Platanus Rzedowskii
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''Platanus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
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 The Northern Hemisphere is the half of Earth that is north of the Equator. For other planets in the Solar System, north is defined as being in the same celestial hemisphere relative to the invariable plane of the solar system as Earth's Nort ...
. They are the sole living members of the family Platanaceae. 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, aft ...
, and most are found in riparian 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 Drought tolerance is the ability to which a plant maintains its biomass production during arid or drought conditions. Some plants are naturally adapted to dry conditions'','' surviving with protection mechanisms such as desiccation tolerance, detox ...
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 se ...
(''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''), 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 peri ...
, which referred to '' Platanus orientalis''.


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 b ...
s may be fused at the base, and the petals are 3–7 and are spatulate. 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 is conne ...
). 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 o ...
) 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 filame ...
s; the female has a superior ovary with 3–7 carpels. 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 ope ...
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. 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 Johnn ...
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 in ...
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 eukaryote ...
sequences suggests that the ''P. racemosa'' species complex in Western North America (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, 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 co ...
, ''P. oaxacana; P. occidentalis''
s.l. SL may refer to: Arts and entertainment * SL (rapper), a rapper from London * ''Second Life'', a multi-user 3D virtual world * Sensei's Library, an Internet site dedicated to the game of Go * Subdominant leittonwechselklänge * Leica SL, a mirror ...
with two ubpecies: ''P. occidentalis, P. palmeri''; '' P. rzedowskii''). The two groups form genetically and morphologically distinct
evolutionary lineages An evolutionary lineage is a temporal series of populations, organisms, cells, or genes connected by a continuous line of descent from ancestor to descendant.The Oxford English Dictionary defines biological lineage as "a sequence of species each of ...
( 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 evolutionary 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 together ...
hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
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. Intr ...
): * ''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. I ...
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 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 language, Otomi: ''Hyodi Ndämxei''), is one of the Political divisions of Mexico, 32 federal entities of Mexico. I ...
and northern Hidalgo; ''P. mexicana'' var. ''interior'' according Nixon & Poole) mark the hybrid zone 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 several ...
of ''P. mexicana'' is from this zone and shows the characteristical intermediate morphology, ''P. mexicana'' s.str. would represent a
nothospecies #REDIRECT Hybrid name In botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name, but there is no requirement that a hybrid name should be created for plants that are believed to be of hybrid origin. ...
: ''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 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 ( cladogenesis), 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 History ...
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 normal ...
: 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 (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
. 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 dif ...
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 co ...
(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 the e ...
''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 Republic ...
in Nagorno-Karabakh. File:Fossil Platanus leaf.jpg, Fossil ''Platanus'' leaf from the
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
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 (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...
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 th ...
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'', 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 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. The ...
e of some
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
species including ''
Phyllonorycter platani ''Phyllonorycter platani'' is a moth of the family Gracillariidae. The moth is found in Europe, the Near East and the eastern Palearctic realm, as well as California in the United States. The wingspan is 8–10 mm. It flies in two gener ...
'' and Setaceous Hebrew Character. 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 se ...
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 bi ...
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'' in the Old World. The tree is an important part of the literary scenery of Plato's dialogue
Phaedrus Phaedrus may refer to: People * Phaedrus (Athenian) (c. 444 BC – 393 BC), an Athenian aristocrat depicted in Plato's dialogues * Phaedrus (fabulist) (c. 15 BC – c. AD 50), a Roman fabulist * Phaedrus the Epicurean (138 BC – c. 70 BC), an Epic ...
. 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 ''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, du ...
. 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 phil ...
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 training i ...
's opera Serse features a famous aria, " Ombra mai fu," 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 as an embodiment of sorrowful sentiments due to its autumnal shedding of leaves. The legendary Dry tree first recorded by
Marco Polo Marco Polo (, , ; 8 January 1324) was a Venetian merchant, explorer and writer who travelled through Asia along the Silk Road between 1271 and 1295. His travels are recorded in ''The Travels of Marco Polo'' (also known as ''Book of the Marv ...
was possibly a platanus. According to the legend, it marked the site of the battle between
Alexander the Great Alexander III of Macedon ( grc, wikt:Ἀλέξανδρος, Ἀλέξανδρος, Alexandros; 20/21 July 356 BC – 10/11 June 323 BC), commonly known as Alexander the Great, was a king of the Ancient Greece, ancient Greek kingdom of Maced ...
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