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''Ginkgo'' is a genus of non-flowering seed plants. The scientific name is also used as the English name. The order to which it belongs,
Ginkgoales Ginkgoales are a gymnosperm order containing only one extant species: ''Ginkgo biloba'', the ginkgo tree. It is monotypic, (the only taxon) within the class Ginkgoopsida, which itself is monotypic within the division Ginkgophyta . The order incl ...
, first appeared in the Permian, 270 million years ago, and is now the only living genus within the order. The rate of evolution within the genus has been slow, and almost all its species had become extinct by the end of the Pliocene. The sole surviving species, '' Ginkgo biloba'' is only found in the wild in China, but is cultivated around the world. The relationships between ginkgos and other groups of plants are not fully resolved.


Prehistory

The ginkgo (''Ginkgo biloba'') is a living fossil, with fossils similar to the modern plant dating back to the Permian, 270 million years ago. The closest living relatives of the clade are the cycads, which share with the extant ''G. biloba'' the characteristic of motile sperm. The ginkgo and cycad lineages are thought to have an extremely ancient divergence dating to the early
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
. Fossils attributable to the genus ''Ginkgo'' with reproductive organs similar to the modern species first appeared in the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
, and the genus diversified and spread throughout
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
during the Jurassic and
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 ...
. It declined in diversity as the Cretaceous progressed with the extinction of species such as ''
Ginkgo huolinhensis ''Ginkgo huolinhensis'' is an extinct species of seed plant in the family Ginkgoaceae The Ginkgoaceae is a family of gymnosperms which appeared during the Mesozoic Era, of which the only extant representative is ''Ginkgo biloba'', which is fo ...
'', and by the
Palaeocene 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 ''palaiĂ ...
, only a few ''Ginkgo'' species, '' Ginkgo cranei'' and '' Ginkgo adiantoides'', remained in the Northern Hemisphere, while a markedly different (and poorly documented) form persisted in the Southern Hemisphere. At the end of the Pliocene, ''Ginkgo'' fossils disappeared from the fossil record everywhere except in a small area of central China, where the modern species survived. It is doubtful whether the Northern Hemisphere fossil species of ''Ginkgo'' can be reliably distinguished. Given the slow pace of evolution and morphological similarity between members of the genus, there may have been only one or two species existing in the Northern Hemisphere through the entirety of the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
: present-day ''G. biloba'' (including ''G. adiantoides'') and '' G. gardneri'' from the Palaeocene of Scotland. At least morphologically, ''G. gardneri'' and the Southern Hemisphere species are the only known post-Jurassic taxa that can be unequivocally recognised. The remainder may have been ecotypes or
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
. The implications would be that ''G. biloba'' had occurred over an extremely wide range, had remarkable genetic flexibility and, though evolving genetically, never showed much
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. While it may seem improbable that a species may exist as a contiguous entity for many millions of years, many of the ginkgo's life-history parameters fit. These are: extreme longevity; slow reproduction rate; (in Cenozoic and later times) a wide, apparently contiguous, but steadily contracting distribution coupled with, as far as can be demonstrated from the fossil record, extreme ecological conservatism (restriction to disturbed streamside environments). Royer et al. (2003), p. 91. Modern-day ''G. biloba'' grows best in well-watered and drained environments, and the extremely similar fossil ''Ginkgo'' favoured similar environments. The sediment records at the majority of fossil ''Ginkgo'' localities indicate it grew primarily in disturbed environments along streams and
levee A levee (), dike (American English), dyke (English in the Commonwealth of Nations, Commonwealth English), embankment, floodbank, or stop bank is a structure that is usually soil, earthen and that often runs parallel (geometry), parallel to ...
s. ''Ginkgo'' therefore presents an "ecological paradox" because, while it possesses some favourable traits for living in disturbed environments (clonal reproduction), many of its other life-history traits (slow growth, large seed size, late reproductive maturity) are the opposite of those exhibited by modern plants that thrive in disturbed settings. Given the slow rate of evolution of the genus, it is possible that ''Ginkgo'' represents a pre-
angiosperm 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 ...
strategy for survival in disturbed streamside environments. ''Ginkgo'' evolved in an era before flowering plants, when ferns, cycads, and cycadeoids dominated disturbed streamside environments, forming a low, open, shrubby canopy. The large seeds of ''Ginkgo'' and its habit of "bolting"—growing to a height of before elongating its side branches—may be adaptations to such an environment. Diversity in the genus ''Ginkgo'' dropped through the Cretaceous (along with that of ferns, cycads, and cycadeoids) at the same time the flowering plants were on the rise, which supports the notion that flowering plants, with their better adaptations to disturbance, displaced ''Ginkgo'' and its associates over time. '' Trichopitys heteromorpha'' from the earliest Permian of France, is one of the earliest fossils ascribed to the Ginkgophyta. It had multiple-forked non-laminar leaves with cylindrical, thread-like ultimate divisions. '' Sphenobaiera'' (early Permian-Cretaceous) had wedge-shaped leaves divided into narrow dichotomously-veined lobes, lacking distinct petioles (leaf stalks). ''Baiera'' (Triassic-Jurassic) had similar multiple-lobed leaves but with petioles.


Phylogeny

, molecular phylogenetic studies have produced at least six different placements of ''Ginkgo'' relative to cycads,
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
,
gnetophytes Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three rel ...
and angiosperms. The two most common are that ''Ginkgo'' is a sister to a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
composed of conifers and gnetophytes and that ''Ginkgo'' and cycads form a clade within the gymnosperms. A 2013 study examined the reasons for the discrepant results and concluded that the best support was for the monophyly of ''Ginkgo'' and cycads.


Etymology

The genus name is regarded as a misspelling of the
Japanese pronunciation The phonology of Japanese features about 15 consonant phonemes, the cross-linguistically typical five-vowel system of , and a relatively simple phonotactic distribution of phonemes allowing few consonant clusters. It is traditionally described ...
''gin kyo'' for the kanji 銀杏 meaning "silver apricot", which is found in Chinese herbology literature such as (Daily Use Materia Medica) (1329) and '' Compendium of Materia Medica'' published in 1578.T. Hori, A historical survey of Ginkgo biloba based on Japanese and Chinese classical literatures, Plant Morphology, 2001, 31, 31-40 Despite its spelling, which is due to a complicated etymology including a transcription error, "ginkgo" is usually pronounced , which has given rise to the common alternative spelling "gingko". The spelling pronunciation is also documented in some dictionaries. Engelbert Kaempfer first introduced the spelling ''ginkgo'' in his book ''Amoenitatum Exoticarum''. It is considered that he may have misspelled "Ginkyo" as "Ginkgo". This misspelling was included by Carl Linnaeus in his book ''Mantissa plantarum II'' and has become the name of the tree's genus.


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{{Authority control Extant Permian first appearances Gymnosperm genera Permian plants Prehistoric gymnosperm genera Taxa named by Carl Linnaeus