Page semi-protected

Venus flytrap

From Wikipedia, the free encyclopedia
Jump to navigation Jump to search

Venus flytrap
Venus Flytrap showing trigger hairs.jpg
Leaf
Scientific classification edit
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Droseraceae
Genus: Dionaea
Sol. ex J.Ellis 1768
Species:
D. muscipula
Binomial name
Dionaea muscipula
J.Ellis
Dionaea distribution (revised).svg
Distribution
Synonyms[2]
  • Dionea Raf., spelling variant
  • Dionaea corymbosa
    (Raf.) Steud. (1840)
  • Dionaea crinita
    Sol. (1990) as synonym
  • Dionaea dentata
    D'Amato (1998) name published without description
  • Dionaea heterodoxa
    D'Amato (1998) nom.nud.
  • Dionaea muscicapa
    St.Hil. (1824) sphalm.typogr.
  • Dionaea sensitiva
    Salisb. (1796)
  • Dionaea sessiliflora
    (Raf.) Steud. (1840)
  • Dionaea uniflora
    (Raf.) Steud. (1840)
  • Drosera corymbosa
    Raf. (1833)
  • Drosera sessiliflora
    Raf. (1833)
  • Drosera uniflora
    Raf. (1833)

The Venus flytrap (Dionaea muscipula) is a carnivorous plant native to subtropical wetlands on the East Coast of the United States in North Carolina and South Carolina.[3] It catches its prey—chiefly insects and arachnids—with a trapping structure formed by the terminal portion of each of the plant's leaves, which is triggered by tiny hairs (called "trigger hairs" or "sensitive hairs") on their inner surfaces.

When an insect or spider crawling along the leaves contacts a hair, the trap prepares to close, snapping shut only if another contact occurs within approximately twenty seconds of the first strike. Triggers may occur if one-tenth of the insect is within contact.[4] The requirement of redundant triggering in this mechanism serves as a safeguard against wasting energy by trapping objects with no nutritional value, and the plant will only begin digestion after five more stimuli to ensure it has caught a live bug worthy of consumption.

Dionaea is a monotypic genus closely related to the waterwheel plant (Aldrovanda vesiculosa) and sundews (Drosera), all of which belong to the family Droseraceae.

Although widely cultivated for sale, the population of the Venus flytrap has been rapidly declining in its native range.[5] The species is currently under Endangered Species Act review by the U.S. Fish & Wildlife Service.[6]

Etymology

The plant's common name refers to Venus, the Roman goddess of love. The genus name, Dionaea ("daughter of Dione"), refers to the Greek goddess Aphrodite, while the species name, muscipula, is Latin for both "mousetrap" and "flytrap".[7][8] The Latin word muscipula ("mousetrap") is derived from mus ("mouse") and decipula ("trap"), while the homonym word muscipula ("flytrap") is derived from musca ("fly") and decipula ("trap").[9][10][8]

Historically, the plant was also known by the slang term "tipitiwitchet" or "tippity twitchet", possibly an oblique reference to the plant's resemblance to human female genitalia.[7][11] The term is similar to the term tippet-de-witchet which derives from tippet and witchet (archaic term for vagina).[12][13] In contrast, the English botanist John Ellis, who gave the plant its scientific name in 1768, wrote that the plant name tippitywichit was an indigenous word from either Cherokee or Catawba.[8][14] The plant name according to the Handbook of American Indians derives from the Renape word titipiwitshik ("they (leaves) which wind around (or involve)").[15][16]

Discovery

On 2 April 1759, the North Carolina colonial governor, Arthur Dobbs, penned the first written description of the plant in a letter to English botanist Peter Collinson.[17] In the letter he wrote: "We have a kind of Catch Fly Sensitive which closes upon anything that touches it. It grows in Latitude 34 but not in 35. I will try to save the seed here."[14][18] A year later, Dobbs went into greater detail about the plant in a letter to Collinson dated Brunswick, 24 January 1760.[19][20][21]

The great wonder of the vegetable kingdom is a very curious unknown species of Sensitive. It is a dwarf plant. The leaves are like a narrow segment of a sphere, consisting of two parts, like the cap of a spring purse, the concave part outwards, each of which falls back with indented edges (like an iron spring fox-trap); upon anything touching the leaves, or falling between them, they instantly close like a spring trap, and confine any insect or anything that falls between them. It bears a white flower. To this surprising plant I have given the name of Fly trap Sensitive.

— Arthur Dobbs

This was the first detailed recorded notice of the plant by Europeans. The description was before John Ellis' letter to The London Magazine on 1 September 1768,[8] and his letter to Carl Linnaeus on 23 September 1768,[22] in which he described the plant and proposed its English name Venus's Flytrap and scientific name Dionaea muscipula.[23]

Description

The Venus flytrap is a small plant whose structure can be described as a rosette of four to seven leaves, which arise from a short subterranean stem that is actually a bulb-like object. Each stem reaches a maximum size of about three to ten centimeters, depending on the time of year;[24] longer leaves with robust traps are usually formed after flowering. Flytraps that have more than seven leaves are colonies formed by rosettes that have divided beneath the ground.

Illustration from Curtis's Botanical Magazine by William Curtis (1746–1799)

The leaf blade is divided into two regions: a flat, heart-shaped photosynthesis-capable petiole, and a pair of terminal lobes hinged at the midrib, forming the trap which is the true leaf. The upper surface of these lobes contains red anthocyanin pigments and its edges secrete mucilage. The lobes exhibit rapid plant movements, snapping shut when stimulated by prey. The trapping mechanism is tripped when prey contacts one of the three hair-like trichomes that are found on the upper surface of each of the lobes. The mechanism is so highly specialized that it can distinguish between living prey and non-prey stimuli, such as falling raindrops;[25] two trigger hairs must be touched in succession within 20 seconds of each other or one hair touched twice in rapid succession,[25] whereupon the lobes of the trap will snap shut, typically in about one-tenth of a second.[26] The edges of the lobes are fringed by stiff hair-like protrusions or cilia, which mesh together and prevent large prey from escaping. These protrusions, and the trigger hairs (also known as sensitive hairs) are likely homologous with the tentacles found in this plant's close relatives, the sundews. Scientists have concluded that the snap trap evolved from a fly-paper trap similar to that of Drosera.[27]

The holes in the meshwork allow small prey to escape, presumably because the benefit that would be obtained from them would be less than the cost of digesting them. If the prey is too small and escapes, the trap will usually reopen within 12 hours. If the prey moves around in the trap, it tightens and digestion begins more quickly.

Speed of closing can vary depending on the amount of humidity, light, size of prey, and general growing conditions. The speed with which traps close can be used as an indicator of a plant's general health. Venus flytraps are not as humidity-dependent as are some other carnivorous plants, such as Nepenthes, Cephalotus, most Heliamphora, and some Drosera.

The Venus flytrap exhibits variations in petiole shape and length and whether the leaf lies flat on the ground or extends up at an angle of about 40–60 degrees. The four major forms are: 'typica', the most common, with broad decumbent petioles; 'erecta', with leaves at a 45-degree angle; 'linearis', with narrow petioles and leaves at 45 degrees; and 'filiformis', with extremely narrow or linear petioles. Except for 'filiformis', all of these can be stages in leaf production of any plant depending on season (decumbent in summer versus short versus semi-erect in spring), length of photoperiod (long petioles in spring versus short in summer), and intensity of light (wide petioles in low light intensity versus narrow in brighter light).[citation needed]

Habitat and distribution

Habitat

Map of the original distribution of the Venus flytrap

The Venus flytrap is found in nitrogen- and phosphorus-poor environments, such as bogs and wet savannahs. Small in stature and slow-growing, the Venus flytrap tolerates fire well and depends on periodic burning to suppress its competition.[28] Fire suppression threatens its future in the wild.[29] It survives in wet sandy and peaty soils. Although it has been successfully transplanted and grown in many locales around the world, it is native only to the coastal bogs of North and South Carolina in the United States, specifically within a 100-kilometer (60 mi) radius of Wilmington, North Carolina.[30] One such place is North Carolina's Green Swamp. There also appears to be a naturalized population of Venus flytraps in northern Florida as well as an introduced population in western Washington.[31][32] The nutritional poverty of the soil is the reason it relies on such elaborate traps: insect prey provide the nitrogen for protein formation that the soil cannot. They tolerate mild winters, and Venus flytraps that do not go through a period of winter dormancy will weaken and die after a period of time.[33]

They are full sun plants, usually found only in areas with less than 10% canopy cover.[5] The microhabitat where it thrives is typically sparse with grasses, herbs, sphagnum, and often bare patches where there aren't enough nutrients for noncarnivorous plants to survive, or where fires regularly clear competition and prevent cover from forming. Thus, natural fires are an important part of its habitat, required every 3–5 years in most places for D. muscipula to thrive. After fire, D. muscipula seeds germinate well in ash and sandy soil, with seedlings growing well in the open post-fire conditions. The seeds germinate immediately without a dormant period.[5]

Distribution

Dionaea muscipula occurs naturally only along the coastal plain of North and South Carolina in the U.S, with all known current sites within 90 km of Wilmington, North Carolina.[34] A 1958 survey of herbaria specimens and old documents found 259 sites where the historical record documented the presence of D. muscipula, within 21 counties in North and South Carolina.[35] As of 2019, it was considered extirpated in North Carolina in the inland counties of Moore, Robeson, and Lenoir, as well as the South Carolina coastal counties of Charleston and Georgetown. Remaining extant populations exist in North Carolina in Beaufort, Craven, Pamlico, Carteret, Jones, Onslow, Duplin, Pender, New Hanover, Brunswick, Columbus, Bladen, Sampson, Cumberland, and Hoke counties, and in South Carolina in Horry county.[34]

Population

A large-scale survey in 2019, conducted by the North Carolina Natural Heritage Program, counted a total of 163,951 individual Venus flytraps in North Carolina and 4,876 in South Carolina, estimating a total of 302,000 individuals remaining in the wild in its native range.[36] This represents a reduction of more than 93% from a 1979 estimate of approximately 4,500,000 individuals.[5] A 1958 study found 259 confirmed extant or historic sites.[35] As of 2016, there were 71 known sites where the plant could be found in the wild. Of these 71 sites, only 20 were classified as having excellent or good long-term viability.[6]

Carnivory

A closing trap

Prey selectivity

A time lapse showing Venus flytrap catching prey

Most carnivorous plants selectively feed on specific prey. This selection is due to the available prey and the type of trap used by the organism. With the Venus flytrap, prey is limited to beetles, spiders and other crawling arthropods. The Dionaea diet is 33% ants, 30% spiders, 10% beetles, and 10% grasshoppers, with fewer than 5% flying insects.[37]

Given that Dionaea evolved from an ancestral form of Drosera (carnivorous plants that use a sticky trap instead of a snap trap) the reason for this evolutionary branching becomes clear. Drosera consume smaller, aerial insects, whereas Dionaea consume larger terrestrial bugs. Dionaea are able to extract more nutrients from these larger bugs. This gives Dionaea an evolutionary advantage over their ancestral sticky trap form.[38]

Mechanism of trapping

Closeup of one of the hinged trigger hairs

The Venus flytrap is one of a very small group of plants capable of rapid movement, such as Mimosa pudica, the Telegraph plant, sundews and bladderworts.

The mechanism by which the trap snaps shut involves a complex interaction between elasticity, turgor and growth. The trap only shuts when there have been two stimulations of the trigger hairs; this is to avoid inadvertent triggering of the mechanism by dust and other wind-borne debris. In the open, untripped state, the lobes are convex (bent outwards), but in the closed state, the lobes are concave (forming a cavity). It is the rapid flipping of this bistable state that closes the trap,[26] but the mechanism by which this occurs is still poorly understood. When the trigger hairs are stimulated, an action potential (mostly involving calcium ions—see calcium in biology) is generated, which propagates across the lobes and stimulates cells in the lobes and in the midrib between them.[39]

It is hypothesized that there is a threshold of ion buildup for the Venus flytrap to react to stimulation.[40] The acid growth theory states that individual cells in the outer layers of the lobes and midrib rapidly move 1H+ (hydrogen ions) into their cell walls, lowering the pH and loosening the extracellular components, which allows them to swell rapidly by osmosis, thus elongating and changing the shape of the trap lobe. Alternatively, cells in the inner layers of the lobes and midrib may rapidly secrete other ions, allowing water to follow by osmosis, and the cells to collapse. Both of these mechanisms may play a role and have some experimental evidence to support them.[41][42] Flytraps show a clear example of memory in plants, the plant knows if its hair has been touched, and remembers that for a few seconds. The plant then recalls that incident if a second touch happens during that time frame, and closes.[43] After closing, the flytrap counts additional stimulations of the trigger hairs, to five total, to start the production of digesting enzymes.[44]

Chrysomelid beetle, Paria.

Digestion

If the prey is unable to escape, it will continue to stimulate the inner surface of the lobes, and this causes a further growth response that forces the edges of the lobes together, eventually sealing the trap hermetically and forming a "stomach" in which digestion occurs. Release of the digestive enzymes is controlled by the hormone jasmonic acid, the same hormone that triggers the release of toxins as an anti-herbivore defense mechanism in non-carnivorous plants. (See Evolution below)[44][45] Once the digestive glands in the leaf lobes have been activated, digestion is catalysed by hydrolase enzymes secreted by the glands.

Oxidative protein modification is likely to be a pre-digestive mechanism used by Dionaea muscipula. Aqueous leaf extracts have been found to contain quinones such as the naphthoquinone plumbagin that couples to different NADH-dependent diaphorases to produce superoxide and hydrogen peroxide upon autoxidation.[46] Such oxidative modification could rupture animal cell membranes. Plumbagin is known to induce apoptosis, associated with the regulation of the Bcl-2 family of proteins.[47] When the Dionaea extracts were pre-incubated with diaphorases and NADH in the presence of serum albumin (SA), subsequent tryptic digestion of SA was facilitated.[46] Since the secretory glands of Droseraceae contain proteases and possibly other degradative enzymes, it may be that the presence of oxygen-activating redox cofactors function as extracellular pre-digestive oxidants to render membrane-bound proteins of the prey (insects) more susceptible to proteolytic attacks.[46]

Digestion takes about ten days, after which the prey is reduced to a husk of chitin. The trap then reopens, and is ready for reuse.[48]

Evolution

Drosera falconeri, with short, wide, sticky leaf traps

Carnivory in plants is a very specialized form of foliar feeding, and is an adaptation found in several plants that grow in nutrient-poor soil. Carnivorous traps were naturally selected to allow these organisms to compensate for the nutrient deficiencies of their harsh environments and compensate for the reduced photosynthetic benefit.[49] Phylogenetic studies have shown that carnivory in plants is a common adaptation in habitats with abundant sunlight and water but scarce nutrients.[38] Carnivory has evolved independently six times in the angiosperms based on extant species, with likely many more carnivorous plant lineages now extinct.[50]

The "snap trap" mechanism characteristic of Dionaea is shared with only one other carnivorous plant genus, Aldrovanda. For most of the 20th century, this relationship was thought to be coincidental, more precisely an example of convergent evolution. Some phylogenetic studies even suggested that the closest living relatives of Aldrovanda were the sundews.[51] It was not until 2002 that a molecular evolutionary study, by analyzing combined nuclear and chloroplast DNA sequences, indicated that Dionaea and Aldrovanda were closely related and that the snap trap mechanism evolved only once in a common ancestor of the two genera.[52][53]

A 2009 study[51] presented evidence for the evolution of snap traps of Dionaea and Aldrovanda from a flypaper trap like Drosera regia, based on molecular data. The molecular and physiological data imply that Dionaea and Aldrovanda snap traps evolved from the flypaper traps of a common ancestor with Drosera. Pre-adaptations to the evolution of snap traps were identified in several species of Drosera, such as rapid leaf and tentacle movement. The model proposes that plant carnivory by snap trap evolved from the flypaper traps, driven by increasing prey size. Bigger prey provides greater nutritional value, but large insects can easily escape the sticky mucilage of flypaper traps; the evolution of snap traps would therefore prevent escape and kleptoparasitism (theft of prey captured by the plant before it can derive benefit from it), and would also permit a more complete digestion.[51][52]

In 2016, a study of the expression of genes in the plant's leaves as they captured and digested prey was published in the journal, Genome Research. The gene activation observed in the leaves of the plants gives support to the hypothesis that the carnivorous mechanisms present in the flytrap are a specially adapted version of mechanisms used by non-carnivorous plants to defend against herbivorous insects.[45][54] In many non-carnivorous plants, jasmonic acid serves as a signaling molecule for the activation of defense mechanisms, such as the production of hydrolases, which can destroy chitin and other molecular components of insect and microbial pests.[55] In the Venus flytrap, this same molecule has been found to be responsible for the activation of the plant's digestive glands. A few hours after the capture of prey, another set of genes is activated inside the glands, the same set of genes that is active in the roots of other plants, allowing them to absorb nutrients. The use of similar biological pathways in the traps as non-carnivorous plants use for other purposes indicates that somewhere in its evolutionary history, the Venus flytrap repurposed these genes to facilitate carnivory.

Proposed evolutionary history

Carnivorous plants are generally herbaceous, and their traps the result of primary growth. They generally do not form readily fossilizable structures such as thick bark or wood. As a result, there is no fossil evidence of the steps that might link Dionaea and Aldrovanda, or either genus with their common ancestor, Drosera. Nevertheless, it is possible to infer an evolutionary history based on phylogenetic studies of both genera. Researchers have proposed a series of steps that would ultimately result in the complex snap-trap mechanism:[51][52]

  • Larger insects usually walk over the plant, instead of flying to it,[56] and are more likely to break free from sticky glands alone. Therefore, a plant with wider leaves, like Drosera falconeri,[51] must have adapted to move the trap and its stalks in directions that maximized its chance of capturing and retaining such prey—in this particular case, longitudinally. Once adequately "wrapped", escape would be more difficult.[56]
  • Evolutionary pressure then selected for plants with shorter response time, in a manner similar to Drosera burmannii or Drosera glanduligera. The faster the closing, the less reliant on the flypaper model the plant would be.
  • As the trap became more and more active, the energy required to "wrap" the prey increased. Plants that could somehow differentiate between actual insects and random detritus/rain droplets would have an advantage, thus explaining the specialization of inner tentacles into trigger hairs.
  • Ultimately, as the plant relied more on closing around the insect rather than gluing them to the leaf surface, the tentacles so evident in Drosera would lose their original function altogether, becoming the "teeth" and trigger hairs—an example of natural selection utilizing pre-existing structures for new functions.
  • Completing the transition, the plant eventually developed the depressed digestive glands found inside the trap, rather than using the dews in the stalks, further differentiating it from genus Drosera.

Phylogenetic studies using molecular characters place the emergence of carnivory in the ancestors of Dionaea muscipula to 85.6 million years ago, and the development of the snap-trap tn the ancestors of Dionaea and its sister genus Aldrovanda to approximately 48 million years ago.[57]

Cultivation

Dionaea muscipula 'Akai Ryu', Japanese for 'Red Dragon', in cultivation

Plants can be propagated by seed, taking around four to five years to reach maturity. More commonly, they are propagated by clonal division in spring or summer. Venus flytraps can also be propagated in vitro using plant tissue culture.[58] Most Venus flytraps found for sale in nurseries garden centers have been produced using this method, as this is the most cost-effective way to propagate them on a large scale. Regardless of the propagation method used, the plants will live for 20 to 30 years if cultivated in the right conditions.[59]

Cultivars

Venus flytraps are by far the most commonly recognized and cultivated carnivorous plant, and they are frequently sold as houseplants. Various cultivars (cultivated varieties) have come into the market through tissue culture of selected genetic mutations, and these plants are raised in large quantities for commercial markets.

Conservation

Although widely cultivated for sale as a houseplant, D. muscipula has suffered a significant decline in its population in the wild. The population in its native range is estimated to have decreased 93% since 1979.[5][36]

Status

The species is under Endangered Species Act review by the U.S. Fish & Wildlife Service.[60] The current review commenced in 2018, after an initial "90-day" review found that action may be warranted. A previous review in 1993 resulted in a determination that the plant was a "Potential candidate without sufficient data on vulnerability".[61] The IUCN Red List classifies the species as "vulnerable".[62] The State of North Carolina lists Dionaea muscipula as a species of "Special Concern–Vulnerable".[63] In 2010, CITES listed it as an Appendix II species.[64]  NatureServe classified it as "Imperiled" (G2) in a 2018 review.[65]

The U.S. Fish and Wildlife Service has not indicated a timeline to conclude its current review of Dionaea muscipula. The Endangered Species Act specifies a two-year timeline for a species review. However, the species listing process takes 12.1 years on average.[66]

Threats

The Venus flytrap is only found in the wild in a very particular set of conditions, requiring flat land with moist, acidic, nutrient-poor soils that receive full sun and burn frequently in forest fires, and is therefore sensitive to many types of disturbance.[5] A 2011 review identified five categories of threats for the species: agriculture, road-building, biological resource use (poaching and lumber activities), natural systems modifications (drainage and fire suppression), and pollution (fertilizer).[67]  

Habitat loss is a major threat to the species. The human population of the coastal Carolinas is rapidly expanding. For example, Brunswick County, North Carolina, which has the largest number of Venus flytrap populations, has seen a 27% increase in its human population from 2010 to 2018.[68] As the population grows, residential and commercial development and road building directly eliminate flytrap habitat, while site preparation that entails ditching and draining can dry out soil in surrounding areas, destroying the viability of the species.[69][65] Additionally, increased recreational use of natural areas in populated areas directly destroys the plants by crushing or uprooting them.[5]

Fire suppression is another threat to the Venus flytrap. In the absence of regular fires, shrubs and trees encroach, outcompeting the species and leading to local extirpations.[28][70] D. muscipola requires fire every 3–5 years, and best thrives with annual brush fires.[71] Although flytraps and their seeds are typically killed alongside their competition in fires, seeds from flytraps adjacent to the burnt zone propagate quickly in the ash and full sun conditions that occur post a fire disturbance.[72] Because the mature plants and new seedlings are typically destroyed in the regular fires that are necessary to maintain their habitat, D. muscipula's survival relies upon adequate seed production and dispersal from outside the burnt patches back into the burnt habitat, requiring a critical mass of populations, and exposing the success of any one population to metapopulation dynamics. These dynamics make small, isolated populations particularly vulnerable to extirpation, for if there are no mature plants adjacent to the fire zone, there is no source of seeds post-fire.[5]

Poaching has been another cause of population decline. Harvesting Venus flytraps on public land became illegal in North Carolina in 1958, and since then a legal cultivation industry has formed, growing tens of thousands of flytraps in commercial greenhouses for sale as household plants. Yet in 2016, the NY Times reported that demand for wild plants still exists, which "has led to a 'Venus flytrap crime ring.'"[73] In 2014, the state of North Carolina made Venus flytrap poaching a felony.[74] Since then, several poachers have been charged, with one man receiving 17 months in prison for poaching 970 Venus flytraps,[75] and another man charged with 73 felony counts in 2019.[76] Poachers may do greater harm to the wild populations than a simple count of individuals taken would indicate, as they may selectively harvest the largest plants at a site, which have more flowers and fruit and therefore generate more seeds than smaller plants.[5]

Additionally, the species is particularly vulnerable to catastrophic climate events.  Most Venus flytrap sites are only 2–4 meters (6.5 –13 feet) above sea level and are located in a region prone to hurricanes, making storm surges and rising sea levels a long-term threat.[5]

Designations

In 2005, the Venus flytrap was designated as the state carnivorous plant of North Carolina.[77]

In alternative medicine

Venus flytrap extract is available on the market as an herbal remedy, sometimes as the prime ingredient of a patent medicine named "Carnivora". According to the American Cancer Society, these products are promoted in alternative medicine as a treatment for a variety of human ailments including HIV, Crohn's disease and skin cancer, but "available scientific evidence does not support the health claims made for Venus flytrap extract".[78]

See also

References

  1. ^ Schnell, D.; Catling, P.; Folkerts, G.; Frost, C.; Gardner, R.; et al. (2000). "Dionaea muscipula". IUCN Red List of Threatened Species. 2000: e.T39636A10253384. doi:10.2305/IUCN.UK.2000.RLTS.T39636A10253384.en. Listed as Vulnerable (VU A1acd, B1+2c v2.3)
  2. ^ Schlauer, J. (N.d.) Dionaea muscipula. Carnivorous Plant Database.
  3. ^ Kew World Checklist of Selected Plant Families
  4. ^ Sumner, Thomas (20 November 2012). "Investigating the Venus Flytrap". insidescience.com. Retrieved 18 March 2018.
  5. ^ a b c d e f g h i j Waller, Donald (21 October 2016). "Petition to list the Venus flytrap (Dionaea muscipula Ellis) as Endangered under the 1973 Endangered Species Act". USFWS. Retrieved 14 December 2019.
  6. ^ a b "Venus Flytrap: Under Endangered Species Act review" (PDF). U.S. Fish & Wildlife Service. June 2017. Retrieved 15 December 2019.
  7. ^ a b "Background Information on Venus Fly Traps – Venus Fly Trap naming and history". FlyTrapCare.com. 4 April 2008. Archived from the original on 17 December 2008.
  8. ^ a b c d Ellis, John (1768). The London Magazine, Or, Gentleman's Monthly Intelligencer. October 1768. R. Baldwin. p. 523.
  9. ^ Donaldson, John William (1852). Varronianus: A Critical and Historical Introduction to the Ethnography of Ancient Italy and to the Philological Study of the Latin Language. J. W. Parker & Son. pp. 431. cipula.
  10. ^ Wase, Christopher (1662). Dictionarium Minus: A Compendious Dictionary English-Latin and Latin-English. Maxwell.
  11. ^ Rice, Barry (January 2007). "How did the Venus flytrap get its name?". The Carnivorous Plant FAQ.
  12. ^ "tippet-de-witchet, n. – Green's Dictionary of Slang". greensdictofslang.com. Retrieved 21 January 2020.
  13. ^ Williams, Gordon (2001). "wicket". A Dictionary of Sexual Language and Imagery in Shakespearean and Stuart Literature: Three Volume Set Volume I A–F Volume II G–P Volume III Q–Z. A&C Black. p. 1533. ISBN 978-0-485-11393-8.
  14. ^ a b Mabey, Richard (2016). "The Challenge of Carnivorous Plants: The Tipitiwitchet (Chapter 16)". The Cabaret of Plants: Forty Thousand Years of Plant Life and the Human Imagination. W. W. Norton & Company. ISBN 978-0-393-24877-7.
  15. ^ Mabey, Richard (2016). The Cabaret of Plants: Forty Thousand Years of Plant Life and the Human Imagination. W. W. Norton & Company. ISBN 978-0-393-24877-7.
  16. ^ Hodge, Frederick Webb (1912). Handbook of American Indians North of Mexico: N–Z. U.S. Government Printing Office. pp. 759. titipiwitshik
  17. ^ Irmscher, Christoph (1999). The Poetics of Natural History: From John Bartram to William James. Rutgers University Press. p. 31. ISBN 978-0-8135-2615-7.
  18. ^ Bartram, John (1942). Diary of a journey through the Carolinas, Georgia and Florida : From July 1, 1765 to April 10, 1766. American Philosophical Society. p. 104. hdl:2027/uc1.32106020417272.
  19. ^ Lewis Weston Dillwyn; Peter Collinson (1843). Hortus Collinsonianus. An account of the plants cultivated by the late Peter Collinson. W. C. Murray & D. Rees. p. 18.
  20. ^ Gardeners Chronicle & New Horticulturist. Volumes 3–4. Haymarket Publishing. 1875. p. 306.
  21. ^ The Nature Conservancy Archived 29 August 2017 at the Wayback Machine – Venus Flytrap
  22. ^ Ellis, John (23 September 1768). Letter 23 September 1768, London to Carl Linnaeus.
  23. ^ Directions for Bringing over Seeds and Plants, from the East Indies and Other Distant Countries, in a State of Vegetation: Together with a Catalogue of Such Foreign Plants as Are Worthy of Being Encouraged in Our American Colonies, for the Purposes of Medicine, Agriculture, and Commerce. To Which is Added, the Figure and Botanical Description of a New Sensitive Plant, Called Dionæa muscipula: or, Venus's Fly-trap – (London, printed and sold by L. Davis, 1770).
  24. ^ "Venus flytraps". The Carnivorous Plant FAQ. Retrieved 13 June 2005.
  25. ^ a b Raven, Peter H.; Evert, Ray Franklin; Eichhorn, Susan E. (2005). Biology of Plants (7th ed.). W.H. Freeman and Company. ISBN 978-0-7167-1007-3.
  26. ^ a b Forterre, Yoël; Skotheim, Jan M.; Dumais, Jacques; Mahadevan, L. (27 January 2005). "How the Venus flytrap snaps" (PDF). Nature. 433 (7024): 421–425. Bibcode:2005Natur.433..421F. doi:10.1038/nature03185. PMID 15674293. Archived from the original (PDF) on 2 December 2007.
  27. ^ Cameron, Kenneth M.; Wurdack, Kenneth J.; Jobson, Richard W. (2002). "Molecular evidence for the common origin of snap-traps among carnivorous plants". American Journal of Botany. 89 (9): 1503–1509. doi:10.3732/ajb.89.9.1503. PMID 21665752.
  28. ^ a b W. Schulze; E.D. Schulze; I. Schulze & R. Oren (2001). "Quantification of insect nitrogen utilization by the Venus flytrap Dionaea muscipula catching prey with highly variable isotope signatures". Journal of Experimental Botany. 52 (358): 1041–1049. doi:10.1093/jexbot/52.358.1041. PMID 11432920.
  29. ^ Leege, Lissa. "How does the Venus flytrap digest flies?". Scientific American. Retrieved 20 August 2008.
  30. ^ Darwin, C. R. 1875. Insectivorous Plants.
  31. ^ Schnell, D. E. (2002). Carnivorous Plants of the United States and Canada (2nd ed.). Timber Press. ISBN 978-0-88192-540-1.
  32. ^ Giblin, D. Nd. Dionaea muscipula. Burke Museum of Natural History and Culture.
  33. ^ "International Carnivorous Plant Society". Carnivorousplants.org. Archived from the original on 28 July 2014. Retrieved 26 August 2013.
  34. ^ a b "Venus Flytrap: Under Endangered Species Act review" (PDF). U.S. Fish & Wildlife Service. June 2017. Retrieved 15 December 2019.
  35. ^ a b Roberts, Patricia R.; Oosting, H. J. (1958). "Responses of Venus Fly Trap (Dionaea muscipula) to Factors Involved in Its Endemism". Ecological Monographs. 28 (2): 193–218. doi:10.2307/1942208. ISSN 1557-7015. JSTOR 1942208.
  36. ^ a b "Biodiversity Day: Venus Flytrap Preservation | NC DNCR". www.ncdcr.gov. Retrieved 14 December 2019.
  37. ^ Ellison, DM; Gotelli, NJ (2009). "Energetics and the evolution of carnivorous plants – Darwin's 'Most Wonderful plants in the world'". Journal of Experimental Botany. 60 (1): 19–42. doi:10.1093/jxb/ern179. PMID 19213724.
  38. ^ a b Gibson, TC; Waller, DM (2009). "Evolving Darwin's 'most wonderful' plant: ecological steps to a snap-trap". New Phytologist. 183 (1): 575–587. doi:10.1111/j.1469-8137.2009.02935.x. PMID 19573135.
  39. ^ Hodick, Dieter; Sievers, Andreas (1989). "The action potential of Dionaea muscipula Ellis". Planta. 174 (1): 8–18. doi:10.1007/BF00394867. PMID 24221411.
  40. ^ "Press release: The Trap Snaps Shut". Wiley ChemBioChem. January 2010. doi:10.1002/(issn)1439-7633/asset/homepages/press/201001press (inactive 18 March 2020). Cite journal requires |journal= (help)
  41. ^ Williams, S. E. 2002. Comparative physiology of the Droseraceae sensu stricto – How do tentacles bend and traps close? Proceedings of the 4th International Carnivorous Plant Society Conference. Tokyo, Japan. pp. 77–81.
  42. ^ Hodick, Dieter; Sievers, Andreas (1988). "On the mechanism of closure of Venus flytrap (Dionaea muscipula Ellis)". Planta. 179 (1): 32–42. doi:10.1007/BF00395768. PMID 24201419.
  43. ^ Chamovitz, Daniel (2012). What a Plant Knows. United States: Scientific American / Farrar, Straus and Giroux. p. 117. ISBN 9780374533885.
  44. ^ a b Böhm, J.; Scherzer, S.; Krol, E.; Kreuzer, I.; von Meyer, K.; Lorey, C.; Mueller, T. D.; Shabala, L.; Monte, I.; Solano, R.; Al-Rasheid, K. A. S.; Rennenberg, H.; Shabala, S.; Neher, E.; Hedrich, R. (2016). "The Venus Flytrap Dionaea muscipula Counts Prey-Induced Action Potentials to Induce Sodium Uptake". Current Biology. 26 (3): 286–295. doi:10.1016/j.cub.2015.11.057. ISSN 0960-9822. PMC 4751343. PMID 26804557.
  45. ^ a b Bemm, Felix; Becker, Dirk; Larisch, Christina; Kreuzer, Ines; Escalante-Perez, Maria; Schulze, Waltraud X.; Ankenbrand, Markus; Weyer, Anna-Lena Van de; Krol, Elzbieta; Al-Rasheid, Khaled A.; Mithöfer, Axel; Weber, Andreas P.; Schultz, Jörg; Hedrich, Rainer (4 May 2016). "Venus flytrap carnivorous lifestyle builds on herbivore defense strategies". Genome Research. 26 (6): 812–825. doi:10.1101/gr.202200.115. ISSN 1088-9051. PMC 4889972. PMID 27197216. Retrieved 26 May 2016.
  46. ^ a b c Galek H, Osswald WF, Elstner EF (1990). "Oxidative protein modification as predigestive mechanism of the carnivorous plant Dionaea muscipula: an hypothesis based on in vitro experiments". Free Radic Biol Med. 9 (5): 427–434. doi:10.1016/0891-5849(90)90020-J. PMID 2292436.
  47. ^ Hsu YL, Cho CY, Kuo PL, Huang YT, Lin CC (August 2006). "Plumbagin (5-Hydroxy-2-methyl-1,4-naphthoquinone) Induces Apoptosis and Cell Cycle Arrest in A549 Cells through p53 Accumulation via c-Jun NH2-Terminal Kinase-Mediated Phosphorylation at Serine 15 in Vitro and in Vivo". J Pharmacol Exp Ther. 318 (2): 484–494. doi:10.1124/jpet.105.098863. PMID 16632641.
  48. ^ Produced by Neil Lucas (7 December 2009). "Plants". Life. BBC One.
  49. ^ AM Ellison (2006). "Nutrient limitation and stoichiometry of carnivorous plants" (PDF). Plant Biology. 8 (6): 740–747. doi:10.1055/s-2006-923956. PMID 17203429.
  50. ^ Albert, V. A.; Williams, S. E.; Chase, M. W. (11 September 1992). "Carnivorous plants: phylogeny and structural evolution". Science. 257 (5076): 1491–1495. Bibcode:1992Sci...257.1491A. doi:10.1126/science.1523408. ISSN 0036-8075. PMID 1523408.
  51. ^ a b c d e Gibson, T. C.; Waller, D. M. (2009). "Evolving Darwin's 'most wonderful' plant: ecological steps to a snap-trap". New Phytologist. 183 (3): 575–587. doi:10.1111/j.1469-8137.2009.02935.x. PMID 19573135.
  52. ^ a b c Cameron, K. M.; Wurdack, K. J.; Jobson, R. W. (2002). "Molecular evidence for the common origin of snap-traps among carnivorous plants". American Journal of Botany. 89 (9): 1503–1509. doi:10.3732/ajb.89.9.1503. PMID 21665752.
  53. ^ Rivadavia, F.; K. Kondo; M. Kato & M. Hasebe (2003). "Phylogeny of the sundews, Drosera (Droseraceae), based on chloroplast rbcL and nuclear 18S ribosomal DNA Sequences". American Journal of Botany. 90 (1): 123–130. doi:10.3732/ajb.90.1.123. PMID 21659087.
  54. ^ Stokstad, E. (12 May 2016). "How the Venus flytrap acquired its taste for meat". Science. 352 (6287): 756. Bibcode:2016Sci...352..756S. doi:10.1126/science.352.6287.756. PMID 27174967.
  55. ^ Turner, John G.; Ellis, Christine; Devoto, Alessandra (1 May 2002). "The Jasmonate Signal Pathway". The Plant Cell. 14 (suppl 1): S153–S164. doi:10.1105/tpc.000679. ISSN 1532-298X. PMC 151253. PMID 12045275.
  56. ^ a b "Venus flytrap origins uncovered". BBC News. 2009.
  57. ^ Ellison, Aaron M.; Adamec, Lubomír (2018). Carnivorous Plants: Physiology, Ecology, and Evolution. Oxford University Press. ISBN 978-0-19-877984-1.
  58. ^ Jang, Gi-Won; Kim, Kwang-Soo; Park, Ro-Dong (2003). "Micropropagation of Venus fly trap by shoot culture". Plant Cell, Tissue and Organ Culture. 72 (1): 95–98. doi:10.1023/A:1021203811457.
  59. ^ D'Amato, Peter (1998). The Savage Garden: Cultivating Carnivorous Plants. Berkeley, California: Ten Speed Press. ISBN 978-0-89815-915-8.
  60. ^ "Federal Register / Vol. 82 / December 20, 2017 / Proposed Rules" (PDF). Government Publishing Office (US). 20 December 2017. Retrieved 14 December 2019.
  61. ^ "Federal Register / Vol. 58, No. 188" (PDF). cdn.loc.gov. 30 September 1993. Retrieved 14 December 2019.
  62. ^ "The IUCN Red List of Threatened Species". IUCN Red List of Threatened Species. Retrieved 15 December 2019.
  63. ^ "Threatened Search Results". plants.usda.gov. Retrieved 15 December 2019.
  64. ^ "Appendices | CITES". www.cites.org. Retrieved 15 December 2019.
  65. ^ a b "Comprehensive Report Species – Dionaea muscipula". explorer.natureserve.org. Retrieved 15 December 2019.
  66. ^ "Many endangered species face long waits for protection: Scientists concerned about decreasing global diversity as a result". ScienceDaily. Retrieved 15 December 2019.
  67. ^ Jennings, David E.; Rohr, Jason R. (1 May 2011). "A review of the conservation threats to carnivorous plants". Biological Conservation. Ecoregional-scale monitoring within conservation areas, in a rapidly changing climate. 144 (5): 1356–1363. doi:10.1016/j.biocon.2011.03.013. ISSN 0006-3207.
  68. ^ "U.S. Census Bureau QuickFacts: Brunswick County, North Carolina". www.census.gov. Retrieved 15 December 2019.
  69. ^ Luken, James O. (2012). "Long-Term Outcomes of Venus Flytrap (Dionaea muscipula) Establishment". Restoration Ecology. 20 (6): 669–670. doi:10.1111/j.1526-100X.2012.00888.x. ISSN 1526-100X.
  70. ^ Luken, James (2007). "Performance of Dionaea muscipula as influenced by developing vegetation". The Journal of the Torrey Botanical Society. 134 (1): 45–52. doi:10.3159/1095-5674(2007)134[45:PODMAI]2.0.CO;2 – via bioone.org.
  71. ^ Yearsley, Connor (May 2017). "The Venus Flytrap: Conserving the Carnivorous Curiosity". Journal of the American Botanical Council. 14 (5).
  72. ^ "Venus Flytraps in Peril: Why Everyone's Favorite Carnivorous Houseplant Is Under Threat". www.mentalfloss.com. 30 September 2019. Retrieved 15 December 2019.
  73. ^ Mele, Christopher (28 November 2016). "Venus Flytraps Need Protection From Poachers in North Carolina". The New York Times. ISSN 0362-4331. Retrieved 15 December 2019.
  74. ^ Evans, Jon (18 September 2014). "Stealing Venus Flytrap plants now a felony". www.wect.com. Retrieved 21 July 2017.
  75. ^ Love, Shayla (28 July 2016). "A poacher who stole 970 venus flytraps in N.C. is sentenced to prison". Washington Post. Archived from the original on 29 July 2016. Retrieved 15 December 2019.
  76. ^ Still, Johanna F. (18 March 2019). "Man charged with 73 counts of poaching Venus Flytrap, bond at $750,000". Port City Daily. Retrieved 15 December 2019.
  77. ^ "Venus Flytrap State Carnivorous Plant | State Symbols USA". statesymbolsusa.org. Retrieved 17 October 2019.
  78. ^ "Venus Flytrap". American Cancer Society. November 2008. Archived from the original on 18 February 2015. Retrieved 22 September 2013.

External links