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Araceae

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Araceae
Temporal range:115–0MaEarly Cretaceous[1]- Recent
Inflorescence ofSpathiphyllum cochlearispathum
Scientific classificationEdit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Order: Alismatales
Family: Araceae
Juss.[2]
Subfamilies
Snake lily (Dracunculus vulgaris) inCrete
Arrowleaf elephant ear (Xanthosoma sagittifolium), clearly showing the characteristic spadix and spathe

TheAraceaeare afamilyofmonocotyledonousflowering plantsin whichflowersare borne on a type ofinflorescencecalled aspadix.The spadix is usually accompanied by, and sometimes partially enclosed in, aspathe(or leaf-likebract). Also known as thearum family,members are often colloquially known asaroids.This family of 114generaand about 3,750 known species[3]is most diverse in theNew Worldtropics, although also distributed in theOld Worldtropics and northerntemperate regions.

Description

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Within the Araceae, species are oftenrhizomatousortuberous;many areepiphytic,creepinglianasorviningplants, and theleavesand tissues of the entire plant nearly always contains irritatingcalcium oxalatecrystals orraphides,in varying degrees.[4][5]The foliage can vary considerably from species to species. The majority of species produce aninflorescenceconsisting of a spadix (which some compare to acorn cob,in appearance), which is nearly always surrounded by a modified leaf bract called aspathe.[6]Inmonoeciousaroids, possessing separate male and female flowers (but with both flowers present on one plant), the spadix is usually organized with female flowers towards the bottom and male flowers at the top. In aroids withperfect flowers,thestigmais no longer receptive when thepollenis released, thus preventing self-fertilization.Some species aredioecious.[7]

Many plants in this family arethermogenic(heat-producing).[8]Their flowers can reach up to 45 °C, even if the surrounding air temperature is much lower. One reason for this unusually high temperature is to attract insects (usuallybeetles) to pollinate the plant, rewarding the beetles with heat energy, in addition to preventing tissue damage in colder regions. Some examples of thermogenic aroids areSymplocarpus foetidus(eastern skunk-cabbage),Amorphophallus titanum(titan arum),Amorphophallus paeoniifolius(elephant-foot yam),Helicodiceros muscivorus(dead-horse arum lily), andSauromatum venosum(voodoo lily). Some species, such asA.titanumandH.muscivorus,give off a very pungent smell akin to rotten meat, which serves to attract flies for pollination. The heat produced by the plant helps to convey the scent further.

Toxicity

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Within the Araceae family, the majority of species produce calcium oxalate crystals in the form ofraphides.While it is possible to consume the cooked foliage of certain genera, such asAlocasia,Colocasia,andXanthosoma,as well as the ripened fruits ofMonstera deliciosa,these raphide compounds are irritating (and even dangerous) for many animals, including humans. Consumption of raw aroid vegetation may causeedema,vesicleformation ordysphagia,accompanied by a painful stinging and burning in the mouth and throat, with symptoms occurring for up to two weeks, depending on amount consumed. In smaller amounts, patients report feeling a mild to extreme sensation of sand or glass in the esophagus and mouth, lasting up to 48 hours.[9]Additionally, in heavier instances of ingestion,anaphylactic shockcould cause swelling of the throat, restricting breathing. The genusDieffenbachiais famously known as "dumb-cane" for this reason; however, given the presence of irritating compounds across the family, this nickname may be applied to virtually any genera within the Araceae.

Taxonomy

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Phylogeny

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Phylogeny based on theAngiosperm Phylogeny Website.[10]

Araceae

GymnostachydoideaeBogner & Nicolson 1991

OrontioideaeBrown ex Müller 1860

Lemnoideae

PothoideaeEngler 1876

MonsteroideaeEngler 1876

LasioideaeEngler 1876

ZamioculcadoideaeBogner & Hesse 2005

AroideaeArnott 1832

Classification

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One of the earliest observations of species in the Araceae was conducted byTheophrastusin his workEnquiry into Plants.[11]The Araceae were not recognized as a distinct group of plants until the 16th century. In 1789,Antoine Laurent de Jussieuclassified all climbing aroids asPothosand all terrestrial aroids as eitherArumorDracontiumin his bookFamilles des Plantes.[citation needed]

The first major system of classification for the family was produced byHeinrich Wilhelm Schott,who publishedGenera Aroidearumin 1858 andProdromus Systematis Aroidearumin 1860. Schott's system was based on floral characteristics, and used a narrow conception of a genus.Adolf Englerproduced a classification in 1876, which was steadily refined up to 1920. His system is significantly different from Schott's, being based more on vegetative characters and anatomy. The two systems were to some extent rivals, with Engler's having more adherents before the advent ofmolecular phylogeneticsbrought new approaches.[12]

A comprehensive taxonomy of Araceae was published by Mayo et al. in 1997.[13]

Modern studies based on gene sequences show the Araceae (including theLemnoideae,duckweeds) to bemonophyletic,and the first diverging group within theAlismatales.[14]TheAPG III systemof 2009 recognizes the family, including the genera formerly segregated in the Lemnaceae.[15]The sinking of the Lemnaceae into the Araceae was not immediately universally accepted. For example, the 2010New Flora of the British Islesused aparaphyleticAraceae and a separate Lemnaceae.[16]HoweverLemnaand its allies were incorporated in Araceae in the 2019 edition.[17]: 872 A comprehensivegenomicstudy ofSpirodela polyrhizawas published in February 2014.[18]

Genera

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The cuckoo-pint or lords and ladies (Arum maculatum) is a common arum in British woodlands.
Arisaema triphyllum

143 genera are accepted within the Araceae.[19]Anthurium,Epipremnum,Monstera,PhilodendronandZantedeschiaare some of the most well-known genera of the family, as are theColocasia(taro,arbi) andXanthosoma('elephant-ear', ‘ape), which are both cultivated for human consumption. The largest unbranched inflorescence in the world is that of the arumAmorphophallus titanum(titan arum).[20]

The Araceae includes many ornamental genera of global economic importance:Aglaonema,Alocasia,Anthurium,Caladium,Dieffenbachia,Epipremnum,Homalomena,Monstera,Nephthytis,Rhaphidophora,Scindapsus,Spathiphyllum,Syngonium,andZamioculcas,to name but a few. The aquatic generaAnubias,BucephalandraandCryptocoryneare highly prized and cultivated aquarium plants; other, recently-described genera, such as theLagenandraof India, are gradually becoming more known in theaquascapingworld.[21]Philodendronis an important genus in the ecosystems ofneotropicalrainforests,and is widely used in home and interior decorating.Symplocarpusfoetidus(skunk cabbage) is a common eastern North American species. An interesting peculiarity is that this family includes the largest unbranched inflorescence, that of thetitan arum,[20]often erroneously called the "largest flower", and the smallest flowering plant and smallest fruit, in theduckweed,Wolffia.[22]

Fossil record

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The family Araceae has one of the oldestfossilrecord amongangiosperms,with fossil forms first appearing during theEarly Cretaceousepoch.[1][23]Notable fossils from the Early Cretaceous include:Spixiarum kipea,[24]an aroid from the lateAptianof Brazil;[1]Orontiophyllum ferreri,an aroid leaf from the lateAlbianof Spain;[1]andTurolospadix bogneri,an aroid spadix from the late Albian of Spain.[1]

Food plants

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Food plants in the family Araceae includeAmorphophallus paeoniifolius(elephant foot yam),Colocasia esculenta(kochu, taro, dasheen),Xanthosoma(cocoyam, tannia),Typhonium trilobatumandMonstera deliciosa(Mexican breadfruit). While the aroids are little traded, and overlooked by plant breeders to the extent that the Crop Trust calls them "orphan crops", they are widely grown and are important insubsistence agricultureand in local markets. The main food product is thecorm,which is high instarch;leaves and flowers also find culinary use.[25]

See also

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References

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  1. ^abcdeSender, L.M., Doyle, J.A., Upchurch, J.R. Jr., Villanueva-Amadoz, U. and Diez J.B. 2019. Leaf and inflorescence evidence for near-basal Araceae and an unexpected diversity of other monocots from the late Early Cretaceous of Spain. Journal of Systematic Palaeontology, vol. 17, p. 1093–1126.
  2. ^Angiosperm Phylogeny Group (2009), "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III",Botanical Journal of the Linnean Society,161(2): 105–121,doi:10.1111/j.1095-8339.2009.00996.x,hdl:10654/18083
  3. ^Christenhusz, M. J. M. & Byng, J. W. (2016)."The number of known plants species in the world and its annual increase".Phytotaxa.261(3). Magnolia Press: 201–217.doi:10.11646/phytotaxa.261.3.1.
  4. ^"Araceae in Flora of North America @ efloras.org".Efloras.org.Retrieved17 January2014.
  5. ^"Araceae".The Encyclopedia of House Plants.Retrieved30 June2024.
  6. ^"Araceae - Flowering Plant Families, UH Botany".University of Hawaii.Retrieved17 January2014.
  7. ^"Araceae"(PDF).Iowa State University.Archived fromthe original(PDF)on 3 May 2013.Retrieved17 January2014.
  8. ^Korotkova, Nadja; Barthlott, Wilhelm (2009)."On the thermogenesis of the Titan arum (Amorphophallus titanum) ".Plant Signaling & Behavior.4(11): 1096–1098.doi:10.4161/psb.4.11.9872.PMC2819525.PMID19838070.
  9. ^Watson, John T.; Jones, Roderick C.; Siston, Alicia M.; Diaz, Pamela S.; Gerber, Susan I.; Crowe, John B.; Satzger, R. Duane (2005). "Outbreak of Food-borne Illness Associated with Plant Material Containing Raphides".Clinical Toxicology.43(1): 17–21.doi:10.1081/CLT-44721.PMID15732442.S2CID388923.
  10. ^Stevens, P.F. (2001)."Araceae".Angiosperm Phylogeny Website.13.Retrieved30 December2017.
  11. ^Bown, Deni (2000).Aroids: plants of the Arum family.Timber Press. p. 46.ISBN0881924857.
  12. ^Grayum, Michael H. (1990), "Evolution and Phylogeny of the Araceae",Annals of the Missouri Botanical Garden,77(4): 628–697,doi:10.2307/2399668,JSTOR2399668
  13. ^Mayo, S. J.; Bogner, J.; Boyce, P. C. (1997).The genera of Araceae.London: Royal Botanic Gardens, Kew.ISBN1-900347-22-9.OCLC60140655.
  14. ^Stevens, P.F."Araceae".Angiosperm Phylogeny Website.
  15. ^Angiosperm Phylogeny Group III (2009)."An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III".Botanical Journal of the Linnean Society.161(2): 105–121.doi:10.1111/j.1095-8339.2009.00996.x.hdl:10654/18083.
  16. ^Stace, C.A.(2010),New Flora of the British Isles(Third ed.), Cambridge, UK: Cambridge University Press,ISBN978-0-521-70772-5pp. 830–834.
  17. ^Stace, C. A.(2019).New Flora of the British Isles(Fourth ed.). Middlewood Green, Suffolk, U.K.: C & M Floristics.ISBN978-1-5272-2630-2.
  18. ^Wang, W.; Haberer, G.; Gundlach, H.; Gläßer, C.; Nussbaumer, T.; Luo, M. C.; Lomsadze, A.; Borodovsky, M.; Kerstetter, R. A.; Shanklin, J.; Byrant, D. W.; Mockler, T. C.; Appenroth, K. J.; Grimwood, J.; Jenkins, J.; Chow, J.; Choi, C.; Adam, C.; Cao, X.-H.; Fuchs, J.; Schubert, I.; Rokhsar, D.; Schmutz, J.; Michael, T. P.; Mayer, K. F. X.; Messing, J (2014)."TheSpirodela polyrhizagenome reveals insights into its neotenous reduction fast growth and aquatic lifestyle ".Nature Communications.5:3311.Bibcode:2014NatCo...5.3311W.doi:10.1038/ncomms4311.PMC3948053.PMID24548928.
  19. ^Araceae Juss.Plants of the World Online.Retrieved 28 April 2024.
  20. ^ab"Titan Arum FAQs | Biological Sciences Greenhouse".Ohio State University.2012-05-16.Retrieved17 January2014.
  21. ^"Aquarium Cryptocoryne Plants".Aquarium Fish International.Retrieved17 January2014.
  22. ^"What is the smallest flower in the world?".Library of Congress.Retrieved17 January2014.
  23. ^Nauheimer, L., Metzler, D. and Renner, S.S. 2012. Global history of the ancient monocot family Araceae inferred with models accounting for past continental positions and previous ranges based on fossils. New Phytologist, vol. 195, p. 938-950.
  24. ^Coiffard, C., Mohr, B.A.R. and Bernardes de Oliveira, M.E.C. 2013. The Early Cretaceous aroid,Spixiarum kipeagen. et sp. nov., and implications on early dispersal and ecology of basal monocots. Taxon, vol. 62. p. 997-1008.
  25. ^"Aroids. Colocasia Xanthosoma".The Crop Trust.Retrieved2019-04-06.

Further reading

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  • Bown, Deni (2000). Aroids: Plants of the Arum Family [ILLUSTRATED]. Timber Press.ISBN0-88192-485-7
  • Croat, Thomas B (1998). "History and Current Status of Systematic Research with Araceae".Aroideana.21.online
  • Grayum, Michael H (1990). "Evolution and Phylogeny of the Araceae".Annals of the Missouri Botanical Garden.77(4): 628–697.doi:10.2307/2399668.JSTOR2399668.
  • Keating R C (2004). "Vegetative anatomical data and its relationship to a revised classification of the genera of Araceae".Annals of the Missouri Botanical Garden.91(3): 485–494.JSTOR3298625.
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