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An upper pitcher of Nepenthes lowii, a tropical pitcher plant thatsupplements its carnivorous diet with tree shrew droppings.[1][2][3]

Carnivorous plantFrom Wikipedia, the free encyclopedia

"Insectivorous Plants" redirects here. For the book by Charles Darwin, see Insectivorous Plants(book).

Carnivorous plants are plants thatderive some or most of their nutrients(but not energy) from trapping andconsuming animals or protozoans,typically insects and other arthropods.Carnivorous plants have adapted togrow in places where the soil is thin orpoor in nutrients, especially nitrogen,such as acidic bogs and rockoutcroppings. Charles Darwin wroteInsectivorous Plants, the first well­known treatise on carnivorous plants, in1875.[4]

True carnivory is thought to haveevolved independently six times in fivedifferent orders of flowering plants,[5][6]and these are now represented by morethan a dozen genera. These include about 630 species that attract and trap prey, produce digestive enzymes,and absorb the resulting available nutrients.[7] Additionally, over 300 protocarnivorous plant species inseveral genera show some but not all of these characteristics.

Contents

1 Trapping mechanisms1.1 Pitfall traps1.2 Flypaper traps1.3 Snap traps1.4 Bladder traps1.5 Lobster­pot traps1.6 Combination traps1.7 Borderline carnivores

2 Evolution3 Ecology and modelling of carnivory4 Classification

4.1 Dicots4.2 Monocots

5 Cultivation6 Medicinal uses7 Cultural depictions

8 See also

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The pitchers of Heliamphorachimantensis are an example of pitfalltraps.

8 See also9 References10 Further reading

Trapping mechanisms

Five basic trapping mechanisms are found in carnivorous plants.

1. Pitfall traps (pitcher plants) trap prey in a rolled leaf thatcontains a pool of digestive enzymes or bacteria.

2. Flypaper traps use a sticky mucilage.3. Snap traps utilize rapid leaf movements.4. Bladder traps suck in prey with a bladder that generates an

internal vacuum.5. Lobster­pot traps force prey to move towards a digestive

organ with inward­pointing hairs.

These traps may be active or passive, depending on whethermovement aids the capture of prey. For example, Triphyophyllum isa passive flypaper that secretes mucilage, but whose leaves do notgrow or move in response to prey capture. Meanwhile, sundews areactive flypaper traps whose leaves undergo rapid acid growth, whichis an expansion of individual cells as opposed to cell division. Therapid acid growth allows the sundew tentacles to bend, aiding in theretention and digestion of prey.[8]

The sundew species Drosera glanduligera employs a uniquetrapping mechanism with features of both flypaper and snap traps;this has been termed a catapult­flypaper trap.[9]

Pitfall traps

Main article: Pitcher plant

Pitfall traps are thought to have evolved independently on at least four occasions. In general they arephytotelmata, water bodies collected or secreted into specialised containers, and ultimately held by plantsfor various functions such as in particular, the trapping and digestion of prey. The simplest ones areprobably those of Heliamphora, the marsh pitcher plant. In this genus, the traps are clearly derivedevolutionarily from a simple rolled leaf whose margins have sealed together. These plants live in areas ofhigh rainfall in South America such as Mount Roraima and consequently have a problem ensuring theirpitchers do not overflow. To counteract this problem, natural selection has favoured the evolution of anoverflow similar to that of a bathroom sink—a small gap in the zipped­up leaf margins allows excess waterto flow out of the pitcher.

Heliamphora is a member of the Sarraceniaceae, a New World family in the order Ericales (heathers andallies). Heliamphora is limited to South America, but the family contains two other genera, Sarracenia andDarlingtonia, which are endemic to the Southeastern United States (with the exception of one species) and

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Darlingtonia californica: note thesmall entrance to the trap underneaththe swollen "balloon" and thecolourless patches that confuse preytrapped inside.

Brocchinia reducta: a carnivorousbromeliad.

California respectively. Sarracenia purpurea subsp. purpurea (the northern pitcher plant) can be found asfar north as Canada. Sarracenia is the pitcher plant genus most commonly encountered in cultivation,because it is relatively hardy and easy to grow.

In the genus Sarracenia, the problem of pitcher overflow is solvedby an operculum, which is essentially a flared leaflet that covers theopening of the rolled­leaf tube and protects it from rain. Possiblybecause of this improved waterproofing, Sarracenia species secreteenzymes such as proteases and phosphatases into the digestive fluidat the bottom of the pitcher; Heliamphora relies on bacterialdigestion alone. The enzymes digest the proteins and nucleic acidsin the prey, releasing amino acids and phosphate ions, which theplant absorbs.

Darlingtonia californica, the cobra plant, possesses an adaptationalso found in Sarracenia psittacina and, to a lesser extent, inSarracenia minor: the operculum is balloon­like and almost sealsthe opening to the tube. This balloon­like chamber is pitted withareolae, chlorophyll­free patches through which light can penetrate.Insects, mostly ants, enter the chamber via the opening underneath

the balloon. Once inside, they tire themselves trying to escape from these false exits, until they eventuallyfall into the tube. Prey access is increased by the "fish tails", outgrowths of the operculum that give theplant its name. Some seedling Sarracenia species also have long, overhanging opercular outgrowths;Darlingtonia may therefore represent an example of neoteny.

The second major group of pitcher plants are the monkey cups ortropical pitcher plants of the genus Nepenthes. In the hundred or sospecies of this genus, the pitcher is borne at the end of a tendril,which grows as an extension to the midrib of the leaf. Most speciescatch insects, although the larger ones, such as Nepenthes rajah,also occasionally take small mammals and reptiles. Nepenthesbicalcarata possesses two sharp thorns that project from the base ofthe operculum over the entrance to the pitcher. These likely serve tolure insects into a precarious position over the pitcher mouth, wherethey may lose their footing and fall into the fluid within.[10]

The pitfall trap has evolved independently in at least two othergroups. The Albany pitcher plant Cephalotus follicularis is a smallpitcher plant from Western Australia, with moccasin­like pitchers.The rim of its pitcher's opening (the peristome) is particularlypronounced (both secrete nectar) and provides a thorny overhang tothe opening, preventing trapped insects from climbing out. Thelining of most pitcher plants is covered in a loose coating of waxyflakes, which are slippery for insects, prey that are often attracted bynectar bribes secreted by the peristome and by bright flower­likeanthocyanin patterning. In at least one species, Sarracenia flava, thenectar bribe is laced with coniine, a toxic alkaloid also found inhemlock, which probably increases the efficiency of the traps by intoxicating prey.[11]

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Pinguicula gigantea with prey. Theinsect was too large and was strongenough to escape from the stickysurface.

The leaf of a Drosera capensisbending in response to the trapping ofan insect.

The final carnivore with a pitfall­like trap is the bromeliad Brocchinia reducta. Like most relatives of thepineapple, the tightly packed, waxy leaf bases of the strap­like leaves of this species form an urn. In mostbromeliads, water collects readily in this urn and may provide habitats for frogs, insects and, more usefulfor the plant, diazotrophic (nitrogen­fixing) bacteria. In Brocchinia, the urn is a specialised insect trap, witha loose, waxy lining and a population of digestive bacteria.

Flypaper traps

The flypaper trap is based ona sticky mucilage, or glue.The leaf of flypaper traps isstudded with mucilage­secreting glands, which maybe short (like those of thebutterworts), or long andmobile (like those of manysundews). Flypapers haveevolved independently atleast five times.

In the genus Pinguicula, themucilage glands are quite

short (sessile), and the leaf, while shiny (giving the genus itscommon name of 'butterwort'), does not appear carnivorous.However, this belies the fact that the leaf is an extremely effectivetrap of small flying insects (such as fungus gnats), and its surfaceresponds to prey by relatively rapid growth. This thigmotropicgrowth may involve rolling of the leaf blade (to prevent rain fromsplashing the prey off the leaf surface) or dishing of the surfaceunder the prey to form a shallow digestive pit.

The sundew genus (Drosera) consists of over 100 species of active flypapers whose mucilage glands areborne at the end of long tentacles, which frequently grow fast enough in response to prey (thigmotropism)to aid the trapping process. The tentacles of D. burmanii can bend 180° in a minute or so. Sundews areextremely cosmopolitan and are found on all the continents except the Antarctic mainland. They are mostdiverse in Australia, the home to the large subgroup of pygmy sundews such as D. pygmaea and to anumber of tuberous sundews such as D. peltata, which form tubers that aestivate during the dry summermonths. These species are so dependent on insect sources of nitrogen that they generally lack the enzymenitrate reductase, which most plants require to assimilate soil­borne nitrate into organic forms.

Closely related to Drosera is the Portuguese dewy pine, Drosophyllum, which differs from the sundews inbeing passive. Its leaves are incapable of rapid movement or growth. Unrelated, but similar in habit, are theAustralian rainbow plants (Byblis). Drosophyllum is unusual in that it grows under near­desert conditions;almost all other carnivores are either bog plants or grow in moist tropical areas.

Recent molecular data (particularly the production of plumbagin) indicate that the remaining flypaper,Triphyophyllum peltatum, a member of the Dioncophyllaceae, is closely related to Drosophyllum and formspart of a larger clade of carnivorous and non­carnivorous plants with the Droseraceae, Nepenthaceae,

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Drosera capensis responding tocaptured prey. This scene is about 4hours in real­time.

The snap traps of Dionaea muscipulaclose rapidly when the sensitive hairson the leaf lobes are triggered.

Ancistrocladaceae and Plumbaginaceae. This plant is usuallyencountered as a liana, but in its juvenile phase, the plant iscarnivorous. This may be related to a requirement for specificnutrients for flowering.

Snap traps

The only two active snaptraps—the Venus flytrap(Dionaea muscipula) and thewaterwheel plant(Aldrovanda vesiculosa)—

are believed to have had a common ancestor with similaradaptations. Their trapping mechanism has also been described as a"mouse trap", "bear trap" or "man trap", based on their shape andrapid movement. However, the term snap trap is preferred as otherdesignations are misleading, particularly with respect to the intendedprey. Aldrovanda is aquatic and specialised in catching smallinvertebrates; Dionaea is terrestrial and catches a variety ofarthropods, including spiders.[12]

The traps are very similar, with leaves whose terminal section isdivided into two lobes, hinged along the midrib. Trigger hairs (threeon each lobe in Dionaea muscipula, many more in the case ofAldrovanda) inside the trap lobes are sensitive to touch. When atrigger hair is bent, stretch­gated ion channels in the membranes of cells at the base of the trigger hair open,generating an action potential that propagates to cells in the midrib.[13] These cells respond by pumping outions, which may either cause water to follow by osmosis (collapsing the cells in the midrib) or cause rapidacid growth.[14] The mechanism is still debated, but in any case, changes in the shape of cells in the midriballow the lobes, held under tension, to snap shut,[13] flipping rapidly from convex to concave[15] andinterring the prey. This whole process takes less than a second. In the Venus flytrap, closure in response toraindrops and blown­in debris is prevented by the leaves having a simple memory: for the lobes to shut, twostimuli are required, 0.5 to 30 seconds apart.[16][17]

The snapping of the leaves is a case of thigmonasty (undirected movement in response to touch). Furtherstimulation of the lobe's internal surfaces by the struggling insects causes the lobes to close even tighter(thigmotropism), sealing the lobes hermetically and forming a stomach in which digestion occurs over aperiod of one to two weeks. Leaves can be reused three or four times before they become unresponsive tostimulation, depending on the growing conditions.

Bladder traps

Bladder traps are exclusive to the genus Utricularia, or bladderworts. The bladders (vesicula) pump ionsout of their interiors. Water follows by osmosis, generating a partial vacuum inside the bladder. The bladderhas a small opening, sealed by a hinged door. In aquatic species, the door has a pair of long trigger hairs.Aquatic invertebrates such as Daphnia touch these hairs and deform the door by lever action, releasing thevacuum. The invertebrate is sucked into the bladder, where it is digested. Many species of Utricularia (such

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The tip of one stolon of Utriculariavulgaris, showing stolon, branchingleaf­shoots, and transparent bladdertraps

Genlisea violacea traps and leaves

as U. sandersonii) are terrestrial, growing on waterlogged soil, and their trapping mechanism is triggered ina slightly different manner. Bladderworts lack roots, but terrestrial species have anchoring stems thatresemble roots. Temperate aquatic bladderworts generally die back to a resting turion during the wintermonths, and U. macrorhiza appears to regulate the number of bladders it bears in response to the prevailingnutrient content of its habitat.

Lobster­pot traps

A lobster­pot trap is achamber that is easy to enter,and whose exit is eitherdifficult to find or obstructedby inward­pointing bristles.Lobster pots are the trappingmechanism in Genlisea, thecorkscrew plants. Theseplants appear to specialise inaquatic protozoa. A Y­shaped modified leaf allowsprey to enter but not exit.Inward­pointing hairs forcethe prey to move in a

particular direction. Prey entering the spiral entrance that coilsaround the upper two arms of the Y are forced to move inexorably towards a stomach in the lower arm ofthe Y, where they are digested. Prey movement is also thought to be encouraged by water movementthrough the trap, produced in a similar way to the vacuum in bladder traps, and probably evolutionarilyrelated to it.

Outside of Genlisea, features reminiscent of lobster­pot traps can be seen in Sarracenia psittacina,Darlingtonia californica, and, some horticulturalists argue, Nepenthes aristolochioides.

Combination traps

The trapping mechanism of the sundew Drosera glanduligera combines features of both flypaper and snaptraps; it has been termed a catapult­flypaper trap.[18]

Borderline carnivores

Main article: Protocarnivorous plant

To be a fully fledged carnivore, a plant must attract, kill, and digest prey;[5][19] and it must benefit fromabsorbing the products of the digestion (mostly amino acids and ammonium ions).[20] To manyhorticulturalists, these distinctions are a matter of taste. There is a spectrum of carnivory found in plants:from completely non­carnivorous plants like cabbages, to borderline carnivores, to unspecialised andsimple traps, like Heliamphora, to extremely specialised and complex traps, like that of the Venus flytrap.

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Roridula gorgonias: a borderlinecarnivore that gains nutrients fromits "prey" via the droppings of apredatory bug

The borderline carnivores include Roridula and Catopsis berteroniana. Catopsis is a borderline carnivorousbromeliad, like Brocchinia reducta. However, unlike B. reducta, whichproduces the enzyme phosphatase, C. berteroniana has not been shownto produce any digestive enzymes at all.[21] In these pitfall traps, preysimply fall into the urn, assisted by the waxy scales located on the rim.Roridula has a more intricate relationship with its prey. The plants inthis genus produce sticky leaves with resin­tipped glands and lookextremely similar to some of the larger sundews. However, they do notdirectly benefit from the insects they catch. Instead, they form amutualistic symbiosis with species of assassin bug (genus Pameridea),which eat the trapped insects. The plant benefits from the nutrients inthe bugs' faeces.[22]

A number of species in the Martyniaceae (previously Pedaliaceae),such as Ibicella lutea, have sticky leaves that trap insects. However,

these plants have not been shown conclusively to be carnivorous.[23] Likewise, the seeds of Shepherd'sPurse,[23] urns of Paepalanthus bromelioides,[24] bracts of Passiflora foetida,[25] and flower stalks andsepals of triggerplants (Stylidium)[26] appear to trap and kill insects, but their classification as carnivores iscontentious.

The production of specific prey­digesting enzymes (proteases, ribonucleases, phosphatases, etc.) issometimes used as a criterion for carnivory. However, this would probably discount Heliamphora[27] andDarlingtonia,[28] both of which appear to rely on the enzymes of symbiotic bacteria to break down theirprey but are generally considered as carnivores. However, discounting the enzyme­based definition leavesopen the question of Roridula.

Evolution

The evolution of carnivorous plants is obscured by the paucity of their fossil record. Very few fossils havebeen found, and then usually only as seed or pollen. Carnivorous plants are generally herbs, and their trapsare produced by primary growth. They generally do not form readily fossilisable structures such as thickbark or wood. The traps themselves would probably not be preserved in any case.

Still, much can be deduced from the structure of current traps. Pitfall traps are quite clearly derived fromrolled leaves. The vascular tissues of Sarracenia is a case in point. The keel along the front of the trapcontains a mixture of leftward­ and rightward­facing vascular bundles, as would be predicted from thefusion of the edges of an adaxial (stem­facing) leaf surface. Flypapers also show a simple evolutionarygradient from sticky, non­carnivorous leaves, through passive flypapers to active forms. Molecular datashow the Dionaea–Aldrovanda clade is closely related to Drosera,[29] but the traps are so dissimilar that thetheory of their origin—very fast­moving flypapers became less reliant on glue—remains rather speculative.

There are over a quarter of a million species of flowering plants. Of these, only around 630 are known to becarnivorous. True carnivory has probably evolved independently at least six times;[5] however, some ofthese "independent" groups probably descended from a recent common ancestor with a predisposition tocarnivory. Some groups (the Ericales and Caryophyllales) seem particularly fertile ground for carnivorous

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Artist's restoration of Archaeamphoralongicervia, the earliest knowncarnivorous plant

preadaptation, although in the former case, this may be more to dowith the ecology of the group than its morphology, as most of themembers of this group grow in low­nutrient habitats such as heathand bog.

It has been suggested that all trap types are modifications of asimilar basic structure—the hairy leaf.[30] Hairy (or morespecifically, stalked­glandular) leaves can catch and retain drops ofrainwater, especially if shield­shaped or peltate, thus promotingbacteria growth. Insects land on the leaf, become mired by thesurface tension of the water, and suffocate. Bacteria jumpstartdecay, releasing from the corpse nutrients that the plant can absorbthrough its leaves. This foliar feeding can be observed in most non­carnivorous plants. Plants that were better at retaining insects orwater therefore had a selective advantage. Rainwater can be retainedby cupping the leaf, leading to pitfall traps. Alternatively, insectscan be retained by making the leaf stickier by the production ofmucilage, leading to flypaper traps.

The pitfall traps may have evolved simply by selection pressure forthe production of more deeply cupped leaves, followed by "zippingup" of the margins and subsequent loss of most of the hairs, exceptat the bottom, where they help retain prey.

The lobster­pot traps of Genlisea are difficult to interpret. They may have developed from bifurcatedpitchers that later specialised on ground­dwelling prey; or, perhaps, the prey­guiding protrusions of bladdertraps became more substantial than the net­like funnel found in most aquatic bladderworts. Whatever theirorigin, the helical shape of the lobster pot is an adaptation that displays as much trapping surface as possiblein all directions when buried in moss.

The traps of the bladderworts may have derived from pitchers that specialised in aquatic prey when flooded,like Sarracenia psittacina does today. Escaping prey in terrestrial pitchers have to climb or fly out of a trap,and both of these can be prevented by wax, gravity and narrow tubes. However, a flooded trap can be swumout of, so in Utricularia, a one­way lid may have developed to form the door of a proto­bladder. Later, thismay have become active by the evolution of a partial vacuum inside the bladder, tripped by prey brushingagainst trigger hairs on the door of the bladder.

Flypaper traps include the various true flypapers and the snap traps of Aldrovanda and Dionaea. Theproduction of sticky mucilage is found in many non­carnivorous genera, and the passive glue traps in Byblisand Drosophyllum could easily have evolved.

The active glue traps use rapid plant movements to trap their prey. Rapid plant movement can result fromactual growth, or from rapid changes in cell turgor, which allow cells to expand or contract by quicklyaltering their water content. Slow­moving flypapers like Pinguicula exploit growth, but the Venus flytrapuses such rapid turgor changes that glue became unnecessary. The stalked glands that once made it andwhich are so evident in Drosera have become the teeth and trigger hairs—an example of natural selection

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The traps of Catopsis berteronianaare unlikely to have descended from ahairy leaf or sepal.

hijacking preexisting structures for new functions. Also, the nerve­like sensory system of the Venus Fly­trap evolved about 135 millionyears ago in the Cretaceous period. It has an active steel trap, likethat of Aldrovanda, the water wheel plant.[31]

Recent taxonomic analysis[32] of the relationships within theCaryophyllales indicate that the Droseraceae, Triphyophyllum,Nepenthaceae and Drosophyllum, while closely related, areembedded within a larger clade that includes non­carnivorousgroups such as the tamarisks, Ancistrocladaceae, Polygonaceae andPlumbaginaceae. Interestingly, the tamarisks possess specialisedsalt­excreting glands on their leaves, as do several of thePlumbaginaceae (such as the sea lavender, Limonium), which mayhave been co­opted for the excretion of other chemicals, such asproteases and mucilage. Some of the Plumbaginaceae (e.g.Ceratostigma) also have stalked, vascularised glands that secretemucilage on their calyces and aid in seed dispersal and possibly inprotecting the flowers from crawling parasitic insects. These areprobably homologous with the tentacles of the carnivorous genera.Perhaps carnivory evolved from a protective function, rather than anutritional one. The balsams (such as Impatiens), which are closelyrelated to the Sarraceniaceae and Roridula, similarly possess stalkedglands.

The only traps that are unlikely to have descended from a hairy leaf or sepal are the carnivorous bromeliads(Brocchinia and Catopsis). These plants use the urn—a fundamental part of a bromeliad—for a newpurpose and build on it by the production of wax and the other paraphernalia of carnivory.

Ecology and modelling of carnivory

Carnivorous plants are widespread but rather rare. They are almost entirely restricted to habitats such asbogs, where soil nutrients are extremely limiting, but where sunlight and water are readily available. Onlyunder such extreme conditions is carnivory favoured to an extent that makes the adaptations obvious.

The archetypal carnivore, the Venus flytrap, grows in soils with almost immeasurable nitrate and calciumlevels. Plants need nitrogen for protein synthesis, calcium for cell wall stiffening, phosphate for nucleic acidsynthesis, and iron for chlorophyll synthesis. The soil is often waterlogged, which favours the production oftoxic ions such as ammonium, and its pH is an acidic 4 to 5. Ammonium can be used as a source ofnitrogen by plants, but its high toxicity means that concentrations high enough to fertilise are also highenough to cause damage.

However, the habitat is warm, sunny, constantly moist, and the plant experiences relatively littlecompetition from low growing Sphagnum moss. Still, carnivores are also found in very atypical habitats.Drosophyllum lusitanicum is found around desert edges and Pinguicula valisneriifolia on limestone(calcium­rich) cliffs.[33]

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Drosophyllum lusitanicum is one of thefew carnivorous plants to grow in dry,alkaline soil.

In all the studied cases, carnivory allows plants to grow and reproduce using animals as a source ofnitrogen, phosphorus and possibly potassium.[34][35][36] However, there is a spectrum of dependency onanimal prey. Pygmy sundews are unable to use nitrate from soil because they lack the necessary enzymes(nitrate reductase in particular).[37] Common butterworts (Pinguicula vulgaris) can use inorganic sources ofnitrogen better than organic sources, but a mixture of both is preferred.[34] European bladderworts seem touse both sources equally well. Animal prey makes up fordiffering deficiencies in soil nutrients.

Plants use their leaves to intercept sunlight. The energy is usedto reduce carbon dioxide from the air with electrons from waterto make sugars (and other biomass) and a waste product,oxygen, in the process of photosynthesis. Leaves also respire, ina similar way to animals, by burning their biomass to generatechemical energy. This energy is temporarily stored in the formof ATP (adenosine triphosphate), which acts as an energycurrency for metabolism in all living things. As a wasteproduct, respiration produces carbon dioxide.

For a plant to grow, it must photosynthesise more than itrespires. Otherwise, it will eventually exhaust its biomass anddie. The potential for plant growth is net photosynthesis, thetotal gross gain of biomass by photosynthesis, minus thebiomass lost by respiration. Understanding carnivory requires a cost­benefit analysis of these factors.[20]

In carnivorous plants, the leaf is not just used to photosynthesise, but also as a trap. Changing the leaf shapeto make it a better trap generally makes it less efficient at photosynthesis. For example, pitchers have to beheld upright, so that only their opercula directly intercept light. The plant also has to expend extra energyon non­photosynthetic structures like glands, hairs, glue and digestive enzymes.[38] To produce suchstructures, the plant requires ATP and respires more of its biomass. Hence, a carnivorous plant will haveboth decreased photosynthesis and increased respiration, making the potential for growth small and the costof carnivory high.

Being carnivorous allows the plant to grow better when the soil contains little nitrate or phosphate. Inparticular, an increased supply of nitrogen and phosphorus makes photosynthesis more efficient, becausephotosynthesis depends on the plant being able to synthesise very large amounts of the nitrogen­richenzyme RuBisCO (ribulose­1,5­bis­phosphate carboxylase/oxygenase), the most abundant protein on Earth.

It is intuitively clear that the Venus flytrap is more carnivorous than Triphyophyllum peltatum. The formeris a full­time moving snap­trap; the latter is a part­time, non­moving flypaper. The energy "wasted" by theplant in building and fuelling its trap is a suitable measure of the carnivory of the trap.

Using this measure of investment in carnivory, a model can be proposed.[20] Above is a graph of carbondioxide uptake (potential for growth) against trap respiration (investment in carnivory) for a leaf in a sunnyhabitat containing no soil nutrients at all. Respiration is a straight line sloping down under the horizontalaxis (respiration produces carbon dioxide). Gross photosynthesis is a curved line above the horizontal axis:as investment increases, so too does the photosynthesis of the trap, as the leaf receives a better supply ofnitrogen and phosphorus. Eventually another factor (such as light intensity or carbon dioxide concentration)

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Modelling carnivory in plants: gross photosynthesis, respiration andnet photosynthesis as a function of the plant's investment incarnivorous adaptations. Non­zero optimum carnivory occurs inbrightly lit habitats with very limiting soil nutrients.

Modelling carnivory in plants: gross photosynthesis, respiration andnet photosynthesis as a function of the plant's investment incarnivorous adaptations. An optimum carnivory of zero occurs inpoorly lit habitats with abundant soil nutrients.

will become more limiting to photosynthesis than nitrogen or phosphorus supply. As a result, increasing theinvestment will not make the plant grow better. The net uptake of carbon dioxide, and therefore, the plant'spotential for growth, must be positive for the plant to survive. There is a broad span of investment wherethis is the case, and there is also a non­zero optimum. Plants investing more or less than this optimum willtake up less carbon dioxide than an optimal plant, and hence growing less well. These plants will be at aselective disadvantage. At zero investment the growth is zero, because a non­carnivorous plant cannotsurvive in a habitat with absolutely nosoil­borne nutrients. Such habitats donot exist, so for example, Sphagnumabsorbs the tiny amounts of nitrates andphosphates in rain very efficiently andalso forms symbioses with diazotrophiccyanobacteria.

In a habitat with abundant soil nutrientsbut little light (as shown above), thegross photosynthesis curve will belower and flatter, because light will bemore limiting than nutrients. A plantcan grow at zero investment incarnivory; this is also the optimuminvestment for a plant, as anyinvestment in traps reduces netphotosynthesis (growth) to less than thenet photosynthesis of a plant thatobtains its nutrients from soil alone.

Carnivorous plants exist between thesetwo extremes: the less limiting light andwater are, and the more limiting soilnutrients are, the higher the optimuminvestment in carnivory, and hence themore obvious the adaptations will be tothe casual observer.

The most obvious evidence for thismodel is that carnivorous plants tend togrow in habitats where water and lightare abundant and where competition isrelatively low: the typical bog. Thosethat do not tend to be even morefastidious in some other way.Drosophyllum lusitanicum grows wherethere is little water, but it is even moreextreme in its requirement for brightlight and low disturbance than most

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Part­time carnivory in Triphyophyllumpeltatum may be due to an unusually highneed for potassium at a certain point in thelife cycle, just before flowering.

other carnivores. Pinguicula valisneriifolia grows in soils with high levels of calcium but requires strongillumination and lower competition than many butterworts.[39]

In general, carnivorous plants are poor competitors, because they invest too heavily in structures that haveno selective advantage in nutrient­rich habitats. They succeed only where other plants fail. Carnivores are tonutrients what cacti are to water. Carnivory only pays off when the nutrient stress is high and where light isabundant.[40] When these conditions are not met, some plants give up carnivory temporarily. Sarraceniaspp. produce flat, non­carnivorous leaves (phyllodes) in winter. Light levels are lower than in summer, solight is more limiting than nutrients, and carnivory does not pay. The lack of insects in winter exacerbatesthe problem. Damage to growing pitcher leaves prevent them from forming proper pitchers, and again, theplant produces a phyllode instead.

Many other carnivores shut down in some seasons. Tuberoussundews die back to tubers in the dry season, bladderworts toturions in winter, and non­carnivorous leaves are made by mostbutterworts and Cephalotus in the less favourable seasons.Utricularia macrorhiza varies the number of bladders itproduces based on the expected density of prey.[41] Part­timecarnivory in Triphyophyllum peltatum may be due to anunusually high need for potassium at a certain point in the lifecycle, just before flowering.

The more carnivorous a plant is, the less conventional itshabitat is likely to be. Venus flytraps live in a very specialisedhabitat, whereas less carnivorous plants (Byblis, Pinguicula) arefound in less unusual habitats (i.e., those typical for non­carnivores). Byblis and Drosophyllum both come fromrelatively arid regions and are both passive flypapers, arguablythe lowest maintenance form of trap. Venus flytraps filter their prey using the teeth around the trap's edge,so as not to waste energy on hard­to­digest prey. In evolution, laziness pays, because energy can be used forreproduction, and short­term benefits in reproduction will outweigh long­term benefits in anything else.

Carnivory rarely pays, so even carnivorous plants avoid it when there is too little light or an easier source ofnutrients, and they use as few carnivorous features as are required at a given time or for a given prey item.There are very few habitats stressful enough to make investing biomass and energy in trigger hairs andenzymes worthwhile. Many plants occasionally benefit from animal protein rotting on their leaves, butcarnivory that is obvious enough for the casual observer to notice is rare.

Bromeliads seem very well preadapted to carnivory, but only one or two species can be classified as trulycarnivorous. By their very shape, bromeliads will benefit from increased prey­derived nutrient input. In thissense, bromeliads are probably carnivorous, but their habitats are too dark for more extreme, recognisablecarnivory to evolve. Most bromeliads are epiphytes, and most epiphytes grow in partial shade on treebranches. Brocchinia reducta, on the other hand, is a ground dweller.

Many carnivorous plants are not strongly competitive and rely on circumstances to suppress dominatingvegetation. Accordingly, some of them rely on fire ecology for their continued survival.

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Nepenthes mirabilis in a road cut inPalau. Showing habit and habitat.

For the most part carnivorous plant populations are not dominantenough to be dramatically significant, ecologically speaking, butthere is an impressive variety of organisms that interact with variouscarnivorous plants in sundry relationships of kleptoparasitism,commensalism, and mutualism. For example, small insectivoressuch as tree frogs often exploit the supply of prey to be found inpitcher plants, and the frog Microhyla nepenthicola actuallyspecialises in such habitats. Certain crab spiders such asMisumenops nepenthicola live largely on the prey of Nepenthes, andother, less specialised, spiders may build webs where they trapinsects attracted by the smell or appearance of the traps; somescavengers, detritivores, and also organisms that harvest or exploitthose in turn, such as the mosquito Wyeomyia smithii are largely ortotally dependent on particular carnivorous plants. Plants such asRoridula species combine with specialised bugs (Pameridearoridulae) in benefiting from insects trapped on their leaves.

Associations with species of pitcher plants are so many and variedthat the study of Nepenthes infauna is something of a discipline inits own right. Camponotus schmitzi, the diving ant, has an intimatedegree of mutualism with the pitcher plant Nepenthes bicalcarata; itnot only retrieves prey and detritus from beneath the surface of theliquid in the pitchers, but repels herbivores, and cleans the pitcher peristome, maintaining its slipperynature. The ants have been reported to attack struggling prey, hindering their escape, so there might be anelement of myrmecotrophy to the relationship. Numerous species of mosquitoes lay their eggs in the liquid,where their larvae play various roles, depending on species; some eat microbes and detritus, as is commonamong mosquito larvae, whereas some species of Toxorhynchites also breed in pitchers, and their larvae arepredators of other species of mosquito larvae. Apart from the crab spiders on pitchers, an actual small, redcrab Geosesarma malayanum will enter the fluid, robbing and scavenging, though reputedly it does so atsome risk of being captured and digested itself.

Nepenthes rajah has a remarkable mutualism with two unrelated small mammals, the tree shrew Tupaiamontana and the Summit Rat, Rattus baluensis. The tree shrews and the rats defecate into the plant's trapswhile visiting them to feed on sweet, fruity secretions from glands on the pitcher lids.[42] The tree shrewalso has a similar relationship with at least two other giant species of Nepenthes. More subtly, Hardwicke'sWoolly Bat (Kerivoula hardwickii), a small species, roosts beneath the operculum (lid) of Nepentheshemsleyana.[43] The bat's excretions that land in the pitcher pay for the shelter, as it were. To the plant theexcreta are more readily assimilable than intact insects would be.

There also is a considerable list of Nepenthes endophytes; these are microbes other than pathogens that livein the tissues of pitcher plants, often apparently harmlessly.

Classification

See also: List of carnivorous plants

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Stylidium turbinatum

Aldrovanda vesiculosa

The classification of all flowering plants is currently in a state of flux. In the Cronquist system, theDroseraceae and Nepenthaceae were placed in the order Nepenthales, based on the radial symmetry of theirflowers and their possession of insect traps. The Sarraceniaceae was placed either in the Nepenthales, or inits own order, the Sarraceniales. The Byblidaceae, Cephalotaceae, and Roridulaceae were placed in theSaxifragales; and the Lentibulariaceae in the Scrophulariales (now subsumed into the Lamiales[44]).

In more modern classification, such as that of the Angiosperm Phylogeny Group, the families have beenretained, but they have been redistributed amongst several disparate orders. It is also recommended thatDrosophyllum be considered in a monotypic family outside the rest of the Droseraceae, probably moreclosely allied to the Dioncophyllaceae. The current recommendations are shown below (only carnivorousgenera are listed):

Dicots

Asterales (sunflower and daisy order)Stylidiaceae

Stylidium (trigger plants, a borderline carnivore)Caryophyllales, (carnation order)

DioncophyllaceaeTriphyophyllum (a tropical liana)

DrosophyllaceaeDrosophyllum (Portuguese dewy pine)

Droseraceae (sundew family)Aldrovanda (waterwheel plant)Dionaea (Venus Flytrap)Drosera (sundews)†Droserapollis†Droserapites†Droseridites†Fischeripollis†Palaeoaldrovanda†Saxonipollis

Nepenthaceae (tropical pitcher­plant family)Nepenthes (tropical pitcher plants or monkey­cups, including Anurosperma)

Ericales (heather order)Roridulaceae

Roridula (a borderline carnivore)Sarraceniaceae (trumpet pitcher family)

†ArchaeamphoraSarracenia (North American trumpet pitchers)Darlingtonia (cobra plant/lily)Heliamphora (sun or marsh pitchers)

Lamiales (mint order)Byblidaceae

Byblis (rainbow plants)Lentibulariaceae (bladderwort family)

Pinguicula (butterworts)Genlisea (corkscrew plant)

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Byblis liniflora

Cephalotus follicularis

Utricularia (bladderworts, including Polypompholyx, the fairy aprons or pink petticoatsand Biovularia an obsolete genus)

Martyniaceae (all borderline carnivores, related to the sesame plant)Ibicella

Oxalidales (wood sorrel order)Cephalotus (Albany pitcher plant)

Monocots

Poales (grass order)Bromeliaceae (bromeliad or pineapple family)

Brocchinia (a terrestrial bromeliad)Catopsis (a borderline carnivore)

Eriocaulaceae (pipewort family)Paepalanthus bromelioides (a borderlinecarnivore)

Cultivation

In horticulture, carnivorous plants are considered a curiosity or ararity, but are becoming more common in cultivation with theadvent of mass­production tissue­culture propagation techniques.Venus flytraps are still the most commonly grown, usually availableat garden centers and hardware stores, sometimes offered alongsideother easy to grow varieties. Nurseries that specialize in growingcarnivorous plants exclusively also exist, more uncommon ordemanding varieties of carnivorous plants can be obtained fromspecialist nursies. California Carnivores is a notable example ofsuch a nursery that specializes in the cultivation of carnivorousplants. It is owned and operated by horticulturalist, Peter D'Amato.

Although different species of carnivorous plants have differentcultivation requirements in terms of sunlight, humidity, soilmoisture, etc., there are commonalities. Most carnivorous plantsrequire rainwater, or water that has been distilled, deionised byreverse osmosis, or acidified to around pH 6.5 using sulfuric acid.Common tap or drinking water contains minerals (particularlycalcium salts) that will quickly build up and kill the plant. This is because most carnivorous plants haveevolved in nutrient­poor, acidic soils and are consequently extreme calcifuges. They are therefore verysensitive to excessive soil­borne nutrients. Since most of these plants are found in bogs, almost all are veryintolerant of drying. There are exceptions: tuberous sundews require a dry (summer) dormancy period, andDrosophyllum requires much drier conditions than most.

Outdoor­grown carnivorous plants generally catch more than enough insects to keep themselves properlyfed. Insects may be fed to the plants by hand to supplement their diet; however, carnivorous plants aregenerally unable to digest large non­insect food items; bits of hamburger, for example, will simply rot, andthis may cause the trap, or even the whole plant, to die.

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Many Sarracenia hybrids are easy togrow.

Cultivated Nepenthes rajah and a fewother species.

A carnivorous plant that catches no insects at all will rarely die,although its growth may be impaired. In general, these plants arebest left to their own devices: after underwatering with tap­water,the most common cause of Venus flytrap death is prodding the trapsto watch them close and feeding them inappropriate items.

Most carnivorous plants require bright light, and most will lookbetter under such conditions, as this encourages them to synthesisered and purple anthocyanin pigments. Nepenthes and Pinguiculawill do better out of full sun, but most other species are happy indirect sunlight.

Carnivores mostly live in bogs, and those that do not are generallytropical. Hence, most require high humidity. On a small scale, thiscan be achieved by placing the plant in a wide saucer containingpebbles that are kept permanently wet. Small Nepenthes species grow well in large terraria.

Many carnivores are native to cold temperate regions and can be grown outside in a bog garden year­round.Most Sarracenia can tolerate temperatures well below freezing, despite most species being native to thesoutheastern United States. Species of Drosera and Pinguicula also tolerate subfreezing temperatures.Nepenthes species, which are tropical, require temperatures from 20 to 30 °C to thrive.

Carnivorous plants require appropriate nutrient­poor soil. Mostappreciate a 3:1 mixture of Sphagnum peat to sharp horticulturalsand (coir is an acceptable, and more ecofriendly substitute forpeat). Nepenthes will grow in orchid compost or in pure Sphagnummoss.

Ironically, carnivorous plants are themselves susceptible toinfestation by parasites such as aphids or mealybugs. Althoughsmall infestations can be removed by hand, larger infestationsnecessitate use of an insecticide.

Isopropyl alcohol (rubbing alcohol) is effective as a topicalinsecticide, particularly on scale insects. Diazinon is an excellentsystemic insecticide that is tolerated by most carnivorous plants.Malathion and Acephate (Orthene) have also been reported astolerable by carnivorous plants.

Although insects can be a problem, by far the biggest killer ofcarnivorous plants (besides human maltreatment) is grey mold(Botrytis cinerea). This thrives under warm, humid conditions and

can be a real problem in winter. To some extent, temperate carnivorous plants can be protected from thispathogen by ensuring that they are kept cool and well ventilated in winter and that any dead leaves areremoved promptly. If this fails, a fungicide is in order.

The easiest carnivorous plants for beginners are those from the cool temperate zone. These plants will dowell under cool greenhouse conditions (minimum 5 °C in winter, maximum 25 °C in summer) if kept inwide trays of acidified or rain water during summer and kept moist during winter:

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Depiction of a native being consumedby a Ya­te­veo ("I see you")carnivorous tree of Central America,from Land and Sea by J.W. Buel,1887

Audrey Junior, the man­eating plantfrom the cult film The Little Shop ofHorrors

Drosera capensis, the Cape sundew: attractive strap­leaved sundew, pink flowers, very tolerant ofmaltreatment.Drosera binata, the fork­leaved sundew: large, Y­shaped leaves.Sarracenia flava, the yellow trumpet pitcher: yellow, attractively veined leaves, yellow flowers inspring.Pinguicula grandiflora, the common butterwort: purple flowers in spring, hibernates as a bud(hibernaculum) in winter. Fully hardy.Pinguicula moranensis, the Mexican butterwort: pink flowers, non­carnivorous leaves in winter.

Venus flytraps will do well under these conditions but are actually rather difficult to grow: even if treatedwell, they will often succumb to grey mold in winter unless well ventilated. Some of the lowland Nepenthesare very easy to grow as long as they are provided with relatively constant, hot and humid conditions.

Medicinal uses

A study published in 2009 by researchers from Tel Aviv Universityindicates that secretions produced by carnivorous plants containcompounds that have anti­fungal properties and may lead to thedevelopment of a new class of anti­fungal drugs that will beeffective against infections that are resistant to current anti­fungaldrugs.[45][46]

Cultural depictions

Carnivorous plants have longbeen the subject of popularinterest and exposition,much of it highly inaccurate.Fictional plants have beenfeatured in a number ofbooks, movies, televisionseries, and video games.Typically, these fictionaldepictions includeexaggerated characteristics,such as enormous size orpossession of abilitiesbeyond the realm of reality,

and can be viewed as a kind of artistic license. Two of the mostfamous examples of fictional carnivorous plants in popular cultureare the 1960s black comedy The Little Shop of Horrors and the triffids of John Wyndham's The Day of theTriffids. Other movies, such as The Hellstrom Chronicle (1971), and television series utilize accuratedepictions of carnivorous plants for cinematic purposes.

The earliest known depiction of carnivorous plants in popular culture was a case wherein a large man­eating tree was reported to have consumed a young woman in Madagascar in 1878, as witnessed by Dr.Carl Liche. Liche reported the events in the South Australian Register in 1881. The woman, pictured in an

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accompanying artwork, was supposed to have been a member of the Mkodos, a "little known but crueltribe". The account has been debunked as pure myth as it appears Dr. Liche, the Mkodos, and the tree wereall fabrications.[47]

See also

Aggressive mimicryCarnivorous fungusPredatory dinoflagellate

References1. Clarke C.M., Bauer U., Lee C.C., Tuen A.A., Rembold K., Moran J.A. (2009). "Tree shrew lavatories: a novel

nitrogen sequestration strategy in a tropical pitcher plant". Biology Letters 5 (5): 632–635.doi:10.1098/rsbl.2009.0311 (https://dx.doi.org/10.1098%2Frsbl.2009.0311).

2. Chin L., Moran J.A., Clarke C. (2010). "Trap geometry in three giant montane pitcher plant species from Borneois a function of tree shrew body size". New Phytologist 186 (2): 461–470. doi:10.1111/j.1469­8137.2009.03166.x(https://dx.doi.org/10.1111%2Fj.1469­8137.2009.03166.x).

3. Clarke C., Moran J.A., Chin L. (2010). "Mutualism between tree shrews and pitcher plants: perspectives andavenues for future research". Plant Signaling & Behavior 5 (10): 1187–1189. doi:10.4161/psb.5.10.12807(https://dx.doi.org/10.4161%2Fpsb.5.10.12807).

4. Darwin C (1875). Insectivorous plants(http://web.archive.org/web/20061023071735/http://pages.britishlibrary.net/charles.darwin3/insectivorous/insect01.htm). London: John Murray. ISBN 1­4102­0174­0. Archived from the original(http://pages.britishlibrary.net/charles.darwin3/insectivorous/insect01.htm) on 2006­10­23.

5. Albert, V.A., Williams, S.E., and Chase, M.W. (1992). "Carnivorous plants: Phylogeny and structuralevolution". Science 257 (5076): 1491–1495. doi:10.1126/science.1523408(https://dx.doi.org/10.1126%2Fscience.1523408). PMID 1523408(https://www.ncbi.nlm.nih.gov/pubmed/1523408).

6. Ellison, A.M., and Gotelli, N.J. (2009). "Energetics and the evolution of carnivorous plants—Darwin's 'mostwonderful plants in the world'.". Journal of Experimental Botany 60 (1): 19–42. doi:10.1093/jxb/ern179(https://dx.doi.org/10.1093%2Fjxb%2Fern179). PMID 19213724(https://www.ncbi.nlm.nih.gov/pubmed/19213724).

7. Barthlott, W., S. Porembski, R. Seine & I. Theisen (translated by M. Ashdown) 2007. The Curious World ofCarnivorous Plants: A Comprehensive Guide to Their Biology and Cultivation (http://books.google.co.uk/books?id=vYwnc_L6el0C). Timber Press, Portland.

8. Williams, S. E. 2002. Comparative physiology of the Droseraceae sensu stricto—How do tentacles bend andtraps close? (http://carnivorousplants.org/cpn/articles/ICPS2002confp77_81.pdf) Proceedings of the 4thInternational Carnivorous Plant Society Conference. Tokyo, Japan. pp. 77­81.

9. Poppinga S., Hartmeyer S.R.H., Seidel R., Masselter T., Hartmeyer I., Speck T. (2012). "Catapulting tentacles ina sticky carnivorous plant" (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3458893). PLoS ONE 7 (9): e45735.doi:10.1371/journal.pone.0045735 (https://dx.doi.org/10.1371%2Fjournal.pone.0045735). PMC 3458893(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3458893). PMID 23049849(https://www.ncbi.nlm.nih.gov/pubmed/23049849).

10. Clarke (1993). "The possible functions of the thorns of Nepenthes bicalcarata (Hook.f.) pitchers"(http://www.carnivorousplants.org/cpn/articles/CPNv22n1_2p27_28.pdf) (PDF). Carnivorous Plant Newsletter 22(1–2): 27–28.

11. Mody N. V., Henson R., Hedin P. A., Kokpol U., Miles D. H. (1976). "Isolation of the insect paralyzing agentconiine from Sarracenia flava". Cellular and Molecular Life Sciences 32 (7): 829–830. doi:10.1007/BF02003710(https://dx.doi.org/10.1007%2FBF02003710).

9/15/2015 Carnivorous plant ­ Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Carnivorous_plant 19/21

12. Famous Insect Eating Plant Catches Many Spiders(http://www.sciencenews.org/view/generic/id/6007/title/From_the_March_23,_1935,_issue), The ScienceNewsletter, March 23, 1935, issue

13. Hodick D, Sievers A (1989). "The action potential of Dionaea muscipula Ellis"(http://www.springerlink.com/index/KL80VV1327508844.pdf) (PDF). Planta 174 (1): 8–18.doi:10.1007/BF00394867 (https://dx.doi.org/10.1007%2FBF00394867).

14. Hodick D, Sievers A (1988). "On the mechanism of closure of Venus flytrap (Dionaea muscipula Ellis)"(http://www.springerlink.com/index/QPK061437U675H10.pdf) (PDF). Planta 179 (1): 32–42.doi:10.1007/BF00395768 (https://dx.doi.org/10.1007%2FBF00395768).

15. Forterre Y, Skotheim JM, Dumais J, Mahadevan L (2005). "How the Venus flytrap snaps"(http://www.nature.com/nature/journal/v433/n7024/abs/nature03185.html). Nature 433 (7024): 421–5.doi:10.1038/nature03185 (https://dx.doi.org/10.1038%2Fnature03185). PMID 15674293(https://www.ncbi.nlm.nih.gov/pubmed/15674293).

16. http://www.carnivorousplants.org/cpn/articles/CPNv09n3p65_75_78.pdf17. http://www.carnivorousplants.org/cpn/articles/CPNv09n4p91_100.pdf18. Poppinga, S., Hartmeyer, S.R.H., Seidel, R., Masselter, T., Hartmeyer, I. & Speck, T. (September 2012).

"Catapulting tentacles in a sticky carnivorous plant"(http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0045735). PLoS ONE 7 (9): e45735.doi:10.1371/journal.pone.0045735 (https://dx.doi.org/10.1371%2Fjournal.pone.0045735). PMC 3458893(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3458893). PMID 23049849(https://www.ncbi.nlm.nih.gov/pubmed/23049849).

19. Juniper, B. E.; Robbins, R.J.; Joel, D.M. (1989). The Carnivorous Plants. Academic Press. ISBN 0­12­392170­8.

20. Givnish TJ, Burkhardt EL, Happel RE, Weintraub JD (1984). "Carnivory in the bromeliad Brocchinia reducta,with a cost­benefit model for the general restriction of carnivorous plants to sunny, moist, nutrient­poor habitats".American Naturalist 124 (4): 479–497. doi:10.1086/284289 (https://dx.doi.org/10.1086%2F284289).JSTOR 00030147 (https://www.jstor.org/stable/00030147). (Requires JSTOR subscription)

21. Plachno, B.J.; Jankun, A. (2005). "Phosphatase activity in glandular structures of carnivorous plant traps". Proc.of the International Botanical Congress. The Jagiellonian University, Krakow, Poland. p. 1716.

22. Hartmeyer, S. (1998). "Carnivory in Byblis revisited II: The phenomenon of symbiosis on insect trapping plants"(http://www.carnivorousplants.org/cpn/samples/Science274Byblis2.htm). Carnivorous Plant Newsletter 27 (4):110–113.

23. Schnell, Donald E. (2002). Carnivorous plants of the United States and Canada. Timber Press. ISBN 0­88192­540­3.

24. Rice, Barry A. (2006). Growing Carnivorous Plants. Timber Press. ISBN 0­88192­807­0.25. Radhamani, T.R., Sudarshana, L., and Krishnan, R. (1995). "Defence and carnivory: Dual role of bracts in

Passiflora foetida" (http://medind.nic.in/imvw/imvw14993.html). Journal of Biosciences 20 (5): 657–664.doi:10.1007/BF02703305 (https://dx.doi.org/10.1007%2FBF02703305).

26. Darnowski, D.W., Carroll, D.M., Płachno, B., Kabanoff, E., and Cinnamon, E. (2006). "Evidence ofprotocarnivory in triggerplants (Stylidium spp.; Stylidiaceae)" (http://www.thieme­connect.com/ejournals/abstract/plantbiology/doi/10.1055/s­2006­924472). Plant Biology (Stuttgart) 8 (6): 805–812. doi:10.1055/s­2006­924472 (https://dx.doi.org/10.1055%2Fs­2006­924472). PMID 17058181(https://www.ncbi.nlm.nih.gov/pubmed/17058181).

27. Jaffe, K., Michelangeli, F., Gonzalez, J.M., Miras, B., and Ruiz, M.C. (1992). "Carnivory in pitcher plants ofthe genus Heliamphora (Sarraceniaceae)". New Phytologist 122 (4): 733–744. doi:10.1111/j.1469­8137.1992.tb00102.x (https://dx.doi.org/10.1111%2Fj.1469­8137.1992.tb00102.x). JSTOR 2557442(https://www.jstor.org/stable/2557442).

28. Ellison, A.M. and Farnsworth, E.J. (2005). "The cost of carnivory for Darlingtonia californica (Sarraceniaceae):Evidence from relationships among leaf traits". American Journal of Botany 92 (7): 1085–1093.doi:10.3732/ajb.92.7.1085 (https://dx.doi.org/10.3732%2Fajb.92.7.1085).

9/15/2015 Carnivorous plant ­ Wikipedia, the free encyclopedia

https://en.wikipedia.org/wiki/Carnivorous_plant 20/21

29. Cameron K, Wurdack KJ, Jobson RW (2002). "Molecular evidence for the common origin of snap­traps amongcarnivorous plants". American Journal of Botany 89 (9): 1503–1509. doi:10.3732/ajb.89.9.1503(https://dx.doi.org/10.3732%2Fajb.89.9.1503). PMID 21665752(https://www.ncbi.nlm.nih.gov/pubmed/21665752).

30. Slack A (1988). Carnivorous plants. London: Alphabooks. pp. 18–19. ISBN 0­7136­3079­5.31. Williams, S. E. (June 15, 1976). Comparative Sensory Physiology of the Droseraceae­The Evolution of a Plant

Sensory System. Proceedings of the American Philosophical Society, 120, 3, 187­204.32. Cameron KM, Chase MW, Swensen SM (1995). "Molecular evidence for the relationships of Triphyophyllum

and Ancistrocladus". American Journal of Botany 83 (6): 117–118. doi:10.2307/2445804(https://dx.doi.org/10.2307%2F2445804). Discussion of this paper(http://www.carnivorousplants.org/cpn/samples/Science262Evol.htm) at the International carnivorous plantsociety website (original paper requires JSTOR subscription).

33. Zamora R, Gomez JM, Hodar JA (1997). "Responses of a carnivorous plant to prey and inorganic nutrients in aMediterranean environment". Oecologia 111 (4): 443–451. doi:10.1007/s004420050257(https://dx.doi.org/10.1007%2Fs004420050257).

34. Thoren LM, Karlsson PS (1998). "Effects of supplementary feeding on growth and reproduction of threecarnivorous plant species in a subarctic environment". Journal of Ecology 86 (3): 501–510. doi:10.1046/j.1365­2745.1998.00276.x (https://dx.doi.org/10.1046%2Fj.1365­2745.1998.00276.x).

35. Hanslin HM, Karlsson PS (1996). "Nitrogen uptake from prey and substrate as affected by prey capture level andplant reproductive status in four carnivorous plant species". Oecologia 106 (3): 370–375.doi:10.1007/BF00334564 (https://dx.doi.org/10.1007%2FBF00334564).

36. Deridder F, Dhondt AA (1992). "A positive correlation between naturally captured prey, growth and flowering inDrosera intermedia in two contrasting habitats". Belgian Journal of Botany 125: 30–44.

37. Karlsson PS, Pate JS (1992). "Contrasting effects of supplementary feeding of insects or mineral nutrients on thegrowth and nitrogen and phosphorus economy of pygmy species of Drosera". Oecologia 92 (1): 8–13.doi:10.1007/BF00317256 (https://dx.doi.org/10.1007%2FBF00317256).

38. Gallie, D. R. & Chang, S. C. (1997). "Signal transduction in the carnivorous plant Sarracenia purpurea ­regulation of secretory hydrolase expression during development and in response to resources"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC158611). Plant Physiology 115 (4): 1461–1471.doi:10.1104/pp.115.4.1461 (https://dx.doi.org/10.1104%2Fpp.115.4.1461). PMC 158611(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC158611). PMID 9414556(https://www.ncbi.nlm.nih.gov/pubmed/9414556).

39. Zamora R, Gomez JM, Hodar JA (1988). "Fitness responses of a carnivorous plant in contrasting ecologicalscenarios". Ecology 79 (5): 1630–1644. doi:10.1890/0012­9658(1998)079[1630:FROACP]2.0.CO;2(https://dx.doi.org/10.1890%2F0012­9658%281998%29079%5B1630%3AFROACP%5D2.0.CO%3B2).ISSN 0012­9658 (https://www.worldcat.org/issn/0012­9658).

40. Brewer JS (2002). "Why don't carnivorous pitcher plants compete with non­carnivorous plants for nutrients?"(http://www.esajournals.org/doi/pdf/10.1890/0012­9658(2003)084%5B0451:WDTCPP%5D2.0.CO%3B2).Ecology 84 (2): 451–462. doi:10.1890/0012­9658(2003)084[0451:WDTCPP]2.0.CO;2(https://dx.doi.org/10.1890%2F0012­9658%282003%29084%5B0451%3AWDTCPP%5D2.0.CO%3B2).ISSN 0012­9658 (https://www.worldcat.org/issn/0012­9658).

41. Knight SE, Frost TM (1991). "Bladder control in Utricularia macrorhiza ­ lake­specific variation in plantinvestment in carnivory". Ecology (Ecological Society of America) 72 (2): 728–734. doi:10.2307/2937212(https://dx.doi.org/10.2307%2F2937212). JSTOR 2937212 (https://www.jstor.org/stable/2937212).

42. Greenwood M., Clarke C., Lee C.C., Gunsalam A., Clarke R.H. (2011). "A unique resource mutualism betweenthe giant Bornean pitcher plant, Nepenthes rajah, and members of a small mammal community". PLoS ONE 6(6): e21114. doi:10.1371/journal.pone.0021114 (https://dx.doi.org/10.1371%2Fjournal.pone.0021114).

43. Scharmann, M. & T.U. Grafe 2013. Reinstatement of Nepenthes hemsleyana (Nepenthaceae), an endemic pitcherplant from Borneo, with a discussion of associated Nepenthes taxa. Blumea, published online on May 8, 2013.doi:10.3767/000651913X668465 (https://dx.doi.org/10.3767%2F000651913X668465)

9/15/2015 Carnivorous plant ­ Wikipedia, the free encyclopedia

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44. Muller K, Borsch T, Legendre L, Porembski S, Theisen I, Barthlott W (2004). "Evolution of carnivory inLentibulariaceae and the Lamiales" (http://www.thieme­connect.com/DOI/DOI?10.1055/s­2004­817909). PlantBiology (Stuttgart) 6 (4): 477–490. doi:10.1055/s­2004­817909 (https://dx.doi.org/10.1055%2Fs­2004­817909).PMID 15248131 (https://www.ncbi.nlm.nih.gov/pubmed/15248131).

45. Eilenberg, Haviva; Pnini­Cohen, Smadar; Rahamim, Yocheved; Sionov, Edward; Segal, Esther; Carmeli,Shmuel; Zilberstein, Aviah (December 2009). "Induced production of antifungal naphthoquinones in the pitchersof the carnivorous plant Nepenthes khasiana" (http://jxb.oxfordjournals.org/cgi/content/full/61/3/911). Journal ofExperimental Botany (Oxford University Press) 61 (3): 911–922. doi:10.1093/jxb/erp359(https://dx.doi.org/10.1093%2Fjxb%2Ferp359). PMC 2814117(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2814117). PMID 20018905(https://www.ncbi.nlm.nih.gov/pubmed/20018905). Retrieved 2010­04­22.

46. "Carnivorous plants may save people" (http://www.israel21c.org/201004117868/briefs/carnivorous­plants­may­save­people). Israel 21c Innovation News Service (http://www.israel21c.org/). April 11, 2010. Retrieved2010­04­13.

47. Ron Sullivan and Joe Eaton (2007­10­27). "The Dirt: Myths about man­eating plants ­ something to chew on"(http://www.sfgate.com/cgi­bin/article.cgi?f=/c/a/2007/10/27/HOIASVF6T.DTL). San Francisco Chronicle.Retrieved 2007­10­26.

Further reading

Slack A (1986). Insect­eating Plants and How to Grow Them.Sherborne UK: Alphabooks. ISBN 0­906670­42­X.Juniper BE, Robins RJ, Joel DM (1989). The CarnivorousPlants. Academic Press, San Diego.Carnivorous Plant Database(http://www.omnisterra.com/bot/cp_home.cgi) provides anup­to­date, searchable database of all the published species ofcarnivorous plants.Carnivorous Plant FAQ (http://www.sarracenia.com/faq.html) at Sarracenia.comList of films and TV shows that feature carnivorous plants­most of them fictional(http://www.sarracenia.com/faq/faq1395.html)Botanical Society of America ­ Carnivorous Plants Online(http://www.botany.org/carnivorous_plants/)Inner World of Carnivorous Plants from the John Innes Centre(https://wikis.nbi.ac.uk/InnerWorlds/index.php/Inner_World_of_Carnivorous_Plants#.UThnmRyePAk)Ellison, A.M. 2006. Nutrient Limitation and Stoichiometry of Carnivorous Plants.(http://harvardforest.fas.harvard.edu/publications/pdfs/ellison_PlantBio_2006.pdf) PDF (334 KB) PlantBiol. 8: 740–747.

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