Ginkgo a living fossil
Kingdom Plantae -- Plants
Subkingdom Tracheobionta -- Vascular plants
Superdivision Spermatophyta -- Seed plants
Division/phylum Ginkgophyta -- Ginkgo
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.
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.
oldest fossils of the genus Ginkgo (Ginkgo digitata) are
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.
click to enlarge
Middle Jurassic of China, Shunosaurus under a
Ginkgo yimaensis with chichi
© Brian Engh
Upper Triassic, Stage: Rhaetian,
Bed 2, Astartekloft, East Greenland
photo courtesy The Field Museum, GEO86387d, Photographer John Weinstein
Middle Jurassic, Henan Province, China
(redrawn from Zhou and Zhang, 1989a)
Maximum diversity in species
existed during the Cretaceous period in the
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:
Maps of Ginkgo fossil spread.
Ginkgo evolution (click to enlarge)
From: Zhou Z, Zheng S , 2003
|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.
Ginkgo apodes with Sinornis santensis
picture © Alexander Lang
Scalby, North Yorkshire, UK
photo © David Scarboro
Late Jurassic, Oregon
National Museum of Natural History, Washington D.C.
photo © Nico Bal
Eocene, MacAbee Formation,
Cache Creek, British Columbia, Canada
photo © David Scarboro
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.
picture Heinrich Harder
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
but so far it is not well documented (Ginkgo patagonica).
Ginkgo adiantoides, Paleocene, Scotland
photo © Cor Kwant
Video: Different shapes
of leaves on Ginkgo biloba.
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
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
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.
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.
In 2016 fossil wood -Ginkgoxylon
liaoningense- was collected from the Tiaojishan Formation in western
Liaoning, northeastern China.
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).
wood, 17 cm diam., Miocene
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.
|Life history traits of the genus
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!
picture © Atsuko Kato
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thanks to David Scarboro from Reading (UK), Open University tutor,
for providing many pictures for this page.
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© Cor Kwant