.<ref name=P&L />
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| − | [[File:Nepenthes distillatoria - Thesaurus zeylanicus (1737).jpg|thumb|right|Illustration of ''Bandura zeylanica'' (''N. distillatoria'') from Burmann's ''Thesaurus Zeylanicus'' of 1737]]
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| − | ''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''.<ref>Burmann, J. 1737. ''Thesaurus Zeylanicus''. Amsterdam.</ref>
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| − | The next mention of tropical pitcher plants was made in 1790, when Portuguese priest [[João de Loureiro]] described ''Phyllamphora mirabilis'', or the "marvellous urn-shaped leaf", from [[Vietnam]]. Despite living in the country for around 35 years, it seems unlikely that Loureiro observed living plants of this species, as he stated the lid is a moving part, [[rapid plant movement|actively opening and closing]]. In his most celebrated work, ''Flora Cochinchinensis'', he writes:<ref>de Loureiro, J. 1790. ''Flora Cochinchinensis'' 2: 606–607.</ref>
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| − | <blockquote>[...] (the) leaf-tip ends in a long hanging tendril, twisted spirally in the middle, from which hangs a sort of vase, oblong, pot-bellied, with a smooth lip with a projecting margin and a lid affixed to one side, which of its own nature freely opens and closes in order to receive the dew and store it. A marvellous work of the Lord! [translated from French in ''[[Pitcher-Plants of Borneo]]'']<ref name=P&L /></blockquote>
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| − | ''Phyllamphora mirabilis'' was eventually transferred to the genus ''Nepenthes'' by Rafarin in 1869.<ref>Schlauer, J. {{Tooltip| N.d. | No date}} [http://www.omnisterra.com/bot/cp_home.cgi?name=Nepenthes+mirabilis ''Nepenthes mirabilis'']. Carnivorous Plant Database.</ref> As such, ''P. mirabilis'' is the [[basionym]] of this most cosmopolitan of tropical pitcher plant species.<ref name="Clarke 1997"/>
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| − | Loureiro's description of a moving lid was repeated by [[Jean Louis Marie Poiret]] in 1797. Poiret described two of the four ''Nepenthes'' species known at the time: ''N. madagascariensis'' and ''N. distillatoria''. He gave the former its current name and called the latter ''Nepente de l'Inde'', or simply "''Nepenthes'' of India", although this species is absent from the mainland. In [[Jean-Baptiste Lamarck]]'s ''Encyclopédie Méthodique Botanique'', he included the following account:<ref name=Poiret />
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| − | <blockquote>This urn is hollow, as I have just said, usually full of soft, clear water, and then closed. It opens during the day and more than half the liquid disappears, but this loss is repaired during the night, and the next day the urn is full again and closed by its lid. This is its sustenance, and enough for more than one day because it is always about half-full at the approach of night. [translated from French in ''[[Pitcher-Plants of Borneo]]'']<ref name=P&L /></blockquote>
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| − | [[File:Veitch Nepenthes house.jpg|thumb|right|The ''Nepenthes'' house of the [[Veitch Nurseries]] as illustrated in ''[[The Gardeners' Chronicle]]'', 1872]]
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| − | 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.<ref name="Barthlott 2007">Barthlott, W., Porembski, S., Seine, R., and Theisen, I. 2007. ''The Curious World of Carnivorous Plants.'' Portland, Oregon: Timber Press.</ref><ref name=P&L /> 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 [[Japan]]ese botanist [[Shigeo Kurata]], whose work in the 1960s and 1970s did much to bring attention to these plants.<ref name=Sarawak>Clarke, C.M. & C.C. Lee 2004. ''[[Pitcher Plants of Sarawak]]''. Natural History Publications (Borneo), Kota Kinabalu.</ref>
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| − | ==Cultivation==
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| − | [[File:2005-12-18 N rajah 034.jpg|thumb|Cultivated ''[[Nepenthes rajah]], ''[[Nepenthes aristolochioides]]'' and other species]]
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| − | ''Nepenthes'' may be cultivated in greenhouses. Easier species include ''[[Nepenthes alata|N. alata]]'', ''[[Nepenthes ventricosa|N. ventricosa]]'', ''[[Nepenthes khasiana|N. khasiana]]'', and ''[[Nepenthes sanguinea|N. sanguinea]]''. These four species are highlanders (''N. alata'' has both lowland and highland forms), some easy lowlander species are ''[[Nepenthes rafflesiana|N. rafflesiana]]'', ''[[Nepenthes bicalcarata|N. bicalcarata]]'', ''[[Nepenthes mirabilis|N. mirabilis]]'', and ''[[Nepenthes hirsuta|N. hirsuta]]''.<ref name="PietropaoloPietropaolo1986">{{cite book|author1=James Pietropaolo|author2=Patricia Ann Pietropaolo|title=Carnivorous Plants of the World|url=https://books.google.com/books?id=yQ5IAAAAYAAJ|year=1986|publisher=Timber Press|isbn=978-0-88192-066-6|page=49}}</ref>
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| − | 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 (insect)|cricket]]s may be beneficial. Terrarium culture of smaller plants, such as ''[[Nepenthes bellii|N. bellii]]'', [[Nepenthes × trichocarpa|''N.'' × ''trichocarpa'']] and ''[[Nepenthes ampullaria|N. ampullaria]]'', is possible, but most plants will get too large over time.<ref name="D'Amato2013">{{cite book|author=Peter D'Amato|title=The Savage Garden, Revised: Cultivating Carnivorous Plants|url=https://books.google.com/books?id=dOA2pFHQG4AC&pg=PT89|date=2 July 2013|publisher=Ten Speed Press|isbn=978-1-60774-411-5|page=89}}</ref><ref name="Rice2006">{{cite book|author=Barry A. Rice|title=Growing carnivorous plants|url=https://books.google.com/books?id=NJfuAAAAMAAJ|year=2006|publisher=Timber Press, Incorporated|isbn=978-0-88192-807-5}}</ref>
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| − | Plants can be propagated by seed, cuttings, and [[Plant tissue culture|tissue culture]]. Seeds are usually sown on damp chopped ''[[Sphagnum]]'' [[moss]], or on sterile plant tissue culture media once they have been properly disinfected. The seeds generally become nonviable soon after harvesting, so seed are not usually the preferred method of propagation. A 1:1 mixture of orchid medium with moss or [[perlite]] has been used for germination and culture. Seed may take two months to germinate, and two years or more to yield mature plants. Cuttings may be rooted in damp ''Sphagnum'' moss in a plastic bag or tank with high humidity and moderate light. They can begin to root in one to two months and start to form pitchers in about six months. [[Plant tissue culture|Tissue culture]] is now used commercially and helps reduce collection of wild plants, as well as making many rare species available to hobbyists at reasonable prices. ''Nepenthes'' species are considered threatened or endangered plants and all of them are listed in [[CITES]] appendices 2, with the exception of ''[[Nepenthes rajah|N. rajah]]'' and ''[[Nepenthes khasiana|N. khasiana]]'' which are listed in CITES appendix 1.<ref name="D'Amato2013"/>{{rp|353}}
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| − | ==Hybrids and cultivars==
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| − | [[File:VentricosaXtrusmadensis-1.jpg|thumb|200px|The complex man-made hybrid ''N. ventricosa'' × (''N. lowii'' × ''N. macrophylla'')]]
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| − | {{See also|List of Nepenthes natural hybrids|List of Nepenthes cultivars}}
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| − | There are many [[Hybrid (biology)|hybrid]] ''Nepenthes'' and numerous named [[cultivar]]s. Some of the more well-known, artificially produced hybrids and cultivars include:{{citation needed|date=February 2017}}
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| − | | |
| − | *''N.'' × ''coccinea'' ((''N. rafflesiana'' × ''N. ampullaria'') × ''N. mirabilis'')
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| − | *''N.'' × ''ventrata'' (''N. ventricosa'' × ''N. alata'')
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| − | *''N.'' × 'Bloody Mary' (''N. ventricosa'' × ''N. ampullaria'')
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| − | *''N.'' 'D'mato' (''N. lowii'' × ''N. ventricosa'')
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| − | *''N.'' × ''mixta'' (''N. northiana'' × ''N. maxima'')
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| − | *''N.'' 'Syurga' (''N. ventricosa'' × ''N. northiana'')
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| − | *''N.'' 'Menarik' (''N. rafflesiana'' × ''N. veitchii'')
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| − | *''N.'' 'Emmarene' (''N. khasiana'' × ''N. ventricosa'')
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| − | *''N.'' 'Judith Finn' (''N. spathulata'' × ''N. veitchii'')
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| − | ==See also==
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| − | * [[Nepenthes classification|''Nepenthes'' classification]]
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| − | * [[Nepenthes infauna|''Nepenthes'' infauna]]
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| | | | |
| | ==References== | | ==References== |
| − | {{Reflist|2}}
| + | <references/> |
| − | * [http://www.omnisterra.com/botany/cp/pictures/nepenthe/dansermg/dans10.htm Danser's Monograph on ''Nepenthes''] (covers species from Malaysia, Indonesia and New Guinea, but not elsewhere)
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| − | * [http://delta-intkey.com/angio/www/nepentha.htm Nepenthaceae] in: Watson, L., and M. J. Dallwitz (1992 onwards). [http://delta-intkey.com/angio The Families of Flowering Plants]. Descriptions, Illustrations, Identification, Information Retrieval.
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| − | | |
| − | ==Further reading==
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| − | {{refbegin|2}}
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| − | * {{cite journal | last1 = Amagase | first1 = S. | last2 = Nakayama | first2 = S. | last3 = Tsugita | first3 = A. | year = 1969 | title = Acid protease in ''Nepenthes''. II. Study on the specificity of nepenthesin | url = | journal = The Journal of Biochemistry | volume = 66 | issue = 4| pages = 431–439 }}
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| − | * {{cite journal | last1 = Athauda | first1 = S.B.P. | last2 = Matsumoto | first2 = K. | last3 = Rajapakshe | first3 = S. | last4 = Kuribayashi | first4 = M. | last5 = Kojima | first5 = M. | last6 = Kubomura-Yoshida | first6 = N. | last7 = Iwamatsu | first7 = A. | last8 = Shibata | first8 = C. | last9 = Inoue | first9 = H. | last10 = Takahashi | first10 = K. | year = 2004 | title = Enzymatic and structural characterization of nepenthesin, a unique member of a novel subfamily of aspartic proteinases | url = http://www.biochemj.org/bj/imps_x/pdf/BJ20031575.pdf | format = PDF | journal = Biochemical Journal | volume = 381 | issue = 1| pages = 295–306 | doi = 10.1042/BJ20031575 | pmid=15035659 | pmc=1133788}}
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| − | * {{cite journal | last1 = Bauer | first1 = U. | last2 = Bohn | first2 = H.F. | last3 = Federle | first3 = W. | year = 2008 | title = Harmless nectar source or deadly trap: ''Nepenthes'' pitchers are activated by rain, condensation and nectar | journal = Proceedings of the Royal Society B | volume = 275 | issue = 1632| pages = 259–265 | doi=10.1098/rspb.2007.1402 | pmid=18048280 | pmc=2593725}}
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| − | * {{cite journal | last1 = Beaver | first1 = R.A. | year = 1979 | title = Biological studies of the fauna of pitcher plants ''Nepenthes'' in west Malaysia | url = | journal = Annales de la Société Entomologique de France | volume = 15 | issue = | pages = 3–17 }}
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| − | * 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.
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| − | * {{cite journal | last1 = Beaver | first1 = R.A. | year = 1985 | title = Geographical variation in food web structure in ''Nepenthes'' pitcher plants | url = | journal = Ecological Entomology | volume = 10 | issue = 3| pages = 241–248 | doi=10.1111/j.1365-2311.1985.tb00720.x}}
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| − | * {{cite journal | last1 = Beekman | first1 = E.M. | year = 2004 | title = A Note on the Priority of Rumphius' Observation of Decapod Crustacea Living In ''Nepenthes'' | url = | journal = Crustaceana | volume = 77 | issue = 8| pages = 1019–1021 | doi = 10.1163/1568540042781748 }}
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| − | * {{cite journal | last1 = Bohn | first1 = H.F. | last2 = Federle | first2 = W. | year = 2004 | title = Insect aquaplaning: ''Nepenthes'' pitcher plants capture prey with the peristome, a fully wettable water-lubricated anisotropic surface | url = http://www.pnas.org/cgi/reprint/101/39/14138.pdf | format = PDF | journal = [[Proceedings of the National Academy of Sciences]] | volume = 101 | issue = 39| pages = 14138–14143 | doi = 10.1073/pnas.0405885101 | pmid=15383667 | pmc=521129}}
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| − | * {{fr icon}} Boulay, J. 1997. [http://dionee.gr.free.fr/bulletin/txt/d_38_c.htm Les ''Nepenthes'']. ''[[Dionée]]'' '''38'''.
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| − | * {{cite journal | last1 = Carlquist | first1 = S | year = 1981 | title = Wood Anatomy of Nepenthaceae | url = | journal = Bulletin of the Torrey Botanical Club | volume = 108 | issue = 3| pages = 324–330 | doi = 10.2307/2484711 }}
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| − | * {{cite journal | last1 = Chia | first1 = T.F. | last2 = Aung | first2 = H.H. | last3 = Osipov | first3 = A.N. | last4 = Goh | first4 = N.K. | last5 = Chia | first5 = L.S. | year = 2004 | title = Carnivorous pitcher plant uses free radicals in the digestion of prey | url = | journal = Redox Report | volume = 9 | issue = 5| pages = 255–261 | doi = 10.1179/135100004225006029 | pmid=15606978}}
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| − | * {{cite journal | last1 = Edwards | first1 = P | year = 2005 | title = Growing ''Nepenthes'' – Part 1 | url = https://www.vcps.org/journals/75_Mar_2005Public.pdf | format = PDF | journal = [[Victorian Carnivorous Plant Society Inc.]] | volume = 75 | issue = | pages = 6–8 }}
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| − | * {{cite journal | last1 = Edwards | first1 = P | year = 2005 | title = Growing ''Nepenthes'' – Part 2 | url = https://www.vcps.org/journals/76_Jun_2005Public.pdf | format = PDF | journal = [[Victorian Carnivorous Plant Society Inc.]] | volume = 76 | issue = | pages = 6–9 }}
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| − | * {{cite journal | last1 = Frazier | first1 = C.K. | year = 2000 | title = ''The Enduring Controversies Concerning the Process of Protein Digestion in ''Nepenthes | url = http://www.carnivorousplants.org/cpn/samples/Science292Digest.htm | journal = [[Carnivorous Plant Newsletter]] | volume = 29 | issue = 2| pages = 56–61 }}
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| − | * {{cite journal | last1 = Jenkin | first1 = A | year = 2005 | title = ''Nepenthes'' pollination | url = http://www.vcps.au.com/journals/75_Mar_2005Public.pdf | format = PDF | journal = [[Victorian Carnivorous Plant Society Inc.]] | volume = 75 | issue = | pages = 12–13 | deadurl = yes | archiveurl = https://web.archive.org/web/20110707185305/http://www.vcps.au.com/journals/75_Mar_2005Public.pdf | archivedate = 2011-07-07 | df = }}
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| − | * {{cite journal | last1 = Jennings | first1 = D.E. | last2 = Rohr | first2 = JR. | year = 2011 | title = A review of the conservation threats to carnivorous plants | url = | journal = Biological Conservation | volume = 144| issue = 5| pages = 1356–1363| doi = 10.1016/j.biocon.2011.03.013 }}
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| − | * {{cite journal | last1 = Karagatzides | first1 = J.D. | last2 = Ellison | first2 = A.M. | year = 2009 | title = Construction costs, payback times, and the leaf economics of carnivorous plants | url = | journal = American Journal of Botany | volume = 96 | issue = 9| pages = 1612–1619 | doi = 10.3732/ajb.0900054 | pmid=21622347}}
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| − | * {{cite journal | last1 = Meimberg | first1 = H. | last2 = Wistuba | first2 = A. | last3 = Dittrich | first3 = P. | last4 = Heubl | first4 = G. | year = 2001 | title = Molecular Phylogeny of Nepenthaceae Based on Cladistic Analysis of Plastid trnK Intron Sequence Data | url = | journal = Plant Biology (Stuttgart, Germany) | volume = 3 | issue = 2| pages = 164–175 | doi = 10.1055/s-2001-12897 }}
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| − | * {{cite journal | last1 = Mithöfer | first1 = A | year = 2011 | title = Carnivorous pitcher plants: insights in an old topic | url = | journal = Phytochemistry | volume = 72 | issue = 13| pages = 1678–1682 | doi = 10.1016/j.phytochem.2010.11.024 | pmid=21185041}}
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| − | * {{cite journal | last1 = Mokkamul | first1 = P. | last2 = Chaveerach | first2 = A. | last3 = Sudmoon | first3 = R. | last4 = Tanee | first4 = T. | year = 2007 | title = Species Identification and Sex Determination of the Genus ''Nepenthes'' (Nepenthaceae) | journal = Pakistan Journal of Biological Sciences | volume = 10 | issue = 4| pages = 561–567 | doi = 10.3923/pjbs.2007.561.567 | pmid=19069535}}
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| − | * {{cite journal | last1 = Moran | first1 = J.A. | last2 = Booth | first2 = W.E. | last3 = Charles | first3 = J.K. | year = 1999 | title = Aspects of Pitcher Morphology and Spectral Characteristics of Six Bornean ''Nepenthes'' Pitcher Plant Species: Implications for Prey Capture | url = http://aob.oxfordjournals.org/cgi/reprint/83/5/521.pdf | format = PDF | journal = Annals of Botany | volume = 83 | issue = 5| pages = 521–528 | doi=10.1006/anbo.1999.0857}}
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| − | * {{cite journal | last1 = Nosonovsky | first1 = M | year = 2011 | title = Materials science: slippery when wetted | url = | journal = Nature | volume = 477 | issue = 7365| pages = 412–413 | doi = 10.1038/477412a | pmid=21938059}}
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| − | * {{cite journal | last1 = Osunkoya | first1 = O.O. | last2 = Daud | first2 = S.D. | last3 = Di-Giusto | first3 = B. | last4 = Wimmer | first4 = F.L. | last5 = Holige | first5 = T.M. | year = 2007 | title = Construction Costs and Physico-chemical Properties of the Assimilatory Organs of ''Nepenthes'' Species in Northern Borneo | url = | journal = Annals of Botany | volume = 99 | issue = 5| pages = 895–906 | doi = 10.1093/aob/mcm023 }}
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| − | * {{cite journal | last1 = Pavlovič | first1 = A. | last2 = Masarovičová | first2 = E. | last3 = Hudák | first3 = J. | year = 2007 | title = Carnivorous Syndrome in Asian Pitcher Plants of the Genus ''Nepenthes'' | url = | journal = Annals of Botany | volume = 100 | issue = 3| pages = 527–536 | doi = 10.1093/aob/mcm145 | pmid=17664255 | pmc=2533595}}
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| − | * {{cite journal | last1 = Poppinga | first1 = S. | last2 = Koch | first2 = K. | last3 = Bohn | first3 = H.F. | last4 = Barthlott | first4 = W. | year = 2010 | title = Comparative and functional morphology of hierarchically structured anti-adhesive surfaces in carnivorous plants and kettle trap flowers | url = | journal = Functional Plant Biology | volume = 37 | issue = 10| pages = 952–961 | doi = 10.1071/FP10061 }}
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| − | * {{cite journal | last1 = Riedel | first1 = M. | last2 = Eichner | first2 = A. | last3 = Meimberg | first3 = H. | last4 = Jetter | first4 = R. | year = 2007 | title = Chemical composition of epicuticular wax crystals on the slippery zone in pitchers of five ''Nepenthes'' species and hybrids | url = | journal = Planta | volume = 225 | issue = 6| pages = 1517–1534 | doi = 10.1007/s00425-006-0437-3 | pmid=17109149}}
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| − | * {{cite journal | last1 = Schulze | first1 = W. | last2 = Frommer | first2 = W.B. | last3 = Ward | first3 = J.M. | year = 1999 | title = ''Transporters for ammonium, amino acids and peptides are expressed in pitchers of the carnivorous plant ''Nepenthes | url = http://www.blackwell-synergy.com/doi/pdf/10.1046/j.1365-313x.1999.00414.x | journal = The Plant Journal | volume = 17 | issue = 6| pages = 637–646 | doi = 10.1046/j.1365-313x.1999.00414.x }}
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| − | * {{cite journal | last1 = Vines | first1 = S.H. | year = 1876 | title = ''On the Digestive Ferment of ''Nepenthes | journal = Journal of Anatomy and Physiology | volume = 11 | issue = 1| pages = 124–127 | pmc=1309768 | pmid=17231131}}
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| − | * {{cite journal | last1 = Wong | first1 = T.-S. | last2 = Kang | first2 = S.H. | last3 = Tang | first3 = S.K.Y. | last4 = Smythe | first4 = E.J. | last5 = Hatton | first5 = B.D. | last6 = Grinthal | first6 = A. | last7 = Aizenberg | first7 = J. | year = 2011 | title = Bioinspired self-repairing slippery surfaces with pressure-stable omniphobicity | url = | journal = Nature | volume = 477 | issue = 7365| pages = 443–447 | doi = 10.1038/nature10447 | pmid=21938066}}
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| − | <!--Goddard, M. 1995. Why Latin? ''[[CPS News]]'' 1995(4): 6–7.-->
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| − | {{Refend}}
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| − | | |
| − | ==External links==
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| − | {{Wikispecies}}
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| − | {{Commons}}
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| − | * [http://www.botany.org/Carnivorous_Plants/Nepenthes.php ''Nepenthes'' - the Monkey Cups] from the [[Botanical Society of America]]
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| − | * [https://www.carnivorousplants.co.uk/resources/nepenthes-interactive-guide/ ''Nepenthes'': The Interactive Guide] at CarnivorousPlants.co.uk
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| − | * [http://www.cpphotofinder.com/Nepenthes.html ''Nepenthes'' photographs] at the Carnivorous Plant Photo Finder
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| − | * [https://www.youtube.com/watch?v=trWzDlRvv1M A video about ''Nepenthes rajah''] from ''[[The Private Life of Plants]]''
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| − | * [http://www.sarracenia.com/faq/faq5400.html The Carnivorous Plant FAQ: ''Nepenthes''] by [[Barry Rice (botanist)|Barry Rice]]
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| − | * [http://www.carnivorousplants.org/cp/EvolutionNepenthes.php Evolution -- ''Nepenthes'' Phylogeny] from the [[International Carnivorous Plant Society]]
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| − | * [https://wikis.nbi.ac.uk/InnerWorlds/index.php/Nepenthes_species#.UThsWRyePAk Inner World of ''Nepenthes'' from the John Innes Centre]
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| − | {{Nepenthes}}
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| − | {{CarnivorousPlants}}
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| − | {{Taxonbar|from=Q217530}}
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| − | {{Authority control}}
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| | | | |
| − | [[Category:Nepenthes| ]]
| + | [[Category:Caryophyllales]] |
| − | [[Category:Caryophyllales genera]] | + | [[Category:Plants for Keenan to eat]] |
| − | [[Category:Dioecious plants]]
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| − | [[Category:Taxa named by Carl Linnaeus]]
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| − | [[Category:Articles containing video clips]] | |
