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''Platanus'' is a
genus 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 n ...
consisting of a small number of
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
species native to the Northern Hemisphere. They are the sole living members of the family
Platanaceae Platanaceae, the "plane-tree family", is a family of flowering plants in the order Proteales. The family consists of only a single extant genus ''Platanus'', with eight known species. The plants are tall trees, native to temperate and subtrop ...
. All mature members of ''Platanus'' are tall, reaching in height. All except for '' P. kerrii'' are
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, a ...
, and most are found in
riparian A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks a ...
or other
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
habitats In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
in the wild, though proving drought-tolerant in cultivation. The hybrid
London plane London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a major s ...
(''Platanus ''×'' acerifolia'') has proved particularly tolerant of urban conditions, and has been widely planted in London and elsewhere in the United Kingdom. They are often known in English as ''planes'' or ''plane trees''. A formerly used name that is now rare is ''plantain tree'' (not to be confused with other, unrelated, species with the name). Some North American species are called ''sycamores'' (especially ''
Platanus occidentalis ''Platanus occidentalis'', also known as American sycamore, American planetree, western plane, occidental plane, buttonwood, and water beech, is a species of ''Platanus'' native to the eastern and central United States, the mountains of northeas ...
''), although the term is also used for several unrelated species of trees. The genus name ''Platanus'' comes from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
, which referred to ''
Platanus orientalis ''Platanus orientalis'', the Old World sycamore or Oriental plane, is a large, deciduous tree of the Platanaceae family, growing to or more, and known for its longevity and spreading crown. In autumn its deep green leaves may change to blood red ...
''.


Botany

The flowers are reduced and are borne in balls (globose heads); 3–7 hairy
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined ...
s may be fused at the base, and the petals are 3–7 and are
spatulate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular ...
. Male and female flowers are separate, but borne on the same plant (
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 i ...
). The number of heads in one cluster (
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 ...
) is indicative of the species (see table below). The male flower has 3–8
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
s; the female has a superior ovary with 3–7
carpel Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) '' pistils' ...
s. Plane trees are wind-pollinated. Male flower-heads fall off after shedding their pollen. After being pollinated, the female flowers become
achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not op ...
s that form an aggregate ball. The fruit is a multiple of achenes (plant systematics, Simpson M. G., 2006). Typically, the core of the ball is 1 cm in diameter and is covered with a net of mesh 1 mm, which can be peeled off. The ball is 2.5–4 cm in diameter and contains several hundred achenes, each of which has a single seed and is conical, with the point attached downward to the net at the surface of the ball. There is also a tuft of many thin stiff yellow-green bristle fibers attached to the base of each achene. These bristles help in wind dispersion of the fruits as in the
dandelion ''Taraxacum'' () is a large genus of flowering plants in the family Asteraceae, which consists of species commonly known as dandelions. The scientific and hobby study of the genus is known as taraxacology. The genus is native to Eurasia and Nor ...
. The leaves are
simple Simple or SIMPLE may refer to: * Simplicity, the state or quality of being simple Arts and entertainment * ''Simple'' (album), by Andy Yorke, 2008, and its title track * "Simple" (Florida Georgia Line song), 2018 * "Simple", a song by John ...
and alternate. In the subgenus ''Platanus'' they have a palmate outline. The base of the leaf stalk (petiole) is enlarged and completely wraps around the young stem bud in its axil. The axillary bud is exposed only after the leaf falls off. The mature bark peels off or exfoliates easily in irregularly shaped patches, producing a mottled, scaly appearance. On old trunks, bark may not flake off, but thickens and cracks instead.


Phylogeny

There are two subgenera, subgenus ''Castaneophyllum'' containing the anomalous '' P. kerrii'', and subgenus ''Platanus'', with all the others; recent studies in Mexico have increased the number of accepted species in this subgenus. Within subgenus ''Platanus'', evidence from both
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it ...
and
nuclear gene A nuclear gene is a gene whose physical DNA nucleotide sequence is located in the cell nucleus of a eukaryote. The term is used to distinguish nuclear genes from genes found in mitochondria or chloroplasts. The vast majority of genes in eukaryot ...
sequences suggests that the ''P. racemosa''
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
in
Western North America Western North America is the western edge of the North American continent that borders the Pacific Ocean. It consists of Alaska at the farthest north, down through the western Canadian province of British Columbia, the western U.S. states of ...
(including '' P. racemosa, P. gentryi , P. wrigthii'') is more closely related to the Western Eurasian ''P. orientalis'' than it is to the other North American species ('' P. mexicana''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
, including up to four species: ''P. chiapaensis, P. lindeniana, P. mexicana''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
, ''P. oaxacana; P. occidentalis'' s.l. with two ubpecies: ''P. occidentalis, P. palmeri''; '' P. rzedowskii''). The two groups form genetically and morphologically distinct evolutionary lineages ( sister clades), informally called the “ANA clade” (Atlantic North American lineage) and “PNA-E clade” (Pacific North American-European lineage). Both lineages have been affected by
reticulate evolution Reticulate evolution, or network evolution is the origination of a lineage through the partial merging of two ancestor lineages, leading to relationships better described by a phylogenetic network than a bifurcating tree. Reticulate patterns ca ...
ary processes in the past (ancient or
recent The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
hybridization and
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intro ...
): * ''Platanus palmeri'' (= ''P. occidentalis''
var. In botanical nomenclature, variety (abbreviated var.; in la, varietas) is a taxonomic rank below that of species and subspecies, but above that of form. As such, it gets a three-part infraspecific name. It is sometimes recommended that the ...
''palmeri'') – forming the southwesternmost populations of ''P. occidentalis'' s.l. – carries nuclear
intron An intron is any nucleotide sequence within a gene that is not expressed or operative in the final RNA product. The word ''intron'' is derived from the term ''intragenic region'', i.e. a region inside a gene."The notion of the cistron .e., gene ...
sequences (second intron of the Leafy gene) of PNA-E origin. It lacks the plastid haplotype specific for the northeastern populations (''P. occidentalis'' s.str.) * The
internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...
s of the nuclear-encoded rRNA genes of ''P. occidentalis'' s.l. and ''P. rzedowskii'' include ANA-specific variants with functional 5.8S rDNA as well as PNA-E-specific variants showing signs of pseudogeny. The latter are shared with ''P. gentryi,'' the PNA-E species closest to the ANA clade area and the northern/ interior populations of ''P. mexicana'' s.l. This indicates that already the
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
of ''P. rzedowskii'' and ''P. occidentalis'' s.l. had been in contact with a member of the PNA-E clade. * Likewise, '' P. rzedowskii'' from Nueva León is a genetic mosaic, and may have originated from earlier hybridization within the ANA clade, between southernmost ''P. occidentalis'' s.l. (''P. palmeri'') and ''P. mexicana'' s.l., or their ancestors. Today the ranges of ''P. occidentalis'' s.l. and ''P. mexicana'' s.l. are mutually exclusive. ''Platanus rzedowskii'' is geographically and morphologically intermediate between ''P. occidentalis'' s.l. and ''P. mexicana'' s.l. * Morphological reinvestigation including the originally collected material revealed that the interior populations of ''P. mexicana'' (northern
Querétaro Querétaro (), officially the Free and Sovereign State of Querétaro ( es, Estado Libre y Soberano de Querétaro, links=no; Otomi: ''Hyodi Ndämxei''), is one of the 32 federal entities of Mexico. It is divided into 18 municipalities. Its capi ...
and northern
Hidalgo Hidalgo may refer to: People * Hidalgo (nobility), members of the Spanish nobility * Hidalgo (surname) Places Mexico * Hidalgo (state), in central Mexico * Hidalgo, Coahuila, a town in the north Mexican state of Coahuila * Hidalgo, Nuevo Le� ...
; ''P. mexicana'' var. ''interior'' according Nixon & Poole) mark the
hybrid zone A hybrid zone exists where the ranges of two interbreeding species or diverged intraspecific lineages meet and cross-fertilize. Hybrid zones can form ''in situ'' due to the evolution of a new lineage but generally they result from secondary cont ...
between ''P. rzedowskii'' and ''P. mexicana'' s.l. and the (former) contact zone to the species of the PNA-E clade (''P. gentryi'', ''P. wrightii''). Since the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
of ''P. mexicana'' is from this zone and shows the characteristical intermediate morphology, ''P. mexicana'' s.str. would represent a nothospecies: ''P. × mexicana''. The remaining populations of ''P. mexicana'' s.l., ''P. lindeniana,'' show no sign of introgression from either ''P. rzedowskii, P. occidentalis-palmeri'' or the Western North American species (''P. racemosa'' species aggregate), with the exception of one
heterozygotic Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
''P. oaxacana'' population from northcentral
Oaxaca ) , population_note = , population_rank = 10th , timezone1 = CST , utc_offset1 = −6 , timezone1_DST = CDT , utc_offset1_DST = −5 , postal_code_type = Postal ...
. The genus ''Platanus'' exemplarily illustrates the concept of a
Coral of Life The coral of life is a metaphor or a mathematical model useful to illustrate evolution of life or phylogeny at various levels of resolution, including individual organisms, populations, species and large taxonomic groups. Its use in biology resolves ...
, a species network. Its modern-day species are not only the product of evolutionary
dichotomies A dichotomy is a partition of a whole (or a set) into two parts (subsets). In other words, this couple of parts must be * jointly exhaustive: everything must belong to one part or the other, and * mutually exclusive: nothing can belong simul ...
(
cladogenesis Cladogenesis is an evolutionary splitting of a parent species into two distinct species, forming a clade. This event usually occurs when a few organisms end up in new, often distant areas or when environmental changes cause several extinctions ...
), the splitting of an ancestral lineage into two (
Tree of Life The tree of life is a fundamental archetype in many of the world's mythological, religious, and philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The Assyrian Sacred Tree: A Histo ...
metaphor) but also evolutionary
anastomoses An anastomosis (, plural anastomoses) is a connection or opening between two things (especially cavities or passages) that are normally diverging or branching, such as between blood vessels, leaf veins, or streams. Such a connection may be norm ...
: hybridization and introgression. The fossil record of leaves and fruit identifiable to ''Platanus'' begins in the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
. Despite the geographic separation between North America and Old World, species from these continents will cross readily resulting in fertile
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
s such as the London plane, which is an anthropogenic hybrid (
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
) between the North American ''P. occidentalis''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
(ANA clade) and the
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 th ...
''P. orientalis'' (PNA-E clade). Widely used as a park tree across Europe, it frequently backcrosses with its native parent. File:Skhtorashen176.jpg, The -year-old ''Platanus orientalis'' tree
Tnjri Tnjri ( hy, Տնջրի , from տնջրի ''tnǰri'', which in the Karabakh dialect means ‘plane tree’) is a -year-old giant Oriental plane tree situated nearby the village of Skhtorashen, ''de facto'' in the Martuni Province of the Republ ...
in
Nagorno-Karabakh Nagorno-Karabakh ( ) is a landlocked region in the South Caucasus, within the mountainous range of Karabakh, lying between Lower Karabakh and Syunik, and covering the southeastern range of the Lesser Caucasus mountains. The region is mos ...
. File:Fossil Platanus leaf.jpg, Fossil ''Platanus'' leaf from the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
of
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
, Canada. File:Fossil Platanus Fruit 0444.jpg, Fossil ''Platanus'' fruit from the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
of
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest Ter ...
, Canada.


Species

The following are recognized species of plane trees:


Diseases

Planes are susceptible to plane anthracnose (''Apiognomonia veneta''), a
fungal A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
disease that can defoliate the trees in some years. The most severe infections are associated with cold, wet spring weather. ''P. occidentalis'' and the other American species are the most susceptible, with ''P. orientalis'' the most resistant. The hybrid London plane is intermediate in resistance. ''
Ceratocystis platani ''Ceratocystis platani'' is a fungus that causes a disease on plane trees in the genus ''Platanus''. Description ''Ceratocystis platani'' is believed to be native to southeastern USA. The fungus was previously considered to be part of the '' Ce ...
'', a wilt disease, has become a significant problem in recent years in much of Europe. The North American species are mostly resistant to the disease, with which they probably coevolved, while the old world species are highly sensitive. Other diseases such as
powdery mildew Powdery mildew is a fungal disease that affects a wide range of plants. Powdery mildew diseases are caused by many different species of ascomycete fungi in the order Erysiphales. Powdery mildew is one of the easier plant diseases to identify, as ...
occur frequently, but are of lesser importance. ''Platanus'' species are used as food plants by the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...
e of some Lepidoptera species including '' Phyllonorycter platani'' and
Setaceous Hebrew Character The setaceous Hebrew character (''Xestia c-nigrum'') is a moth of the family Noctuidae. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. It is found in the Palearctic realm. It is a common specie ...
. In the 21st century a disease, commonly known as Massaria disease, has attacked plane trees across Europe. It is caused by the fungus ''
Splanchnonema platani ''Splanchnonema platani'' is a fungus in the genus ''Splanchnonema''. It was formerly known under the name ''Massaria platani''. The anamorph of the fungus is known as ''Macrodiplodiopsis desmazieresii''. The fungus has caused serious damage to ...
'', and causes large lesions on the upper sides of branches.


Uses

The principal use of these trees is as ornamental trees, especially in urban areas and by roadsides. The
London plane London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a major s ...
is particularly popular for this purpose. The
American plane ''Platanus occidentalis'', also known as American sycamore, American planetree, western plane, occidental plane, buttonwood, and water beech, is a species of ''Platanus'' native to the eastern and central United States, the mountains of northeas ...
is cultivated sometimes for timber and investigations have been made into its use as a
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms biom ...
crop. The oriental plane is widely used as an ornamental and also has a number of minor medicinal uses.


Cultural history

Most significant aspects of cultural history apply to ''
Platanus orientalis ''Platanus orientalis'', the Old World sycamore or Oriental plane, is a large, deciduous tree of the Platanaceae family, growing to or more, and known for its longevity and spreading crown. In autumn its deep green leaves may change to blood red ...
'' in the Old World. The tree is an important part of the literary scenery of Plato's dialogue Phaedrus. Because of
Plato Plato ( ; grc-gre, Πλάτων ; 428/427 or 424/423 – 348/347 BC) was a Greek philosopher born in Athens during the Classical period in Ancient Greece. He founded the Platonist school of thought and the Academy, the first institutio ...
, the tree also played an important role in the scenery of Cicero's
De Oratore ''De Oratore'' (''On the Orator''; not to be confused with '' Orator'') is a dialogue written by Cicero in 55 BC. It is set in 91 BC, when Lucius Licinius Crassus dies, just before the Social War and the civil war between Marius and Sulla, d ...
. The trees also provided the shade under which
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical Greece, Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatet ...
and Plato’s famed philosophical schools were held.
Handel George Frideric (or Frederick) Handel (; baptised , ; 23 February 1685 – 14 April 1759) was a German-British Baroque composer well known for his operas, oratorios, anthems, concerti grossi, and organ concertos. Handel received his train ...
's opera Serse features a famous aria, "
Ombra mai fu "" ("Never was a shade…"), also known as "Largo from ''Xerxes''", is the opening aria from the 1738 opera '' Serse'' by George Frideric Handel. Context The opera was a commercial failure, lasting only five performances in London after its pre ...
," which the title character sings in praise of his favorite plane tree. The plane tree has been a frequent motif featured in
Classical Chinese poetry Classical Chinese poetry is traditional Chinese poetry written in Classical Chinese and typified by certain traditional forms, or modes; traditional genres; and connections with particular historical periods, such as the poetry of the Tang dyn ...
as an embodiment of sorrowful sentiments due to its autumnal shedding of leaves. The legendary Dry tree first recorded by Marco Polo was possibly a platanus. According to the legend, it marked the site of the battle between
Alexander the Great Alexander III of Macedon ( grc, Ἀλέξανδρος, Alexandros; 20/21 July 356 BC – 10/11 June 323 BC), commonly known as Alexander the Great, was a king of the ancient Greek kingdom of Macedon. He succeeded his father Philip II to ...
and
Darius III Darius III ( peo, 𐎭𐎠𐎼𐎹𐎺𐎢𐏁 ; grc-gre, Δαρεῖος ; c. 380 – 330 BC) was the last Achaemenid King of Kings of Persia, reigning from 336 BC to his death in 330 BC. Contrary to his predecessor Artaxerxes IV Arses, Dar ...
. The German camouflage pattern ''Platanenmuster'' ("plane tree pattern"), designed in 1937–1942 by Johann Georg Otto Schick, was the first dotted camouflage pattern.


Footnotes


References

;Books * * ;Journals * * ;Web * *


External links


Botany of plane trees


concentrating on the North American species *
A developmental and evolutionary analysis of embryology in Platanus (Platanaceae), a basal eudicot
', abstract of article by Sandra K. Floyd et al. in ''American Journal of Botany'', 1999;86:1523–1537.
Photos, measurements, and location details of large plane trees worldwide
{{Authority control Platanaceae Eudicot genera