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It consists of approximately 180 species in three [[Extant taxon|extant]] genera, the vast majority being in the sundew genus ''[[Drosera]]''. The family also contains the well-known [[Venus flytrap]] (''Dionaea muscipula'') and the more obscure [[waterwheel plant]] (''Aldrovanda vesiculosa''), both of which are the only living species of their respective genera.<ref name="Christenhusz-Byng2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. |name-list-style=amp | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 | publisher = Magnolia Press | doi-access = free }}</ref> Representatives of the Droseraceae are found on all continents except Antarctica. ==Description== Droseraceae are [[carnivorous plant|carnivorous herbaceous plants]] that may be annuals or perennials. Their leaves are alternate and [[Glossary of botanical terms#adaxial|adaxially]] [[Glossary of botanical terms#circinate|circinate]], with at least one leaf surface containing hairs with mucilage-producing glands at the tip. Their flowers are bisexual, usually with three [[carpels]] and five [[sepal]]s, [[petal]]s and [[stamen]]s. Their pollen grains are [[Glossary of botanical terms#triporate|triporate]] or multiporate and released in [[Glossary of botanical terms#tetrad|tetrads]]. Despite being carnivorous, their flowers are insect-pollinated, typically with white to purple flowers that close at night. They produce small seeds that are dispersed by wind and water.<ref name="Judd-2018">{{Cite book|title=Plant Systematics: A Phylogenetic Approach|last1=Judd|first1=Walter|last2=Campbell|first2=Christopher|last3=Kellogg|first3=Elizabeth|last4=Stevens|first4=Peter|last5=Donoghue|first5=Michael|publisher=Sinauer Associates|year=2018|isbn=978-1-60535-389-0|location=MA, USA|pages=459}}</ref> Most of the members of Droseraceae are contained in the genus ''[[Drosera]]'', the sundews. Both ''[[Dionaea (plant)|Dionaea]]'' and ''[[Aldrovanda]]'' have only one extant species. ''Drosera'' species trap prey by secreting a sticky substance from hairs on their leaves. ''Dionaea'' and ''Aldrovanda'' both use snap-traps that close rapidly when the leaves are disturbed. ''Dionaea'' is terrestrial, while ''Aldrovanda'' is strictly aquatic. Like carnivorous plants of other families, the Droseraceae are able to supplement their nutrient intake, especially that of nitrogen, by capturing and digesting small animals such as insects. In this way, these plants are able to thrive in nutrient-deficient areas, such as [[sphagnum bog]]s. ===''Drosera''=== ''Drosera'' is one of the largest genera of carnivorous plants, and individual species vary extensively in their specific morphology. Common to all members of ''Drosera'' are highly modified leaves lined with tentacle-like glandular [[trichome]]s. At the end of each trichome, a bead of highly viscous [[mucilage]] is secreted, which resembles a drop of dew. The mucilage is a fairly pure aqueous solution of acidic polysaccharides with high molecular weights, which makes the mucilage not only highly viscous, but also very sticky,<ref name ="rost">{{ citation | title = Physical and chemical properties of the mucilage secreted by ''Drosera capensis'' | year = 1977 | author = Rost, K. | journal = Phytochemistry | volume = 16 | issue = 9 | pages = 1635–1638 |doi=10.1016/s0031-9422(00)88783-x | last2 = Schauer | first2 = R. }}</ref> so much so, a single drop of mucilage may be stretched to lengths of up to a meter and cover one million times its original surface area.<ref name ="rost"/><ref name="zhang">{{citation | doi=10.1186/1477-3155-8-20 | pmid = 20718990 | title = Nanofibers and nanoparticles from the insect capturing adhesive of the Sundew (''Drosera'') for cell attachment | year = 2010 | author = Zhang, M. | journal = Journal of Nanobiotechnology | volume = 8 | issue = 20 | pages = 20 | last2 = Lenaghan | first2 = S.C. | last3 = Xia | first3 = L. | last4 = Dong | first4 = L. | last5 = He | first5 = W. | last6 = Henson | first6 = W.R. | last7 = Fan | first7 = X. | pmc = 2931452 | doi-access = free }}</ref> Insects and other prey animals are attracted by the smell of this mucilage and become stuck in it. Such snares are termed "flypaper traps", but the trapping mechanism of sundews is often erroneously described as "passive". In fact, sundew traps are quite active and sensitive, and the disturbance of one or a few trichomes quickly triggers an action potential that stimulates the rapid movement of other trichomes toward the prey. The leaf then curls in on itself, enveloping the prey for digestion.<ref name="williams">{{ citation | title = Comparative physiology of the Droseraceae sensu stricto – How do tentacles bend and traps close? | year = 2002 | author = Williams, S.E. | journal = Proceedings of the 4th International Carnivorous Plant Conference | pages = 77–81 }}</ref> Four ''Drosera'' subgenera are recognized today: subgenus ''Regiae'' and subgenus ''Arcturia'' are each monotypic (''D. regia'' and ''D. arcturi,'' respectively), and the remaining ''Drosera'' are divided into two clades, subgenus ''Ergaleium'' and subgenus ''Drosera''.<ref name="Fleischmann-2018">{{Citation|last1=Fleischmann|first1=A.|title=Systematics and evolution of droseraceae|date=2018|url=https://espace.curtin.edu.au/handle/20.500.11937/70040|work=Carnivorous Plants: Physiology, Ecology, and Evolution|pages=45–57|language=en|isbn=978-0-19-877984-1|access-date=2019-12-13|last2=Cross|first2=Adam|last3=Gibson|first3=R.|last4=Gonella|first4=P.|last5=Dixon|first5=Kingsley}}</ref> ===''Dionaea''=== ''[[Dionaea muscipula]]'', better known as the [[Venus flytrap]], is a globally famous carnivorous plant and according to [[Charles Darwin]], "one of the most wonderful in the world."<ref name ="darwin">{{citation | title = Insectivorous Plants | year = 1875 | author = Darwin, C. | publisher = John Murray | location = London, UK | url=http://darwin-online.org.uk }}</ref> The leaves of ''Dionaea'' are also highly modified and form a "snap-trap" that quickly shuts when a stimulus is detected. Three large trichomes extend outward on the inner surface of the trap. Two of these three hairs must be stimulated within a certain amount of time to trigger the trap. The trap closes as the result of a flipping of the trap lobes from a position where the exterior of the trap is concave to one where the exterior is convex. This movement can begin as soon as 0.4 seconds after stimulation and can be completed after one second.<ref name="williams"/> ===''Aldrovanda''=== ''[[Aldrovanda vesiculosa]]'', also called the waterwheel plant, is a free-floating, rootless, aquatic plant.<ref>{{Cite web|url=https://www.iucnredlist.org/species/162346/901031|title=Aldrovanda vesiculosa, Waterwheel|last=Cross|first=A.|date=2012|website=IUCN Red List|access-date=December 13, 2019}}</ref> It is less well-known than its relative ''Dionaea muscipula'', but the two have similar trap structures. In 1875, Darwin described ''Aldrovanda'' as "a miniature aquatic ''Dionaea''".<ref name="darwin" /> The trap of ''Aldrovanda'' is aquatic and is smaller and faster than that of ''Dionaea''.<ref name="williams"/> In addition, while two stimuli are required to close a trap in ''Dionaea'', only one is required in ''Aldrovanda''. The trap of ''Aldrovanda'' closes about ten times faster than that of ''Dionaea''.<ref name="williams"/> ==Etymology== The type genus for the Droseraceae is ''Drosera'', which was described and named by [[Carl Linnaeus|Linnaeus]] in 1753. The name was derived from the Greek word "droseros", meaning "dewy" or "drops of water". The ''Principia Botanica'', published in 1787, states "Sun-dew (''Drosera'') derives its name from small drops of a liquor-like dew, hanging on its fringed leaves, and continuing in the hottest part of the day, exposed to the sun."<ref>{{Cite book|url=https://books.google.com/books?id=ZM4gQwAACAAJ|title="Principia Botanica", Or a Concise and Easy Introduction to the Sexual Botany of Linnaeus, with the "genera", Their Mode of Growth ... the Number of "species" to Each "genus", where Principally Native and the Number Indigenous to the British Isles ... Together with Three Indexes: I. of the Linnaean "genera" Accented, with the British Names ; II. of Such Trivial Names as Were the "genera" of Old Authors ; III. of the British Names, with the Linnaean "genera", to which are Added Many of the Specific Names, Also, a Table of Several Vegetable Drugs Not in the Indexes ...|last=Linné|first=Carl von|date=1787|publisher=G.G.J. and J. Robinson|language=en}}</ref> ==Phylogeny== In 1867, Bentham and Hooker placed six genera in the Droseraceae: ''Dionaea, Aldrovanda, Drosera, Drosophyllum, Byblis,'' and ''Roridula''.<ref name="Ellison-2009">{{Cite journal|last1=Ellison|first1=Aaron M.|last2=Gotelli|first2=Nicholas J.|date=2009-01-01|title=Energetics and the evolution of carnivorous plants—Darwin's 'most wonderful plants in the world'|url=https://academic.oup.com/jxb/article/60/1/19/567619|journal=Journal of Experimental Botany|language=en|volume=60|issue=1|pages=19–42|doi=10.1093/jxb/ern179|pmid=19213724|issn=0022-0957|doi-access=free}}</ref> Although these genera had significant differences in leaf and flower morphologies, they were grouped together on the basis of insect traps that appeared to be homologous.<ref name="Ellison-2018">{{Cite book|url=https://books.google.com/books?id=l_pADwAAQBAJ|title=Carnivorous Plants: Physiology, Ecology, and Evolution|last1=Ellison|first1=Aaron M.|last2=Adamec|first2=Lubomír|date=2018|publisher=Oxford University Press|isbn=978-0-19-877984-1|language=en}}</ref> In 1922, ''Byblis'' and ''Roridula'' were moved into a new family, the Byblidaceae (and later further split out, forming the Roridulaceae).<ref name="Ellison-2018" /> In the 1990s, both morphological and molecular evidence began to build that ''Drosophyllum'' differed from the other genera in the Droseraceae,<ref name="williams94" /> ''Drosophyllum'', another monotypic genus (''[[Drosophyllum lusitanicum]]'' being the only species'')'', exhibits a flypaper-type trap similar to those of ''Drosera'', but ''Drosophyllum'' does not actively curl its leaves to envelop captured prey animals. This important morphological distinction led researchers to question the validity of this taxon's placement in Droseraceae. Other significant trait differences in ''Drosophyllum'' include [[pollen]] structure, trichome anatomy, and a woody stem with a deep taproot.<ref name="williams94" /> Ultimately, ''Drosophyllum'' was shown to be more closely related to the carnivorous [[liana]] ''[[Triphyophyllum]]'' and the noncarnivorous liana ''[[Ancistrocladus]]'', and is, thus, classified elsewhere (to be specific, its own monotypic family [[Drosophyllaceae]]).<ref name="cameron" /> and APG III (2009) placed it into its own family, the Drosophyllaceae.<ref name="Ellison-2018" /> This left only the three genera (''Dionaea, Aldrovanda, Drosera'') that are classified as Droseraceae today. Despite some debate, taxonomists have tended to include at least two of these three genera, and, in general, all three, in this family since at least 1906.<ref name="cameron">{{citation | doi = 10.3732/ajb.89.9.1503 | title = Molecular evidence for the common origin of snap-traps among carnivorous plants | year = 2002 | author = Cameron, K. M. | journal = American Journal of Botany | volume = 89 | issue = 9 | pages = 1503–9 | last2 = Wurdack | first2 = K. J. | last3 = Jobson | first3 = R. W. | pmid=21665752}}</ref> Separate families for ''Dionaea'' and ''Aldrovanda'' have been proposed in the past. These were Dionaecae, proposed in 1933, and Aldrovandaceae, proposed in 1949.<ref name="williams94">{{citation | title = Relationships of Droseraceae: a cladistic analysis of ''rbcL'' sequence and morphological data | year = 1994 | author = Williams, S.E. | journal = American Journal of Botany | volume = 81 | issue = 8 | pages = 1027–1037 | last2 = Albert | first2 = V.A. | last3 = Chase | first3 = M.W. | doi=10.2307/2445297| jstor = 2445297 }}</ref> Ultimately, molecular and morphological evidence support the inclusion of all three, confirming that the Droseraceae are a [[Monophyly|monophyletic]] group.<ref name="cameron" /> Molecular evidence also shows that the two genera with traps that snap shut (''Dionaea'' and ''Aldrovanda'') are more closely related to each other than to ''Drosera'', suggesting snap traps evolved only once.<ref name="cameron" /> The family Droseraceae is part of the order [[Caryophyllales]] in the Superasterid clade within the [[core eudicots]].<ref name="cameron" /><ref name="Judd-2018" /> The family totals nearly 200 species. Caryophyllales are divided into two major suborders: [[Caryophyllineae]], which contains the "core" Caryophyllales, such as [[Cactaceae]] and [[Amaranthaceae]] and is sister to the [[Polygonineae]] – the "non-core" Caryophyllales. This non-core clade is where Droseraceae is placed.<ref name="cuenoud">{{citation | title = Molecular phylogenetics of Caryophyllales based on nuclear 18S rDNA and plastid rbcL, atpB, and matK DNA sequences | year = 2002 | author = Cuénoud, P. | journal = American Journal of Botany | volume = 89 | issue = 1 | pages = 132–144 | last2 = Savolainen | first2 = V. | last3 = Chatrou | first3 = L.W. | last4 = Powell | first4 = M. | last5 = Grayer | first5 = R.J. | last6 = Chase | first6 = M.W. | doi=10.3732/ajb.89.1.132 | pmid=21669721}}</ref> Recent molecular and biochemical evidence<ref>{{Cite web|url=http://www.mobot.org/MOBOT/Research/APweb/welcome.html|title=ANGIOSPERM PHYLOGENY WEBSITE, version 14}}</ref> suggests the carnivorous taxa in the order [[Caryophyllales]] (the families Droseraceae, [[Drosophyllaceae]], [[Nepenthaceae]], and the species ''[[Triphyophyllum peltatum]]'') all belong to the same [[clade]], which does not consist only of carnivorous plants, but also of some noncarnivorous plants such as those in the family [[Ancistrocladaceae]]. The fossil record of Droseraceae is the richest of any carnivorous plant family. Fossil [[pollen]] has been attributed to several [[Extant taxon|extant]], as well as [[extinction|extinct]], genera, although some are of questionable validity. ==Evolution== Darwin concluded that carnivory in plants was [[Convergent evolution|convergent]], writing in 1875 that ''[[Utricularia]]'' and ''[[Nepenthes]]'' were not "at all related to the Droseraceae".<ref name="darwin" /> This remained a subject of debate for over a century. In 1960, [[Léon Croizat|Leon Croizat]] concluded that carnivory was monophyletic, and placed all the carnivorous plants together at the base of the angiosperms.<ref name="Ellison-2009" /> Molecular studies over the past 30 years have led to a wide consensus that Darwin was correct, with studies showing that carnivory evolved at least six times in the angiosperms, and that trap designs such as pitcher traps and flypaper traps are analogous rather than homologous.<ref>{{Cite journal|last1=Albert|first1=V. A.|last2=Williams|first2=S. E.|last3=Chase|first3=M. W.|date=1992-09-11|title=Carnivorous plants: phylogeny and structural evolution|journal=Science|language=en|volume=257|issue=5076|pages=1491–1495|doi=10.1126/science.1523408|issn=0036-8075|pmid=1523408|bibcode=1992Sci...257.1491A }}</ref> The origin of carnivory within the ancestors of the Droseraceae has been dated to 85.6 million years ago, with the evolution of snap-traps dated to 48 million years ago.<ref name="Ellison-2018" /> Researchers have hypothesized that carnivory in the Droseraceae began with simple flypaper traps, followed by movement of tentacles in some ''Drosera''-like species, followed by movement of leaves, leading eventually to the development of snap-traps in ''Dionaea'' and ''Aldrovanda'' by increasing the speed of the leaf movements and altering the morphology of the leaves.<ref>{{Cite journal|last1=Heubl|first1=G.|last2=Bringman|first2=G.|last3=Meimberg|first3=H.|date=November 2006|title=Molecular Phylogeny and Character Evolution of Carnivorous Plant Families in Caryophyllales - Revisited|journal=Plant Biology|volume=8|issue=6|pages=821–830|doi=10.1055/s-2006-924460|pmid=17066364|bibcode=2006PlBio...8..821H }}</ref> <ref>{{Cite journal|last1=Poppinga|first1=Simon|last2=Hartmeyer|first2=Siegfried R. H.|last3=Masselter|first3=Tom|last4=Hartmeyer|first4=Irmgard|last5=Speck|first5=Thomas|date=2013-07-01|title=Trap diversity and evolution in the family Droseraceae|journal=Plant Signaling & Behavior|volume=8|issue=7|pages=e24685|doi=10.4161/psb.24685|pmc=3907454|pmid=23603942|bibcode=2013PlSiB...8E4685P }}</ref> Due to the sister relationship of ''Dionaea'' and ''Aldrovanda,'' it is likely that the snap-trap mechanism only evolved once, but it is unknown if the common ancestor was terrestrial or aquatic.<ref>{{Cite journal|last1=Rivadavia|first1=Fernando|last2=Kondo|first2=Katsuhiko|last3=Kato|first3=Masahiro|last4=Hasebe|first4=Mitsuyasu|date=2003|title=Phylogeny of the sundews, Drosera (Droseraceae), based on chloroplast rbcL and nuclear 18S ribosomal DNA Sequences|journal=American Journal of Botany|language=en|volume=90|issue=1|pages=123–130|doi=10.3732/ajb.90.1.123|issn=1537-2197|pmid=21659087|doi-access=free}}</ref> ==Gallery== <gallery perrow="5"> File:Drosera communis.jpg|''Drosera communis'' File:Drosera rotundifolia leaf1.jpg|''Drosera rotundifolia'' File:Drosera filiformis leaf Darwiniana.jpg|''Drosera filiformis'' File:AldrovandaVesiculosa4.jpg|''Aldrovanda vesiculosa'' File:Dionaea, muscoid fly.jpg|''Dionaea muscipula'' </gallery> ==References== {{Reflist}} ==External links== * {{Commons category-inline}} {{Angiosperm families}} {{Taxonbar|from=Q156185}} {{Authority control}} [[Category:Droseraceae| ]] [[Category:Caryophyllales families]] </textarea><div class="templatesUsed"><div class="mw-templatesUsedExplanation"><p><span id="templatesused">Pages transcluded onto the current version of this page<span class="posteditwindowhelplinks"> (<a href="/wiki/Help:Transclusion" title="Help:Transclusion">help</a>)</span>:</span> </p></div><ul> <li><a href="/wiki/Template:-" class="mw-redirect" title="Template:-">Template:-</a> (<a href="/w/index.php?title=Template:-&action=edit" class="mw-redirect" title="Template:-">view source</a>) (template 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