Editing Pitcher plant

{{short description|Carnivorous plant}}{{More citations needed|date=July 2024}}[[File:Pitcher plant diagram ency brit.png|thumb|right|upright=1.36|Pitcher of ''[[Nepenthes distillatoria]]''. '''A:''' Honey-glands from attractive surface of lid. '''B:''' Digestive fluid from interior of pitcher, in pocket-like depression of epidermis, opening downwards. '''C:''' Transverse section of the same.]]
[[File:Pitcher plant trichomes pollen.jpg|thumb|right|[[Scanning electron micrograph]] inner surface of pitcher plant 

]]
[[File:Pitcher plants in the Linn Run bog..jpg|thumb|right|Pitcher plants growing in a bog in Pennsylvania]]

'''Pitcher plants''' are  [[carnivorous plant]]s 
known as '''pitfall traps'''—a prey-trapping mechanism featuring a deep cavity filled with digestive [[liquid]]. The traps of pitcher plant are considered to be "true" pitcher plants and are formed by specialized leaves. The plants attract and drown the prey with [[nectar]].<ref name="Krol-etal-2011">{{Cite journal | year = 2011
 | last1 = Krol | first1 = E.
 | last2 = Plancho | first2 = B. J.
 | last3 = Adamec | first3 = L.
 | last4 = Stolarz | first4 = M. | last5 = Dziubinska | first5 = H. | last6 =Trebacz | first6 = Kgv.
 | title = Quite a few reasons for calling carnivores 'the most wonderful plants in the world'
 | journal = Annals of Botany | volume = 109 | issue = 1 | pages = 47–64
 | doi =  10.1093/aob/mcr249 | pmid=21937485 | pmc=3241575}}</ref>

== Types ==
The term "pitcher plant" generally refers to members of the [[Nepenthaceae]] and [[Sarraceniaceae]] families, but similar pitfall traps are employed by the monotypic [[Cephalotaceae]] and some members of the [[Bromeliaceae]]. The families [[Nepenthaceae]] and [[Sarraceniaceae]] are the most species-rich families of pitcher plants.<ref>{{Cite web |title=The evolution of carnivorous pitcher plants |url=https://www.obga.ox.ac.uk/evolution-carnivorous-pitcher-plants |access-date=2025-02-14 |website=www.obga.ox.ac.uk |language=en}}</ref>

=== Nepenthaceae ===
The Nepenthaceae contains a single [[genus]], ''[[Nepenthes]]'', containing over 100 species and numerous hybrids and cultivars. In this genus of [[Old World]] pitcher plants, the pitchers are borne at the end of [[tendril]]s that extend from the [[midrib]] of an otherwise unexceptional leaf. Old World pitcher plants are typically characterized as having reduced and symmetrical pitchers with a comprehensive waxy coating on the surface of the inner pitcher wall. The plants themselves are often climbers, accessing the canopy of their habitats using the aforementioned tendrils, although others are found on the ground in forest clearings, or as [[epiphyte]]s on trees.<ref>{{cite journal |last1=Clarke |first1=Charles |last2=Moran |first2=Jonathan |date=2015 |title=Climate, soils and vicariance - their roles in shaping the diversity and distribution of Nepenthes in Southeast Asia |url=https://link.springer.com/article/10.1007/s11104-015-2696-x |access-date= |journal=Plant and Soil |volume=403 |issue=1–2 |pages=37–51 |publisher=Springer International Publishing|doi=10.1007/s11104-015-2696-x }}</ref>
=== Sarraceniaceae ===

The New World pitcher plants (Sarraceniaceae), which comprise three genera, are ground-dwelling herbs whose pitchers arise from a horizontal [[rhizome]]. In this family, the entire leaf forms the pitcher, as opposed to Nepenthaceae, where the pitcher arises from the terminal portion of the leaf. The species of the genus ''[[Heliamphora]]'', which are popularly known as marsh pitchers (or erroneously as sun pitchers), have a simple rolled-leaf pitcher, at the tip of which is a spoon-like structure that secretes nectar. They are restricted to areas of high rainfall in [[South America]]. The [[North America]]n genus ''[[Sarracenia]]'' are the trumpet pitchers, which have a more complex trap than ''Heliamphora'', with an [[Operculum (Botany)|operculum]], which prevents excess accumulation of rainwater in most of the species. The single species in the [[California]] genus ''[[Darlingtonia (plant)|Darlingtonia]]'' is popularly known as the cobra plant, due to its possession of an inflated "lid" with elegant false-exits, and a forked "tongue", which serves to ferry ants and other prey to the entrance of the pitcher. The species in the genus ''Sarracenia'' readily hybridize, making their classification a complex matter.{{citation needed|date=October 2023}}

The purple pitcher plant, ''[[Sarracenia purpurea]]'', is the [[floral emblem]] of the province of [[Newfoundland and Labrador]], Canada.

=== Cephalotaceae ===
The [[Cephalotaceae]] is a monotypic family with but one genus and species, ''[[Cephalotus|Cephalotus follicularis]]''. This species has a small (2–5&nbsp;cm) pitcher similar in form to those of ''Nepenthes''. Unlike in ''Nepenthes'', in ''Cephalotus'' ''follicularis'' the petiole is attached to the rear of the upper trap rim rather than to the base of the pitcher.<ref name=":0">{{Cite book |author=Wilhelm Barthlott |translator=Michael Ashdown  |url=https://www.worldcat.org/oclc/71189676 |title=The curious world of carnivorous plants : a comprehensive guide to their biology and cultivation |date=2007 |publisher=Timber Press |isbn=978-0-88192-792-4 |edition=English language |location=Portland, Ore. |oclc=71189676}}</ref> The species occurs in only one location in southwestern [[Australia]].{{citation needed|date=October 2023}}

=== Bromeliaceae ===
A few species of bromeliads ([[Bromeliaceae]]), such as ''[[Brocchinia reducta]]''  and ''[[Catopsis berteroniana]]'', are known or suspected to be carnivorous.<ref>{{Cite journal |last1=Givnish |first1=Thomas J. |last2=Burkhardt |first2=Elizabeth L. |last3=Happel |first3=Ruth E. |last4=Weintraub |first4=Jason D. |date=1984 |title=Carnivory in the Bromeliad Brocchinia reducta, with a Cost/Benefit Model for the General Restriction of Carnivorous Plants to Sunny, Moist, Nutrient-Poor Habitats |url=https://doi.org/10.1086/284289 |journal=The American Naturalist |volume=124 |issue=4 |pages=479–497|doi=10.1086/284289 |s2cid=84947503 }}</ref>

{{Gallery| File:Nepenthes muluensis.jpg|''[[Nepenthes muluensis]]'' pitchers hang from tendrils. (This specimen is cultivated.)
 | File:kew.gardens.pitcher.plant.sarracenia.arp.jpg|Most North American pitcher plants belong to the genus ''[[Sarracenia]]''.
 | File:Darlingtonia californica ne8.JPG|Cobra lilies (''[[Darlingtonia californica]]'') use window-like areolae to lure insects into their hollow leaves.
 | File:Cephalotus follicularis001.jpg|The Australian pitcher plant is the only member of the Australian genus ''[[Cephalotus]]''.
 | File:Nepenthes truncata Exhibition of Carnivorous Plants Prague 2016 2.jpg|''[[Nepenthes truncata]]''
 | File:Nepenthes villosa.jpg|Pitcher of ''[[Nepenthes villosa]]''
}}

==Feeding behavior==

=== Attraction ===
Foraging, flying, or crawling [[insect]]s such as flies are attracted to a cavity formed by the cupped leaf, often by visual lures such as [[anthocyanin]] pigments, and [[nectar]]. Many pitcher plants exhibit patterns of ultraviolet coloration which may play a role in attracting insects.<ref name=":0" /> Some species, such as ''Cephalotus follicularis'', likely use camouflage to trap insects, as their coloration matches that of the surrounding environment and the plants are often embedded in the substrate such that the traps are flush with the ground.<ref name=":0" />

Olfactory cues can also play a role in attraction. For example, ''Nepenthes rafflesian''a uses flower-scent mimicry to attract insects to its pitchers.<ref>{{Cite journal |last1=Di Giusto |first1=Bruno |last2=Bessière |first2=Jean-Marie |last3=Guéroult |first3=Michaël |last4=Lim |first4=Linda B. L. |last5=Marshall |first5=David J. |last6=Hossaert-McKey |first6=Martine |last7=Gaume |first7=Laurence |date=2010-04-28 |title=Flower-scent mimicry masks a deadly trap in the carnivorous plant Nepenthes rafflesiana: Flower-scent mimicry in Nepenthes rafflesiana |journal=Journal of Ecology |language=en |volume=98 |issue=4 |pages=845–856 |doi=10.1111/j.1365-2745.2010.01665.x|s2cid=82299236 |doi-access=free }}</ref>

=== Capture ===
The rim of the pitcher (peristome) is slippery when moistened by condensation or nectar, causing insects to fall into the trap. The walls of the pitfall may be covered with waxy scales, protruding [[aldehyde]] crystals, [[Plant cuticle|cuticular]] folds, downward-pointing hairs, or guard-cell-originating [[wikt:lunate|lunate]] cells, to help prevent escape.<ref name="Krol-etal-2011" /> The small bodies of liquid contained within the pitcher traps are called [[phytotelmata]]. They drown the insect, whose body is gradually dissolved. This may occur by bacterial action (the bacteria being washed into the pitcher by rainfall), or by digestive [[enzyme]]s secreted by the plant itself. Pitcher trap fluids largely vary in their viscoelasticity and acidity, which then dictates which type of prey they can target. For example, increased viscoelasticity is associated with increased insect retention to help capture flying insects such as flies, whereas increased fluid acidity can decrease insect killing-time, which can help capture crawling insects such as ants.<ref>{{Cite journal |last1=Bazile |first1=Vincent |last2=Le Moguédec |first2=Gilles |last3=Marshall |first3=David J. |last4=Gaume |first4=Laurence |date=2015-03-01 |title=Fluid physico-chemical properties influence capture and diet in Nepenthes pitcher plants |url=https://doi.org/10.1093/aob/mcu266 |journal=Annals of Botany |volume=115 |issue=4 |pages=705–716 |doi=10.1093/aob/mcu266 |issn=0305-7364 |pmc=4343297 |pmid=25672361}}</ref> Some pitcher plants contain [[Mutualism (biology)|mutualistic]] insect [[larva]]e, which feed on trapped prey, and whose [[excreta]] the plant absorbs.<ref>{{Cite journal | year = 1998
 | last = McAlpine | first = D.K.
 | title = Review of the Australian stilt flies (Diptera: Micropezidae) with a phylogenetic analysis of the family
 | journal = Invertebrate Taxonomy | volume = 12 | issue = 1 | pages = 55–134
 | doi = 10.1071/IT96018}}</ref>

=== Digestion ===
Whatever the mechanism of digestion, the prey items are converted into a solution of [[amino acid]]s, [[peptide]]s, [[phosphate]]s, [[ammonium]] and [[urea]], from which the plant obtains its mineral nutrition (particularly [[nitrogen]] and [[phosphorus]]). Like all carnivorous plants, pitcher plants all grow in locations where the [[soil]] is too poor in minerals and/or too acidic for most plants to survive. Pitcher plants supplement available nutrients and minerals (which plants normally obtain through their roots) with the constituents of their insect prey.{{citation needed|date=October 2023}}

=== Feces-trapping symbiosis===
Mature plants of ''[[Nepenthes lowii#Carnivory|Nepenthes lowii]]'' attract [[tree shrew]]s (''[[Tupaia montana]]''), which feed on nectar that the plant produces but also defecate into the pitcher, providing nitrates and other nutrients. The plant and tree shrew have a [[symbiosis|symbiotic relationship]]. The rim of ''N. lowii'' is not slippery so that tree shrews can easily get in and out; it provides more nectar than other pitcher plants. The shape of the pitcher rim and the position of the nectar ensure that the animal's hindquarters are over the rim while it feeds.<ref>[http://www.livescience.com/9666-pitcher-plant-doubles-toilet.html Pitcher Plant Doubles as Toilet]</ref>

''Nepenthes rafflesiana'' var. ''elongata'' has a similar relationship with [[Hardwicke's woolly bat]]s (''Kerivoula hardwickii'').<ref>{{Cite journal |last1=Grafe |first1=T. Ulmar |last2=Schöner |first2=Caroline R. |last3=Kerth |first3=Gerald |last4=Junaidi |first4=Anissa |last5=Schöner |first5=Michael G. |date=2011-06-23 |title=A novel resource–service mutualism between bats and pitcher plants |journal=Biology Letters |language=en |volume=7 |issue=3 |pages=436–439 |doi=10.1098/rsbl.2010.1141 |issn=1744-9561 |pmc=3097880 |pmid=21270023}}</ref> The bats roost inside the pitchers and the plants derive much of their foliar nitrogen from the feces of the bats. Compared to other varieties of ''Nepenthes rafflesiana'' that do not exhibit this form of mutualism, ''N. rafflesiana'' var. ''elongata'' has elongated pitchers that can accommodate both single bats and mother-juvenile pairs. As well as its elongated shape, ''N. rafflesiana'' var. ''elongata'' has reduced volumes of pitcher fluid compared to other species, leaving more space to accommodate the bats.

==Evolution of the form==
It is widely assumed pitfall traps evolved by epiascidiation (infolding of the leaf with the adaxial or upper surface becoming the inside of the pitcher),<ref name="Albert-etal-1992">{{Cite journal | year = 1992
 | last1 = Albert | first1 = V.A.
 | last2 = Williams | first2 = S.E.
 | last3 = Chase | first3 = M.W.
 | title = Carnivorous Plants: Phylogeny and Structural Evolution
 | journal = Science | volume = 257 | issue = 5076 | pages = 1491–1495
 | pmid = 1523408 | doi=10.1126/science.1523408| bibcode = 1992Sci...257.1491A }}</ref><ref>{{Cite journal | year = 1999
 | last1 = Owen Jr | first1 = T.P.
 | last2 = Lennon | first2 = K.A.
 | title = Structure and Development of Pitchers from the Carnivorous Plant ''Nepenthes alta'' (Nepenthaceae)
 | journal = American Journal of Botany | volume = 86 | issue = 10 | pages = 1382–1390
 | pmid = 10523280 | doi=10.2307/2656921| jstor = 2656921 }}</ref> with [[natural selection|selection]] pressure favouring more deeply cupped leaves over [[evolution]]ary time.  The pitcher trap evolved independently in three [[eudicot]] lineages and one [[monocot]] lineage, representing a case of [[convergent evolution]].<ref name="Albert-etal-1992" /> Some pitcher plant families (such as Nepenthaceae) are placed within clades consisting mostly of [[Carnivorous plant#Flypaper traps|flypaper traps]], indicating that some pitchers may have evolved from the common ancestors of today's flypaper traps by loss of mucilage.<ref>{{cite journal |last1=Pritchard|display-authors=etal|title=Evolutionary adaptations in pitcher plants |journal=International Journal of Evolutionary Biology |date=2002 |volume=12 |issue=3 |pages=62–81}}</ref>

==See also==
* [[Darlingtonia State Natural Site]] – A nature preserve for pitcher plants in [[Oregon]], [[United States]].
* [[Baghmara Pitcher Plant Wildlife Sanctuary]]

==References==

{{reflist}}
* {{Cite EB1911 |wstitle=Pitcher Plants |volume=21 |pages=663–665}}
* {{cite journal |last1=Bazile |first1=Vincent |last2=Le Moguédec |first2=Gilles |last3=Marshall |first3=David J. |last4=Gaume |first4=Laurence |date=2015-03-01 |title=Fluid physico-chemical properties influence capture and diet in Nepenthes pitcher plants |journal=Annals of Botany |volume=115 |issue=4 |pages=705–716 |doi=10.1093/aob/mcu266 |issn=0305-7364 |pmc=4343297 |pmid=25672361}}

==Further reading==
* Juniper, B.E., R.J. Robins & D.M. Joel (1989). ''[[The Carnivorous Plants (1989 book)|The Carnivorous Plants]]''. Academic Press, London. ISBN 9780123921703. OCLC 490279526.
* Schnell, D. (2003). ''Carnivorous Plants of the United States and Canada''. Second Edition. Timber Press, Portland, Ore., ISBN 9780881925401.

==External links==
* [http://gardenofeaden.blogspot.co.uk/2010/02/how-does-pitcher-plant-attract-catch.html How does a pitcher plant attract, catch and trap insects]
* [http://www.thenakedscientists.com/HTML/questions/question/2797/ Carnivorous plants can photosynthesise, so why eat flies?]

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{{CarnivorousPlants}}

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[[Category:Carnivorous plants]]
[[Category:Plant common names]]