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Ginkgoales are a
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, Î³Ï…Î¼Î½ÏŒÏ ...
order containing only one
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
species: ''
Ginkgo biloba ''Ginkgo biloba'', commonly known as ginkgo or gingko ( ), also known as the maidenhair tree, is a species of tree native to China. It is the last living species in the order Ginkgoales, which first appeared over 290 million years ago. Fossils ...
'', the ginkgo tree. It is
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
, (the only taxon) within the class Ginkgoopsida, which itself is monotypic within the division Ginkgophyta . The order includes five
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideall ...
, of which only
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 for this reason sometimes regarded as a living fossil A living fossil is an extant tax ...
remains extant.


History

Ginkgophyta and
Cycadophyta Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
have a very 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 ...
. The earliest representative of the group in the fossil record is probably '' Trichopitys'' from the
Asselian In the geologic timescale, the Asselian is the earliest geochronologic age or lowermost chronostratigraphic stage of the Permian. It is a subdivision of the Cisuralian Epoch or Series. The Asselian lasted between and million years ago (Ma) ...
(299-293 million years ago) of France. The earliest representatives of ''
Ginkgo ''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, first appeared in the Permian, 270 million years ago, and is now the only living genus within ...
'', represented by reproductive organs similar to the living species, first appear in the Middle Jurassic, alongside other, related forms such as ''
Yimaia ''Yimaia'' is a extinct genus of Ginkgoalean tree, and the only member of the family Yimaiaceae. In botanical form classification, its a form taxon for ginkgoalean ovulate organs. ''Yimaia'' species are distinguished from other Ginkgoales by ...
'' and ''
Karkenia Karkeniaceae is an extinct family in the order Ginkgoales. It contains the single genus ''Karkenia''. It is distinguished by "Ovulate organs consisting of a peduncle and helically arranged, up to about 100 small, orthotropous but incurved ovule ...
'', which have differently arranged reproductive structures and seeds associated with ''Ginkgo''-like leaves. The diversity of Ginkgoales declined during the Late Cretaceous and Cenozoic, coincident with the rise of flowering plants, with all Ginkgophytes aside from ''Ginkgo'' being extinct by the end of the Cretaceous. The only remaining Ginkgophyte was ''Ginkgo adiantoides'' – a polymorphic species. Modern ''Ginkgo'' trees are native to China.


Reproduction

''Ginkgo'' trees produce ovulate and pollen-bearing structures. These structures 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 ...
, in that male and female structures come from different ''Ginkgo'' plants. The pollen organs are very similar to angiospermous
catkins A catkin or ament is a slim, cylindrical flower cluster (a spike), with inconspicuous or no petals, usually wind-pollinated (anemophilous) but sometimes insect-pollinated (as in ''Salix''). They contain many, usually unisexual flowers, arranged cl ...
. They come from the
axils A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
of the bud scales, and the leaves from the ''Ginkgo'' tree spur shoots. Pollen is contained in sacs of two to four at the tips of sporophylls on the strobiloid. Ovules of ''Ginkgo'' trees come from stalks from leaf axils on the short shoots, each containing two ovules. The ovule is fertilized by the flagellated male gametes, which can move about freely. This fertilization process begins on the tree itself in the spring. The swollen fruit-like
ovules In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...
, about 2–3 cm in diameter, fall from the tree in the fall, and fertilization continues into the winter/spring. This ovule contains a single large seed, similar to that of a
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
.


Morphology


Ginkgophyte wood

Fossils that appear ''Ginkgo''-like are filed under a
morphogenus Form classification is the classification of organisms based on their morphology (biology), morphology, which does not necessarily reflect their biological relationships. Form classification, generally restricted to palaeontology, reflects uncer ...
called ''Ginkgoxylon, Ginkgomyeloxylon,'' or ''Protoginkgoxylon''. Fossilized ginkgophyte wood is not commonly found in the record, possibly because it degrades easily, and possibly because it is difficult to tell apart from the much more pervasive
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
samples. Like conifer wood, it has secondary thin-walled xylem and a primary vascular system composed of eustele and bifacial vascular
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from w ...
. The tracheids in the secondary
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
rays have pitting that occurs only on the walls and is circularly bordered.


Ginkgophyte foliage

''Ginkgophyte''
foliage A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
has stayed largely consistent since the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
. Its historically wide territory makes it an important leaf morphology, and its unique
stomata In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
and isotopic profile give it a key role in recreations of the Mesozoic and
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 ...
. Leaf fossils that resemble the Ginkgophytes are known as ''
Ginkgoites ''Ginkgoites'' is a genus that refers to extinct plants belonging to Ginkgoaceae. Fossils of these plants have been found around the globe during the Triassic, Jurassic, Cretaceous, with fossils also known from the Paleogene. The name was created ...
''. There are similar, now extinct, morphogens, such as ''Sphenobaiera'', which describes fan-shaped, deeply divided leaves without clear petioles. The distinctive shape of the modern ''Ginkgo biloba'' gives the impression of a very narrow leaf morphology, but the group is varied and diverse. The genus ''Ginkgo'' by itself contains a range of morphologies. ''Ginkgo digitata'', from the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
, has long, wedge-shaped
lamina Lamina may refer to: Science and technology * Planar lamina, a two-dimensional planar closed surface with mass and density, in mathematics * Laminar flow, (or streamline flow) occurs when a fluid flows in parallel layers, with no disruption betwee ...
e with the intercostal regions covered in stomata and resin bodies, while ''G. pluripartita'' has at most 2 cm-long leaves and is intercostally hypostomatic.


Fossil gallery

File:Ginkgo biloba leaf 01.jpg, A 6.7 cm tall ''Ginkgo biloba'' leaf, with insect herbivory.
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington state. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Wash ...
, Republic, Ferry County, Washington, USA,
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
,
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
, 49 million years old File:Ginkgo biloba 01 SR 87-36-02 A.jpg, A 70 mm-wide ''Ginkgo biloba'' leaf. Klondike Mountain Formation, Republic, Ferry County, Washington, USA, Eocene, Ypresian, 49 million years old File:Ginkgo biloba MacAbee BC.jpg, ''Ginkgo biloba''
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
fossil leaf from the Tranquille Shale of MacAbee, British Columbia, Canada File:Ginkgo huttoni.jpg, Fossil of ''Ginkgo huttoni''. Photo taken at Naturalis Museum in
Leiden Leiden (; in English and archaic Dutch also Leyden) is a city and municipality in the province of South Holland, Netherlands. The municipality of Leiden has a population of 119,713, but the city forms one densely connected agglomeration wit ...
, The Netherlands. File:Gingkoites huttoni 1.jpg, Fossil of ''
Ginkgoites ''Ginkgoites'' is a genus that refers to extinct plants belonging to Ginkgoaceae. Fossils of these plants have been found around the globe during the Triassic, Jurassic, Cretaceous, with fossils also known from the Paleogene. The name was created ...
huttoni''


References

{{Taxonbar, from=Q835097 . Plant orders Jurassic plants Extant Permian first appearances