Ginkgo: The ‘Living Fossil’ That Alone Survives From a Lineage Older Than the Dinosaurs

Every autumn, fan-shaped yellow leaves carpet the streets. It is such a common street tree that we pass it without a second look, yet the ginkgo is the only species surviving today from an ancient plant lineage, the order Ginkgoales, which goes back to the Permian, before the dinosaurs appeared. That is why it is nicknamed a “living fossil.” This nickname, however, bundles several different times together. If we separate the time when the lineage began, the time when leaves resembling today’s appeared, and the time when today’s species, Ginkgo biloba, appears in the fossil record, it becomes clearer what the ginkgo has kept, and for how long.

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A Lineage Born in the Permian, a Single Surviving Species

The earliest fossil usually counted in the ginkgo’s lineage is Trichopitys, from earliest Permian (Asselian, about 299–293 million years ago) strata in France. The Ginkgoales were a common group in Permian and Triassic floras, and in the Jurassic and Early Cretaceous the genus Ginkgo spread across the continents of the Northern Hemisphere. The start of this lineage thus comes before the Triassic, when the first dinosaurs appeared.

Then things changed. From the Late Cretaceous into the Cenozoic, the diversity of the Ginkgoales declined, and by the end of the Cretaceous every group except the genus Ginkgo had disappeared. Because this decline overlaps with the rise of flowering plants (angiosperms), one hypothesis holds that flowering plants, better suited to disturbed environments, pushed the ginkgo group aside. By the end of the Pliocene, the ginkgo had almost vanished from the fossil record except for a small area of central China.

What remains today is a single species, the ginkgo (Ginkgo biloba). It is the only living species in the division Ginkgophyta, the class Ginkgoopsida, the order Ginkgoales, the family Ginkgoaceae and the genus Ginkgo. A single species stands alone for an entire division, and few living things in taxonomy have been left this solitary.

How Far Does “Living Fossil” Hold True?

A 2019 study of 545 ginkgo genomes published in Nature Communications introduces the ginkgo as a living fossil “with almost no morphological change for at least 200 million years.” To understand this claim accurately, it helps to look at the fossil record stage by stage.

First, the genus Ginkgo. Fossils of the genus with reproductive organs similar to those of today’s species appear in the Jurassic. The University of California Museum of Paleontology (UCMP) at Berkeley traces the genus’s line back to the Early Jurassic, about 190 million years ago, while Wikipedia and other sources place its first appearance in the Middle Jurassic. Ginkgo huttonii, common in Jurassic strata in Yorkshire, England, had simple leaves divided into four lobes with dense, radiating veins. They resemble today’s ginkgo leaves but are not the same.

Leaves almost identical to today’s are confirmed in much later fossils. According to UCMP, by the Paleocene (about 66–56 million years ago) the genus Ginkgo had narrowed to what was effectively a single species, the group usually called Ginkgo adiantoides, whose leaves resemble modern ginkgo leaves so closely as to be “virtually indistinguishable.” The fossil record of today’s species, G. biloba, itself runs from the early Eocene, about 51.5 million years ago. In short, the lineage began about 290 million years ago, leaves resembling today’s are found in Jurassic fossils, and nearly identical leaves are confirmed in Paleocene fossils at the latest. “200 million years” is the phrase the researchers used to introduce the ginkgo, and the nickname is read accurately only if we also remember that each fossil resembles today’s ginkgo to a different degree.

It is not only the leaves. A 2013 study in BMC Evolutionary Biology analyzed 20 fossils of ginkgophyte seedlings (young plants) from the early Middle Triassic, collected from a single bedding plane of the Grès à Voltzia sandstone in France. Four stages were preserved in sequence, from the stage in which seed leaves form inside the seed and it germinates to the stage in which the taproot thickens and properly formed leaves appear, and the researchers concluded that germination and seedling development were “nearly identical” to those of the living ginkgo. The main difference was that the first leaves were larger than those of today’s ginkgo and divided in two. The way the seed sprouts was, in other words, already in place more than 240 million years ago.

Fan-Shaped Leaves and Forking Veins: A Group Apart Even Among Gymnosperms

Hold a ginkgo leaf up to the light and you can see the rule its veins follow. Two veins enter the leaf blade through the leaf stalk, then fork in two again and again as they run to the edge of the fan. This is called dichotomous venation. The forked veins never meet again to form a net. It is a structure entirely different from the net venation seen in cherry or maple leaves.

The ginkgo is a gymnosperm, a plant whose seeds are exposed, yet it forms its own division, distinct from both the conifers and the cycads. Among living plants, the cycads are considered its closest relatives. Different from the needles of conifers and from the feather-like leaves of cycads, this fan-shaped leaf is the last form left by a lineage that otherwise vanished long ago. In Flora Japonica, published by Philipp Franz von Siebold and Zuccarini, the ginkgo’s leaves and reproductive organs appeared in a detailed plate. Beneath the plate is written Salisburia adiantifolia, an old name some scholars used at the time.

Swimming Sperm and a Seed That Is “Not a Fruit”

The ginkgo’s reproduction keeps the ways of ancient plants. The ginkgo is dioecious, with separate female and male trees. Male trees produce small pollen cones, and female trees produce two ovules at the tip of a stalk. The pollen reaches the ovules on the wind.

In 1896, Sakugoro Hirase (平瀬作五郎), a Japanese botanical illustrator, became the first to discover swimming sperm in the ginkgo. According to Harvard University’s Arnold Arboretum, about four months after the pollen is shed in spring, sperm form in the pollen tube inside the ovule, still attached to the tree, and swim to the egg cell. The sperm are about 70–90 micrometers across, large for plant sperm, and the number of flagella covering them is given as anywhere from about 1,000 to several thousand, depending on the source. They look nothing like human sperm, which swim with a single tail. Among living seed plants, the only groups that fertilize with swimming sperm like this are the ginkgo and the cycads.

The yellow autumn balls commonly called “ginkgo fruit” are not fruit in the botanical sense. As a gymnosperm, the ginkgo has no ovary enclosing the ovule and cannot bear true fruit. That yellow structure is a seed, not a fruit. Of the layers that wrap the seed (the seed coat), the outermost is the soft, fleshy sarcotesta; beneath it, exposed once the soft layer is removed, is the hard sclerotesta, and inside that hard shell lie a thin inner layer, nutritive tissue and the embryo.

The source of the foul smell on autumn streets is butyric acid in the sarcotesta. Fallen seeds smell like rancid butter or vomit. The kernel inside the hard shell, the “ginkgo nut” we roast and eat, is widely used in Asian cooking. Eating many at once or eating them over a long period, however, can cause poisoning from ginkgotoxin (4′-O-methylpyridoxine), and cases of seizures have been reported. This compound is heat-stable, so it does not go away with cooking, and it is wise to limit how many you eat.

Names That Remember a Slow-Growing Tree

The Korean name eunhaeng (銀杏) means “silver apricot.” The name comes from the seed’s resemblance to an apricot and its surface coated with a silvery white powder. Another name, gongsonsu (公孫樹), reflects how long the tree takes to bear seeds. Because a female tree can take nearly 30 years of growth before it produces seeds, it was called “the tree whose seeds come only in the grandchild’s generation.” It is also called apgaksu (鴨脚樹), “duck-foot tree,” because its leaves resemble a duck’s foot.

The scientific name Ginkgo has a story too. Engelbert Kaempfer, the first Westerner to study the tree, in Nagasaki, Japan, in 1690, wrote the name as “Ginkgo” in his book Amoenitates Exoticae, published in 1712. If he had been transcribing the Japanese “ginkyo,” “ginkio” or “ginkjo” would have been more natural, so this spelling was long regarded as Kaempfer’s mistake. Later research, however, suggested that he followed the spelling of his interpreter, Imamura Gen’emon, who spoke the Nagasaki dialect. Linnaeus adopted the spelling as it was from Kaempfer’s record, and it became fixed as the genus name.

Trees That Have Endured a Thousand Years

Slow-growing ginkgos live long. Some trees have been measured at more than 1,600 years old, and there are estimates that the largest trees exceed 3,500 years. In Korea, the ginkgo at Yongmunsa Temple in Yangpyeong, Gyeonggi Province, is the best-known example. Designated a Natural Monument in 1962, the tree is estimated to be about 1,100 years old as of 2024, and at 38.8 m it is the tallest ginkgo in Korea. Legend says that Crown Prince Maui of Silla planted it on his way to Geumgangsan, and it was prized so highly that it is said to have been granted the court rank of dangsanggwan (senior third grade) under King Sejong of Joseon.

The ginkgo’s vitality also showed itself under the most extreme ordeal. When the atomic bomb fell on Hiroshima on August 6, 1945, six ginkgo trees about 1,130–2,160 m from the hypocenter are widely reported to have survived and sprouted again. In Japan, trees that survived the atomic bombing are called hibakujumoku (被爆樹木), “A-bombed trees,” and the ginkgo is sometimes called a “bearer of hope.” Of the six, the Hosenbo ginkgo, the closest to the hypocenter at about 1,130 m, sprouted again even after the temple was destroyed. When the temple was rebuilt after the war, moving or cutting down the tree was considered, but in 1994 it was decided to leave the tree where it was and build around it, so today the steps in front of the main hall split to the left and right with the tree between them.

Endangered in the Wild, a Street Tree in the City

This tree that looks so common is listed as Endangered (EN) on the Red List of the International Union for Conservation of Nature (IUCN). For centuries the ginkgo was thought to have already disappeared from the wild. Whether natural populations survived was long debated, and a 2012 study in the American Journal of Botany examined ginkgo forests in the Dalou Mountains (大婁山) of southwestern China. Pointing out that the makeup of the plants growing alongside the ginkgo matched the ancient floras in which ginkgo fossils are found, and that the ages of several trees went back to before people settled the region, the researchers regarded these populations as fragments of the tree’s original natural range.

The “semi-wild” ginkgos of Tianmu Mountain (天目山) in Zhejiang grow along streams, on rocky slopes and at cliff edges. Some Chinese scholars, however, hold that the ginkgo has been planted and preserved by monks for about 1,000 years, so how much of it is wild is still unsettled. The 2019 study mentioned above analyzed the genomes of 545 trees chosen from 51 populations around the world, found three glacial refugia within China, and confirmed traces of ginkgos from eastern China being carried by people to other continents several times. The researchers presented the ginkgo as a case of “how people have saved a species from extinction and assisted its resurgence,” and argued that people’s protection of ginkgo forests, rooted in traditional culture and faith, supported the tree’s survival and spread.

The ginkgos in cities today are trees that people planted and raised. The ginkgo tolerates pollution and cramped root space well, rarely gets diseases even in cities and has few pests, so many cities in Korea, China and Japan have planted it as a street tree. It is familiar enough that Sungkyunkwan University and the University of Tokyo use the ginkgo leaf as their emblem. Behind the way a tree that nearly vanished from the wild became the most common sight on our streets lies the long time people and this tree have spent together.

A Single Leaf That Has Endured Time

The ginkgo shows us two things together. One is that a lineage born in the Permian passed through countless changes until only a single living species remained, and that the form of its leaves and seeds was kept for a very long time. The other is that human hands carried that life, imperiled in the wild, to temple courtyards and city streets. Each yellow leaf among those that pile up at our feet every autumn holds both the memory of unimaginably distant geological ages and a thousand-year story of living alongside people. Being able to look closely at a life so deep and ancient is a quiet wonder the created world offers us.

References

• Ginkgoales – Wikipedia
• Ginkgo biloba – Wikipedia
• Ginkgo huttonii – Wikipedia
• Ginkgoales: Fossil Record – UCMP Berkeley
• Bauer et al. (2013), Fossil ginkgophyte seedlings from the Triassic of France resemble modern Ginkgo biloba – BMC Evolutionary Biology
• Zhao et al. (2019), Resequencing 545 ginkgo genomes across the world reveals the evolutionary history of the living fossil – Nature Communications
• Tang et al. (2012), Evidence for the persistence of wild Ginkgo biloba populations in the Dalou Mountains, southwestern China – American Journal of Botany
• The Swimming of the Ginkgo Sperm – Arnold Arboretum of Harvard University
• Ginkgo (Eunhaengnamu) – Korean Wikipedia (in Korean)
• Ginkgo Tree of Yongmunsa Temple, Yangpyeong – Korean Wikipedia (in Korean)
• A-bombed Ginkgo trees in Hiroshima, Japan – The Ginkgo Pages
• Hibakujumoku – Wikipedia