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Pitcher plants are several different
carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants still generate some of their energy from photosynthesis. Car ...
s which have modified leaves known as pitfall traps—a prey-trapping mechanism featuring a deep cavity filled with digestive
liquid A liquid is a nearly incompressible fluid that conforms to the shape of its container but retains a (nearly) constant volume independent of pressure. As such, it is one of the four fundamental states of matter (the others being solid, gas, an ...
. The traps of what are considered to be "true" pitcher plants are formed by specialized leaves. The plants attract and drown their prey with nectar.


Types

The term "pitcher plant" generally refers to members of the
Nepenthaceae ''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mo ...
and
Sarraceniaceae Sarraceniaceae are a family of pitcher plants, belonging to order Ericales (previously Nepenthales). The family comprises three extant genera: '' Sarracenia'' (North American pitcher plants), '' Darlingtonia'' (the cobra lily or California pi ...
families, but similar pitfall traps are employed by the monotypic
Cephalotaceae ''Cephalotus'' ( or ; Greek: ''κεφαλή'' "head", and ''οὔς''/''ὠτός'' "ear", to describe the head of the anthers) is a genus which contains one species, ''Cephalotus follicularis'' the Albany pitcher plant, a small carnivorous pit ...
and some members of the
Bromeliaceae The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...
. The families
Nepenthaceae ''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mo ...
and
Sarraceniaceae Sarraceniaceae are a family of pitcher plants, belonging to order Ericales (previously Nepenthales). The family comprises three extant genera: '' Sarracenia'' (North American pitcher plants), '' Darlingtonia'' (the cobra lily or California pi ...
are the most species-rich families of pitcher plants. The Nepenthaceae contains a single
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencla ...
, ''
Nepenthes ''Nepenthes'' () is a genus of carnivorous plants, also known as tropical pitcher plants, or monkey cups, in the monotypic family Nepenthaceae. The genus includes about 170 species, and numerous natural and many cultivated hybrids. They are mos ...
'', containing over 100 species and numerous hybrids and cultivars. In this genus of
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by th ...
pitcher plants, the pitchers are borne at the end of
tendril In botany, a tendril is a specialized stem, leaf or petiole with a threadlike shape used by climbing plants for support and attachment, as well as cellular invasion by parasitic plants such as ''Cuscuta''. There are many plants that have tend ...
s that extend from the
midrib This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary ...
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 An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s on trees. The New World pitcher plants (Sarraceniaceae), which comprise three genera, are ground-dwelling herbs whose pitchers arise from a horizontal
rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow ho ...
. 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 The genus ''Heliamphora'' ( or ; Greek: ''helos'' "marsh" and ''amphoreus'' " amphora") contains 23 species of pitcher plants endemic to South America.McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. '' Sarraceniaceae of South America'' ...
'', 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 North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the C ...
n genus ''
Sarracenia ''Sarracenia'' ( or ) is a genus comprising 8 to 11 species of North American pitcher plants, commonly called trumpet pitchers. The genus belongs to the family Sarraceniaceae, which also contain the closely allied genera '' Darlingtonia'' and ...
'' are the trumpet pitchers, which have a more complex trap than ''Heliamphora'', with an operculum, which prevents excess accumulation of rainwater in most of the species. The single species in the Californian genus '' 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 damn pitcher. The species in the genus ''Sarracenia'' readily hybridize, making their classification a complex matter. The
Cephalotaceae ''Cephalotus'' ( or ; Greek: ''κεφαλή'' "head", and ''οὔς''/''ὠτός'' "ear", to describe the head of the anthers) is a genus which contains one species, ''Cephalotus follicularis'' the Albany pitcher plant, a small carnivorous pit ...
is a monotypic family with but one genus and species, ''
Cephalotus follicularis ''Cephalotus'' ( or ; Greek: ''κεφαλή'' "head", and ''οὔς''/''ὠτός'' "ear", to describe the head of the anthers) is a genus which contains one species, ''Cephalotus follicularis'' the Albany pitcher plant, a small carnivorous pit ...
''. This species has a small (2–5  cm) pitcher similar in form to those of ''Nepenthes''. It occurs in only one location in southwestern Australia. A few species of bromeliads (
Bromeliaceae The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...
), such as ''
Brocchinia reducta ''Brocchinia reducta'' is one of a few carnivorous bromeliads. It is native to southern Venezuela, Brazil, Colombia, and Guyana, and is found in nutrient-poor soil. ''B. reducta'' adapts to different environments, when growing on rocks it uses ...
'' and ''
Catopsis berteroniana ''Catopsis berteroniana'' , commonly known as the powdery strap airplant or the lantern of the forest, is an epiphytic bromeliad thought to be a possible carnivorous plant, similar to '' Brocchinia reducta'', although the evidence is equivocal. I ...
'', are known or suspected to be carnivorous. Bromeliads are
monocots Monocotyledons (), commonly referred to as monocots, ( Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
, and given that they all naturally collect water where their leaves meet each other, and that many collect detritus, it is not surprising that a few should have been naturally selected to develop the habit into carnivory by the addition of
wax Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
and downward-pointing
hair Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and fin ...
s. The purple pitcher plant, ''
Sarracenia purpurea ''Sarracenia purpurea'', the purple pitcher plant, northern pitcher plant, turtle socks, or side-saddle flower, is a carnivorous plant in the family Sarraceniaceae. Description Like other species of '' Sarracenia'', ''S. purpurea'' obtains mos ...
'', is the
floral emblem In a number of countries, plants have been chosen as symbols to represent specific geographic areas. Some countries have a country-wide floral emblem; others in addition have symbols representing subdivisions. Different processes have been used to ...
of the province of
Newfoundland and Labrador Newfoundland and Labrador (; french: Terre-Neuve-et-Labrador; frequently abbreviated as NL) is the easternmost province of Canada, in the country's Atlantic region. The province comprises the island of Newfoundland and the continental region ...
, Canada.


Feeding behavior

Foraging, flying, or crawling
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of ...
s such as flies are attracted to a cavity formed by the cupped leaf, often by visual lures such as
anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical com ...
pigments, and
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualis ...
. 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 In organic chemistry, an aldehyde () is an organic compound containing a functional group with the structure . The functional group itself (without the "R" side chain) can be referred to as an aldehyde but can also be classified as a formyl group ...
crystals,
cuticular A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
folds, downward-pointing hairs, or guard-cell-originating
lunate Lunate is a crescent or moon-shaped microlith. In the specialized terminology of lithic reduction, a lunate flake is a small, crescent-shaped flake removed from a stone tool during the process of pressure flaking. In the Natufian period, a luna ...
cells, to help prevent escape. The small bodies of liquid contained within the pitcher traps are called
phytotelmata Phytotelma (plural phytotelmata) is a small water-filled cavity in a terrestrial plant. The water accumulated within these plants may serve as the habitat for associated fauna and flora. A rich literature in German summarised by Thienemann (19 ...
. 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 Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. ...
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. Some pitcher plants contain mutualistic insect
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...
e, which feed on trapped prey, and whose
excreta Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks after lea ...
the plant absorbs. Whatever the mechanism of digestion, the prey items are converted into a solution of
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
s,
peptide Peptides (, ) are short chains of amino acids linked by peptide bonds. Long chains of amino acids are called proteins. Chains of fewer than twenty amino acids are called oligopeptides, and include dipeptides, tripeptides, and tetrapeptides. ...
s,
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
s,
ammonium The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaterna ...
and
urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important ...
, from which the plant obtains its mineral nutrition (particularly
nitrogen Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at se ...
and
phosphorus Phosphorus is a chemical element with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ea ...
). Like all carnivorous plants, they all grow in locations where the
soil Soil, also commonly referred to as earth or dirt, is a mixture of organic matter, minerals, gases, liquids, and organisms that together support life. Some scientific definitions distinguish ''dirt'' from ''soil'' by restricting the former ...
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. Mature plants of ''
Nepenthes lowii ''Nepenthes lowii'' , or Low's pitcher-plant,Phillipps, A. & A. Lamb 1996. '' Pitcher-Plants of Borneo''. Natural History Publications (Borneo), Kota Kinabalu. is a tropical pitcher plant endemic to Borneo. It is named after Hugh Low, who discove ...
'' attract
tree shrew The treeshrews (or tree shrews or banxrings) are small mammals native to the tropical forests of South and Southeast Asia. They make up the entire order Scandentia, which split into two families: the Tupaiidae (19 species, "ordinary" treeshrews ...
s (
Tupaia montana The mountain treeshrew (''Tupaia montana'') is a treeshrew species within the Tupaiidae. It is endemic to Borneo and inhabits montane forests in Sarawak and Sabah. The first specimen was described by Oldfield Thomas and was part of a zoolog ...
), which feed on nectar that the plant produces but also defecate into the pitcher, providing nitrates and other nutrients. The plant and tree shrew have a
symbiotic relationship Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
. 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.


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), with
selection Selection may refer to: Science * Selection (biology), also called natural selection, selection in evolution ** Sex selection, in genetics ** Mate selection, in mating ** Sexual selection in humans, in human sexuality ** Human mating strateg ...
pressure favouring more deeply cupped leaves over
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary time. The pitcher trap evolved independently in three
eudicot The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dico ...
lineages and one
monocot Monocotyledons (), commonly referred to as monocots, (Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one ...
lineage, representing a case of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Some pitcher plant families (such as Nepenthaceae) are placed within clades consisting mostly of flypaper traps, indicating that some pitchers may have evolved from the common ancestors of today's flypaper traps by loss of mucilage.


See also

* Darlingtonia State Natural Site – A nature preserve for pitcher plants in Oregon, United States


References

* 10. * 11. Bazile, Vincent; Le Moguédec, Gilles; Marshall, David J.; Gaume, Laurence (2015-03-01). "Fluid physico-chemical properties influence capture and diet in Nepenthes pitcher plants". ''Annals of Botany''. 115 (4): 705–716. doi:10.1093/aob/mcu266.
ISSN An International Standard Serial Number (ISSN) is an eight-digit serial number used to uniquely identify a serial publication, such as a magazine. The ISSN is especially helpful in distinguishing between serials with the same title. ISSNs a ...
 0305-7364. PMC 4343297.
PMID PubMed is a free search engine accessing primarily the MEDLINE database of references and abstracts on life sciences and biomedical topics. The United States National Library of Medicine (NLM) at the National Institutes of Health maintain the ...
 25672361.


Further reading

* Juniper, B.E., R.J. Robins & D.M. Joel (1989). '' The Carnivorous Plants''. Academic Press, London. * Schnell, D. (2003). ''Carnivorous Plants of the United States and Canada''. Second Edition. Timber Press, Oregon, U.S.A.


External links


How does a pitcher plant attract, catch and trap insects

Carnivorous plants can photosynthesise, so why eat flies?
{{DEFAULTSORT:Pitcher Plant Carnivorous plants Plant common names