Ginkgo biloba
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Ginkgo a living fossil

. Classification:
Kingdom  Plantae -- Plants
    Subkingdom  Tracheobionta -- Vascular plants
        Superdivision  Spermatophyta -- Seed plants
            Division/phylum  Ginkgophyta -- Ginkgo
                Class  Ginkgoopsida
                    Order  Ginkgoales
                        Family  Ginkgoaceae -- Ginkgo family
                           Genus  Ginkgo L.
The only living representative of the order Ginkgoales is the Ginkgo biloba.
Source: National PLANTS Database -USDA, NRCS. 2001. The PLANTS Database, Version 3.1

The genus Ginkgo is one of the major groups in the phylum Ginkgophytes. Fossils of Ginkgo are recognized by its leaves, reproductive organs and wood. Its leaf remains are the most often found.

Its earliest leaf fossils date back to 270 million years ago in the Permian period of the Paleozoic when seed ferns and ferns dominated and Ginkgo, cycads and conifers came into prominence. 
So in the era of the dinosaurs (Jurassic 213 million years ago) it already existed!

Ginkgo fossil Perm Germany (photo Cor Kwant)
Ginkgo Baiera 
Lower Upper Permian (Lower Thuringian)
Zechstein / Kupferschiefer, Harz area, Central Germany
Museum für Naturkunde der Humboldt-Universität zu Berlin
photo © Cor Kwant

Sphenobaiera (photo Prof. R.A. Spicer)
photo © Prof. R.A. Spicer

ginkgo and dinos

Ginkgoales is a group of gymnosperms composed of the family Ginkgoaceae traditionally described as consisting of six “families” and possibly 19 genera, for instance Baiera, Ginkgoites, Ginkgoidium, Ginkgo, Arctobaiera, Sphenobaiera, Windwardia, Trichopitys. However this classification is ill-defined, for instance variations in Baiera, Sphenobaiera and Ginkgoites are now considered to be Ginkgo as well.

Its maximum diversity was achieved in the Jurassic and Early Cretaceous periods of the Mesozoic. The pteridosperms (‘‘seed ferns’’), and especially the Peltaspermales, are at present thought to be the most plausible ancestral group. 
The Ginkgoales are so isolated evolutionarily that efforts to establish its closest extant sister group have remained controversial. Nevertheless, there is a growing belief, supported by genetic research, that cycads are the most plausible closest relative (both have flagellated motile sperm and pollen tubes) rather than conifers.

Undoubted oldest fossils of the genus Ginkgo (Ginkgo digitata) are from 
the Early Jurassic period in the Mesozoic from a single location (Fergana) in what is now the Asiatic part of the former USSR, and therefore it is considered to be the oldest extant genus among seed plants. 

During this time Ginkgo, Bennettitales, ferns, cycads and conifers dominated the vegetation of the world. Mesozoic climate: no polar ice, wet and warm climatic conditions followed after aridity in the early Mesozoic.

During the Middle Jurassic there was an increase in species, Ginkgo digitata, Ginkgo huttoni, Ginkgo yimaensis, throughout the northern parts of the Laurasian supercontinent.

Shunosaurus under Ginkgo yimaensis (picture Brian Engh)
click to enlarge
Middle Jurassic of China, Shunosaurus under a
Ginkgo yimaensis with chichi
© Brian Engh

Ginkgo digitata

Ginkgo digitata
(from A.C. Seward, 1919)

Fossils picture gallery.

Ginkgo sp. Upper Triassic, East Greenland (photo The Field Museum)
Ginkgo sp.
Upper Triassic,  Stage: Rhaetian,
Bed 2, Astartekloft, East Greenland
photo courtesy The Field Museum, GEO86387d, Photographer John Weinstein
Ginkgo yimaensis
Ginkgo yimaensis
Middle Jurassic, Henan Province, China
 (redrawn from Zhou and Zhang, 1989a)

Maximum diversity in species existed during the Cretaceous period in the Northern Hemisphere, areas now known as Asia, Europe and North America, for instance Ginkgo coriacea, Ginkgo adiantoides, Ginkgo tigrensis, Ginkgo apodes, Ginkgo digitata, Ginkgo yimaensis, Ginkgo gardneri. Ginkgo was also present in the Southern Hemisphere:  Gondwana.

Maps of Ginkgo fossil spread.


Ginkgo evolution scheme

Ginkgo evolution (click to enlarge)

From: Zhou Z, Zheng S , 2003
 picture © Z. Zhou

Most species are mainly distinguishable on the basis of leaf anatomy and geographical distribution. It is well known that leaves on a living Ginkgo tree have a considerable variation in form, therefore the fossil record is supposed to include many species close to Ginkgo biloba (thus certain species names are adopted only as a matter of convenience). 

However fossils of Ginkgo yimaensis (170 mya) and Ginkgo apodes (121 mya) are found with Ginkgo-like reproductive organs and these discoveries leave no doubt about their belonging to the genus Ginkgo.

Sinornis santensis with Ginkgo apodes (image Alexander Lang)
 Ginkgo apodes with Sinornis santensis
Cretaceous, China
picture © Alexander Lang

Ginkgo huttoni
Ginkgo huttoni
Middle Jurassic,
Scalby, North Yorkshire, UK
photo © David Scarboro

More: Fossils picture gallery.

Ginkgo siberica
Ginkgo sibirica
Late Jurassic, Oregon
National Museum of Natural History, Washington D.C.
photo © Nico Bal
ginkgo dissecta
Ginkgo dissecta
Eocene, MacAbee Formation,
Cache Creek, British Columbia, Canada
photo ©  David Scarboro

Cretaceous landscape
The World of Dinosaurs, Cretaceous landscape North America (left: Ginkgo tree)
picture from The United States Postal Service

In structure and form (morphologically) indistinguishable forms of Ginkgo biloba extend back to the Early Cretaceous, therefore many researchers consider the fossil and modern forms to belong to the same species. This fossil form is most commonly identified as Ginkgo adiantoides, however many other designations are considered essentially equal.
It was common and widespread for a long time. 

Click here for Ginkgo fossils picture gallery: 
Paleozoic, Mesozoic, Tertiary and more.

Rhamphorhynchus, Jurassic
picture Heinrich Harder
dinos asteroid
At the end of the Cretaceous period all dinosaurs,  marine reptiles, and about 75 percent of all other species on earth became extinct.

Only three (or four) species were left in the Tertiary (65 million years ago): Ginkgo adiantoides (mainly Northern Hemisphere) with leaves virtually indistinguishable from modern Ginkgo biloba, andGinkgo gardneri (late Paleocene, only found on the Isle of Mull, Scotland).
Ginkgo jiayinensis (Northeast China, Wuyun Formation of Jiayin) differs from other Cenozoic ginkgos in having amphistomatic leaves (stomata  (pores) on both surfaces of the leaf) due to environmental adaptation.

In the Southern Hemisphere a different, more strongly digitate type of Ginkgo leaf persists into the Eocene, but so far it is not well documented (Ginkgo patagonica).

Ginkgo adiantoides (photo Cor Kwant)
Ginkgo adiantoides, Paleocene, Scotland
Naturalis, Leiden
 photo © Cor Kwant

Video: Different shapes of leaves on Ginkgo biloba. 
More videos here.
video © Cor Kwant

The decline is possibly the result of the extensive cooling that occurs throughout the Northern Hemisphere and rainfall gradually shifting from one of the summer-wet to one of summer-dry. The rise of the angiosperms might have played a role as well. The extinction of the dinosaurs  as potential seed dispersers of the large seeds may also have influenced this decline, which is in line with the fossil records. Read more about seed dispersers on my Propagation-page. Climate of the Cenozoic: very warm to glacial period.
About 7 million years ago Ginkgo disappeared from the fossil record of North America.
In Eurasia only very limited numbers of fossils are found from the Pliocene, and for the Pleistocene, the only known occurences of Ginkgo are from southwestern Japan. It was gone from Europe by about 2.5 million years ago.

Ginkgo biloba L. survived in some areas of China where the impact of glaciation was minimal.
DNA analyses have demonstrated  (Zhao et al., 2019) that Ginkgo refugia occur in southwestern, eastern and southern China:

- A refugium occurs in southwestern China especially around the southern slopes of Jinfo Mountain (Jinfo Shan) of  Nanchuan County at the boundary of Chongqing Municipality and Guizhou Province (see area map; 28°53'N; 107°27'E). Ecological work in this area, as well as in adjacent parts of Guizhou Province, has identified numerous small populations, for instance in Wuchuan County and Tuole (Panxian), that can be considered to be either wild or remnants of wild plants. Evidence for the persistance of wild Ginkgo biloba (Ginkgoaceae) populations in the valley and lower mountain slopes of the Dalou mountains was published in 2012 (Tang et al.) and in 2019 (Zhao et al.).
More info and photos here.

- Ginkgos in eastern China, e.g. on West Tianmu Mountain, Zhejiang province:  this nature reserve is an another refugium for wild Ginkgo biloba communities.
Non-Chinese populations are all genetically close to this eastern lineage, indicating multiple human-mediated introductions of Ginkgo from eastern China into North America and Europe. More info: article Zhao et al.

- A third refugium has been genetically classified in southern China e.g. in Fujian province.
Interesting is that nearly all fossil finds of Ginkgo lie poleward of 40 oN latitude, during the Tertiary more toward 40 o due to cooling and increased seasonality. Its survival in China is deviating from the long-term history of Ginkgo.

The Chinese Ginkgo biloba survived essentially unchanged. From the Jurassic till now there was a reduction of individual ovule-stalks and a decrease in the number of ovules. The size of the ovules increased. The ovulate organs may find their origin in the Middle JurassicGinkgo yimaensis (170 million years ago). The embryos of Ginkgo biloba seeds possess a temperature-dependent developmental-delay mechanism that allows seeds to survive winter by preventing premature germination in fall. This and other cold-climate adaptations appear to have evolved within the genus Ginkgo during the early Cretaceous, when the Northern Hemisphere was undergoing dramatic cooling after a long period of stable, warm conditions. Fossilized leaves and reproductive organs from the Lower Cretaceous period (Ginkgo apodes found in the 121 million years old Yixian Formation in N.E. China) show that their morphology has changed little until now.

Ginkgo adiantoides from the Tertiary period (56 million years ago) is remarkably similar to the modern Ginkgo biloba.

Fossil woods of the Ginkgoales are rare. A well-known example is Ginkgo beckii named after prof. George Beck, who discovered the petrified wood in the 1930s near Columbia River Gorge, Vantage, central Washington, North America.

Dating from the Miocene period, 15 million-year-old petrified logs of Ginkgo beckii and many other tree species have been uncovered from flood sediments and lava flows near this region. Ginkgo logs were carried by raging flood waters to the vantage area, where they lay burried in lake bed sediments covered by lava flows for millions of years.

Ginkgo beckii shows a striking similarity to Ginkgo biloba wood, with the possible exception of fewer pits per unit length on the radial walls of its tracheids.

petrified Ginkgo wood Arizona (photo Cor Kwant)
larger image
petrified Ginkgo wood, 6 x 7 cm, limb cross section, Triassic
Arizona, north of Canyon de Chelly, Chinle formation
 photo © Cor Kwant

In 2016 fossil wood -Ginkgoxylon liaoningense- was collected from the Tiaojishan Formation in western Liaoning, northeastern China. 
It probably dates from the Middle to Late Jurassic.
Its anatomy departs slightly from that of modern Ginkgo, it displays all its characteristic features. It differs only in having a more mixed type of radial pitting, which also occurs, albeit locally, in modern Ginkgo wood. The xylem structures of Ginkgoxylon liaoningense illustrates the basal state of Ginkgo wood anatomy and thus contributes to the understanding of Ginkgo evolution.

Ginkgoxylon liaoningense
Ginkgoxylon liaoningense

Believed to be fossil wood is known from the Lower Permian (Hueco Formation, New Mexico) and  Late Triassic (Holbrook area, Chinle Formation, Petrified Forest National Park, Arizona). 
From the Eocene period a single specimen is known as Ginkgo bonesii, Clarno Formation, Oregon. 

Ginkgo petrified wood (photo Lewis Goodman)

petrified Ginkgo wood, 17 cm diam., Miocene 
Vantage, North America
photo © Lewis Goodman

Fossil ginkgoalean wood is probably scarce because of lack of chemical resistance of the wood to degradation of the cell walls after deposition in sediments.

More photos and info about Ginkgo wood.

Life history traits of the genus Ginkgo are for instance:
- highly conservative morphologically
- slow space of evolution
- slow reproduction rate (late sexual maturity)
- large seeds
- long life span
- conservative in its ecological tolerance: well-drained disturbed habitats; now representing an ecological paradox because the living Ginkgo biloba L. has few of the life-history traits typical for plants that prosper in modern disturbed settings. 
But it now also thrives in for instance New York City Center, it does not suffer from air pollution and pests!
video CreatureCast

Click here for fossils picture gallery.
Maps of Ginkgo fossil spread.

Spuren picture
picture © Atsuko Kato
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Special thanks to David Scarboro from Reading  (UK), Open University tutor,
for providing many pictures for this page.

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©  Cor Kwant 
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