Platanus
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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 nom ...
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 subtropic ...
. 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, ...
, 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 p ...
habitats in the wild, though proving drought-tolerant in cultivation. The hybrid London plane (''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 coine ...
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). The number of heads in one cluster (
inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
) is indicative of the species (see table below). The male flower has 3–8 stamens; 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 achenes 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 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 and nuclear gene 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 W ...
(including '' P. racemosa, P. gentryi , P. wrigthii'') is more closely related to the
Western Eurasian Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago an ...
''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, ''P. oaxacana; P. occidentalis'' s.l. with two ubpecies: ''P. occidentalis, P. palmeri''; '' P. rzedowskii''). The two groups form
genetically Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar working i ...
and morphologically distinct evolutionary lineages (
sister clades In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
), 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 hybridization and introgression): * ''Platanus palmeri'' (= ''P. occidentalis'' var. ''palmeri'') – forming the southwesternmost populations of ''P. occidentalis'' s.l. – carries nuclear intron 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 spacers 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 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 contact ...
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 sever ...
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 ''P. oaxacana'' population from northcentral
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the political divisions of Mexico, Federative Entities of Mexico. It is ...
. 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 simult ...
(
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 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 hybrids 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 (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 Europe, Western and Southern Europe and Anatolia, on the south by North Africa ...
''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 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 m ...
. 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 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 A wilt disease is any number of diseases that affect the vascular system of plants. Attacks by fungi, bacteria, and nematodes can cause rapid killing of plants, large tree branches or even entire trees. Wilt diseases in woody plants tend to fal ...
, 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, a ...
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. ...
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'', 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 is particularly popular for this purpose. The American plane is cultivated sometimes for timber and investigations have been made into its use as a biomass crop. The
oriental plane ''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 ...
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 institution ...
, the tree also played an important role in the scenery of Cicero's De Oratore. 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 period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ph ...
and Plato’s famed philosophical schools were held. Handel's opera
Serse ''Serse'' (; English title: ''Xerxes''; HWV 40) is an opera seria in three acts by George Frideric Handel. It was first performed in London on 15 April 1738. The Italian libretto was adapted by an unknown hand from that by Silvio Stampiglia (1 ...
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 dy ...
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. 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