Ginkgoales

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Ginkgoales
Temporal range: Permianpresent
Fossil Ginkgoales.jpg
Leaves and seeds of Ginkgo yimaensis (left) Yimaia recurva (top right) and Karkenia henanensis (bottom right)
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Gymnospermae
Division: Ginkgophyta
Class: Ginkgoopsida
Order: Ginkgoales
Gorozhankin, 1904
Families

Ginkgoales are a gymnosperm order containing only one extant species: Ginkgo biloba , the ginkgo tree. [1] The order has a long fossil record extending back to the Early Permian around 300 million years ago from fossils found worldwide. The order was a common component of Triassic flora before the super dominance of conifers.

Contents

Evolution

Ginkgophyta and Cycadophyta have a very ancient divergence dating to the early Carboniferous. [2] The earliest representative of the group in the fossil record is probably Trichopitys from the Asselian (299-293 million years ago) of France. The earliest representatives of Ginkgo , represented by reproductive organs similar to the living species, first appear in the Middle Jurassic, alongside other, related forms such as Yimaia and Karkenia , which have differently arranged reproductive structures and seeds associated with Ginkgo-like leaves. [3] [4] 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. [5] [4] The only remaining Ginkgophyte was Ginkgo adiantoides – a polymorphic species. [6] Modern Ginkgo trees are native to China. [5]

Reproduction

Ginkgo trees produce ovulate and pollen-bearing structures. These structures are dioecious, in that male and female structures come from different Ginkgo plants. [5] The pollen organs are very similar to angiospermous catkins. They come from the axils 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, about 2–3 cm in diameter, fall from the tree in the fall, and fertilization continues into the winter/spring. [4] [6] This ovule contains a single large seed, similar to that of a cycad. [4]

Morphology

Ginkgophyte wood

Fossils that appear Ginkgo-like are filed under a morphogenus 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 samples. [4] Like conifer wood, it has secondary thin-walled xylem and a primary vascular system composed of eustele and bifacial vascular cambium. The tracheids in the secondary xylem rays have pitting that occurs only on the walls and is circularly bordered. [5]

This recreation displays one example of early Gingkophyte foliage. Live reconstruction of Cretophasmomima melanogramma - journal.pone.0091290.g007.png
This recreation displays one example of early Gingkophyte foliage.

Ginkgophyte foliage

Ginkgophyte foliage has stayed largely consistent since the Mesozoic. Its historically wide territory makes it an important leaf morphology, and its unique stomata and isotopic profile give it a key role in recreations of the Mesozoic and Cenozoic. Leaf fossils that resemble the Ginkgophytes are known as Ginkgoites . 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, has long, wedge-shaped laminae with the intercostal regions covered in stomata and resin bodies, while G. pluripartita has at most 2 cm-long leaves and is intercostally hypostomatic. [4]

Related Research Articles

<span class="mw-page-title-main">Gametophyte</span> Haploid stage in the life cycle of plants and algae

A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

<span class="mw-page-title-main">Vascular plant</span> Clade of plants with xylem and phloem

Vascular plants, also called tracheophytes or collectively tracheophyta, are plants that have lignified tissues for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue to conduct products of photosynthesis. The group includes most land plants other than mosses.

<i>Ginkgo biloba</i> Species of tree

Ginkgo biloba, commonly known as ginkgo or gingko, also known as the maidenhair tree, is a species of gymnosperm tree native to East Asia. It is the last living species in the order Ginkgoales, which first appeared over 290 million years ago, and fossils very similar to the living species, belonging to the genus Ginkgo, extend back to the Middle Jurassic epoch approximately 170 million years ago. The tree was cultivated early in human history and remains commonly planted, and is widely regarded as a living fossil.

<span class="mw-page-title-main">Conifer</span> Group of cone-bearing seed plants

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 extant conifers are perennial woody plants with secondary growth. The great majority are trees, though a few are shrubs. Examples include cedars, Douglas-firs, cypresses, firs, junipers, kauri, larches, pines, hemlocks, redwoods, spruces, and yews. As of 2002, Pinophyta contained seven families, 60 to 65 genera, and more than 600 living species.

<span class="mw-page-title-main">Gnetophyta</span> Division of plants containing three genera of gymnosperms

Gnetophyta is a division of plants, grouped within the gymnosperms, that consists of some 70 species across the three relict genera: Gnetum, Welwitschia, and Ephedra. The earliest unambiguous records of the group date to the Jurassic, and they achieved their highest diversity during the Early Cretaceous. The primary difference between gnetophytes and other gymnosperms is the presence of vessel elements, a system of small tubes (xylem) that transport water within the plant, similar to those found in flowering plants. Because of this, gnetophytes were once thought to be the closest gymnosperm relatives to flowering plants, but more recent molecular studies have brought this hypothesis into question, with many recent phylogenies finding them to be nested within the conifers.

<span class="mw-page-title-main">Gymnosperm</span> Clade of non-flowering, naked-seeded vascular plants

The gymnosperms are a group of seed-producing plants that include conifers, cycads, Ginkgo, and gnetophytes, forming the clade Gymnospermae. The term gymnosperm comes from the composite word in Greek: γυμνόσπερμος, and literally means 'naked seeds'. The name is based on the unenclosed condition of their seeds. The non-encased condition of their seeds contrasts with the seeds and ovules of flowering plants (angiosperms), which are enclosed within an ovary. Gymnosperm seeds develop either on the surface of scales or leaves, which are often modified to form cones, or on their own as in yew, Torreya, and Ginkgo. The life cycle of a gymnosperm involves alternation of generations, with a dominant diploid sporophyte phase, and a reduced haploid gametophyte phase, which is dependent on the sporophytic phase. The term "gymnosperm" is often used in paleobotany to refer to all non-angiosperm seed plants. In that case, to specify the modern monophyletic group of gymnosperms, the term Acrogymnospermae is sometimes used.

<i>Nageia</i> Genus of conifers in the family Podocarpaceae

Nageia is a genus of conifers belonging to the podocarp family Podocarpaceae. Nageia includes evergreen shrubs and trees, from one to 54 meters in height. A 2009 treatment of the genus recognized five species. Some authors consider Nageia formosensis to be a separate species from Nageia nagi, thus recognizing six species. The podocarp genera have been reshuffled by various botanists. Most recently, several species formerly classed as Nageia were moved to the new genus Retrophyllum, while Nageia falcata and Nageia mannii were moved to the new genus Afrocarpus.

<i>Afrocarpus</i> Genus of conifers

Afrocarpus is a genus of conifer of the family Podocarpaceae. Two to six species are recognized. They are evergreen trees native to Africa. Afrocarpus was designated a genus in 1989, when several species formerly classified in Podocarpus and Nageia were reclassified.

<span class="mw-page-title-main">Bennettitales</span> Extinct order of seed plants

Bennettitales is an extinct order of seed plants that first appeared in the Permian period and became extinct in most areas toward the end of the Cretaceous. Bennettitales were amongst the most common seed plants of the Mesozoic, and had morphologies including shrub and cycad-like forms. The foliage of bennettitaleans is superficially nearly indistinguishable from that of cycads, but they are distinguished from cycads by their more complex flower-like reproductive organs, at least some of which were likely pollinated by insects.

<i>Torreya</i> Genus of conifers

Torreya is a genus of conifers comprising six or seven species placed in the family Taxaceae, though sometimes formerly placed in Cephalotaxaceae. Four species are native to eastern Asia; the other two are native to North America. They are small to medium-sized evergreen trees reaching 5–20 m, rarely 25 m, tall. Common names include nutmeg yew.

<i>Retrophyllum</i> Genus of conifers

Retrophyllum is a genus of conifers in the family Podocarpaceae. It contains five generally recognized extant species with a disjunct distribution in the Southern Hemisphere, found in Papuasia and also in South America. Retrophyllum are evergreen trees typically occurring in tropical rainforests and cloud forests.

<i>Cephalotaxus</i> Genus of conifers

Cephalotaxus, commonly called plum yew or cowtail pine, is a genus of conifers comprising 11 species, either considered the only member of the family Cephalotaxaceae, or in the Taxaceae when that family is considered in a broad sense. The genus is endemic to eastern Asia, though fossil evidence shows it had a wider Northern Hemisphere distribution in the past. The species are evergreen shrubs and small trees reaching 1.0–10 metres (3–33 ft) tall.

<span class="mw-page-title-main">Trochodendraceae</span> Family of flowering plants

Trochodendraceae is the only family of flowering plants in the order Trochodendrales. It comprises two extant genera, each with a single species along with up to five additional extinct genera and a number of extinct species. The living species are native to south east Asia. The two living species both have secondary xylem without vessel elements, which is quite rare in angiosperms. As the vessel-free wood suggests primitiveness, these two species have attracted much taxonomic attention.

<i>Dioon</i> Genus of cycads in the family Zamiaceae

Dioon is a genus of cycads in the family Zamiaceae. It is native to Mexico and Central America. Their habitats include tropical forests, pine-oak forest, and dry hillsides, canyons and coastal dunes.

A strobilus is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to the woody seed strobili of conifers. Strobili are characterized by a central axis surrounded by spirally arranged or decussate structures that may be modified leaves or modified stems.

<i>Ginkgo</i> Genus of ancient seed plants with a single surviving species

Ginkgo is a genus of non-flowering seed plants, assigned to the gymnosperms. The scientific name is also used as the English common name. The order to which the genus belongs, Ginkgoales, first appeared in the Permian, 270 million years ago, and Ginkgo 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 found in the wild only in China, but is cultivated around the world. The relationships between ginkgos and other groups of plants are not fully resolved.

<i>Ginkgo dissecta</i> Extinct species of tree

Ginkgo dissecta is an extinct ginkgo species in the family Ginkgoaceae described from a series of isolated fossil leaves. The species is known from Early Eocene sediments exposed in the province of British Columbia, Canada, and Washington, US. It is one of two Ginkgo species found at the Washington and British Columbia sites.

<i>Ginkgo huttonii</i> Extinct species of tree

Ginkgo huttonii is an extinct Ginkgo species in the family Ginkgoaceae from the Jurassic of England. The fossil is also known by the name, Ginkgoites huttonii, the genus, Ginkgoites, referring to a group of extinct members of the Ginkgoaceae. G. huttonii was a broad-leaved, deciduous gymnosperm bearing resemblance to the only living member of the Ginkgoaceae, Ginkgo biloba.

<i>Yimaia</i> Extinct genus of seed-bearing plants

Yimaia is an 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 the presence of "Ovulate organs consisting of a peduncle and up to eight or nine terminal, sessile, contiguous and orthotropous ovules." The ovules are associated with leaves of either Baiera or Ginkgoites leaf morphospecies. Fossils have been found in Middle Jurassic deposits in China.

<i>Baiera</i> Extinct genus of seed plants in the family Ginkgoaceae

Baiera is a genus of prehistoric gymnosperms in the order Ginkgoales. It is one of the oldest fossil foliage types of Ginkgoales, and is related to the genera Ginkgo and Ginkgoites. Fossils of Baiera are found worldwide, and have been known from the Permian to the Cretaceous.

References

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  2. Stull, Gregory W.; Qu, Xiao-Jian; Parins-Fukuchi, Caroline; Yang, Ying-Ying; Yang, Jun-Bo; Yang, Zhi-Yun; Hu, Yi; Ma, Hong; Soltis, Pamela S.; Soltis, Douglas E.; Li, De-Zhu (19 July 2021). "Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms". Nature Plants. 7 (8): 1015–1025. doi:10.1038/s41477-021-00964-4. ISSN   2055-0278. PMID   34282286. S2CID   236141481.
  3. Zhou, Zhi-Yan (March 2009). "An overview of fossil Ginkgoales". Palaeoworld. 18 (1): 1–22. doi:10.1016/j.palwor.2009.01.001.
  4. 1 2 3 4 5 6 Taylor, Thomas N.; Taylor, Edith L.; Krings, Michael (29 December 2008). Paleobotany, Second Edition: The Biology and Evolution of Fossil Plants (2nd ed.). Academic Press. ISBN   9780123739728.
  5. 1 2 3 4 Beck, Charles (2014). "Ginkgoales". Access Science. doi:10.1036/1097-8542.289700 . Retrieved 13 April 2017.
  6. 1 2 Jalalpour, Julie; Malkin, Matt; Poon, Peter; Rehrmann, Liz; Yu, Jerry (1997). "Introduction to the Ginkgoales". www.ucmp.berkeley.edu. Retrieved 20 April 2017.