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The Araceae are a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
monocotyledon Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
ous
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s in which
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s are borne on a type of
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a
spathe In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
(or leaf-like
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
). Also known as the arum family, members are often colloquially known as aroids. This family of 140
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and about 4,075 known species is most diverse in the
New World The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...
tropics, although also distributed in the
Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by the ...
tropics and northern
temperate regions In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
. Many species display very decorative leaves and flowers, and they are widely used for gardening; popular as indoor plants and also outdoor plants where climates are mild, and winter freezes will not generally occur. However, some temperate species are also very popular in
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the e ...
-climate gardening, or in moderately cool temperate zones, such as ''
Zantedeschia ''Zantedeschia'' is a genus of eight species of herbaceous, perennial, flowering plants in the family Araceae, native to southern Africa from South Africa north to Malawi. The genus has been introduced on all continents except Antarctica. Com ...
''.


Description

Species within Araceae are often
rhizomatous In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow ho ...
or
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
ous, and the leaves nearly always contain
calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrate ...
crystals or
raphide Raphides (pronounced /ˈræfɪˌdiz/, singular raphide /ˈreɪfʌɪd/ or raphis) are needle-shaped crystals of calcium oxalate monohydrate (prismatic monoclinic crystals) or calcium carbonate as aragonite ( dipyramidal orthorhombic crystals), fo ...
s, in varying degrees. The
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
can vary considerably from species to species. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
is composed of a spadix, which is almost always surrounded by a modified leaf called a
spathe In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or of ...
. In
monoecious Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system alongside gynomonoecy, andromonoecy and trimonoecy. Monoecy is conne ...
aroids (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 towards the top. In aroids with
perfect flower Plant reproductive morphology is the study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction. Among all living organisms, flowers, which are the reproductive s ...
s, the stigma is no longer receptive when the
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
is released, thus preventing self-
fertilization Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a new individual organism or offspring and initiate its development. Proce ...
. Some species are
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
. Many plants in this family are
thermogenic Thermogenic means tending to produce heat, and the term is commonly applied to drugs which increase heat through metabolic stimulation, or to microorganisms which create heat within organic waste. Approximately all enzymatic reaction in the huma ...
(heat-producing). Their flowers can reach up to 45 °C even when the surrounding air temperature is much lower. One reason for this unusually high temperature is to attract insects (usually beetles) to pollinate the plant, rewarding the beetles with heat energy. Another reason is to prevent tissue damage in cold regions. Some examples of thermogenic Araceae are:'' Symplocarpus foetidus'' (eastern skunk cabbage),''
Amorphophallus titanum ''Amorphophallus'' (from Ancient Greek , "without form, misshapen" + ''phallos'', "penis", referring to the shape of the prominent spadix) is a large genus of some 200 tropical and subtropical tuberous herbaceous plants from the ''Arum'' family ...
'' (titan arum),''
Amorphophallus paeoniifolius ''Amorphophallus paeoniifolius'', the elephant foot yam or whitespot giant arum, is a tropical tuber crop grown primarily in Africa, South Asia, Southeast Asia and the tropical Pacific islands. Because of its production potential and popularit ...
'' (elephant foot yam), '' Helicodiceros muscivorus'' (dead horse arum lily), and ''
Sauromatum venosum ''Sauromatum venosum'' (syn. ''Typhonium venosum'')''Sauromatum venosum''.
Pl ...
'' (voodoo lily). Some species, such as titan arum and the dead horse arum, 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

Within the Araceae, genera such as ''
Alocasia ''Alocasia'' is a genus of rhizomatous or tuberous, broad-leaved, perennial, flowering plants from the family Araceae. There are about 90 accepted species native to tropical and subtropical Asia and eastern Australia. Around the world, many growe ...
'', ''
Arisaema ''Arisaema'' is a large and diverse genus of the flowering plant family Araceae. The largest concentration of species is in China and Japan, with other species native to other parts of southern Asia as well as eastern and central Africa, Mexico ...
'', ''
Caladium ''Caladium'' is a genus of flowering plants in the family Araceae. They are often known by the common name elephant ear (which they share with the closely related genera '' Alocasia'', ''Colocasia'', and ''Xanthosoma''), heart of Jesus, and ange ...
'', ''
Colocasia ''Colocasia'' is a genus of flowering plants in the family Araceae, native to southeastern Asia and the Indian subcontinent. Some species are widely cultivated and naturalized in other tropical and subtropical regions. The names elephant-ear ...
'', ''
Dieffenbachia ''Dieffenbachia'' , commonly known as dumb cane or leopard lily, is a genus of tropical flowering plants in the family Araceae. It is native to the New World Tropics from Mexico and the West Indies south to Argentina. Some species are widely cul ...
'', '' Syngonium'', ''
Epipremnum ''Epipremnum'' is a genus of flowering plants in the family (biology), family Araceae, found in tropics, tropical forests from China, the Himalayas, and Southeast Asia to Australia the western Pacific. They are evergreen perennial plant, perennia ...
'', ''
Aglaonema ''Aglaonema'' is a genus of flowering plants in the arum family, Araceae. They are native to tropical and subtropical regions of Asia and New Guinea.
'' and ''
Philodendron ''Philodendron'' is a large genus of flowering plants in the family (biology), family Araceae. , the World Checklist of Selected Plant Families accepted 489 species; other sources accept different numbers. Regardless of number of species, the gen ...
'' contain calcium oxalate crystals in the form of raphides. When accidentally consumed, these may cause
edema Edema, also spelled oedema, and also known as fluid retention, dropsy, hydropsy and swelling, is the build-up of fluid in the body's Tissue (biology), tissue. Most commonly, the legs or arms are affected. Symptoms may include skin which feels t ...
, vesicle formation, and
dysphagia Dysphagia is difficulty in swallowing. Although classified under "symptoms and signs" in ICD-10, in some contexts it is classified as a disease#Terminology, condition in its own right. It may be a sensation that suggests difficulty in the passag ...
, accompanied by painful stinging and burning in the mouth and throat, with symptoms occurring for up to two weeks after ingestion. Patients report feeling a mild to extreme sensation of sand or glass in the esophagus and mouth, depending on amount ingested.


Taxonomy


Phylogeny

Phylogeny based on the
Angiosperm Phylogeny Website The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis ...
.


Classification

One of the earliest observations of species in the Araceae was conducted by
Theophrastus Theophrastus (; grc-gre, Θεόφραστος ; c. 371c. 287 BC), a Greek philosopher and the successor to Aristotle in the Peripatetic school. He was a native of Eresos in Lesbos.Gavin Hardy and Laurence Totelin, ''Ancient Botany'', Routledge ...
in his work ''
Enquiry into Plants Theophrastus's ''Enquiry into Plants'' or ''Historia Plantarum'' ( grc-gre, Περὶ φυτῶν ἱστορία, ''Peri phyton historia'') was, along with his mentor Aristotle's ''History of Animals'', Pliny the Elder's '' Natural History'' a ...
''. The Araceae were not recognized as a distinct group of plants until the 16th century. In 1789,
Antoine Laurent de Jussieu Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an e ...
classified all climbing aroids as ''Pothos'' and all terrestrial aroids as either ''Arum'' or ''Dracontium'' in his book ''Familles des Plantes''. The first major system of classification for the family was produced by
Heinrich Wilhelm Schott Heinrich Wilhelm Schott (7 January 1794 in Brünn (Brno), Moravia – 5 March 1865 at Schönbrunn Palace, Vienna) was an Austrian botanist well known for his extensive work on aroids (Araceae). He studied botany, agriculture and chemistry at ...
, who published ''Genera Aroidearum'' in 1858 and ''Prodromus Systematis Aroidearum'' in 1860. Schott's system was based on floral characteristics, and used a narrow conception of a genus.
Adolf Engler Heinrich Gustav Adolf Engler (25 March 1844 – 10 October 1930) was a German botanist. He is notable for his work on plant taxonomy and phytogeography, such as ''Die natürlichen Pflanzenfamilien'' (''The Natural Plant Families''), edited with ...
produced 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 of
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
brought new approaches. A comprehensive taxonomy of Araceae was published by Mayo et al. in 1997. Modern studies based on gene sequences show the Araceae (including the
Lemnoideae Lemnoideae is a subfamily of flowering aquatic plants, known as duckweeds, water lentils, or water lenses. They float on or just beneath the surface of still or slow-moving bodies of fresh water and wetlands. Also known as bayroot, they arose fr ...
, duckweeds) to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, and the first diverging group within the
Alismatales The Alismatales (alismatids) are an order of flowering plants including about 4,500 species. Plants assigned to this order are mostly tropical or aquatic. Some grow in fresh water, some in marine habitats. Description The Alismatales compri ...
. The
APG III system The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a fur ...
of 2009 recognizes the family, including the genera formerly segregated in the Lemnaceae. The sinking of the Lemnaceae into the Araceae was not immediately universally accepted. For example, the 2010 ''New Flora of the British Isles'' used a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
Araceae and a separate Lemnaceae. pp. 830–834. However ''Lemna'' and its allies were incorporated in Araceae in the 2019 edition. A comprehensive
genomic Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...
study of ''
Spirodela polyrhiza ''Spirodela polyrhiza'' ( ''S. polyrrhiza'') is a species of duckweed known by the common names common duckmeat, greater duckweed, great duckmeat, common duckweed, and duckmeat. It can be found nearly worldwide in many types of freshwater habitat ...
'' was published in February 2014.


Genera

''
Anthurium ''Anthurium'' (; Schott, 1829) is a genus of about 1,000Mantovani, A. and T. E. Pereira. (2005)''Anthurium'' (section ''Urospadix''; subsection ''Flavescentiviridia'').''Rodriguesia'' 56(88), 145–60. species of flowering plants, the largest ...
'' and ''
Zantedeschia ''Zantedeschia'' is a genus of eight species of herbaceous, perennial, flowering plants in the family Araceae, native to southern Africa from South Africa north to Malawi. The genus has been introduced on all continents except Antarctica. Com ...
'' are two well-known members of this family, as are ''Colocasia esculenta'' (
taro Taro () (''Colocasia esculenta)'' is a root vegetable. It is the most widely cultivated species of several plants in the family Araceae that are used as vegetables for their corms, leaves, and petioles. Taro corms are a food staple in Africa ...
) and ''
Xanthosoma ''Xanthosoma'' is a genus of flowering plants in the arum family, Araceae. The genus is native to tropical America but widely cultivated and naturalized in other tropical regions. Several are grown for their starchy corms, an important food stap ...
roseum'' (elephant ear or ‘ape). The largest unbranched inflorescence in the world is that of the arum ''Amorphophallus titanum'' (titan arum). The family includes many ornamental plants: ''
Dieffenbachia ''Dieffenbachia'' , commonly known as dumb cane or leopard lily, is a genus of tropical flowering plants in the family Araceae. It is native to the New World Tropics from Mexico and the West Indies south to Argentina. Some species are widely cul ...
'', ''
Aglaonema ''Aglaonema'' is a genus of flowering plants in the arum family, Araceae. They are native to tropical and subtropical regions of Asia and New Guinea.
'', ''
Caladium ''Caladium'' is a genus of flowering plants in the family Araceae. They are often known by the common name elephant ear (which they share with the closely related genera '' Alocasia'', ''Colocasia'', and ''Xanthosoma''), heart of Jesus, and ange ...
'', '' Nephthytis'', and ''
Epipremnum ''Epipremnum'' is a genus of flowering plants in the family (biology), family Araceae, found in tropics, tropical forests from China, the Himalayas, and Southeast Asia to Australia the western Pacific. They are evergreen perennial plant, perennia ...
'', to name a few. The genera '' Cryptocoryne'', ''
Anubias ''Anubias'' is a genus of aquatic and semi-aquatic flowering plants in the family Araceae, native to tropical central and western Africa. They primarily grow in rivers and streams, but can also be found in marshes. They are characterized by broad ...
'' and ''
Bucephalandra ''Bucephalandra'' is a genus of flowering plants in the family Araceae. There are 30 species of ''Bucephalandra'' which have been discovered in Borneo and have been formally described by S.Y. Wong and P.C. Boyce. Most of the species are found i ...
'' are many popular aquarium plants. ''
Philodendron ''Philodendron'' is a large genus of flowering plants in the family (biology), family Araceae. , the World Checklist of Selected Plant Families accepted 489 species; other sources accept different numbers. Regardless of number of species, the gen ...
'' is an important plant in the ecosystems of the
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainfores ...
s and is often used in home and interior decorating. ''Symplocarpus foetidus'' (skunk cabbage) is a common eastern North American species. An interesting peculiarity is that this family includes the largest unbranched inflorescence, that of the
titan arum Titan most often refers to: * Titan (moon), the largest moon of Saturn * Titans, a race of deities in Greek mythology Titan or Titans may also refer to: Arts and entertainment Fictional entities Fictional locations * Titan in fiction, fictio ...
, often erroneously called the "largest flower" and the smallest flowering plant and smallest fruit, found in the duckweed, ''
Wolffia ''Wolffia'' is a genus of aquatic plants with a cosmopolitan distribution. They include the smallest flowering plants on Earth. Commonly called watermeal or duckweed, these aquatic plants resemble specks of cornmeal floating on the water. Indivi ...
''.


Fossil record

The family Araceae has one of the oldest
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record among
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
, with fossil forms first appearing during the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
epoch.Sender, 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.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. Notable fossils from the Early Cretaceous include: ''Spixiarum kipea'',Coiffard, C., Mohr, B.A.R. and Bernardes de Oliveira, M.E.C. 2013. The Early Cretaceous aroid, ''Spixiarum kipea'' gen. et sp. nov., and implications on early dispersal and ecology of basal monocots. Taxon, vol. 62. p. 997-1008. an aroid from the late
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
of Brazil; ''Orontiophyllum ferreri'', an aroid leaf from the late
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0.9 M ...
of Spain; and ''Turolospadix bogneri'', an aroid spadix from the late Albian of Spain.


Food plants

Food plants in the family Araceae include ''
Amorphophallus paeoniifolius ''Amorphophallus paeoniifolius'', the elephant foot yam or whitespot giant arum, is a tropical tuber crop grown primarily in Africa, South Asia, Southeast Asia and the tropical Pacific islands. Because of its production potential and popularit ...
'' (elephant foot yam), ''
Colocasia esculenta Taro () (''Colocasia esculenta)'' is a root vegetable. It is the most widely cultivated species of several plants in the family Araceae that are used as vegetables for their corms, leaves, and petioles. Taro corms are a food staple in African ...
'' (kochu, taro, dasheen), ''Xanthosoma'' (cocoyam, tannia), '' Typhonium trilobatum'' and ''
Monstera deliciosa ''Monstera deliciosa'', the Swiss cheese plant or split-leaf philodendron is a species of flowering plant native to tropical forests of southern Mexico, south to Panama. It has been introduced to many tropical areas, and has become a mildly invas ...
'' (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 in
subsistence agriculture Subsistence agriculture occurs when farmers grow food crops to meet the needs of themselves and their families on smallholdings. Subsistence agriculturalists target farm output for survival and for mostly local requirements, with little or no su ...
and in local markets. The main food product is the
corm A corm, bulbo-tuber, or bulbotuber is a short, vertical, swollen underground plant stem that serves as a storage organ that some plants use to survive winter or other adverse conditions such as summer drought and heat (perennation). The word ' ...
, which is high in
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
; leaves and flowers also find culinary use.


See also

*
List of foliage plant diseases (Araceae) This is a list of diseases of foliage plants belonging to the Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, ...


References


Further reading

* Bown, Deni (2000). Aroids: Plants of the Arum Family LLUSTRATED Timber Press. *
online
* *


External links


International Aroid Society

The Uberlist of Araceae (pdf)




i


Plants of the World Online
b
Royal Botanic Gardens, Kew
{{Authority control Alismatales families Extant Maastrichtian first appearances