Ginkgo

New Haven and London

 Ginkgo
the tree that time forgot

Peter Crane
W I T H A F O R E W O R D BY PET E R R AV E N
 Published with assistance from the foundation established in memory of
Philip Hamilton McMillan of the Class of 1894, Yale College.

Copyright © 2013 by Peter Crane. All rights reserved. This book may not be reproduced,
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Library of Congress Cataloging-in-Publication Data

Crane, Peter R.
Ginkgo : the tree that time forgot / Peter Crane.
p.cm.
Includes bibliographical references and index.
ISBN 978-0-300-18751-9 (cloth : alk. paper)
1. Ginkgo. I. Title.
QK494.5.G48C73 2013
585′.7—dc232012032831

A catalogue record for this book is available from the British Library.

This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

10 9 8 7 6 5 4 3 2 1
 FOR EMILY AND SAM

with hope for their generation’s wider view of the world

Ginkgo biloba

Dieses Baum’s Blatt, der von Osten

Meinem Garten anvertraut,
Giebt geheimen Sinn zu kosten,
Wie’s den Wissenden erbaut.

Ist es ein lebendig Wesen?

Das sich in sich selbst getrennt;
Sind es zwey? die sich erlesen,
Dass man sie als eines kennt.

Solche Frage zu erwiedern

Fand ich wohl den rechten Sinn;
Fühlst du nicht an meinen Liedern,
Dass ich Eins und doppelt bin?

—Johann Wolfgang von Goethe, September 15, 1815

Ginkgo biloba

This tree’s leaf which from the Orient

Is entrusted to my garden
Lets us savor a secret meaning
As to how it edifies the learned man.

Is it one living being?

That divides itself into itself
Are there two who have chosen each other,
So that they are known as one?

To reply to such a question

I found, I think, the condign sense.
Do you not feel that in my poems
I am single and twofold?

—English translation by Kenneth Northcott, 2006

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 Contents

ForewordPeter H. Raven, xiii

Preface, xvii

pa rt i Prologue

1 Time, 3
2 Trees, 8
3 Identity, 16

pa rt ii The Living Tree

4 Energy, 27
5 Growth, 35
6 Stature, 42
7 Sex, 53
8 Gender, 60
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contents

9 Seeding, 66
10 Resilience, 73

pa rt iii Origin and Prehistory

11 Origins, 81
12 Ancestry, 89
13 Relationships, 97
14 Recognition, 104
15 Proliferation, 111
16 Winnowing, 119
17 Persistence, 125
18 Prosperity, 132

pa rt i v Decline and Survival

19 Constraint, 143
20 Retreat, 152
21 Extinction, 156
22 Endurance, 162
23 Relic, 168

pa rt v History

24 Antiquity, 177
25 Reprieve, 184
26 Voyages, 190
27 Renewal, 195
28 Naming, 204
29 Resurgence, 209
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contents

pa rt vi Use

30 Gardens, 217
31 Nuts, 226
32 Streets, 234
33 Pharmacy, 242

pa rt vii Future

34 Risk, 253
35 Insurance, 258
36 Gift, 264
37 Legacy, 272

Appendix: List of Common Plant Names

Used in the Text and Latin Equivalents, 279
Notes, 285
Bibliography, 335
Illustration Credits, 362
Index, 365
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 Foreword

Perhaps the best known and most easily recognized of the world’s 100,000 kinds of
trees, ginkgo stands out by virtue of its unique features, amazing history, and long as-
sociation with people. With their distinctive fan-shaped leaves and tall trunks, ginkgo
trees adorn parks, streets, and recreational areas throughout the temperate regions of
the world. When the weather turns sharp, all of the leaves suddenly turn a brilliant yel-
low, dropping soon after to lay a lovely, bright yellow carpet under each tree. Ginkgo
trees are bisexual, some producing seeds and others only pollen-bearing organs. The
seeds are naked, as in other gymnosperms (such as pines, cycads, and cedars); the
fleshy outer coat of the ginkgo seed smells strongly of rancid butter (butyric acid). The
inner “kernel” of the seed is nutlike and eaten widely in the Orient, once the smelly
outer layer and the stony inner one are removed. Among the seed plants, only ginkgo
and cycads form motile sperm within their pollen tubes, a fascinating example of the
survival of an archaic characteristic.
A particularly fascinating feature of ginkgo’s history is that it is extremely rare as
an uncultivated native tree. Once widespread throughout the Northern Hemisphere, it
disappeared at different times in different regions, including most of Eurasia and all of
North America, as the climate changed and new plant communities came into being.
In the Northern Hemisphere, China represents the zone where the most ancient lin-
eages have survived, including many that were once widespread. Only in two moun-

xiii
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foreword

tainous areas of southern China do the varied genetic patterns of the standing trees
make it likely that they or their immediate ancestors were native to the forests where
they still occur. In addition, however, individual trees here and there in southern China
may represent originally wild individuals that survived and eventually were protected
by the people who lived near them, as is the case in the dawn redwood, Metasequoia.
That ginkgo has survived essentially unchanged for as much as 200 million years is a
miracle: virtually all other kinds of plants and animals that occurred with it more than
150 million years ago have become extinct. Those that disappeared included the rela-
tives of the surviving evolutionary lineage to which Ginkgo biloba belongs. Although
defining species from fossil material alone is difficult, ginkgo may legitimately be re-
garded as the oldest surviving kind of plant: the characteristics of the living species
closely resemble those of its Jurassic ancestors!
Starting about a thousand years ago, ginkgo was brought in from the wild, or simply
allowed to survive where native, in the temple gardens and protected forests of China;
it has been nurtured and valued by human beings over the past thousand years or
more. Ginkgo evidently was spread from there to Korea and Japan over the course of
the past 800 years or so. After Europeans discovered it in Japan in the late seventeenth
century, it was brought into cultivation in Europe over the next several decades, even-
tually spreading around the world in areas with a suitable climate. Ginkgo is resistant
to air pollution and to pests and diseases, doing very well as a city tree through tem-
perate zones of both Northern and Southern Hemispheres. Without the attention of
human society, ginkgo would doubtless have become extinct by now, or at best reduced
to a few individuals. In that sense, its history affords an excellent example of how we
must deal with many plant species in this age of change and extinction if we wish to
save them for the future.
As Peter Crane has emphasized in the closing pages of this fine book, human beings
have had a short span of existence on this 4.5 billion–year–old planet, and we live only
a short time as individuals. That we share the Earth with such a venerable organism,
with a history where hundreds and tens of millions of years are relevant, should help to
give us a better perspective from which to think of our own lives and existence here and
thus prepare as well as we are able for the future. Our short-term actions are ruining
the world in which we live much faster than we can imagine, with the world’s sustain-
able capacity sufficient to provide less than two-thirds of what we consume each year,
even though billions of us live hungry and in extreme poverty. The two billion or more
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foreword

additional people who will join our numbers in the next several decades will almost all
be poor, entering a world that we, with our largely short-term view of progress, are de-
stroying rapidly. Might not we be able to learn from the deep past and then redouble
our efforts to sustainably use our planet’s resources and thus live within its productive
capacity while we still have time to do so?

Peter H. Raven, President Emeritus, Missouri Botanical Garden, Saint Louis

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 Preface

The origins of this book trace to an early fondness for the wonderful accounts of
economically important plants written by the late Charlie Heiser in the 1970s and
1980s. These stand in a long tradition of popular science writing, but it was their par-
ticular blend of science and culture, leavened with personal experience, that provided
the inspiration for this book. In focusing on the ginkgo, a tree with such a long and
varied life story, my reach may have exceeded my grasp, but the challenge of trying to
balance my scientific inclination for depth with the need for breadth has brought its
own rewards.
From time to time researching and writing this book has impinged on my other re-
sponsibilities. Therefore, I am especially indebted to the three organizations that have
been my professional homes over the last decade. Work began on this book while I was
director of the Royal Botanic Gardens, Kew; continued at the University of Chicago;
and was finally completed at Yale. I am grateful to all three institutions for allowing me
to devote time to this project, which is not quite a normal piece of scientific research,
had little do with my professional responsibilities, and was always something of an in-
dulgence. Much of the writing was done in Seoul in the summers of 2009, 2010, and
2011, surrounded by ginkgo trees, during my time as a visiting professor in the World
Class Universities program of the National Research Foundation of Korea. I am espe-

xvii
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pre fac e

cially grateful to EWHA University, and my colleagues there, Jae Choe and Yikweon
Jang, for their hospitality and support.
This book would not have happened without the unwavering kindness of Fumiko
Ishizuna in Tokyo, who led me on several incomparable expeditions to key sites for
ginkgo in Japan. Those expeditions, and the opportunities they provided to see truly
spectacular great ginkgos, convinced me that this book was worth writing. I am also
deeply grateful to my friend Zhou Zhiyan in Nanjing for his patience and thoughtful
guidance in many aspects of this work, and not only in those chapters dealing with the
fossil record. Finally, this manuscript probably would have languished forever had it
not been for the dedication and commitment of Ashley DuVal, who more than anyone
else helped drive it through to completion.
Among the many other individuals who have helped me with different aspects of
this story I would especially like to thank Heidi Anderson and Andrew Drinnan in Aus-
tralia; Johanna Eder and Michael Kiehn in Austria; Kevin Aulenback and Spencer Bar-
rett in Canada; Cheng Quan, He Shan-an, Hu Yonghong, and Yungpeng Zhao in China;
Branko M. Begović Bego in Croatia; Kaj Raunsgaard Pedersen in Denmark; Hans Kerp
in Germany; Mitsuyasu Hasebe, Toshiyushi Nagata, Tetsuo Ohi-Toma, and Masamichi
Takahashi in Japan; Gerard Thijsse in the Netherlands; Jolanta Kalisz in Poland; Adrian
Patrut in Romania; Lye Lin Heng in Singapore; John Anderson and Brian Huntley in
South Africa; Hyosig Won in South Korea; Else Marie Friis in Sweden; Julia Buckley,
Eleanor Bunnell, Martin Hamilton, Liz Jaeger, Stephen Jury, Tony Kirkham, Christine
Leon, Brian Mathew, Andrew McRobb, Mark Nesbitt, John Parker, Martin Postle,
Hugh Prendergast, Malin Rivers, Moctar Sacande, Anna Saltmarsh, Wolfgang Stuppy,
and Fiona Wild in the United Kingdom; and Selena Ahmed, Mark Ashton, Bruce Bald-
win, Alona Banai, Jeremy Beaulieu, Graeme Berlyn, Kevin Boyce, Eric Brooks, Gary
Brudvig, Bret Buskirk, Ed Buyarski, Bill Carvell, Jeff Courtney, David Dilcher, Laura
Donnelley, Gerry Donnelly, Michael Donoghue, Ian Glasspool, Chris Haufler, Dave
Hayes, David Heidler, Christie Henry, Pat Herendeen, Nancy Hines, Michelle Hol-
brook, Kirk Johnson, Bill LeFevre, Andrew Leslie, Stefan Little, Chris Liu, Marie Long,
Steve Manchester, Greg McPherson, Herb Meyer, Rachel Meyer, Colleen Murphy-
Dunning, Andrew Newman, Peter Purdue, John Rashford, Laurel Ross, Kemba Sha-
kur, Pamela Soltis, Leroy Squires, Scott Strobel, Gregory Tarver, Douglas Trainor, Mary
Evelyn Tucker, Warren Wagner, Marianne Welch, Elisabeth Wheeler, Mimi Yiengpruk-
sawan, and Qingfu Xiao in the United States. All contributed ideas, information, or ex-
 xix
pre fac e

periences that found their way into the book. I am especially indebted to Bill Chaloner,
Pat Horn, Charles Jarvis, Toshiyushi Nagata, Peter Raven, and Scott Wing for their
sound advice and review of the manuscript, and to Jean Thomson Black, Sara Hoover,
and Dan Heaton at Yale University Press and Al Zuckerman at Writers House for their
kind guidance. The drawings that introduce each section are by Pollyanna von Knor-
ring, who has illustrated my research for almost thirty years.
Research for this book benefited greatly from the availability of a few key resources
that had already drawn together much scattered material on ginkgo, most important
the wonderful Ginkgo Pages Web site by Cor Kwant, the various writings of Peter Del
Tredici, and especially for Western readers the important book edited by Terumitsu
Hori, Robert Ridge, Walter Tulecke, Peter Del Tredici, Jocelyne Trémouillaux-Guiller,
and Hiroshi Tobe, which was produced as part of the celebrations on the centenary of
Hirase’s remarkable discovery of swimming sperm in ginkgo.
Finally, I am deeply grateful to my longtime friend Peter Raven for writing the fore-
word to this book, and to my family, Elinor, Emily, and Sam, for their infinite patience
during my multiyear obsession with this very special tree.
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 part i

Prologue
OVERLEAF Ginkgo leaf decorated with a drawing of the Wang Shi Yuan,
Master of the Nets Garden, Suzhou, China.
 1
Time

One huge ginkgo tree, topping all the others, shot its great limbs
and maidenhair foliage over the fort which we had constructed.
In its shade we continued our discussion . . .
—Sir Arthur Conan Doyle, The Lost World

To most people ginkgo is either the tree with the smelly “fruits” or the plant that is
good for your memory, but that unmistakable acrid aroma, or the leaf extract in the
health food store, is only part of what makes ginkgo unique. Common on city streets
from Beijing to London and Tokyo to New York, ginkgo is an increasingly common
backdrop to the bustle of modern city life. It is hard to imagine that these trees, now
towering above cars and commuters, grew up with the dinosaurs and have come down
to us almost unchanged for 200 million years. Ginkgo is one of the world’s most dis-
tinctive plants and has one of the longest botanical pedigrees; there is no other living
tree with a prehistory so deeply intertwined with that of our planet. This book, an
abridged global biography, sets out to tell ginkgo’s evolutionary and cultural life story.1
My interest in ginkgo stretches back more than thirty-five years, but it grew during
my time, from 1999 to 2006, as director of the Royal Botanic Gardens, Kew. During
those seven years, the oldest and perhaps the most important ginkgo in the United

3
 4
prologue

Kingdom flourished just steps from our family’s home in the garden. We passed it
almost every day; it was a regular stop on tours for distinguished visitors. It was one
of the most beloved of the Old Lions, that handful of trees remaining from the mid-
eighteenth century, when Kew was a royal estate. We watched this tree as it changed
through the seasons, and we worried when a big storm came through. In 2002 it was
designated one of fifty Great British Trees by the Tree Council in its celebration of the
Queen’s Golden Jubilee. It was among the most precious of the nearly fifteen thousand
trees on the Kew landscape.2
I first came across Siegfried Unseld’s delightful little book Goethe und der Ginkgo: Ein
Baum und ein Gedicht through my longtime colleague Else Marie Friis at the Swedish
Museum of Natural History in Stockholm. It led me to track down an English trans-
lation of the famous poem at the heart of Unseld’s book. Then, upon my return to the
United States in late 2006, Christie Henry at the University of Chicago Press presented
me with a copy of Kenneth Northcott’s rendering of Goethe and the Ginkgo. The idea
that those who enjoy Unseld’s book might like to know more about ginkgo was part of
the stimulus for this biographical sketch of a singular tree.
Ginkgo is a botanical oddity, a single species with no close living relatives. Once
regarded as a cousin of pines, yews, and cypresses, it was later recognized as some-
thing quite different. It was first distinguished from conifers in plant classifications of
the early nineteenth century. The evidence that has come to light since—particularly
an astonishing discovery made in Japan in 1896 about the intimate details of its re-
production—has reinforced the isolated position of ginkgo among living plants. So
in the twentieth century, as the world of plants has come into sharper evolutionary
focus, ginkgo has assumed new scientific importance. To borrow a phrase from Dar-
win, ginkgo has become a platypus for the plant kingdom, and plant paleontologists
have traced its lineage millions of years into prehistory. Ginkgo is now the most widely
recognized of all botanical “living fossils”: a tree that time forgot and an increasingly
familiar living link to landscapes of the distant past.3
For almost all of its long tenure on our planet ginkgo inhabited a world without
people, and for much of that time, a world very different from that of today. For tens
of millions of years it lived alongside plants and animals that are long since extinct.
Several different kinds of ginkgolike trees watched as our ancestors transformed from
reptiles to mammals. Fossil ginkgo leaves are known from every continent. The prehis-
 5
time

tory of ginkgo goes back to before the Atlantic Ocean existed and before the southern
continents broke from Antarctica and went their own ways.
As our planet has changed over the past 200 million years, ginkgo has proved re-
markably resilient. It felt the shock as new kinds of plants came to dominate the Earth
about 100 million years ago, but it was oblivious to the traumatic events that eliminated
the dinosaurs a few tens of millions of years later. Soon afterward ginkgo lost its foot-
hold in the southern continents, but in Asia, Europe, and North America it continued
to prosper. It flourished during the great warmth of 50 million years ago and once grew
close to the North Pole. But as the Earth grew cooler, ginkgo found itself in retreat and
was driven from its home in the Arctic.4
For the next forty million years, ginkgo was widespread across the northern conti-
nents. Its distinctive leaves are common in the fossil record. Eventually, though, and
for reasons that are not fully clear, ginkgo began to suffer. By the time our ancestors
diverged from those of other living apes, five million to seven million years ago, ginkgo
was probably already in decline. It was nearly extinguished by the great Ice Ages that
gave birth to our own species. When the last great southward push of the ice had re-
treated, ginkgo was barely hanging on, perhaps only in protected valleys scattered
across eastern and south-central China. By the time modern people arrived in that part
of Asia, perhaps fifty thousand years ago, ginkgo was already a relic.5
Human dominance on our planet could have meant the end for ginkgo, but un-
like many other trees, it has flourished alongside people. In one way or another, it
has proved useful; more unusually, it has become revered. In many cultures, in many
different ways, ginkgo commands unusual respect. Together, these qualities earned
ginkgo a reprieve. The nuts became a delicacy and were used for oil and in medicine.
The tree, with its distinctive leaves and great longevity, also took on symbolic meaning
in Buddhism, Daoism, and Confucianism. From China and Korea, ginkgo spread to
Japan and became incorporated in the indigenous religion of Shintoism. Many of the
great ginkgos of China, Japan, and Korea are in the grounds of Buddhist temples or
Shinto shrines.
From Asia, again with the assistance of people, ginkgo began its renewal. In the
eighteenth century, through the Dutch trading colony of Deshima in southern Japan,
ginkgo became known in Europe. Soon after, seeds found their way back to the Low
Countries and Britain, probably from both China and Japan, but perhaps also from
 6
prologue

A small Buddhist shrine at the base of an ancient ginkgo

at the Heungju Temple, in Taean, Chungnam, South Korea.

Korea. Ginkgo then spread quickly to North America and elsewhere as a horticultural
novelty. The Old Lion at Kew is one of several early ginkgo trees growing in Europe.
By the time of Goethe, and later in the nineteenth century, ginkgo was widely grown as
an unusual and striking tree: a symbol of the East. In just a hundred years, ginkgo re-
turned to many of those places from which it had been extinguished millions of years
before.
In the past fifty years ginkgo has been resurgent; interest in growing ginkgo, what it
stands for scientifically, and the ways in which it might be useful has never been higher.
Ginkgo has become recognized as a valuable street tree that grows well in tough places.
Resistant to disease, tolerant of pollution, and able to withstand extremes of heat and
cold, it is now familiar in urban landscapes over much of the world. Parts of Seoul are
a near ginkgo forest, and ginkgo is among the most common street trees in Manhattan.
 7
time

You can see ginkgo in San Francisco’s Golden Gate Park, in Monet’s garden at Giverny,
and in parks and gardens in all but the warmest and coldest places on our planet. In
recent decades ginkgo has also found its way into the pharmacy. It is among the most
popular of herbal remedies, and its medicinal properties are the subject of advanced
biomedical research. Extracts from ginkgo leaves are the source of a multibillion-dollar
pharmaceutical industry.
Beginning with an introduction to the living tree, this book follows the prehistory of
ginkgo over the past 250 million years from its origin, proliferation, and eventual de-
cline, to its reprieve, renewal, and resurgence through its association with people. The
dominant theme in the biography of ginkgo is survival, and its resilient life story offers
hope for other botanical biographies that are still being written.
 2
Trees

The best time to plant a tree was twenty years ago. The second best time is now.
—Chinese proverb

Among my clearest memories as a student beginning in botany are the giant Atlan-
tic cedars that dominated the center of campus at the University of Reading. They were
part of my life for nearly a decade, and still today every time I see one of these majes-
tic trees, with their sparse flattened branching and truncated crowns, it takes my mind
back to what now seems like a different life. Together with the giant sequoias scattered
across campus, and the gnarled strawberry tree near Whiteknights House, the Atlantic
cedars were planted at the end of the eighteenth century by the Marquess of Blandford,
an ancestor of Winston Churchill. They were his living legacy to thousands of students
who came two hundred years later.1
From the summer of 1974, when I was studying at Kew, it is the magnolias and mon-
key puzzles that stick in my mind. A vivid memory from my time as a new lecturer at
Reading in the late 1970s is of a classically proportioned English oak, long since swept
away, which grew near the Plant Science Laboratories. When I was newly arrived in
Chicago in the early 1980s, the massive American beeches and sugar maples at Warren
Woods, Michigan, a rare tract of midwestern forest spared by the loggers, made an in-

8
 9
trees

delible impression. And everyone in our family remembers the statuesque stone pine
that grew by our garden gate when we lived at Kew; it has watched all fifteen directors
come and go. These encounters with trees punctuate the chapters of my life. They are
the touchstones to many other memories and connect me to the lives of people from
times long before I was born.2
I have been fortunate; a life lived studying plants has given me the chance to inter-
view some of the most spectacular trees on the planet. In 1975 I explored forests of
Atlantic cedars in the Middle Atlas Mountains of Morocco and, in 1981, my first sum-
mer in the United States, I visited the coast redwoods just a few miles north of San
Francisco. It is not hard to see how such miracles of nature, with their cousins the
giant sequoias, galvanized the passion of John Muir and have stirred the hearts of many
others.3
At Humboldt Redwoods State Park, two hundred miles up the coast from San Fran-
cisco, more than a hundred coast redwoods reach 350 feet or more. Few trees can
match them for sheer size, but on a family holiday in Australia in the early 1990s, we
visited equally immense swamp gums in the temperate rain forest of Tasmania. Sev-
eral of those monumental eucalypts top out at more than 330 feet. Thomas Paken-
ham’s spectacular photographs go a long way toward summoning up their grandeur,
but there is no substitute for seeing these miracles of nature close up; you need to smell
them, feel them, and be humbled by the cathedral-like spaces they create.4
For many of us, trees of all kinds, not just the largest and most spectacular, often
take on special meaning. My affinity for the grove of ginkgo trees where we walked with
our baby daughter, and for the black tupelo planted in our garden around the time our
son was born, could not be more different from my feelings for the bricks and concrete
that otherwise define the places where we have lived. Such connections probably go
back deep into our evolutionary past. Our bodies reflect the arboreal lives of our an-
cestors, and in a world without skyscrapers it was trees, like mountains, that reached
up into the sky. It was their roots, like caves, that penetrated deep into the Earth. Trees
are part of us, and they connect heaven and the underworld with primeval power. Our
urge to climb them may not be so different from our urge to climb mountains.5
Until recently my own tree climbing expertise had been pretty much confined to
the small lilac that grew behind my father’s garden shed fifty years ago. Because I have
a tendency to vertigo, scaling tall trees is something I have mainly avoided. But that
 10
prologue

phobia was temporarily banished early one morning in the rain forest of northeastern
Borneo. The promise of sightings of tropical birds, combined with just a hint of peer
pressure, coaxed me onto a ladder strapped to the trunk of an enormous koompassia.
It took me 120 feet, straight up, to the roof of some of Sabah’s last remaining virgin
lowland rain forest. It is hard to grasp the scale of such a giant until you have inched
your way up its trunk and been enveloped by its canopy.6
Trees have been adopted as important symbols that are firmly entwined in human
culture. In Genesis we learn that “out of the ground made the Lord God to grow every
tree that is pleasant to the sight, and good for food; the tree of life also in the midst
of the garden.” It was the fruit of the “tree of the knowledge of good and evil” that
Adam and Eve struggled to resist. The ancient Norse saw their world organized around
Yggdrasil, the World Ash.7
Different cultures in different parts of the world have also been attracted to simi-
lar kinds of trees. Among figs, the banyan and peepal have special significance in the
Hindu faith, and Buddha found enlightenment under one special peepal, the Bodhi
tree. The Mediterranean fig was the tree of life in ancient Egypt, and five species of
fig are at the center of the Afro-Brazilian religion of Candomblé, part of the legacy
of slavery in Brazil. These fig trees have special meaning in the terreiros, the houses
of worship.8
Trees are also living links to our cultural history. Transplanted people carried the
seeds of baobabs from Africa to the New World. Brazil takes its name from Pau Brasil,
the once common tree of the Atlantic rain forest that produced a rich red dye much
prized by early Portuguese traders. The cedars of Lebanon, mentioned many times in
the Bible, are a symbol of nationhood and appear on the Lebanese flag. The palmetto
has similar significance in the state of South Carolina. The bearded fig is part of the
national identity of Barbados, and in China there have been efforts to have ginkgo
adopted officially as the national tree.9
We also connect trees with key people and important moments in history. In Britain
the Royal Oak at Boscobel is a descendant of the tree in which King Charles II report-
edly hid after defeat at the battle of Worcester. At Hatfield House, an English oak marks
the spot where Queen Elizabeth I is said to have learned of her ascent to the throne.
In Sherwood Forest, more than half a million visitors every year flock to see the Major
Oak, the massive tree linked to the legend of Robin Hood.
 11
trees

A stylized ginkgo leaf, carved into the planks of a wooden bridge and painted yellow,
marks the way to the great ginkgo at the Yongmunsa Temple, South Korea.

Oaks have assumed cultural importance on the other side of the Atlantic, too. In
Austin, Texas, the Treaty Oak is the sole survivor of the fourteen Council Oaks, a sacred
meeting place for Native Americans weighing matters of war and peace. The Emanci-
pation Oak at Hampton, Virginia, marks the site of the first southern reading of Lin-
coln’s Emancipation Proclamation. In Chico, California, the Hooker Oak, named for
Joseph Dalton Hooker, one of my predecessors at Kew, is featured in the Errol Flynn
version of the Robin Hood legend. Renamed the Gallows Oak, it is the tree under
whose shade the “merry band” first come together.10
There are many such legends related to the great ginkgo trees of eastern Asia. An
hour and a half drive east of Seoul, South Korea, the massive ginkgo at the Yongmunsa
Temple is one of the largest and most visited in the world. Established late in the first
millennium A.D. in the foothills of Mount Yongmun, the temple is approached by a
 12
prologue

gentle walk uphill through the tranquil mountain forest. As the temple comes into
view, the colossal ginkgo is perfectly placed on a broad plinth just above a rushing
mountain stream. It is approached from below and leaves a strong impression.11
Legends about the origin of the Yongmunsa Ginkgo emphasize its place in Korean
history. According to one, it grew from the staff of Ūisang, the great Buddhist priest
of the Silla Dynasty. Another has it planted in sorrow by Maui, the crown prince of
Gyeongsun, the last Silla king, as he lamented the fall of his nation. It hardly matters
whether either account is true; more important is that this great tree is of deep signifi-
cance to the Yongmunsa Temple and its monks, and has special meaning for the people
of Korea.12
Cultural prominence is just one sign of the importance of the roughly 100,000 dif-
ferent kinds of trees on our planet. From the earliest civilizations trees have been used
for fuel, timber, and food, as well as to make objects of all kinds. They also provide rub-
ber, oils, and medicines, and are indispensable in the day-to-day livelihoods of poor
people in many parts of the world. At Kew we worked on tree conservation with the
Forestry Department in Burkina Faso. Every year Burkina Faso loses tens of thousands
of acres of its scrubby forest to agriculture and the cutting of trees for fuel. At the same
time, its population depends on fuel wood for 90 percent of their household energy.
Understanding how people use trees, and how they can continue to use them sustain-
ably, has practical value.13
Trees are also crucially important in the developed world. The average American
uses a ton of wood every year, equivalent to about forty-three cubic feet of lumber. It
gets turned into everything from baseball bats and baby cribs to xylophones and yo-
yos. A huge amount of wood is also used in buildings, and trees are the main source
of pulp for making card and paper. America consumes around ninety million tons of
paper products every year.14
The OneTree Project took a symbolic approach to try to capture our debt to trees.
An ailing 170-year-old English oak about sixty feet tall was felled in Tatton Park, Chesh-
ire, England, and was turned over to more than seventy designers, artists, and artisans.
Sawdust was burned to make a glaze for pottery, bark was used for tanning, and wood
fibers were used to make paper. The wood itself was turned into an astonishing range
of objects: toys, sculpture, furniture, window frames, ladders, yurts, bowls, and even a
fetal stethoscope.15
Trees are also crucial participants in the ecological processes that determine the
The great ginkgo at the Yongmunsa Temple, South Korea.
 14
prologue

environment on our planet. The fossilized remains of ancient trees are the main com-
ponent of coal, and the carbon of which they were made is an important source of the
carbon dioxide now accumulating in our atmosphere. Forests also affect local and re-
gional climate, and regulate the quantity and quality of water in streams and rivers.
Both above- and belowground, trees help structure different kinds of vegetation and
the communities of animals and microbes that depend on them.
Yet despite the importance of trees, we often remain curiously ambivalent about
them and their future. Trees are easy to take for granted. They seem ubiquitous, and
when they stand in the way of something we really want—a site for a new parking lot,
perhaps—they are easily pushed aside. Bill Vaughan, a columnist for the Kansas City
Star, summed it up with a memorable line: “Suburbia is where the developer bulldozes
out the trees and then names the streets after them.”16
Nevertheless, threats to particular trees can bring out strong emotions. The Wash-
ington Oak in New Jersey survived two battles of Princeton: the first in 1777, when
George Washington rallied his troops to defeat the British; the second in the 1980s,
when it was threatened by a local developer. The citizens of Charleston annexed the
massive Angel Oak to save it from similar threats. Through 2007 and much of 2008
the University of California, Berkeley, became the site of the longest running “urban
tree sit,” a protest to prevent the removal of ninety trees in the campus’s Oak Grove to
make way for a new athletic facility. Despite a lengthy lawsuit, numerous appeals, and,
at one point, a hundred nude protesters perched in the boughs of the threatened oaks,
the Alameda County Superior Court eventually ruled in favor of athletics.17
In the same way, controversies over the logging of old-growth forest or expanding
agriculture into the rain forest can attract national and international political atten-
tion. U.S. President Bill Clinton must have known that he had crafted a fine compro-
mise when neither loggers nor conservationists were happy with his plan for the man-
agement of old-growth forest in the Pacific Northwest. More recently, trees and forests
have featured with increasing prominence in discussions of climate change policy.18
However, as always, it is the immediate and the local that get most of the attention;
longer-term consequences and connections to what is happening elsewhere are too
easily overlooked. Trees, and the ecological systems of which they are part, are assailed
all over the world, all the time. The threats come from all directions; some are direct
and rapid, others are indirect and insidious. Land conversion that removes forest and
replaces it with agriculture, including tree monocultures, such as oil palm or rubber
 15
trees

plantations, is one threat; urbanization is another. Overexploitation for fuel or timber

is also a problem. For commercially valuable trees, such as mahogany and teak, selec-
tive, and sometimes illegal, logging—the botanical equivalent of hunting whales or
poaching tigers—is a persistent threat.
Trees are also caught up in broader environmental changes, such as water or air pol-
lution, as well as invasions of species, including diseases, from elsewhere. They are also
in trouble from changes in soils, water, drainage, and the frequency of fire; rapid cli-
mate change and extreme climate events pose an increasing challenge. The stately life
cycles of most trees are not well suited to the frenetic world that we are creating. The
time it takes for a seed to grow into a mature tree, combined with fragmented land-
scapes that hamper the natural movement of plants and animals, makes it difficult for
new generations to adapt or migrate.19
A large part of what attracts us to trees is their timelessness. Compared to us, and to
much else in our modern world, trees have comforting longevity. They change slowly,
almost imperceptibly. They are beacons of stability that can cross generations. To quote
a well-known proverb, “A society grows great when old men plant trees whose shade
they know that they shall never sit in.” Ecologically speaking, trees find a place to live
and put down roots; in the parlance of ecology they are site occupiers. They arrive,
courtesy of the wind, or perhaps a bird or squirrel, and there they stay. In a densely
populated world filled by restless people with ever escalating demands, such a ponder-
ous lifestyle may not be a winning strategy. Our habits disturb the world on a massive
scale. We create environments that favor weeds: plants that live fast, reproduce early,
and die young. The soothing calm of long-lived trees is easily lost in the turmoil.
With its resilience and longevity ginkgo epitomizes much of what we admire about
trees, and its strong cultural associations underline the strength of that bond. But the
biography of ginkgo also has the dimension of deep time; the link that it provides to
worlds gone by. Many of our familiar trees—ash, hornbeam, magnolia, oak, or wal-
nut—have prehistories measured in millions or even tens of millions of years, but
ginkgo is extreme; it has been ever present for 200 million years or more. And like the
dawn redwood of China and the Wollemi pine of Australia, two other charismatic trees
with biographies that stretch far into the past, ginkgo also has a further attraction; it
very nearly went extinct. Ginkgo is a good news story: a tree that people saved.
 3
Identity

. . . just plant for me a funny looking ginkgo.

—LuEsther Mertz, commemorative plaque, New York Botanical Garden

William Jackson Hooker, the first director of the Royal Botanic Gardens, Kew, was
born in 1785. By then the ginkgo that stands in the historic heart of Kew Gardens was
becoming mature. Half a century later, when Hooker came to Kew to create a na-
tional botanic garden, it was nearly a hundred years old; it had already outlived King
George III, Sir Joseph Banks, and the others from the eighteenth century who had pre-
sided over its planting. Hooker would have seen this tree nearly every day and he would
have recognized it as one of the more unusual and important in his care, just as it was
for me 150 years later.1
William Hooker was the right man, in the right place, at the right time. The son of a
brewer who became professor of botany at the University of Glasgow, he was a superb
artist, gifted scientist, and remarkable visionary. He was tall and thin, with bound-
less energy, as well as patience, tact, and charm; his manners were “easy and urbane.”
Hooker was also an industrious and effective administrator: “He deemed nothing too
small for his notice.”2
Hooker came to Kew in the mid-nineteenth century to take charge of a small part

16
 17
identity

of a country estate in decline. In the eighteenth century Kew had been a royal favorite,
with the landscape of Richmond Gardens created by Capability Brown, and the adja-
cent estate of Frederick and Augusta, Prince and Princess of Wales, studded with mag-
nificent buildings by Sir William Chambers. However, with the death of George III and
also Sir Joseph Banks in 1820, Kew and its collection of living plants fell into decline.3
Appointed director in 1841, William Hooker set Kew on a new trajectory and brought
it back to life. In a little more than two decades he built astonishing greenhouses,
planted grand vistas, unified the eighteenth-century landscapes, and introduced exotic
plants from all around the world. Behind the scenes, using his personal collection as
the core, he began the development of the library and preserved plant collections that
are the foundation of Kew’s global work in plant conservation and science. By the time
Hooker died, through his energy and political genius, he had greatly expanded the
parts of the estate under his control and created a magnificent spectacle for the public.4
Hooker was a scientist, like most directors of Kew have been, but he was also a
teacher. He understood that the way to interest students and others in plants is to show
how they are useful to people. At Kew the collections of useful plants that he used to
illustrate his lectures in Glasgow became the nucleus of what he called the Museum of
Economic Botany, and for the national botanic garden this interest had another dimen-
sion. Hooker knew that the uses of plants were important to his political masters and
to the commercial interests of Britain’s Victorian Empire.5
With more than seventy-six thousand specimens, the Economic Botany collection
at Kew is now the largest of its kind in the world. Tucked away in an air-conditioned
vault, it is a vast miscellany of food plants, medicinal plants, dye plants, and timbers.
There are endless artifacts made from different plant parts: fish traps and a dugout
canoe alongside beautiful necklaces and exquisite fabrics. There is a shirt made from
pineapple fibers and a bowler hat made out of cork. Among the spices, the psycho-
active plants, the tools, the musical instruments, and the many other samples that illus-
trate the interdependence of plants and people, there are some that document the uses
of ginkgo.
There are multiple accessions of ginkgo seeds. In China, ginkgo has been cultivated
for a thousand years for its edible nuts. There are also samples of ginkgo wood, col-
lected by John Quin, one of the first British diplomats in Japan. Between 1867 and 1896,
at the instigation of Kew, he made meticulous observations of the ancient Japanese art
 18
prologue

of creating lacquerware. He recorded how the work was done, and he collected the ma-
terials and tools that were used. Among them is a piece of ginkgo wood. Quin lists it as
used for “such articles as are turned in a lathe, as bowls, rice cups, round trays, etc.”6
Among the other delights of the Economic Botany collection is a single dried ginkgo
leaf acquired not long ago by a Kew botanist visiting China. It is decorated with a
scene from a Chinese garden drawn in fine black lines with an accompanying poem in
minute Chinese characters. To the left a tree, perhaps a pine, stands in the foreground.
In the center is a group of rocks around a small pavilion. Behind, and off in the dis-
tance, are more trees, some of them perhaps willows. On the right a red stamp bears
the name Suzhou in old Chinese characters from the Qin Dynasty.7
To those familiar with Chinese gardens the scene on the ginkgo leaf is well known.
It is from one of the most famous gardens in Suzhou, the Chinese City of Gardens.
It might also be familiar to those who know New York; it is partly re-created in the
Metropolitan Museum of Art. Wang Shi Yuan, the Master of the Nets Garden, is one
of the best examples of a small late-eighteenth-century Chinese private garden. Estab-
lished by a retired bureaucrat on the site of a twelfth- century garden from the southern
Song Dynasty, it is one of nine classical Chinese gardens in Suzhou that are recognized
on the UNESCO list of World Heritage Sites. Several contain ancient ginkgo trees, as
does the oldest of all botanic gardens at the University of Padua in northern Italy.8
The unique fan-shaped leaf has given ginkgo a strong presence all over the world.
The seeds probably first attracted people to this tree, and perhaps saved it from extinc-
tion, but its distinctive leaf has played a large part in transforming ginkgo from a minor
food plant into cultural icon. Ginkgo has the most memorable leaves of any tree, and
its unusual form lies behind much of ginkgo’s rich cultural history.9
The ginkgo leaf has been taken up repeatedly as an instantly recognizable motif.
Ginkgo is native to China and has its longest history of cultivation there. Guo Moruo,
a Chinese revolutionary, a contemporary of Mao, and the first president of the mod-
ern Chinese Academy of Sciences, wrote many poems about plants and flowers, but he
reserved his highest praise for ginkgo. In the depth of the communist struggle against
Chiang Kai-shek he called ginkgo “the Holy One of the East,” a worthy and resounding
symbol of Chinese nationalism.
In South Korea, many famous old ginkgo trees are preserved as natural monuments,
including the ginkgo in Nongso, Seonsan, which villagers honor on the fifteenth day
 19
identity

The ginkgo motif on the gate of the

Sung Kyun Kwan University, which
incorporates the Munmyo Confucian
Shrine, in Seoul, South Korea.

of the lunar New Year. Nearly a hundred feet tall and sixteen feet in diameter, it is im-
posing even in its winter nakedness. It was planted more than four hundred years ago
near a temple and marketplace that are now just a ruin. According to local legend it is
so sacred that birds will not land on it.10
Ginkgo is also deeply embedded in Japanese culture. The ubiquitous T that serves
as the symbol for the prefecture of Tokyo, home to thirteen million people, looks sus-
piciously like a stylized ginkgo leaf. The best sumo wrestlers have their hair tied at the
top in shapes that take the ginkgo name; and when in 1958 two Japanese scientists de-
scribed a new species of beaked whale from the warm waters of the Pacific and Indian
Oceans, they named it Mesoplodon ginkgodens, the “ginkgo-toothed beaked whale.” In
Japan there are also ginkgo crabs, ginkgo mushrooms, ginkgo potatoes, and ginkgo
sharks. Not to mention cupboards, tables, flower vases, farm tools, and musical instru-
ments that all incorporate ginkgo as a part of their name.11
Around the world, governments, businesses, and organizations of all kinds have
taken the ginkgo leaf as part of their identity; Zhejiang Forestry University in China,
Osaka University in Japan, and Sung Kyun Kwan University in South Korea all have
the ginkgo leaf as part of their logo. In the West, when the artist Larry Kirkland was
asked to design the entrance to the new building of the National Academy of Sciences
in Washington, D.C., and to convey the development of human knowledge of the natu-
ral world, he included a sprig of ginkgo leaves and their seeds. Along with Darwin’s
 20
prologue

finches, Mendel’s pea pods, and Morgan’s fruit flies, ginkgo was among the nine images
Kirkland chose to superimpose on the molecular structure of DNA. Appropriately for
researchers working close to the site of the old Cavendish Laboratory, where the struc-
ture of DNA was first worked out, the Department of Plant Sciences at Cambridge Uni-
versity chose a similar juxtaposition; its logo surrounds a ginkgo leaf with a stylized
double helix of DNA.12
The ginkgo identity also extends beyond academia into commerce. You can visit a
Ginkgo café in Melbourne, Australia, or Frankfurt, Germany, and many places in be-
tween. Around the world there are chic Ginkgo spas and Ginkgo restaurants. Almost
all advertise themselves by flourishing the distinctive ginkgo leaf. Marketers and brand
developers link the word and the leaf with an image that is contemporary but timeless,
exotic yet familiar, and, most of all, sublimely elegant.
In the Chicago suburb of Oak Park, a large ginkgo stands outside the former home
and studio of Frank Lloyd Wright. It was probably a young tree when the architect
bought the land. He could easily have cast it aside, but instead he worked around it
as he extended his home. Certainly he would have seen ginkgo on his visits to Japan.
Today, the tens of thousands of visitors each year who come to this shrine of twentieth-
century architecture pass under his great ginkgo and buy their tickets at the Ginkgo
Bookshop. Inside there is ginkgo merchandise of all kinds, from ginkgo dishes to
ginkgo jewelry boxes. An even bigger selection can be found through a few keystrokes
on the Web. The variety of ginkgo products is almost overwhelming, and they continue
a long tradition. For hundreds of years artists and artisans of all kinds, in China, Japan,
and Korea, have incorporated ginkgo into their work.13
The connection between ginkgo and Frank Lloyd Wright seems a natural one. The
influence of the Arts and Crafts movement is especially strong in many contemporary
products that incorporate the ginkgo leaf motif. Its elegance and clean curves connect
easily to an aesthetic that began as a reaction against the machine. Ginkgo was also
taken up in Art Nouveau; there are spectacular renderings of ginkgo twigs and leaves
in the Art Nouveau architecture of Nancy and Prague.14
Ginkgo also remains a source of artistic inspiration. In 2005, in the British Pavilion
at the Fifty-first Venice Biennale, Gilbert and George, artists from the East End of Lon-
don, showed their Ginkgo Pictures: a striking exhibition of twenty-five photo images
created from ginkgo leaves collected in Gramercy Park, New York. Brightly colored,
framed in a black grid, and arranged with portraits of Gilbert and George in repeated,
 21
identity

often symmetrical, patterns, each of the fourteen-foot-high panels is a kind of surreal-

istic stained glass window.15
Ginkgo has made its deepest cultural mark in China, Japan, and Korea, where the
most ancient of all ginkgo trees are beloved. Two large ginkgos stand on either side of
the Temple of the Reclining Buddha in Beijing, in what is now the Beijing Botanic Gar-
den. There are ginkgos at the Confucius Temple in QuFu: just as the Buddha sat be-
neath the Bodhi tree, legend has it that Confucius spent time reading, reflecting, and
teaching beneath a ginkgo tree. A massive old ginkgo in Shanxi Province is said to have
been planted by Lao Tzu, according to legend the founder of Daoism.16
In Korea many of the largest and oldest ginkgos are associated with Buddhist
temples, but there are two large ginkgos at the Munmyo Confucian Shrine on the cam-
pus of Sung Kyun Kwan University in Seoul. My son and I found them there one hot
July day a few years ago. They were in good health, planted symmetrically in the oldest
courtyard of the university. They had stood there for hundreds of years, through good
times and bad.
In Japan there are large ginkgos at some of the country’s most important Buddhist
temples, including the Tamba Kokubunji Temple in Kyoto Prefecture, the Jonichiji
Temple in Toyama Prefecture, the Hida-Kokubunji in Gifu Prefecture, and the Zen-
pukuji Temple in Tokyo. There are also ginkgos at some of the most important Shinto
shrines: the Ubagami Shrine in Miyagi Prefecture, the Katsushika Hachiman Shrine
in Chiba Prefecture, and many others, including the controversial Yasukuni Shrine in
Tokyo.17
In the West, ginkgo is widely used as a commemorative tree. There are several, with
their memorial plaques, in the park close to my home. In Hoboken, New Jersey, just
across the Hudson from where the twin towers of the World Trade Center once stood, a
grove of ginkgo trees provides a living memorial to the victims of the 9/11 terror attack.
In Detroit there is a ginkgo planted by Yoko Ono, and in Caen another planted by the
Dalai Lama. At Kew in 1916, when Sir William Chamber’s eighteenth-century Temple
of the Sun was crushed by a massive cedar of Lebanon that came down in a storm,
Queen Mary marked the site by planting a ginkgo just a few feet from the Old Lion
placed there about 150 years earlier, in the time of her husband’s great-great-great-
grandmother. In Morgantown, Indiana, there is a single large ginkgo planted around
1880, soon after the end of the American Civil War. It commemorates Union prisoners
who survived the notorious Andersonville Prison of the Confederacy.18
 22
prologue

Ginkgo also turns up in celebrated locations around the world. There are ginkgos
on the grounds of the White House in Washington, D.C., and the Imperial Palace in
Tokyo, as well as in Tiananmen Square in Beijing and at the Alamo in San Antonio. In
Ottawa, on the grounds of the residence of the governor general, a ginkgo commemo-
rates the 1985 visit of Chinese President Li Xiannian. Most famous is the ginkgo at
Hiroshima that survived the blast of the first atomic bomb on August 6, 1945. It became
a symbol of endurance in the midst of great destruction and human suffering. On the
other side of the world, there are ginkgos in Missouri near the home of President Harry
Truman, who ordered the bomb to be dropped, and at the University of Chicago there
are ginkgos on Ellis Avenue, a stone’s throw from where Enrico Fermi and his team
ushered in the nuclear age.19
Ginkgo has also become an icon of singularity. Joy Morton, who created the Morton
Salt Company and founded the Morton Arboretum, just west of Chicago, is reputed to
have said: “The Morton Arboretum is a ginkgo and a ginkgo it shall remain.” His vision
was of an organization that was unique and memorable. The Morton Arboretum has
a Ginkgo Way, and its Ginkgo Restaurant is decorated with ginkgo-inspired furniture.
More than seventy ginkgo trees from forty different sources are planted in its beautiful
wooded landscape.20
Joy Morton was right: the inverted deltoid shape and distinctive fan of fine veins
distinguish the leaf of ginkgo from those of every other plant. Engelbert Kaempfer, the
first westerner to take note of the tree during his time in Japan at the end of the seven-
teenth century, saw similarities with leaflets of the maidenhair fern. From Kaempfer’s
comparison comes one of the English common names of ginkgo, the maidenhair tree.
But while the leaves of the maidenhair fern do have a hint of the ginkgoesque, there is
nothing with which the leaf of a real ginkgo is easily confused.
The Swedish naturalist Linnaeus, who sought to name and catalogue all the world’s
plants in the eighteenth century, gave ginkgo its formal scientific name. He took it from
the name that Kaempfer transliterated from the Japanese. Despite the awkward combi-
nation of consonants, Linnaeus was evidently content; he simply added the epithet bi-
loba, referring to a characteristic feature of some ginkgo leaves: a blade that is notched
along its leading edge, or sometimes deeply divided into two.
Only a few decades later Goethe took Linnaeus’s designation, Ginkgo biloba, for his
famous poem in the West-East Divan. In the middle stanza he asks:
 23
identity

Is it one living being?

That divides itself into itself
Are there two who have chosen each other,
So that they are known as one?

Goethe turned to ginkgo to express his feelings for Marianne Willemer, his muse
and the young wife of a close friend, but he also knew that his question had deeper
significance. He struggled not only with his attraction to Marianne but also to find
meaning behind the structure of plants. The scientific study of plant form begins with
Goethe. He was the first to use the term morphology, as the scientific study of biological
form is now called.21
Modern science provides no clear answer to Goethe’s question about the ginkgo
leaf, but his broader point has been taken up. The overwhelming variety of plant life,
of which ginkgo is a key part, still challenges us to search for a fundamental organiza-
tional theme, a Bauplan, through which it all can be linked. Goethe once said: “From
top to bottom a plant is all leaf.” How leaves arose, and how they have been modified
over thousands of millennia, is the key to much that we would like to understand about
the evolution of plants.22
This page intentionally left blank
 part ii

The Living Tree

OVERLEAF Extensive stalactite-like zhōng-rǔ, or chi-chi, on the
Kitakanegasawa ginkgo, northern Honshu, Japan.
 4
Energy

All growth depends upon activity. There is no development physically

or intellectually without effort, and effort means work.
—Calvin Coolidge, “Have Faith in Massachusetts”

The elegance of a ginkgo leaf begins with its stalk; it is long, sometimes a little longer
than seems quite right for the length of the blade, but the two flow easily together. As
the blade expands, the two nerves, which pass concealed through the leaf stalk, emerge
as fine veins: each supplies one half of the leaf. The veins divide and diverge relentlessly
within the blade; only rarely is the pattern disturbed by veins that rejoin.1
Leaves are of vital importance to a living ginkgo tree; they provide energy indepen-
dence. Leaves are clean- energy factories; natural, mass-produced solar panels, each
packed with sophisticated biochemical machinery capable of transforming the sun’s
energy into chemical energy that plants and animals can use. This miracle of natural al-
chemy, the process of photosynthesis, has been probed by some of the world’s greatest
scientific minds. We understand a great deal about how it works, down to the level of
molecules and atoms, but much still remains elusive. My Yale colleague Gary Brudvig
investigates the subatomic transfers of energy that lie at the heart of photosynthesis.
Yet this process, first accomplished by simple organisms more than two billion years

27
 28
the living tree


 

Scattered stomata, the breathing pores on the underside of a ginkgo leaf, seen from the outside, showing
the projections from nearby cells that arch over the entrance. Carbon dioxide enters the leaf through
the stomata, and the density of stomata on fossil ginkgo leaves has been used to estimate the amount
of carbon dioxide in the Earth’s atmosphere in the distant past: more stomata are needed to sustain the
same supply of carbon dioxide when the concentration of carbon dioxide in the atmosphere is low.

ago, is one we still struggle to re-create in even the world’s most advanced chemistry
laboratories.2
Leaves are where the chemical building blocks that make and sustain a ginkgo tree
are created, and they are assembled from the simplest of materials. Water comes from
the soil, drawn up the trunk from the roots, and brings with it key nutrients, especially
nitrogen and phosphorous. It is distributed across each fan-shaped leaf by the diverg-
ing and dichotomizing veins. Carbon dioxide comes from the air, taken into the leaf
through breathing pores, the stomata, adjustable valves that are concentrated on the
lower leaf surface. The carbon that it carries is combined with hydrogen from water to
make sugars. These sugars, and the energy they contain, can be converted into many
different kinds of carbohydrates and can combine and interact with other molecules
to make proteins and the great variety of chemicals needed for life. Oxygen, essential
for us, is a mere by-product.3
This basic function of leaves is fundamental to all plants. The ability to harvest light
and to use it to create chemical energy is the ultimate accomplishment in green chem-
 29
energy

istry, and it could scarcely be more important. Ancient photosynthesis created the re-
serves of coal and oil on which much of modern society is based. Today photosynthesis
builds forests and grasslands. Notwithstanding the other sources of energy available
to us—nuclear energy, wind energy, geothermal energy, and others—photosynthesis
remains the energetic foundation on which most ecosystems, all of agriculture, and
almost all of civilization rest. As ancient people recognized more clearly than we do
today, we all depend on the sun; photosynthesis is what sustains us and most other
living organisms on our planet.4
Over the short term most of the carbon-containing molecules produced by photo-
synthesis are transformed back into carbon dioxide and water as they react with oxy-
gen. This happens in our own bodies, and the bodies of most animals and plants,
as energy in carbohydrates and other compounds is used for a myriad of life pro-
cesses. The results of photosynthesis are also reversed over geologic time as tempo-
rarily buried carbon-containing molecules, produced by once living organisms, come
to the surface and react with oxygen in the air. However, if those carbon compounds
are separated from contact with oxygen, literally taken out of the equation—as, for ex-
ample, when they are buried and deeply locked away in oceans and swamps—then the
oxygen left behind builds up.
Two billion years ago, oxygen produced by ancient photosynthesizing bacteria accu-
mulated in this way, to such a point that it changed the fundamental conditions for
life. Poisonous to the earliest microbes, oxygen now became essential and channeled
evolution in new directions. It created opportunities for new kinds of organisms, and
it allowed new kinds of biological processes to emerge. The long-term impact of photo-
synthesis on our world has been vast. It was one of the key contingent steps in the his-
tory of life to which the origin of ginkgo, and eventually the origin of our own species,
can be traced.5
The magic of photosynthesis resides within chlorophyll, the molecule that makes
plants green. Chlorophyll molecules are packaged together inside tiny green light-
gathering discs, chloroplasts, that are vital to the internal workings of most of the mil-
lions of plant cells that make up a ginkgo leaf. Each cell may contain up to a hundred
chloroplasts; there are hundreds of millions in a single leaf. Inside each chloroplast,
chlorophyll molecules and their complex associated biochemistry are embedded in
membranes less than one–five millionth of an inch thick.6
The clever trick of the chlorophyll molecule, with help from other light-gathering
 30
the living tree

pigments, is to release an electron for every photon of light it absorbs. That electron is
passed through complex intermediaries in less than a thousandth of a second. It pro-
vides the energy that drives the slower process by which carbon dioxide is captured
from the air, combined with hydrogen, and transformed into simple sugars.7
Water is a basic raw material for photosynthesis; the chemical breaking apart of
water molecules provides the hydrogen and replenishes the electrons lost by chloro-
phyll molecules in the light-harvesting step. Water is also indispensable to many other
essential life processes. It was readily available to the earliest plants that lived in the
sea, lakes, ponds, or rivers, but as evolution created opportunities to move on to land,
it also presented a fundamental challenge: the reality of water in short supply. In large
part, the success of plants on land, and everything that has followed from it, including
ourselves, has been a triumph of biological water management.
A fresh ginkgo leaf is stiff, full of water; the leaf stalk is wiry. However, plucked from
the tree, it soon changes character. The leaf blade goes limp. What was once crisp and
firm becomes soft, almost membranous; it is the pressure of water in countless indi-
vidual cells that keeps the leaf stiff. When water is lost and not replenished, the leaf
wilts. For plants, as for animals, a shortage of water can be fatal; to fulfill their energy-
generating and other functions, leaves need to stay hydrated.
In the leaves of most trees big veins supply smaller ones in a hierarchy, and the veins
are connected into a network as they rejoin. Such reticulation builds in redundancy,
a safeguard against damage, and differences in vein sizes allow a single large vein to
supply many smaller ones. However, ginkgo does it differently; the architecture of its
water supply is unique among the world’s trees. The hierarchy of veins is much less pro-
nounced, and all can be traced back to the two slender veins that enter the leaf base
from the leaf stalk. Only one leaf in ten shows any fusion among the veins, and such
reticulations are always sparse. It is an idiosyncratic system, a water supply more remi-
niscent of drip irrigation than a hosepipe; the water leaks from the elongated cells that
make up the veins into the tissues between.8
Keeping the water content of leaves just right is a challenge at the best of times; it is
still more difficult when the soil dries out and water is scarce. Water evaporates from
the leaves of plants and is also lost through the breathing pores. When water is plentiful
and temperatures are warm, this is a good thing; it is the botanical equivalent of sweat-
ing, one way by which the leaves stay cool. Water loss from the leaves also draws water
up the trunk from the roots and distributes it to every branch, every branchlet, every
 31
energy

twig, and every leaf in the crown. But keeping the whole process in balance is crucial;
if too much water is lost and not replenished, then the leaves suffer, and soon the tree
will be in trouble.
In most plants that live on land, the difficulty of staying hydrated is compounded
by the fact that when the breathing pores open to take in carbon dioxide, they also lose
water. Driven by this awkward physiological reality, the upper leaf surface in ginkgo,
and most trees, has a well-developed waterproof covering, the cuticle, which reduces
water loss. Breathing pores are present only on the lower leaf surface, away from the
direct heat and glare of the sun, and they close when water is in short supply.
In mature ginkgo leaves the upper surface is smooth and dark green; the cuticle is
tough, colorless, and almost impermeable. The epidermis, the outermost layer of cells
beneath the upper leaf cuticle, also lacks color, and sunlight streams straight through
to the chlorophyll-rich cells below. Of all the cells in a living ginkgo leaf, these are the
true energy factories: the cells supplied most generously with chloroplasts.
The lower leaf surface is different; its gray-green hue comes from a fine layer of
wax over a much thinner waterproof covering. The covering itself is perforated at the
breathing pores, each of which connects to a well-developed system of air spaces inside
the leaf. This makes it possible for carbon dioxide and water vapor to diffuse rapidly
among the cells that make up the leaf. These are the messy but critical conduits through
which carbon dioxide is supplied for photosynthesis.
Mature ginkgo leaves are tough and resistant to decay, more so than the leaves of
many other trees. The cuticle around the outside, together with strands of mucilage-
like resin inside the leaf between the veins, makes them slow to decompose. Ginkgo
leaves are among the last to succumb to the compost heap; their durability is one
reason why they make such good fossils. Across Japan, the more than half a million
ginkgos planted as street trees produce an overabundance of leaves in the autumn.
In Tokyo, ginkgos make up about 12 percent of all street trees, and their leaves break
down so slowly that for a time a research team for the city was working on how best to
turn them into useful compost.9
As far as I know, no one has ever counted how many leaves there are on a large
ginkgo tree, but even on a small tree it is a very big number. During his Ph.D. studies
at Yale, Kirk Johnson, now director of the U.S. National Museum of Natural History,
cut down a fifty-foot-tall red maple to count the leaves. He was interested in the total
number of leaves produced in a typical patch of forest. Kirk’s red maple was tall but
 32
the living tree

slender, with a trunk just eleven inches in diameter at chest height. It was not what
most people would call a very big tree, but it still took him and a friend eight hours to
count all the leaves. The final total was 99,284. A much bigger ginkgo, perhaps twice as
tall and more profusely branched, might have 300,000 to 500,000 leaves. An old and
truly massive ginkgo, like some of those in China, Japan, or Korea, might bear close to
a million.
Every leaf on a ginkgo tree is a triumph of modular design built through the un-
failing translation of complicated instructions that are written in code in strands of
ginkgo DNA. We have come a long way in learning how to read the language of DNA,
but how it is translated to make a leaf through the carefully choreographed dance of
a multitude of molecules still eludes us. Nevertheless, each leaf is built in a matter
of months, according to complex specifications and with supremely high standards
of quality control. Just as astonishing, after all that effort, is that every leaf is shed as
winter approaches. The creation and disposal of hundreds of thousands of leaves every
year is a testament to the power of photosynthesis, and to the productivity of a large
tree.
In the spring, last year’s hard-won energy is invested to grow and expand the new
leaves. Through the summer each leaf will yield a good return. When light is plentiful,
temperatures are warm, and water is in good supply, leaves produce much more energy
than they consume. However, in the dark and cold of winter, when their delicate bio-
chemical machinery may be damaged, when water may be hard to coax from frozen
soil, and when energy is needed to keep them alive, leaves become a liability. In ginkgo
and many other trees they are quickly and thoroughly discarded. This is the economics
of growth in a deciduous tree. The energy and nutrients used to build the leaves are
partly reclaimed, and the rest is written off.10
Every autumn, just at the moment before the leaves fall, ginkgo is at its most beau-
tiful. By then the leaves are bright yellow; in the low-angled autumn sun they are “bril-
liant as a brimstone butterfly.” Often the yellowing begins at the leaf edge and slowly
expands across the whole blade. It reflects complicated, tightly controlled changes
going on inside each leaf, all apparently triggered by the shorter days rather than the
onset of cold weather. In the process, chloroplasts degenerate, chlorophyll is broken
down, and photosynthesis slows. Valuable nutrients, especially nitrogen and phospho-
rous, are absorbed back into the tree; they are too precious to waste. And as the green
 33
energy

In mid-November, bright yellow leaves cover the lawn beneath a ginkgo tree in the garden
of the President’s House at Yale University in New Haven, Connecticut.

of chlorophyll disappears, the yellow of other light-absorbing pigments, the carote-

noids, come to the fore. Leaves of green turn to leaves of gold.11
For these few short weeks ginkgo is worth its place in any garden. A single ginkgo
can be a dramatic horticultural focal point. In early November, around the time of
the annual Yale-Harvard football game, when the tree peepers are enjoying the last
of New England’s autumn color, the single ginkgo that stands proudly in the garden of
the President’s House at Yale is at its best. Regrettably, the ginkgos at Harvard’s Arnold
Arboretum are still more spectacular.
A massed planting of ginkgos is enough to bring out the sightseers. In the heart of
Tokyo the double allée of more than 140 large ginkgo trees that line the view to the
Meiji Memorial Gallery in the Meiji-Jingu Park is a popular destination. When these
 34
the living tree

trees were planted in the 1920s, they were about twenty feet tall; the largest now ap-
proach a hundred feet. They are meticulously maintained as one of the finest examples
of Western-style landscape architecture in Japan. In the autumn they create a spectacle.
Sightseers come to admire the view, eat roasted ginkgo nuts, and take tea as the late-
afternoon light fades away and winter approaches.12
At the Meiji-Jingu Park, as everywhere else in the world, when the time comes for
ginkgo leaves to be shed they go suddenly, almost all at once, seemingly without provo-
cation. In Monroe, Wisconsin, there is a large ginkgo outside the Dorsch Memorial
Library, planted from a seed reputedly given to Dr. Dorsch by the Chinese ambassador
at Lincoln’s inauguration. For many years there was a competition to guess the date on
which the leaves would fall.13
Ginkgo has the most synchronized leaf drop of any tree I know. In a general way we
understand how trees shed their leaves. It happens because of changes in a layer of cells
right at the point where the stalk attaches to the branch. Ultimately, these cells die, the
walls between them separate, and the leaf falls, but exactly how this happens and why,
in ginkgo, it happens with eerie synchronicity, no one knows. The former U.S. poet lau-
reate Howard Nemerov also wondered.14

Late in November, on a single night

Not even near to freezing, the ginkgo trees
In one consent, and neither to rain nor to wind
But as though to time alone: the golden and the green
Leaves litter the lawn today, that yesterday
Had spread aloft their fluttering fans of light

What signal from the stars? What senses took it in?

What in those wooden motives so decided
To strike their leaves, to down their leaves,
Rebellion or surrender? and if this
Can happen thus, what race shall be exempt?
What use to learn the lessons taught by time,
If a star at any time may tell us: Now.
 5
Growth

The wood will renew the foliage it sheds.

—Irish proverb

Matthew Hargraves, a curator at the Center for British Art at Yale, points out that in
the late eighteenth to mid-nineteenth century, “the tree nearly supplanted the human
figure as the best test of an artist’s mettle.” Any artist looking closely at ginkgo would
quickly have seized upon its unusual form. Even when the leaves are gone, ginkgo is
distinctive; for decades at the beginning of their long lives, ginkgo trees have a sparse
crown, with widely spaced branches that poke out like long, thin, spiky fingers. Their
characteristic silhouette differs from that of any other tree. Unlike the leaves, the archi-
tecture of a ginkgo tree is lacking in grace; it is no American elm.1
Branches of a mature ginkgo are also distinctive in another way. Twigs that are more
than a few years old have sideways-pointing, spurlike, cylindrical pegs: so-called short
shoots, each of which grows by only a fraction of an inch every year. In contrast, the
long shoots on which they are borne may be very long. Each grows rapidly from a bud
at its tip. Growth from a single bud in a single season may produce a foot or more of
new twig. Similar differentiation into long and short shoots occurs in many other trees,
such as apples. It reflects a division of labor between twigs with normal growth and
widely spaced leaves, through which the tree gets bigger, and twigs that are repressed

35
 36
the living tree

Young leaves and pollen cones emerge from short shoots scattered along
the branches of an ancient ginkgo, Kyushu, Japan.

and bear their leaves in a tuft. In ginkgo the difference is especially pronounced. The
long shoots grow straight and are few in number compared with the short shoots. The
spikiness of the ginkgo habit in part reflects this special way of growing.
Short shoots add further character to the ginkgo silhouette. Each can be up to an
inch or more long and bears the crowded scars left by the leaves of previous summers.
On older branches a good-sized short shoot may bear the scars of a hundred or more
leaves borne in a life that may last decades. Botanists in Europe and North America
sometimes pride themselves on being able to identify trees in winter in the absence of
leaves. It is a test of observation, deduction, and memory, which requires looking care-
fully at the bark, the scars of leaves that have been shed, and the pattern of branching.
Only the most inexperienced student would have a problem in recognizing ginkgo. The
characteristic cylindrical short shoots are unique.2
In spring, twigs on ginkgo trees that have been barren since autumn come back to
 37
growth

life. Clusters of leaves, usually about four to six, emerge in miniature from the bud at
the tip of each short shoot. They were formed in the previous growing season and sur-
vived the winter tiny and tightly packed inside their protective buds; each has a short
stalk with a minute leaf blade that is rolled inward on either side. Their precocious
development means that the leaves can be deployed quickly in the spring. With the
coming warmth the bud scales fold back, the leaves are revealed, and the blades unroll.
As the leaf stalk grows in length, the blade expands rapidly.
At bud burst, the leaves are small, lime green, delicate, and vulnerable to a late
freeze. In April 2007, Chicago and the rest of the Midwest suffered a late cold snap as
the wind chill dipped to minus 30° Fahrenheit soon after the leaves had emerged: most
did not survive. Eventually, a few weeks later, the trees summoned the energy to re-
place them, but successive years of such natural abuse would quickly exhaust them.3
On a well-established ginkgo most of the leaves are borne on short shoots, and these
are the leaves that bring in most of the tree’s energy. Short shoots are a neat functional
trick in the economics of tree growth. Each supports a cluster of leaves that yields a
good return without the energetic expense of building a long branch on which to bear
them. With their multiple leaves, short shoots produce a significant amount of energy
with only minimal long-term investment in wood and other tissues. They also help
make the crown open and light, rather than dense and compact. In summer, each long
spiky branch of a young ginkgo is a feathery cylinder of leaf clusters: a single main long
shoot bearing hundreds of short shoots and thousands of leaves.
Short shoots bring in energy but do little to increase the tree’s stature or prevent
it from being overgrown by its neighbors. It is the long shoots that are responsible
for most of the physical growth of the tree. They elongate rapidly during the grow-
ing season and bear leaves alternately, one by one, along their length. In the spring, as
the buds are breaking, new short shoots may start from each of the small buds left in
the axils of last year’s long-shoot leaves. In this way, long shoots extend the length of
branches and expand the size of the crown, while short shoots fill in the crown with
leaves.
The whole system is also beautifully flexible, because the difference between the long
and short shoots is under simple control. The short shoots are held in check by chemi-
cal signals passing back from the dominant thrusting bud at the tip of the long shoot.
With strict discipline controlled by simple chemistry, ginkgo short shoots defer to their
 38
the living tree

leader. However, if signals from the tip of a long shoot are interrupted—for example, if
the bud is damaged or lost—one or more short shoots soon break ranks, start to elon-
gate, and quickly become new leaders.4
The leaves on long and short shoots are slightly different. On short shoots they have
fan-shaped blades that are rarely deeply divided. The upper edge is usually more or less
smooth, or at the most only shallowly notched. The same is true of the first few leaves
that burst from the tip of a long shoot at the beginning of the season. However, leaves
formed on long shoots later in the growing season have a deep central notch that may
be up to two-thirds of the leaf blade. These are the leaves that served Goethe’s poetic
purpose and that Linnaeus had in mind when he coined the epithet biloba.5
The difference between leaves on long and short shoots is beautifully seen in a
nineteenth-century illustration of ginkgo from Siebold and Zuccarini’s Flora Japonica.
The difference probably reflects the different conditions the leaves encounter as they
develop. Leaves on short shoots, and the first-formed leaves on the long shoots, both
begin their development inside tightly closed buds. They are well protected, develop
slowly, and survive in near suspended animation over the winter. The long-shoot leaves
are formed later, and they begin to develop only after growth has started in the spring;
they develop and grow more quickly inside the bud at the tip of a long shoot that is
itself elongating rapidly.
These same factors may also be responsible for the internal differences between the
leaves on long and short shoots, and their different capacities to conduct water. Short-
shoot leaves develop attached to a stubby branch that is already mature, is not grow-
ing rapidly, and that has well-developed tissues capable of providing an ample supply
of water. Long-shoot leaves, on the other hand, are borne on shoots that are imma-
ture, are growing very actively, and have a greater need for water even as their internal
water-conducting tissues are still being formed. Careful measurements show that long-
shoot leaves, perhaps because they are more likely to be short of water as they grow, are
more effective in water conduction.6
The need to keep the leaves fully supplied with water may be the crucial limitation
that prevents ginkgo from growing really tall. All the water needed to keep the leaves
fully hydrated comes from the soil, scavenged molecule by molecule by fine hairs on
young roots. It is then moved to the bigger roots and upward through the outer parts
of the woody column inside the trunk, out through the branches, and eventually to the
leaves. The supply needs to be steady and consistent to replace the water lost through
The illustration of Ginkgo biloba, labeled with a variant of the once-popular name of Salisburia
adiantifolia, from Philipp Franz von Siebold and Joseph Gerhard Zuccarini’s Flora Japonica.
 40
the living tree

the breathing pores, as well as smaller amounts lost by evaporation through the cuticle
and used in photosynthesis.7
What is remarkable is that the entire process of water transport in plants is passive:
there is no hidden microscopic pump and no expenditure of energy on the part of the
tree. The tiny elongated cells through which the water passes are dead and empty, and
except in the early spring, when stored sugars in the trunk and roots begin to be mobi-
lized, and the sap starts to rise, there is little sustained pressure from below. In prin-
ciple, the arrangement could hardly be more simple: as water is lost or used up in the
leaves, it is replaced by water drawn up through the stem all the way from the roots.8
The amount of water needed to keep a big tree fully hydrated varies greatly, de-
pending on the kind of tree and where it is growing, but it can be substantial. A thirty-
year-old sessile oak growing in a plantation in eastern France might use only about a
half-gallon of water a day, while the yevaro, a large tree that dominates certain kinds
of tropical forests in the Venezuelan Amazon, might use five hundred times as much.9
All of this water is literally sucked up from the roots; the taller the tree, the greater
the suction needed, and the greater the internal pressures that are generated. The nega-
tive pressure needed to lift the water against the force exerted by gravity increases by
almost one and a half pounds per square inch for every three feet or so of height. The
resulting tension in the water-filled cells of the wood of a tall tree is enormous. The
elongated cells through which water is drawn have to withstand massive pressures; they
have robust cell walls and specialized internal thickenings that help prevent them from
imploding.
A further hazard created by such pressures is that air inside nearby empty cells may
be sucked into the water column. The result is a kind of botanical embolism; a bubble
of air that creates a blockage and prevents further conduction. In ginkgo, this danger
is minimized by the minute pores that connect adjacent cells; they are just the right
size to allow water through, but too small to let an air bubble pass. In this, as in many
other details, every tree is an extraordinary feat of hydraulic engineering: all created
by natural selection from natural variation over millions of generations.10
The wood that makes up the trunk of ginkgo and other trees not only conducts water
but also provides support. It creates the scaffold on which the leaves are arranged, lift-
ing them up toward the sun and away from the shade created by nearby plants. This
competitive struggle for light was probably the ultimate driver in the evolution of trees.
However, getting bigger has consequences. As trees become larger, water and other
 41
growth

fluids become harder to move around: the distances involved, and the internal pres-
sures on the water-conducting cells, increase. Bigger trees are also heavier, especially
when laden with water after a downpour: the trunk has to be strong enough to carry
the load.
There are also considerations of internal economy. Over the life of a single tree the
more energy diverted to building the trunk and branches, the less energy is available
for reproduction or other kinds of growth. The energetic cost of simple maintenance,
sustaining basic life processes, is also considerable. It makes sense that truly massive
trees live in places where growth never completely shuts down in the winter, and where
water is readily available almost all of the time. For most trees, maintaining a gigantic
branching scaffold and hundreds of thousands of leaves through tough times, when
temperatures are low, the soil water is frozen, and no energy is coming in, may not be
a winning strategy.
So like most things in nature, the height of trees is a trade-off, dictated in part by
the ability of a big tree to conduct water, in part by the energetic costs and benefits of
achieving and maintaining large size, and in part by the physical strength of the wood
and how that relates to the form of the tree. In the case of ginkgo all of these factors
may be at play, but with its broad leaves and their unusual architecture, maintaining
a steady supply of water may be especially important. Ginkgo is perhaps not quite as
efficient in holding onto water and supplying it to the leaves as some of the world’s
most imposing plant giants. Part of the explanation for the great size of the Yongmunsa
Ginkgo, and other truly massive ginkgos, may be that their roots are tapped into a
plentiful water supply.
 6
Stature

. . . as carpenters carve wood, the wise shape their minds . . .

—The Buddha, The Dhammapada

Puccini’s opera Madama Butterfly, set in Japan in 1904, begins in the garden ter-
race of a small house overlooking Nagasaki Harbor. The tragic story of Cho-Cho San,
which Puccini adapted from the novel by John Luther Long, was just one expression
of increased fascination in the West with all things Japanese in the decades after Com-
modore Perry’s Black Ships forced the reclusive nation into contact with the outside
world. Much has been written about the possible models for Long’s story: connections
have been made to the book Madame Chrysanthemum by Pierre Loti, a French naval
officer who was in Nagasaki during the summer of 1885, and to also to Thomas Glover,
a Scottish merchant and key figure in the early industrialization of Japan who lived in
Nagasaki in the late nineteenth century. Another connection is to Philipp Franz von
Siebold, who was in Nagasaki in the 1820s and 1850s. Siebold was no Lieutenant Pinker-
ton, but some parts of his story will sound familiar to those who know Puccini’s opera.1
From the mid-sixteenth century until the time of Commodore Perry, Nagasaki, with
its fine natural harbor, was the only port that linked Japan to the West. Trade was con-
trolled through the monopoly held by the Dutch East India Company. Company ships
brought western luxuries from Holland to Japan and returned laden with fine ceramics

42
 43
s tat u r e

and other goods that helped ignite Europe’s craze for japonism. The elite of Amster-
dam, London, or Paris could order customized sets of Japanese china made in the kilns
of Kyushu. It was into this milieu that Siebold came when he arrived in Nagasaki in
1823 as the doctor to this far-flung outpost of the Dutch trading network. Following in
the footsteps of Engelbert Kaempfer and Carl Peter Thunberg, two of his predecessors
who were also employed by the Dutch East India Company, he became one of the three
great early botanical explorers of Japan and a pioneer of Japanese studies in Europe.
Siebold studied medicine at the University of Würzburg, was a member of the
Senckenberg Natural History Society in Frankfurt, and was much influenced by the
adventures and writings of Alexander Von Humboldt. His dream was to emulate Hum-
boldt’s explorations in far- off countries, and when he was sent to Japan as surgeon
major, Siebold seized the opportunity. He used his time there to gather detailed infor-
mation about the country, its plants and animals, its culture, and the people that he
met.
During the seven years that Siebold spent in Nagasaki in the 1820s, Japan was still
almost entirely closed to the Western world. The Dutch were confined to the island of
Deshima in Nagasaki Harbor, the import and export of goods was strictly monitored,
and there was little contact with the local population. However, as a doctor, and par-
ticularly as an ophthalmologist and obstetrician with expertise in Western medicine,
Siebold had unusual freedom to leave the island to treat Japanese patients and collect
herbs. He also recruited students, who helped him purchase land and establish the first
school in Japan to teach Western medicine.2
During house visits in Nagasaki, Siebold met and fell in love with a local woman,
Kusumoto Taki, who was also given the name Sonogi. Siebold was twenty-seven and
Sonogi sixteen when they met. He wrote to his uncle that he had “become quite at-
tached to a sweet sixteen-year old Japanese girl, who I would not willingly exchange
for a European one.” Sonogi was one of the few Japanese allowed to stay on Deshima.
Siebold called her Otaksa, and they had a daughter together, whom they named Oine.3
Siebold would perhaps have returned to Japan repeatedly except for an incident that
took place in 1826 during the annual visit of the Dutch legation to Edo, modern Tokyo.
Each year, with great ceremony, representatives of the Dutch traders traveled to Edo to
offer gifts in tribute to the Shogun. Kaempfer and Thunberg had made the same jour-
ney and, like them, Siebold valued this rare opportunity to see other parts of the coun-
try and learn more about all aspects of Japanese life. In 1826, as on his earlier journeys
 44
the living tree

Statue of Philipp Franz von Siebold (1796–1866) as a

young man, Siebold Museum, Nagasaki, Japan.

to Edo, Siebold collected many items that helped illuminate Japanese culture, but espe-
cially prized were several detailed maps of Japan and Korea given to him by his friend
Takahashi Sakuzaemon, the court astronomer and supervisor of the Imperial Library.
In return Sakuzaemon received a recently issued Russian map of the world. Siebold’s
motivation was simple curiosity, but for Japan, with its concern about outside influ-
ences, such maps were highly sensitive: possessing one was forbidden, and supplying
one to a foreigner was a capital offense.4
Siebold must have understood the risk he was taking, but Kaempfer and Thun-
berg had returned with similar documents. According to some accounts, Siebold was
already on a boat leaving Nagasaki Harbor when his good fortune ran out. The maps
would probably have gone undetected had it not been for bad weather that brought
his ship back to port. His belongings were unloaded and inspected, and the maps were
discovered, along with another illegal item, a gown bearing the Shogun’s crest. When
 45
s tat u r e

this came to the attention of the Japanese authorities, Siebold was interrogated, as were
about forty Japanese who were in contact with him during the court journey, along with
fifty interpreters and many of his students. Takahashi Sakuzaemon fared the worst. He
was arrested and died in prison soon after.5
The verdict on Siebold was reached more than a year later. The court found no evi-
dence that he was a spy, and also took into account his service to Japan, as well as pleas
from his employers in Europe, but the sentence handed down in October 1829 was
nonetheless severe. Some of his Japanese colleagues were imprisoned or sent to remote
islands. Siebold was banished from Japan for life and left Deshima on December 30,
1829. He said his final goodbyes to Sonogi, Oine, and two of his devoted students, Kō
Ryōsai and Ninomiya Keisaku, on his ship, the Cornelius Houtman, as it left Nagasaki
Harbor. He took with him two small lacquered boxes with portraits of Sonogi and Oine
that contained locks of their hair. Oine was two years and eight months old when her
father left; Siebold entrusted responsibility for her well-being and education to Kō
Ryōsai and Ninomiya Keisaku.6
When Siebold left Nagasaki, Japan was still an isolated country, but more than to
any other Westerner, history gave Siebold the opportunity to view at close quarters the
massive changes wrought as the country opened its doors to the world. Two years after
the arrival of Commodore Perry, Siebold returned to Japan in 1855 with the responsi-
bility of delivering the Netherlands Trading Company’s trade agreement for ratifica-
tion. There was also an emotional reunion with Sonogi and Oine. Siebold had since
married and had five children, Sonogi had married twice, and Oine was on her way to
becoming Japan’s first female doctor.7
Siebold accumulated a vast collection of specimens, books, and artifacts in Japan,
some on his trips to Edo and others in return for medical services. He also preserved
large collections of plants and animals. On his return to Europe he used these to docu-
ment Japanese natural history and culture in a trilogy of important early works on dif-
ferent aspects of Japan. Flora Japonica, Siebold’s work on Japanese plants, illustrated
with lavish plates, remains an influential early work on the plants of Japan.8
The copy of Flora Japonica in the Kew Library was acquired by William Jackson
Hooker, and after his death it was purchased for the nation, along with the rest of
Hooker’s library. As well as Flora Japonica, the books in Hooker’s collection included
superb art by Ehret, the Bauer brothers, and many others. The books became the
nucleus of one of the greatest collections of botanical art anywhere. However, there is
 46
the living tree

also interesting botanical art in the Kew Economic Botany Collection, and particularly
unusual are a series of twenty-six boards, each with an illustration of a plant painted
on its surface. Like Flora Japonica, these also have a link to Siebold.9
Each board is a little more than a foot long and nine inches wide, held in a frame
made of pieces of a small branch. The frames, as well as the boards on which the paint-
ings are made, seem to be made from the plants that are illustrated. One is labeled
“Ginkgo biloba, Linn.,” with the name of the plant written in Chinese Kanji and Japa-
nese Katakana characters below; both read in the old-fashioned way, from right to left.
The records at Kew tell us nothing about when and where this strange xylotheque
was made, or how it came to London, but similar collections of boards at the Koishi-
kawa Botanical Garden in Tokyo, the Berlin-Dahlem Botanic Garden, and the Harvard
University Herbaria, as well as a private collection in Britain, provide some clues. On
the back surface many bear the same red seal of Chikusai Kato, the first plant illustra-
tor employed in the Koishikawa Botanic Garden, which is now part of the University
of Tokyo. The seals incorporate the line “Spring, New creative achievement,” with the
date “11 Meiji,” 1878 in the Western calendar.10
Both the Kew and Berlin collections contain a board illustrating ginkgo. The Chi-
nese characters on the Berlin board read: “Grandfather Grandson tree, more formally,
Silver Apricot.” The two illustrations of ginkgo twigs, with leaves and young seed stalks
attached, are very similar and also closely resemble a painting of ginkgo produced by
Chikusai Kato for the Koishikawa Garden. Parts of the twigs and short shoots in all
three illustrations are almost identical, and the way in which the mature seeds on their
seed stalks are drawn is exactly the same. Only a few details on the Kew board are not
shown in the Berlin illustration, but all are in Kato’s ginkgo painting.11
Chikusai Kato worked for Keisuke Ito, an early professor at Tokyo University Bo-
tanic Garden, who had met Siebold as a young man in 1826. Kato may also have been
influenced by Keiga Kawahara, an artist who worked for Siebold during his time in
Nagasaki. As a result the style of these three ginkgo portraits blends Japanese and
European traditions and reflects the growing early influence of the West on Japan. The
way that the twigs are represented has a clear Japanese influence but the separate illus-
tration of botanical details is adopted from the Western scientific tradition that was
almost certainly introduced by Siebold.
The ginkgo wood used to make the boards in the Kew and Berlin xylotheques has
the same basic structure as that of most woody plants. Cut down a ginkgo and the
A portrait of ginkgo painted onto a board made from ginkgo wood and framed with young ginkgo
branches. The portrait is by Chikusai Kato and was made for Tokyo University in 1878.
 48
the living tree

trunk looks exactly like that of a normal tree; only a specialist would be able to distin-
guish the wood from that of pine or cedar. Around the outside in a narrow band is the
bark, and immediately inside is a zone of softer tissues; the rest of the trunk is made
up of wood.
In its outer part, where water and nutrients are still being transported, the wood is
light in color, but the center of the stem, the heartwood, is dark, and plays no part in
water conduction. Its cells are often blocked by deposits of various kinds that build up
over time. Nevertheless, the heartwood is vitally important: the strength of this dense
central column is crucial for support of the tree.
Almost all of the tissues that make up the twigs, branches, or even the trunk, of a
full-grown ginkgo tree are produced in the same way. Except for the bark, they can all
be traced back to a delicate cylinder of living cells just a single cell thick. In most trees
this cylinder, known technically as the cambium, occurs not far below the bark be-
tween the wood and the softer tissues toward the outside. As they grow, the elongated
cells in this layer divide consistently and regularly at a tangent to the circular cross sec-
tion of the trunk; in the process they form new cells toward both the outside and the
inside.
As soon as the new cells produced by the cambium toward the inside are fully de-
veloped, they die. They become part of the central woody column and add to the mass
of strengthened cells, xylem cells, that are conduits for water on its way from the roots
to the leaves. Toward the outside, on the other side of the cylinder of dividing cells, the
newly formed elongated cells remain alive and form the softer zone of fibrous tissue
immediately inside the bark. It is these phloem cells, the functional counterpoint to
the dead, water-conducting, xylem cells of the wood, that actively transport the sugars
made in the leaves to the lower part of the stem and the roots.
Also produced by the actively dividing layer, but this time running horizontally
rather than vertically through the wood, and scattered at different levels among the
files of dead cells, are thinner-walled, elongated living cells, the so- called rays. Un-
like the cells making up most of the rest of the wood, these cells are alive. They help
transport sap horizontally through wood and are often the cells where starch is stored
through the winter.12
A consequence of the way that the water-conducting cells in the trunk are formed is
that they are arranged in radial files. These come about because each row of elongated
cells, both toward the inside and toward the outside, traces its origin back to one of the
 49
s tat u r e

Close-up of the bark of a young ginkgo

showing the successive layers produced
by a zone of actively dividing cells near
the outer surface of the trunk. In this
tree, which was growing on a busy street
in Seoul, South Korea, the bark has been
partly smoothed by the hundreds of
people who brush against it every year.

cells in the actively dividing layer. Each file preserves a record of the repeated divisions
of that single cell through time. However, just occasionally one file can be seen to have
split into two, and this reflects occasional radial divisions of one of the cells in the ac-
tively dividing cylinder. This is how trees increase in girth. Such divisions increase the
diameter of the cylinder and allow it to expand to keep up with the increasing mass of
wood in the center of the trunk or twig. Ultimately, the increased size of the cylinder
results in an increased circumference of the trunk and, to the outside, the increase in
girth is reflected in the fissures in the bark.
The bark is a layer of corky cells produced by a second and less-well-defined cylin-
der of actively dividing cells. The quantity and texture of bark produced by these cells
varies greatly in different kinds of trees. Smooth bark, such as that of beech trees,
forms where there is little cork and where its production is even around the stem.
Rough and deeply textured bark like that of oak trees results from much more exten-
sive production of corky cells. In ginkgo the bark is intermediate in texture, but a close
 50
the living tree

A cross section through the wood

of living ginkgo, showing the large-
diameter water-conducting cells
produced in the spring and summer and
the smaller-diameter water-conducting
cells produced late in the growing season.
Annual rings are the result of this
rhythmic pattern of growth.

look clearly shows the rhythmic layers produced by the annual growth of an outer
cylinder of actively dividing cells, the cork cambium.13
The individual cells that make up the wood of ginkgo are not much more than a few
hundred thousandths of an inch across but may be up to half an inch long. For many
years it was thought that these kinds of long narrow cells, which make up most of the
trunk in ginkgo and conifers, were inefficient conductors of water; especially com-
pared with what seemed to be more sophisticated systems in most other trees, where
cells are arranged end to end and connected by holes in their end walls to form long
pipes. However, this turns out not to be true; the conducting cells of ginkgo and the
other conifers are equally sophisticated but employ a different system. Each is perfo-
rated with thousands of tiny valves that close when water is in short supply, but that
still permit easy and efficient transport through the wood when water is plentiful and
the valves are open. As often happens, over the eons evolution has invented more than
one solution to the same problem.14
Ginkgo responds to the changing seasons on the outside by dropping its leaves. On
the inside it responds by shutting down the cylinders of dividing cells over the winter.
The effect is most obvious in the wood. New cells of the wood produced in the autumn
have slightly smaller diameters than those produced in spring and summer; then cell
division stops abruptly as the tree becomes dormant. Through the winter no new cells
are produced toward either the inside or the outside. But when growth begins again in
the spring, the new cells produced are of normal size. It is the difference between the
 51
s tat u r e

small-diameter cells produced late in the growing season and the larger-diameter cells
produced early in the next growing season that shows up, even to the unaided eye, as
annual rings: a rhythmic record of the tree’s annual growth. The rings on the young
branches used to make the corners of the Kew ginkgo board show that each had grown
for about eleven years.15
Ginkgo is not grown for timber, but its wood nevertheless has its uses. Technically,
ginkgo is a soft wood like pine or spruce, but its more or less homogenous structure
compared with the wood of most trees, especially hardwoods, makes it resilient. Even
if it regularly gets wet and then dries out, ginkgo wood does not easily shrink, crack,
or warp. As John Quin noted, it can be used as a base for lacquerware where shrinkage
would cause cracking and peeling of the lacquer. In China, when the trees in a ginkgo
orchard cease to be productive, they are cut down, chopped, and shredded to make
particle board. In Japan, larger pieces of timber are used for chopping blocks, furni-
ture, molds, and ornaments.16
The smooth, even texture of ginkgo wood, a result of its relatively simple structure,
also makes it easy to work. Scott Strobel, a colleague at Yale with great skill on a wood
lathe, says that ginkgo wood works more smoothly and easily than any other wood he
has used. The chisel goes through it “like a knife through butter.” Daoist shamans are
said to have engraved their magical spells and seals on old ginkgo wood to communi-
cate with the spirit world. The Vairocana, the Manjusuri, and the Samantabhadra stat-
ues at the Haeinsa Buddhist Temple in South Korea are also carved from ginkgo wood.
Much less ornate, but no less impressive, are the remarkable sculptures housed in a
small temple high above the small town of Ojiya in the mountains of central Honshu.17
The temple is a tiny jewel reached by a steep road that winds up through Japanese
cedars and gives spectacular views to the valley below. Outside it is a modest, square,
wooden building with a peaked roof in traditional Japanese style. In the winter the
outside is protected by thick boards from the heavy snows. Inside it is also plain, but
along the wall opposite the entrance are thirty-five sculptures, each a representation of
the Buddhist Goddess of Mercy with the head centered in a stylized disk representing
the sun. The largest figure, about five feet tall, has its right hand resting on its cheek.
Around the head is an inscription in charcoal. To the right and left are two similar but
smaller figures, and in alcoves on either side others are arranged in two groups of six-
teen; each a little different from the next, and each with its own symbolic meaning.
The sculptures are all the work of Mokujiki Shonin, one of the most famous Bud-
 52
the living tree

dhist monks of eighteenth-century Japan. Born in 1718 and graduating to the priest-
hood in his early twenties, he practiced in the temple at Edo for more than twenty
years before becoming attracted to a particular sect of Buddhism that eschews meat,
fish, and rice for a diet of nuts, leaves, and fruits. From the age of sixty until his death
in 1810 at the age of ninety-three, Mokujiki Shonin traveled from temple to temple in
central Japan and carved more than one thousand wooden Buddha sculptures.18
He was already approaching his eighties when he first visited the temple in Ojiya,
but on his return several years later the old temple was gone, burned down in an acci-
dental fire. He set out to make the new temple unique. The townspeople brought him
pieces of a large ginkgo tree that they had cut from farther down the valley. In a little
over three weeks in August 1803, he crafted them into the remarkable figures that are
the heart of the Ojiya Temple.
 7
Sex

We are survival machines—robot vehicles blindly programmed

to preserve the selfish molecules known as genes.
—Richard Dawkins, The Selfish Gene

All living ginkgos are connected, part of an unbroken chain of genetic continuity
that has survived through thousands of generations. With Darwinian hindsight we
recognize now that this continuity links all living ginkgos to their ancestors that grew
more than 200 million years ago, and that has been sustained, as in all living organ-
isms, by an innate propensity for self-perpetuation through reproduction.1
Reproduction in plants, just like reproduction in humans, involves sex: the bring-
ing together and fusion of reproductive cells, usually but not always from two parents,
to create an embryo that develops into a new individual. Each sex cell—sperm from
the male, egg from the female—carries just a single set of chromosomes, and after
fusion the newly created embryo has two sets: one from the father, the other from the
mother. In this respect ginkgo is not remarkable. But beyond these bare essentials, sex
in ginkgo is a long and complex process.
In ginkgo, as in ourselves, there are separate male and female individuals. In the
tree, pollination—the transfer of pollen from the male to the female, the essential pre-
cursor to sexual fusion—takes place in the spring. Sexual fusion, the initial step in the

53
 54
the living tree

formation of an embryo, follows in the late summer or early autumn, and the result, a
seed containing the embryo of a new young plant, is shed a month or two later. Gen-
erally, the time from pollination to the point at which the seed germinates is about a
year.
A key feature of sexual reproduction, which comes about by combining the chro-
mosomes from each sex cell, is that the resulting offspring are not exactly like either
parent. In a complicated way, through mechanisms mediated at the level of DNA, the
new young plant or animal combines features from both parents. In addition, during
the development of the sex cells on both the maternal and paternal sides, DNA on the
chromosomes is shuffled. As a result, the combination of features specified by the DNA
in every sperm or egg cell ends up being different. These are the main reasons why
brothers and sisters are seldom identical; for a mature male and female ginkgo, this
means that the genetic makeup of every one of their offspring, the embryo inside every
seed that develops on a female tree, is unique.
A further important part of the development of the sex cells, and a necessary pre-
lude to sexual fusion, is that the chromosome number is reduced by half, which en-
sures that the resulting embryo does not end up with four sets of chromosomes, double
the number of its parents. In ginkgo and in all other plants this halving of the chromo-
some number, which occurs at the same time as the shuffling of the DNA, takes place
during the formation of the pollen grains, and also inside the developing seeds as the
eggs are being formed.
The seventeenth-century German scientist Rudolf Jakob Camerarius was the first
to begin to develop a scientific understanding of the crucial importance of the trans-
fer of pollen for the production of seed. Earlier botanists, such as John Ray and Nehe-
miah Grew, had made similar suggestions, but while studying mulberries, Camera-
rius noticed that female plants growing far from males would produce fruits but no
seeds. He also experimented on castor oil plants and maize by removing the pollen-
producing parts, and showed that no seeds were formed.2
From the earliest human encounters with ginkgo it would have been clear that seeds
are produced only on some trees and not on others. Realizing that trees of two different
kinds are needed for seed to be produced would have required a further mental step,
but was nevertheless recognized well before any scientific understanding of the pro-
cess. What may be the earliest written reference to ginkgo in China, from the tenth cen-
 55
sex

tury, includes the statement: “Let male and female ginkgo trees grow near one another,
then fruit will form.”3
Ginkgo is relatively unusual among seed-bearing plants in having male and female
reproductive parts produced on separate trees. Pines, for example, have separate
pollen and seed cones, but both are borne on the same plant and they are therefore
hermaphrodites. It is even more common to have male and female parts closely asso-
ciated in the same structure, as in most flowering trees, like a cherry or a magnolia,
where the pollen-producing male parts surround the seed-producing female parts in
the same flower.
Nevertheless, Darwin was the first to suggest that the complete separation of sexes
in plants might, in the right circumstances, provide advantages through a division of
labor in the production of sex cells. Since then, the conditions under which a switch
from hermaphroditism to having separate males and females might be favored have
been studied in detail, and among flowering plants it is now recognized that this must
have occurred independently more than a hundred times. The basic idea is simple: if
an advantage exists for a plant in terms of passing on more of its genes to the next gen-
eration by specializing as a male or a female, rather than by being both, then natural
selection will tend to push successive populations in that direction.4
Simple differences in the arrangement of the sex organs in plants have important
consequences. In plants like cherries and magnolias, where male and female parts
occur in the same flower, Darwin recognized that self-pollination and self-fertilization
may occur quite often even though many plants have evolved ingenious mechanisms to
prevent it. In pines, which have separate male and female cones on the same plant, the
chances of self-pollination may be reduced, but nevertheless there is still ample oppor-
tunity for self-pollination and potentially for self-fertilization. However, in ginkgo and
plants like it, reproduction always requires a partner. Self-fertilization is a physical im-
possibility.
When male ginkgo trees are mature, the small conelike structures in which the
pollen is formed are produced on normal short shoots at the same time as the young
leaves begin to appear. The cones emerge from the bud at the shoot tip; each has a short
stalk and is borne at the base of one of the tiny developing leaves. On a single short
shoot up to half a dozen pollen cones may be produced among the cluster of young
leaves. At first the pollen cones are slightly fleshy, and just two- or three-tenths of an
 56
the living tree

inch long. They have a central stalk that is completely surrounded by small, tightly
packed side branches, and each side branch has two small, downward-pointing yellow
sacs at the tip in which the pollen is produced.5
On dry warm days in the spring, the central stalk grows quickly, the cones elongate,
the side branches become widely separated, and each pollen sac splits open by a single
lengthwise slit to release the pollen. A triangular bulge at the end of each side branch,
right at the point where the two pollen sacs join, may be involved some way in the
splitting process. For just a few days every year vast quantities of tiny pollen grains are
released into the air and blow away on the wind. In the process the cones dry up, and
soon after, they are shed; desiccated pollen catkins carpet the ground beneath the tree,
their job done for another year. From then until new cones appear the next spring, the
only way to spot a male ginkgo is through negative evidence: the lack of seeds on the
tree or on the ground beneath it.
Each pollen grain is tiny, about a thousandth of an inch long, roughly the shape of
a rugby ball, with a tough, flexible wall except for a single narrowly elliptical region
where the pollen wall is thinner. The grains are not very distinctive and look much like
the pollen of cycads or even some flowering plants. When they are released, the pollen
grains dry out, the thin part of the wall folds inward, the grains become thinner and
slightly longer, and further water loss is slowed.6
The output of pollen grains from a single ginkgo tree over just a few crucial days
every year is extraordinary. Andrew Leslie, a former colleague at the University of Chi-
cago, made a very rough estimate for the thirty-five- to forty-foot ginkgo trees grow-
ing on campus. He calculated that the pollen output from about seven cones on an
individual short shoot might be as many as fifty-nine million individual pollen grains.
With an estimated 17,500 short shoots on a tree of moderate size, this works out to an
annual production of an incredible one trillion pollen grains. This is a vast number,
but it needs to be. Imagine a single microscopic pollen grain blowing in the wind: what
are the odds that it will find its way to exactly the right place on a developing seed on
a female tree?7
At the same time that pollen cones begin to be produced on male trees, or perhaps
just a little later, mature female trees produce ovules. These are the structures in which
the egg cells develop; if pollinated, they will eventually develop into seeds. They are
borne on specialized stalks, and like the pollen cones, each is attached to the short
shoot at the base of one of the young developing leaves. The ovule-bearing structure
 57
sex

of ginkgo is distinctive, quite unlike that of any other living plant; no one would mis-
take it for a flower, or for the woody seed cone of a conifer or cycad. Each has a single,
simple stalk, usually with two young ovules borne at the tip. Each ovule is surrounded
by a collarlike rim and its pointed tip projects upward and outward; if pollinated and
fertilized, each has the potential to develop into a seed.8
At precisely the same time that ginkgo pollen is drifting on the breeze, each young
ovule is tipped by a glistening, watery drop of fluid, the pollination drop. The drop
helps trap passing pollen grains from the air, and any grains that are captured sink in
the fluid. The pollination drop is repeatedly absorbed and restored each day until it has
drawn grains inside the ovule.9
In plants, which are unable to move to find a mate, pollination is the riskiest part of
reproduction, and this is especially the case for ginkgo and other species with separate
male and female plants. The whole sequence of events needs to be beautifully coordi-
nated, with pollen and pollination drop produced at the same time. Timing is every-
thing; if pollination is successful, and if the male sex cells that develop within the pollen
grain manage to fertilize the egg inside the ovule a few months later, then an embryo, a
new young plant, will develop as the seed matures. However, any deviation from exact
synchronization is penalized harshly. Trees with a poor sense of timing will produce
no seeds and leave no offspring for the next generation. It is not hard to see how such
exquisite timing in the reproduction of male and female trees has been coordinated by
the evolutionary process of natural selection.
Normally, only one of the two ovules at the tip of a ginkgo seed stalk develops. It
might be that the other was not pollinated, or loses out in the competition for resources
from the mother tree, or aborts if there are insufficient resources for all the seeds that
are pollinated to develop. Occasionally, however, both of the ovules on a single stalk
mature, and even more rarely there may be three or four mature seeds on each stalk.
In most seeds two or three egg cells are produced, but normally only one is fertilized
or goes on to develop. In about two out of every hundred seeds more than one egg is
fertilized, and more than one embryo develops. In these rare cases, when the seed ger-
minates, two young plants may emerge.10
The mature ginkgo seed is a plumlike structure typically up to about an inch and a
quarter long, and up to about an inch wide, with the embryo developing inside. When
fully ripe, the pale yellow seeds have a distinctive silver sheen which gives the ginkgo
its common name in Chinese, “silver apricot.” Inside, a fleshy yellow pulp surrounds
Clusters of succulent apricot-like fruits on a female ginkgo in the late autumn.
 59
sex

a hard stone, and inside the stone is the developing embryo embedded in the nutritive
tissue that at an earlier stage produced the eggs.
Ginkgo is easy to grow from seed; as long as the fleshy pulp is gone, there is no
built-in dormancy that has to be overcome: during the winter development simply
slows or stops, picking up again as temperatures rise in the spring. From almost the
beginning of its development the embryo has a clear top and bottom. The top, which
points toward the tip of the ovule, eventually develops into the root of the plant, while
at the other end, toward the attachment to the stalk, growth from a zone of actively
dividing cells eventually forms the aboveground parts of the new plant. First to be
produced are a pair of seedling leaves that remain embedded in the nutritive tissue of
the seed. Strangely, the seedling leaves are green, even though they are never released
and never expand. The root grows through the apex of the seed, and as the seedling
leaves elongate, the new shoot apex is pushed out of the seed in the same direction. The
young shoot apex then becomes green and begins to grow upward, toward the light
and against gravity, producing new leaves alternately, one after another on the tree’s
first long shoot.11
Alan Mitchell, one of the great British tree specialists of the late twentieth cen-
tury, estimated that female trees first produce seeds when they are about twenty-five
to thirty years old. Male trees probably reach maturity at about the same time. Before
then it is impossible to tell whether a tree is male or female. But those two or three
decades have not been wasted. The trees have grown in size and built up reserves of
energy. Branching has also created opportunities to produce pollen cones and seeds
in large numbers. When the embryo begins its growth, it has only one group of active
cells that can produce a shoot and new leaves, but every time a new bud is formed a
new group is added. By the time the tree is mature, there are tens of thousands of these
meristems. Every one of them can potentially produce new shoots and leaves. Under
the right circumstances they can also produce pollen grains and ovules with the sex
cells inside. As those sex cells develop, the DNA of the new parent, with its unique set
of genes, is once again prepared for its passage to the next generation.
 8
Gender

We allow our ignorance to prevail upon us and make us think

we can survive alone . . .
—Maya Angelou, address to Centenary College of Louisiana

The most widely accepted explanation for why ginkgo and other trees have separate
male and female plants is that it eliminates negative effects that might come from ex-
treme inbreeding if self-pollination were followed by self-fertilization. This conjecture
is supported by a great deal of evidence that plants produced by self-fertilization are
less successful and leave fewer offspring in the next generation than those arising from
cross-fertilization. Given this principle, it is not surprising, as observed by Darwin,
that where male and female parts occur together in the same flower, there are often
mechanisms or structures to prevent self-fertilization. However, having separate male
and female plants, as in ginkgo, is a particularly blunt and unforgiving mechanism for
ruling out self-fertilization. It makes absolutely certain that offspring are genetically
different from both their parents.1
Ensuring that offspring differ from their parents is likely to be an advantage when
a new generation of plants faces a range of different ecological conditions. Indirectly,
it may also be beneficial over the long haul in a changing world. Variation, after all, as
Darwin was the first to recognize, is the raw material of evolution. Together with the

60
 61
gender

overproduction of offspring and the winnowing effect of selection, minute variations

controlled by genetics create the opportunity for successive populations of ginkgo trees
to respond to changing environmental circumstances. As different variants are favored
by different local conditions, ginkgo populations have the potential to change their ge-
netic makeup through time. This is the essence of natural selection.
However, even acknowledging the advantages in terms of producing new variation,
an inflexible system with separate male and female plants has inherent disadvantages.
In plants that cannot move to find a mate, there is no backup, no fail-safe mecha-
nism that allows seed to be produced if there are no trees of the opposite sex close by.
Many plants have mechanisms to help ensure cross-pollination, but many also have
mechanisms that effectively keep their reproductive options open, just in case cross-
pollination fails to work. Ginkgo does not have such an option. It is also more than
a little ironic that while ginkgo seems to have a mechanism that forces the produc-
tion of new genetic variation, it is also the botanical poster child for stasis, a striking
lack of evolutionary change. Paradoxically, it has stayed much the same across vast
spans of geologic time.2
The seeming simplicity of the male-female distinction in ginkgo made it an obvi-
ous focus of early research to try to understand how sex is controlled in plants. Just
as Camerarius focused on maize and castor oil, which have separate male and female
parts, Joseph von Jacquin, professor of botany at the University of Vienna in the early
nineteenth century, turned to ginkgo to carry out an interesting early experiment into
what determines sex in plants.3
Ginkgo was introduced into cultivation in Europe in the eighteenth century, and,
according to Jacquin, who carefully reviewed these early introductions, the first record
of one of these plants reaching sexual maturity is from 1795, when two trees at Kew,
one of them the Old Lion, produced pollen cones. Cuttings from the ginkgo trees in
England had been sent to the Imperial Palace of the Habsburg Empire at Schönbrunn
in 1781 and were the first ginkgo trees to be grown in Vienna. Perhaps from the same
stock, Jacquin’s father, Nikolaus Joseph von Jacquin, planted the large male ginkgo that
still grows in the botanical garden of the University of Vienna.4
Movement of water through the stems of plants had been well known since the
work of the English clergyman Stephen Hales in the early eighteenth century. Jacquin
wondered therefore whether a female branch grafted onto a male tree might change
sex. He obtained a cutting from a female tree, probably from the first female ginkgo
 62
the living tree

to be recognized in Europe, and made the graft onto the male tree at the University of
Vienna botanical garden.5
The graft was successful; the branch flourished, but it remained female. It continued
to produce seeds, even though it was now attached to a male plant. It also maintained
its independence in another way; every spring its foliage emerged two weeks later than
that of the rest of the tree, and in the autumn its leaves were still green when the rest of
the tree’s leaves had turned yellow.6
Jacquin’s observation was simple, and not unexpected, but it was also profound. It
showed that plants are made of individual parts that are to some extent autonomous,
and that sex and other characteristics are determined not at the level of a whole organ-
ism, as in most animals, but potentially in individual tissues in different parts of the
plant. It highlighted the tension between thinking about a tree as a whole organism,
like a bird or a person, and thinking about a tree as a collection of potentially indepen-
dent organisms, something more like a coral colony. It is a distinction that has practical
consequences. “Dolly the sheep,” the first cloned mammal, was a major scientific ac-
complishment not achieved until the mid-1990s. In contrast, any gardener can estab-
lish a new plant from a cutting; people have been cloning plants for thousands of years.
Not until genetics started to be better understood in the early twentieth century
were the next steps taken toward deeper knowledge of how sex in plants is controlled.
Stimulated by the rediscovery of Gregor Mendel’s work on pea plants, as well as new
research by the Dutch scientist Hugo de Vries in the 1890s, it became clear that the
characteristics of organisms, including “maleness” and “femaleness,” were inherited in
a “particulate” manner. Before then it had been assumed, by Darwin and others, that
inheritance operated by some kind of “blending,” based on the observation that off-
spring were often somewhat intermediate in their features compared with their par-
ents. Writing thirty years after Darwin’s Origin of Species, de Vries had no idea what the
nature of these particles might be. He could not see them and could only infer their
existence from his and Mendel’s experiments. He called them pangenes. We now refer
to them simply as genes.7
De Vries’s breakthrough laid the foundation for the rapid development of the sci-
ence of genetics in the early twentieth century, including the recognition, achieved
independently in 1902 by Theodor Boveri and Walter Sutton, that the “particulate”
hereditary factors were borne on chromosomes. This was later confirmed by Thomas
Hunt Morgan and his team at Columbia University in New York, based in large part on
 63
gender

discoveries made in 1905 by Nettie Stevens and Edmund Beecher Wilson that the two
sexes in insects like the common mealworm also differ in chromosomes in their cells.
This was the first time that physical differences among organisms could be linked to
observable differences in their chromosomes, and it also explained Jacquin’s observa-
tions on ginkgo. The chromosomes in the cells of his graft bore the genes for “female-
ness,” while the chromosomes in the cells on the rest of the tree bore the genes for
“maleness.”8
Nettie Stevens observed that in the female larvae of her mealworms there were
twenty large chromosomes, while the male larvae possessed nineteen large chromo-
somes of equal size, with one of the tenth pair reduced in size. She called the short
chromosome of the unequal pair the Y chromosome and the longer of the two the
X chromosome. Both she and Edmund Beecher Wilson recognized that the XY con-
figuration results in males, while the XX configuration results in females, and we now
know that this same mechanism operates in humans.9
The rediscovery of Mendel’s laws, the recognition of the importance of chromo-
somes, the discovery of sex chromosomes, and Morgan’s work on fruit flies all came at
about the same time as an upsurge in interest in ginkgo. As a result, ginkgo was among
the first plants to be examined for its chromosomes, and it is now well established that
every living cell in a ginkgo plant has twenty-four chromosomes, twelve pairs: a dozen
brought by the male reproductive cell, which develops in pollen grain, the other twelve
contributed by the egg that develops inside the ovule.10
When I was a student, I was taught that just as in humans and mealworms, differ-
ences among one of the pairs of chromosomes in male and female ginkgo trees indi-
cated a system of sex chromosomes, which results in males and females in approxi-
mately equal proportions. However, careful observations over the past few decades
have shown that in ginkgo the supposed X and Y chromosomes are not consistently
different. Furthermore, even though both of these chromosomes have small, so-called
satellite fragments of DNA attached to them, which may be important for sex determi-
nation in other plants, in ginkgo these fragments vary in ways that make them unlikely
determinants of the sex of a mature tree.11
Nevertheless, whether visible or not, there must be real differences in the chromo-
somes of male and female ginkgo trees at the level of genes. This implies that the sex
of a ginkgo embryo is determined at the time that the egg is fertilized, just as it is in
humans. However, there is also evidence that sex determination in ginkgo may not be
 64
the living tree

quite as strongly fixed as it is in many animals, and even that it may not be completely
stable through the life of every tree.12
In the summer of 2006 a small twig on the Old Lion at Kew, a male tree, sponta-
neously produced three seeds. I published a note on this in the Kew Magazine later that
year and had letters and emails from several colleagues suggesting that this was prob-
ably the reemergence of an old graft rather than a sexual switch in part of the tree. This
certainly could have been possible because the practice of grafting female branches
onto male ginkgo trees became widespread in the late nineteenth and early twentieth
centuries, both for reasons of curiosity and to produce viable seeds. We know that such
a graft was made on the Kew tree in 1911, and it apparently fruited copiously before
it was pruned off accidentally by an overenthusiastic arborist. Similar uncertainties
about whether or not a graft was involved also apply to the large, nearly 200-year-old
male tree in the botanical garden in Jena, Germany, that began to produce seeds on a
single branch in the early 1990s.13
However, in other cases there is clear evidence of male ginkgo trees spontaneously
producing seed without human intervention. Martin Hamilton, a native of Kentucky
and one of the many wonderful students trained in horticulture during my time at
Kew, drew my attention to an example that piqued his interest as a young botanist. In
the eastern and southeastern United States some of the best places for marvelous speci-
men trees are historic cemeteries, and the Cave Hill Cemetery in Louisville, Kentucky,
is an excellent example. It boasts a wonderful collection of trees dating from the mid-
1800s, including several massive ginkgos. Among them is a large male tree, one of the
largest and most spectacular in North America. High in the canopy it has a witches’
broom, a dense mass of shoots that have somehow become feminized and that produce
large quantities of seeds every year. These seeds contain perfectly good embryos and
grow normally. Similar spontaneous partial sexual switches have been recorded several
times on old male ginkgo trees in Japan.14
Equally definitive evidence of the same phenomenon comes from the Ginkgo Plan-
tation at the Blandy Experimental Farm in Boyce, Virginia, where more than six hun-
dred ginkgo trees were planted between 1929 and 1947, all from seeds produced by a
large female ginkgo on the University of Virginia campus in Charlottesville. About half
the trees survived and began producing either pollen cones or seeds after twenty or
thirty years. Surveys to determine the sex of each tree were carried out in the late 1970s
 65
gender

and early 1980s, and in May 1982 an especially careful survey was made using a mecha-
nized lift so that the tree canopy could be carefully inspected.15
For all of those trees for which sex could be determined, the ratio was roughly one to
one: 157 female trees to 140 male trees. But the 1982 survey turned up a surprise. Four
trees recorded as females in previous surveys, based on the presence of seeds, turned
out to be predominantly male. These trees were watched especially closely in 1982, and
three of them produced from one to seven seeds each.
Disentangling the factors responsible for such leakiness in sexual expression is not
so easy, but it is perhaps not surprising that a male tree might occasionally produce
a few ovules. Over the long haul, this might be a useful trick if you are a lonely male
without a mate. It also does not completely undermine the normally strict outbreeding
enforced by the rigid separate sex system. At worst, a few ovules on a “leaky male” are
likely to use only a tiny fraction of the pollen grains that would otherwise have gone to
pollinate females, and at best such spontaneous production of ovules, if they mature
into seeds, might provide an intergenerational fail-safe when no mature females are
available.
From an evolutionary point of view, “leaky females”—female trees that occasionally
produce pollen catkins—would be a much less advantageous form of gender modifi-
cation. Pollen is produced in such large quantities that even a few rogue pollen cones
could result in massive self-pollination of the ovules on a mainly female tree and a
large-scale override of the normal system that ensures that sex is possible only be-
tween different individuals. “Leaky females” would also be more difficult to detect
than “leaky males” because pollen catkins are on the tree for only a short time early
in the season, but for the moment, the evidence suggests that while most ginkgo trees
are strictly male or female, when deviations occur, they usually take the form of males
producing seeds rather than the other way round. If this is true, ginkgo would fit the
pattern seen in other plants with separate male and female plants, as well as the theo-
retical predictions: “leaky males” are much more common than “leaky females.”16
 9
Seeding

If you can look into the seeds of time,

And say which grain will grow and which will not,
Speak then to me . . .
—William Shakespeare, Macbeth

I like to think of the old male ginkgo that grows at Kew as one of the celebrity ginkgo
trees of the world: a tree worth making a special effort to meet. As a living connection
to King George III and Sir Joseph Banks, as well as the early introduction of ginkgo
into the West, it would have a rich story to tell if you could engage it in conversation.
However, an equally special celebrity ginkgo grows on the other side of the world in the
Koishikawa Botanical Garden at the University of Tokyo. It is a living link to pre–Meiji
Era Japan. Keisuke Ito, who had been inspired by Siebold, would have known this tree
when he was appointed professor there. Planted more than three hundred years ago,
today it stands over eighty feet tall and has survived many vicissitudes, from the Great
Kanto Earthquake of 1923 to the firebombs that consumed much of Tokyo toward the
end of World War II. It is a precious old female tree, and it has a special connection to
early development of modern science in Japan.1
The Koishikawa Garden was founded on the site of what was once the medicinal

66
 67
seeding

plant garden of the Tokugawa Shogun. In 1868, at a time of great upheaval in Japan, the
land passed to the new imperial Meiji government, and the land became the botanical
garden of the newly founded Tokyo University. The transfer, however, was deeply un-
popular with the samurai, the traditional military class, who opposed the new regime.
They showed their displeasure at the changes that swept Japan in the 1860s in many
ways, but at the Koishikawa Garden one manifestation was the felling of some of the
large trees just before the time of transfer. The Koishikawa Ginkgo was very nearly
among those that were lost.2
Although a time of rapid and difficult social change, the late nineteenth century was
crucially important for the future development of science in Japan. After the installa-
tion of the Meiji government and the creation of a modern system of education, Japa-
nese scientists had their first opportunity to collaborate with colleagues in the West.
In plant science, strong early connections developed with leading botanists working
in Germany. Both Ryokichi Yatabe and Jinzo Matsumura, the first two professors of
botany at Tokyo University, studied in Germany in the late 1880s. A few decades later,
so did Manabu Miyoshi, who returned to found the discipline of plant physiology in
Japan. Also from this group came Kenjiro Fujii, who studied in Munich under Profes-
sor Goebel, did important work on ginkgo, and founded the Japanese scientific journal
Cytologia.3
Since the Meiji Restoration, the ginkgo at the Koishikawa Garden has been visited
by many scientific luminaries, but few have had a life quite as varied and colorful as
the precocious young Englishwoman Marie Stopes, who studied there in 1907 and
1908. Stopes came to Tokyo through her connection to Kenjiro Fujii. They met in 1903,
when both were working in Munich, and they did important research together on fossil
plants from Hokkaido. Both were familiar with the large female ginkgo in the Koishi-
kawa Garden.4
Marie Stopes is best known for her controversial pioneering work promoting family
planning in Britain, and her influential book Married Love, described by some as a sex
manual, was banned as obscene in the United States until 1931. In one survey of the
most significant books of the twentieth century Married Love scored only a little be-
hind Das Kapital and ahead of The Meaning of Relativity. Through her contributions
to women’s rights and her shaping of women’s expectations within marriage, Marie
Stopes is credited with almost single-handedly leading women in Britain out of the re-
pression of the Victorian Age into a more enlightened age of sexual awareness.5
 68
the living tree

However, Marie Stopes began her career focused on a more mundane pursuit: study-
ing plants. At the age of eighteen, she was granted a science scholarship at University
College, London, and she finished a double degree in botany and geology in only two
years. After completing her Ph.D. on reproduction in cycads, she became Britain’s
youngest doctor of science at the age of twenty-five. The University of Manchester
hired her immediately as a lecturer. She did important work on several different kinds
of extinct plants and also wrote a classic paper on different kinds of coals, which is still
widely used. In an era when women faced enormous obstacles in pursuing a career in
science, her geological and paleontological contributions were extraordinary.6
Stopes and Fujii worked under Goebel in Munich, but another important group of
botanists studying plant biology and evolution at that time was in Heidelberg under
the leadership of the great German botanist Eduard Strasburger, who became inter-
ested in the details of reproduction in ginkgo in the 1890s. Richard von Wettstein, pro-
fessor of botany at the University of Vienna, sent him seeds from the graft that Joseph
von Jacquin had made many years earlier. They were dispatched every two weeks from
June through to early September, and based on this material, Strasburger described
many aspects of sexual reproduction in ginkgo. However, he missed a crucial detail; a
detail filled in only later by Sakugoro Hirase, working at the Koishikawa Garden.7
Hirase was an accomplished technician and illustrator. In 1895 he published a
meticulous account of how the embryo developed into a new young plant inside a
ginkgo seed. Through the summer of 1896 Hirase kept up an especially detailed study
of the seeds collected from the Koishikawa tree, and on September 9, 1896, he was the
first to observe the previously overlooked final stage of sexual reproduction in ginkgo.
What Hirase saw was a botanical sensation: pollen grains drawn into the ovule by
the pollination drop produced a branching tube that had grown into the tissues at the
top of the ovule. The bulging base of the tube, hanging down on the inside of the ovule,
contained a pair of large sperm cells. Hirase saw that these two sperm were released
into a cavity in the top of the ovule. Propelled by the synchronized movements of spi-
ral bands of thousands of tiny flexuous hairs, they swam the short distance to fertilize
the egg. He recognized immediately that this curious means of fertilization was unlike
that of any other plant known at that time. Swimming sperm were well known in ferns,
mosses, and similar plants but had never been observed in any seed-producing plant.8
Hirase’s discovery was followed just a few months later by the report of exactly the
same phenomenon in the common Sago Cycad by Seiichiro Ikeno, who was work-
Sakugoro Hirase, the Japanese scientist who was the first to
observe the swimming sperm of ginkgo in 1896.

Close-up photograph of two swimming sperm just before their

release. Each sperm cell is a little less than four thousandths of
an inch long and is propelled by the synchronized movements
of thousands of tiny hairs arranged in a spiral.
 70
the living tree

ing in the College of Agriculture of the Imperial University, Tokyo, and who had been
guided by Hirase’s work. Both discoveries were remarkable. They were also two of the
first research contributions of Japanese scientists that made an impact on the inter-
national stage. In 1897 Ikeno and Hirase summarized their results in a joint paper in
English, and in 1912 they were the first biologists to receive the Imperial Prize of the
Japan Academy.9
From the standpoint of understanding plant evolution, the discoveries of Ikeno
and Hirase were an unanticipated breakthrough. Until their work it had been assumed
that fertilization in ginkgo and cycads was like that in conifers, the male sex cell being
delivered to the egg through a tube growing from the pollen grain. The discovery of
swimming sperm showed that reproduction in ginkgo and cycads was very different
and more like that of ferns: it harked back to an earlier time in plant evolution when
plant reproduction was more reliant on water.
Marie Stopes, who was at Tokyo Imperial University just a few years after Hirase
made his discovery, and who published a diary of her experiences in Japan, refers sev-
eral times to the excitement of repeating Hirase’s observations. Her entry for Septem-
ber 17, 1907, records: “This is just the last day or so for the swimming out of the sper-
matozoids of Ginkgo, and I spent a few delightful hours in the laboratory watching
their infusorian-like movements and the quick vibrations of their spiral crowns of cilia.
Fancy cutting up dozens of juicy Ginkgo seeds!”
A year later, her entry for September 9, 1908, alludes to the effort needed to catch
fertilization in ginkgo in the act: “I went early to the Institute, where there is grand ex-
citement over Ginkgo; the sperms are just swimming out, and they only do it for a day
or two each year. It is no such easy business to catch them, in 100 seeds you can only
get five with sperms at the best of times, and may get one and be thankful. I spent pretty
well the whole day over them, and got three, and several in the pollen tube not yet quite
ripe. It is most entertaining to watch them swimming, their spiral of cilia wave ener-
getically.” She spent the next two days “hunting Ginkgo sperms nearly all the time.”10
The significance of Hirase and Ikeno’s work was further underlined by a paleo-
botanical discovery made just a few years later in Britain. For many years botanists had
puzzled about why many of the large fossil fern leaves discovered in the coal measures
of Europe and North America never had the spore-producing clusters on the under-
side of the leaf that are typical of most modern ferns. In 1903, F. W. Oliver, working at
University College, London, and D. H. Scott, working at Kew, solved the problem by
 71
seeding

The young Marie Stopes working at her desk in 1904, a few years
before she traveled to Japan for her botanical work at Tokyo Imperial University.
Her pioneering work on family planning in Britain came later in her career.

convincingly linking one such leaf with a fossil seed, thus demonstrating that many
so-called fossil fern leaves were not produced by ferns at all. Their work revealed the
existence of several different kinds of extinct plants, that were, in a way, intermediate
between ferns and living seed plants. Such fossil plants came to be called Cycadofili-
cales, or seed ferns. Together, the discoveries made by Hirase and Ikeno on ginkgo and
cycads, and by Oliver and Scott on their fossils, went some way to narrowing the evo-
lutionary gap between living ferns and living seed plants.11
This line of thinking was also reinforced by reports of aberrant ginkgo trees that
occasionally produced seeds and pollen sacs on the margins of otherwise normal-
looking leaves. In 1896 Kenjiro Fujii described three such trees from Kai Prefecture in
Honshu, Japan, and since then others have noted the same phenomenon. Fujii sug-
gested that the formation of seed or pollen sacs on the margins of leaves, together with
 72
the living tree

Hirase’s discovery about the fertilization process, might suggest that ginkgo evolved
from fernlike ancestors.12
An interesting question that has loomed large in the thinking of some botanists is
whether fertilization in ginkgo occurs while the seeds are still attached to the tree, as
is normal in living seed plants, or whether it occurs after the seeds have been shed, as
suggested by the French paleobotanist Louis Emberger. In some ways this would be
more like what happens in ferns, but in ginkgo this is clearly not normal. Fertilization
usually occurs in late summer or early autumn, and most seeds are not shed until a
month or two later, by which time the embryo is already well developed. That said, it is
not impossible that fertilization does occasionally occur on the ground. After pollina-
tion, the developing seeds grow almost to their mature size before fertilization. At the
later stages of their development—for example, in August—young seeds do occasion-
ally fall from the tree, and if that happens just prior to the release of sperm, then viable
embryos might still be produced.13
In Chicago, most ginkgo seeds are on the ground just as the weather is turning cold
in mid- and late November. On a few trees the seeds fall more gradually through the
winter. However, whether it is in the late autumn or after a winter storm or cold snap,
when there is a layer of ginkgo seeds on the ground, no one could mistake them for
apricots, plums, or anything else. The smell is distinctive and powerful. A plant chem-
ist would describe it as rich in butyric acid. Others would recognize a strong similarity
to the unmistakable stench of human vomit. A friend who is a councilor in southwest
London had to convince local residents that the smell about which they were complain-
ing was not the result of binge drinking by hooligans outside her local train station but
the equally malodorous output of a large female ginkgo. And if the smell isn’t enough
incentive to avoid the fallen fruits, the pulp contains ginkgolic acid, an allergy-causing
chemical. Like poison oak or poison ivy, it can produce a nasty rash. If you get it in
your eye it can put you in the hospital.14
 10
Resilience

Strength . . . comes from an indomitable will.

—Mahatma Gandhi, Non-Violent Resistance (Satyagraha)

Brian Mathew is one of those botanical enthusiasts for whom a lifetime at Kew was
an irresistible attraction. People who somehow have plants in their blood often end up
working at Kew for their entire careers, and their connections to the place, through
their friends and through their plants, persist long after retirement. In Brian’s case,
most of his working life was spent in the vast collection of preserved plants at Kew.
Begun by William Jackson Hooker, this massive herbarium now contains about eight
million specimens of dried plants. It may be the largest of its kind in the world, and
Brian’s role was to use that massive reference collection to accurately name the plants
being grown in the gardens by Kew’s horticulturalists.1
For Brian, as for many people at Kew, plants are not just a job but also an avocation,
and his botanical passion finds expression in many different ways. He became an ex-
pert on bulbs, like crocus and snowdrops, and this led him to work on their conserva-
tion in countries like Turkey and Georgia, where bulbs are collected from the wild for
export into the horticultural trade. He also wrote several books, becoming a successful
author; like many at Kew, of course, he also became a passionate gardener.
In 1999, Brian was asked by friends in his village to design and plant a garden on the

73
 74
the living tree

land around his local church. Rarely able to say “no” and always willing to help, he set
to work. No money was available, but some volunteers gave a hand with the planting,
others pitched in with donations of plants. One such donation was a young ginkgo,
growing in a nearby garden that was becoming too big for the space available. It was a
small tree, healthy, well established, and about ten feet tall. Moving a tree, even a rela-
tively small one, is always harder than it looks, but Brian took up the challenge. He
thought that if the root system was reasonably well developed, the tree would stand a
good chance of survival if he dug it up while it was dormant in the winter. Brian’s idea
was to move it first into a pot where it could be cosseted for a season before being re-
planted.
Unfortunately, what seemed straightforward in theory turned out to be harder in
practice. The first problem was that over the years the soil had been mounded up
around the trunk. As Brian dug down, he had difficulty finding any roots at all, let
alone getting to the bottom of the main root mass. Eventually, with the hole around
the tree a couple of feet deep, and deeper digging difficult without uprooting nearby
plants, Brian came across just a few small side roots coming off the main trunk. Below
that the trunk just continued down. At this point, Brian felt that he could go no farther
without impinging too much on the generosity of the donors and disrupting much of
their garden. So, warning that the tree probably would not survive the move, he cut the
trunk below the small side roots and planted it in a large pot. As an experienced gar-
dener Brian knew that in order to maximize the young tree’s slim chances of survival,
he needed to reduce its crown dramatically. If the tree leafed out the next spring, one
or two roots were not going to be enough to keep all the leaves supplied with water. So
Brian pruned the ginkgo severely, placed the pot out of direct sun, and waited.
To his surprise the young tree burst into leaf the following spring. Healthy leaves ap-
peared on the unpruned branches, and the plant looked fine through the summer and
into the autumn. The next winter he found a good place to plant it, in an area used by
children of a preschool. He thought that the ginkgo’s distinctive leaves would make it
an entertaining curiosity for the youngsters. In the pot the roots had developed well,
there was now a nicely developed root mass, and the plant was flourishing despite its
earlier trauma.
The next summer the tree again leafed out beautifully. Unfortunately, however,
sometimes young people and young plants don’t mix. The tree got snapped off, right
in the middle of the trunk, about four feet above the ground. The plant was now just a
 75
resilience

bare trunk, broken at one end with no branches and no leaves. There was nothing for
it; it had to go back into intensive care. Brian dug it up for the second time, planted it
back in its pot, and waited to see whether it would recover. Again it came back from the
dead; this time sprouting at soil level from around the broken stem. Soon there were
several new trunks, and its recovery was rapid and long-lasting.
Brian’s experience with the indomitable resilience of ginkgo is not unusual. Its
tenacity in the face of abuse is one of the reasons that it does well as a street tree. It is
also easy to propagate from small cuttings, which readily take root and establish new
plants. This is probably how ginkgo was spread from garden to garden when it was first
introduced into Europe. And as Brian found out, ginkgos have a peculiarity of their
anatomy that creates new opportunities for growth if the main part of the plant is dam-
aged. Ginkgo, in other words, has an alternative way of reproducing itself that does not
require completing the cycle of sexual reproduction, with all its complexities, risks, and
potential for failure.
In a few trees, such as willows, such mechanisms of self-propagation are especially
well developed. Willows, often streamside plants, perpetuate themselves easily in the
wild. Twigs break off, get washed downstream, become lodged on a bank, and then take
root. A relative of the willow, the aspen, is equally vigorous. It sends up new branches
from its roots that look like separate trees, but they are all connected belowground and
are genetically identical. The establishment of new plants in this way is not common
in ginkgo, but it does have a similar inbuilt means of sustaining itself. This allows it to
occupy a single place for a long period, and as in the case of Brian’s sapling, if the main
branch is damaged, new branches soon form and can quickly take over.2
A characteristic feature of some ginkgo trees, and especially of very old specimens,
is the production of peculiar downward growths from otherwise normal branches.
These develop like stalactites from the roof of a cave. Several may develop along a
single branch, giving the Daliesque impression that the branch is made of wax and has
somehow begun to melt. Some of these downward growths, or burls, may be six feet or
more from top to bottom, and as each reaches the ground, it can resprout and send up
new shoots. Each may eventually develop into a substantial new trunk that can become
separated from the parent tree. In very old trees, where these growths are extensive, the
original main trunk can sometimes be hard to find.
In Japan these downward-growing branches are called chi- chi, literally “breasts,”
from the folklore that mothers who pray to them can increase their breast milk. In
 76
the living tree

China they are called zhōng rǔ, “stalactites.” It is not known what stimulates their
growth, but careful studies show that they form from small buds that become em-
bedded in the wood of a branch as it grows and that somehow are later reactivated.
Every leaf of a ginkgo tree has a small bud, a potential new shoot, in its axil, and as a
branch increases in girth, buds that were once at the surface sometimes become buried
by the tissues around them. While ginkgo trees rarely develop chi-chi until they are
very old, the potential of regrowth from embedded buds seems always to be present in
latent form.3
Peter Del Tredici, one of the world’s experts on ginkgo, who works at the Arnold
Arboretum of Harvard University, has shown that such embedded buds are established
from the seedling stage onward, right at the beginning of the life of a new plant, and are
a normal feature of growth in ginkgo. In all ginkgo seedlings the tiny buds that occur
in the axils of the seedling leaves, the first leaves produced by the embryo, remain in-
active and become embedded in the base of the tree as it grows. If the sapling is then
damaged in some way, as happened to the tree nursed along by Brian Mathew, one of
these leftover buds grows downward from the base of the trunk to produce a woody
basal chi-chi, a so-called lignotuber, which can produce new trunks and branches, as
well as new roots. Del Tredici suggests that these lignotubers are also helpful anchors
on unstable soils.4
The combined effect of extensively developed chi-chi and the associated capacity for
rampant self-propagation is seen most spectacularly in some of the remarkable giant
ginkgos that grow in Aomori Prefecture in northern Honshu, Japan. One of the most
impressive is the Kitakanegasawa Ginkgo, a real giant that grows squeezed between
the crowded houses of Kitakanegasawa village and the main coastal road. It flourishes
right at the point where the steep mountain slopes join the narrow coastal plain. It is
only a few hundred yards from the sea, but its roots draw water from natural fresh-
water springs, and it has a sheltered spot tucked into the lee of the Cape of Odosesaki,
where it is protected from the full force of the winter storms that roll in from the west.
From the small parking lot built for visitors by the local Department of Tourism, the
Kitakanegasawa Ginkgo is a picture of health. The crown is huge, vigorous, and flour-
ishing, with few dead branches or yellow leaves, but inside its personality changes. The
inner crown contains lots of dead wood, and the internal scaffold of the tree reflects its
irrepressible growth. Multiple trunks and branches, many of them snapped off, show
the battering that the tree has taken over centuries from heavy snow, from typhoons,
 77
resilience

The solid trunk of a large ginkgo, showing a mass of thin sucker shoots growing up from the base.
This is one of several large ginkgo trees in and around Shanghai, China.

perhaps even from earthquakes. A tangle of roots shows through where the thin soil
and leaf duff have been worn away by the feet of tourists.
Even the largest ginkgos do not come close to the vast size of a coast redwood, or the
great age of a bristlecone pine, but a truly massive ginkgo, like that at Kitakanegasawa,
is nevertheless a great natural wonder. This is a tree that you walk into, rather than be-
neath, and as you enter you are engulfed by a complex tangle of large upward and hori-
zontal branches, mixed with chi-chi growing downward from the larger limbs. Every-
where there are slender, vigorous, upward-pointing sucker shoots sprouting from the
trunks and larger branches. The Kitakanegasawa Ginkgo is less like a tree and more
like a thicket; the combined circumference of its trunks is more than seventy-two feet.
 78
the living tree

Among them are three substantial new trunks, already clothed by sucker shoots that
are starting life as opportunists from their mother. With their potential for indepen-
dence, they provide good insurance for the future.5
So ginkgo, through evolution, has hedged its bets. It can survive and propagate
itself in different ways. The cycle of pollination, fertilization, dispersal, germination,
and establishment is a seemingly fragile process. It relies on precise synchronization
in the development of pollen and ovules borne on different trees, and benign con-
ditions for seedling development and growth, but evidence from prehistory shows
that it works well. These processes have gone on continuously, and probably in ways
not much different from those of the living species, for more than 200 million years.
Female ginkgo trees readily produce large quantities of seed, even with just a single
male in the vicinity. In ginkgo, as in some other species, males are almost, but not
quite, expendable. Ginkgo also has the fail-safe that at least on the male side it might
very occasionally produce a few seeds. It is not beyond the bounds of possibility that
very, very occasionally a single seed might produce two new plants, one male and one
female.
Ginkgo is also resistant to physical abuse, which means that time is on its side.
Potentially it can wait until its mate arrives. It resprouts vigorously from buds buried
in its underground parts. It is irrepressible; its capacity for self-preservation has helped
it survive through millions of generations.
The Kitakanegasawa Ginkgo exemplifies the resilience of its species. Local people
think that it is the biggest in the world. It was designated as a national treasure only in
September 2004. Somehow, it was overlooked in the initial national surveys of Japan’s
great ginkgos. It is now recognized as the fourth-largest tree in Japan, and it is certainly
the most massive ginkgo. One enthusiast is gathering the evidence that he hopes will
give it a place in the Guinness book of world records. But in a world where so much of
the landscape has been altered by human activity, more important is that the connec-
tion of local people with this marvelous tree is deep. To them, and indeed to anyone
with empathy for nature, this great plant is a testament both to its tenacity and also to
its lasting power.
 part iii

Origin and Prehistory

 OVERLEAF Reconstructions of three different extinct ginkgolike plants discovered
and described by Zhou Zhiyan from the Middle Jurassic, about 170 million years before the present,
at the Yima mine in Henan Province, China. Left: Ginkgo yimaensis; right: Yimaia recurva;
bottom right: Karkenia henanensis.
 11
Origins

To every thing there is a season, and a time to every purpose under the heaven:
a time to be born, and a time to die . . .
—Ecclesiastes 3:1–2

Carl Linnaeus, the eighteenth-century naturalist who coined the name Ginkgo bi-
loba, stands with Arrhenius and Celsius as one of the foremost of all Swedish scientists.
When you arrive at Arlanda Airport in Stockholm, Linnaeus is among the collage of
well-known Swedish citizens who welcome you to their “hometown,” and his image ap-
pears on the one hundred–kronor banknote. Linnaeus spent his entire life in Europe,
but his name is known worldwide, in large part because of his intrepid students who
journeyed all over the globe to collect plants and bring them back for study. Linnaeus
was the leader in efforts to make sense of the rapidly increasing knowledge of plant and
animal life that came from eighteenth-century European exploration in far-flung parts
of the world.
Linnaeus was also an omnipresent force in Sweden. He held sway at the University
of Uppsala for decades and, as physician to the king, had the ear of the royal family.
He was a charismatic teacher who inspired his students. His natural history excursions
into the countryside around Uppsala were legendary. There would be music and lavish
picnics as Linnaeus held forth about the trees, the flowers, the birds, and the insects.

81
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origin and prehistory

Linnaeus could be vain and self-important, but there is no doubting his intellect or his
extraordinary energy.
Linnaeus was among the founding members of the Royal Swedish Academy of Sci-
ences, which today is best known for awarding the Nobel Prizes in physics, chemistry,
and medicine. However, biology remains an important part of the academy’s work,
and its interest in the diversity of plants continues through its support of the nearby
Bergianska Botanical Garden and its links, through history and present-day members,
with the Swedish Museum of Natural History. Each of these three great Swedish institu-
tions—the academy, the Bergianska Garden, and the Museum of Natural History, clus-
tered together in Frescati on the northern edge of Stockholm—has played an impor-
tant role in expanding our understanding of ginkgo and its place in plant evolution.1
The Swedish Museum of Natural History houses one of the world’s great collections
of fossil plants. Successive scientists who have worked there have been leaders in paleo-
botany since the renowned Arctic explorer Alfred Nathorst was appointed as professor
in the newly founded Department of Archegoniates and Fossil Plants in 1884. Much of
what we now know about plants of the past can be traced to the discoveries of Nathorst
and the techniques for studying fossils that he developed.2
My longtime colleague Professor Else Marie Friis now occupies Nathorst’s former
position at the Swedish Museum of Natural History and oversees the massive col-
lections of fossil plants accumulated by him and his successors. Today a quarter of a
million specimens from all over the world are housed in more than seven thousand
drawers and hundreds of cabinets on three floors of a building that Nathorst helped
design. The collections are matched in size, scope, and quality only by those in the
Natural History Museum in London and the U.S. National Museum of Natural History
in the Smithsonian Institution in Washington, D.C. Together with collections in other
museums and universities, these great storehouses of plant fossils are used by scientists
from all over the world and house the physical samples on which our knowledge of the
history of ginkgo and other plants is based.
In the Stockholm collections are ginkgo leaves collected by Nathorst on expeditions
to Greenland and elsewhere. There are also beautifully preserved specimens collected
by him in the late eighteenth century during the heyday of coal mining in southern
Sweden, but among all the ginkgo material in the Stockholm Museum, the most beau-
tiful specimen is a slab of gray siltstone collected in the 1970s from Ishpushta in cen-
tral Afghanistan. Bequeathed by the German paleobotanist Hans Joachim Schweitzer,
 83
origins

Fossil leaves of an ancient ginkgo, Ginkgo cordilobata, from the Early Jurassic of Ishpushta,
Afghanistan, about 190 million years before the present.

it is covered with the beautiful shiny black imprints of more than eight almost com-
plete ginkgo leaves, which Schweitzer and his colleague Martin Kirchner named Ginkgo
cordilobata.3
The great ginkgo slab from Afghanistan comes from rocks of Early Jurassic age that
were laid down near the shores of an ancient sea around 190 million years ago. Each
leaf is divided into six segments, and each segment is itself deeply bilobed. They look
more like the leaves of ginkgo seedlings than like the leaves of mature trees, but there
is no dispute about their identity; it does not take a specialist to immediately see the
link to the familiar tree of our streets and gardens. This slab alone is enough to suggest
the great antiquity of the ginkgo lineage, but it raises a still more fundamental question:
where did ginkgo come from?
Through studies of living and fossil plants we now have a reasonably clear outline
of the main events in the history of plant life and a rough idea of when they took place.
In a few cases, fossil plants have also turned out to be key missing links that help con-
 84
origin and prehistory

nect seemingly separate pieces of the botanical evolutionary puzzle. For example, in
the same way that Archaeopteryx from the Jurassic helps link modern birds to carnivo-
rous dinosaurs, Archaeopteris, a fossil plant with a confusingly similar name, helps link
living seed plants to strange extinct trees that reproduced by spores, like modern ferns,
rather than by seeds. And while our understanding of the early evolution of animals
has been informed by research on exquisitely preserved fossils from the Burgess Shale
in British Columbia, studies of fossil plants from the Rhynie Chert, an equally remark-
able example of exceptional preservation in Scotland, have added new depth to what
we know about the early evolution of plants.
Discovered by the geologist William Mackie in 1912, the Rhynie Chert lies beneath
a grassy slope just outside the small Scottish village of Rhynie, not far from Aberdeen.
The chert is not even visible at the surface, but fortunately Mackie was an alert field
geologist who noticed strange-looking rocks scattered in the field and built into nearby
walls. Their source was later confirmed by digging trenches. More recently a team from
the University of Aberdeen has drilled a series of holes to understand how this unique
deposit was formed.4
The Rhynie Chert preserves the fossilized remains of a series of ancient peat bogs,
one on top of the other, from about 400 million years ago. Different layers preserve en-
tire ecological communities, with plants, algae, and fungi alongside a variety of spider-
like creatures, as well as the oldest known insect. Everything is completely embedded
in hard, glasslike silica exactly where it once lived, and many of the fossils are preserved
with exquisite fidelity.5
The paleontologist Stephen Jay Gould devoted a whole book to the significance of
the Burgess Shale for understanding the early history of animal life. The Rhynie Chert
has had no such booster, but it is equally important for understanding the early history
of plants. The five papers written on the fossil plants of the Rhynie Chert between 1917
and 1921 by Robert Kidston of the British Geological Survey and William H. Lang of
the University of Manchester are classics of plant science. Together with superb work
done toward the end of the twentieth century by others, especially Winfried Remy at
the University of Münster, they revealed peculiar plants and important clues about the
early development of life on land.
The Rhynie Chert provides gloriously detailed evidence that by around 400 million
years ago plants had already begun to move out of the ponds, puddles, and tide pools
in which they had evolved and established a significant foothold—roothold—on the
 85
origins

land. All these plants are small, none more than a foot or so tall, and, compared with
most plants of today, all are very simple in structure. Most consist of slender branches,
sometimes with a capsule at the tip of each branch in which the spores were produced.
A few show features reminiscent of living mosses or hornworts, while others, such as
Asteroxylon, one of the most distinctive of the Rhynie Chert plants, are very like mod-
ern clubmosses. However, there is no hint that any of these small, simple plants was
able to form wood, and none have large, complicated, leaves such as we find today in
ferns, cycads, or ginkgo. The Rhynie Chert captures an early moment in plant evolu-
tion before the origin of trees, large leaves, and many other features that we take for
granted in our modern world.6
Before the Rhynie Chert, even earlier evidence that plants were beginning to make
the transition from living in water to living on land comes from dispersed plant spores
with tough resistant walls. These spores are first recognized in the Middle Ordovician,
about 450 million years ago, and are similar to those of some living liverworts. A little
later, toward the end of the Silurian, about 420 million years ago, minute and very frag-
mentary plant fossils, often no bigger than the head of a pin, provide direct evidence of
the tiny simple plants that produced some of these spores and that were the probable
progenitors of the Rhynie Chert plants.7
The Rhynie Chert seems to have been formed near hot springs like those occurring
today in Yellowstone National Park in the United States, or at Rotorua in New Zealand.
Plants growing in the peat were intermittently flooded with hot water and rapidly im-
pregnated with silica. In places the plants were dead and had already decayed before
they were fossilized. Occasionally, however, living plants must have been overwhelmed,
and fossilization evidently occurred within a few days or even a few hours. The sudden
engulfment preserved delicate structures and ephemeral stages of ancient life cycles
that can be hard to observe even in living plants.
All of the plants from the Rhynie Chert show the basic structural specializations
that help make plant life possible on land. These include a water-resistant, waxy outer
covering, the cuticle, which helps reduce water loss, and breathing pores, stomata,
which regulate the exchange of carbon dioxide, water vapor, and oxygen between the
environment and air spaces inside the plant tissues. Most of the Rhynie Chert plants
also have elongated cells in the center of their stems: by analogy with living plants, the
innermost of these helped transport water from the soil to the aerial parts, while the
outer ones carried the sugars produced by photosynthesis to both the aboveground
 86
origin and prehistory

and belowground parts of the plant. So by 400 million years ago, many of the fun-
damental structures and processes seen today in ginkgo and other living land plants
were already in place, and that these characteristics are shared by almost all land plants
strongly suggests that plants, unlike animals, colonized the land only once.
The evolution of trees from small, simple plants like those preserved in the Rhynie
Chert was probably driven mainly by competition for light. Tall plants have several
potential advantages that may give them an edge in the struggle for survival, such as
improved opportunities for spore dispersal, but light is indispensable for plant growth,
and plants that are taller than their neighbors have access to more of it. During the De-
vonian, several different groups of plants show a tendency toward increased size, aided
by different innovations that provided different kinds of structural support. Several
groups also developed large complex leaves, not too different from those of ginkgo,
cycads, and ferns, probably stimulated by the advantages that came from greater effi-
ciency in harvesting the sun’s energy. Such leaves apparently developed from elabora-
tions of the simple branching seen in the Rhynie Chert plants.
In hindsight, major changes, such as the origin of leaves and the origin of trees, seem
like dramatic evolutionary breakthroughs, but it is not hard to imagine how they could
have come about rather gradually through the process of natural selection envisaged
by Darwin. For example, in Asteroxylon from the Rhynie Chert, the elongated cells in
the center of the stem have characteristic, often more or less spiral, internal thicken-
ings, just like the water-conducting cells of most living plants. Initially, in the lives of
individual plants growing more than 400 million years ago, water may occasionally
have been in short supply; this seemingly minor difference may have been enough to
give plants with weakly developed internal thickenings in their conducting cells a slight
ecological edge over their neighbors. It might have been just sufficient to help the cells
withstand pressures that would otherwise have caused them to collapse.
Over time, providing that the slight difference improved survival and could be in-
herited and passed on to the offspring of the survivors, the internal support of these
cells may have been further strengthened in successive generations. Eventually, the
same specialized cells may have been co- opted by natural selection to support increas-
ingly large stems. The thickenings that initially helped prevent the cells from collapsing
now provided an advantage in structural support. The elaboration of such cells, their
production in large numbers, and their eventual modification for both water conduc-
tion and support allowed plants to grow into trees and also develop large leaves.
 87
origins

Key dates in the early history

of land plants between about
500 million and 300 million
years ago.

Similar kinds of contingent processes, building one upon the other, would also have
been at work as plant reproduction became ever more attuned to life on land. Plants
able to withstand desiccation would have survived preferentially when small freshwater
ponds and damp places dried up, so it is no surprise that all of the land plants found
in the Rhynie Chert, as well as their precursors from the Ordovician and Silurian, pro-
duced spores with tough walls. Resistant to drying out, dispersed by wind through the
air and by water through the soil, these spores would have been the vehicle by which
early plants survived tough times, and they would also have provided the means by
which successive generations moved from place to place.8
When conditions were right, the spores would have germinated to produce a new
free-living plant, but this time one that produced sex organs rather than spores. Re-
markably, a few of these gamete-producing phases in the life cycle have been discov-
ered in the Rhynie Chert, and in some it is just possible to make out the young sperm
developing inside the male sex organs. The sperm cells would have swum through soil
moisture to fertilize the eggs, exactly as happens today in mosses, liverworts, horn-
worts, clubmosses, and ferns. The motile sperm of the Rhynie Chert plants would have
been a holdover from more fully aquatic ancestors, just as they are in ginkgo and in
humans.
The evolution of seeds introduced new levels of complexity to the rather simple
 88
origin and prehistory

kind of sexual reproduction seen in the Rhynie Chert plants. It enabled further inde-
pendence from water in reproduction, allowed for subtle modification of spores into
pollen grains, and ultimately facilitated the de facto transfer of male gametes through
the air. It also meant that the spore, from which the egg cell would ultimately be pro-
duced, could be retained and protected on the parent plant. This opened up new pos-
sibilities to support the growth of the spore, as well as the subsequent growth of the
embryo, from the resources of the parent. These innovations effectively eliminated the
free-living, gamete-producing phase of the life cycle and meant that male gametes no
longer had to swim through soil moisture for fertilization to occur. They also created a
new organ of dispersal, the seed, containing a prepackaged young plant.
Seed plants with this new way of reproducing themselves are first known from about
360 million years ago in the Late Devonian. They foreshadow the kind of sexual repro-
duction seen in ginkgo. Taken at face value, this sets a rough lower limit on the pos-
sible age of the ginkgo lineage. The leaves of Ginkgo cordilobata from Afghanistan set
an upper limit of around 190 million years. One challenge for understanding the origin
of ginkgo is to pinpoint the age of the lineage more precisely. Another is to place the
living tree in its proper evolutionary position in relation to other plants.
 12
Ancestry

Is it on his grandfather’s or his grandmother’s side

that the ape ancestry comes in?
—Samuel Wilberforce, Wilberforce-Huxley debate, June 30, 1860

To connect living ginkgo to the extinct seed plants that were alive between about
190 million and 360 million years ago, the traditional approach is to look for ances-
tors, beginning with those fossils that seem securely related to modern ginkgo, then
working outward and backward to consider other fossils that might provide a con-
nection to other kinds of seed plants. Ginkgo cordilobata from Afghanistan shows that
plants with leaves like those of living ginkgo had already evolved by the Early Jurassic,
between about 175 million and 200 million years ago. Similar leaves are also known,
slightly earlier in the fossil record, from the Late Triassic, and especially interesting are
rich and beautiful collections of fossil plants from rocks of the Molteno Formation in
the Karoo Basin of South Africa.1
The Molteno fossils were collected and studied over many years by John and Heidi
Anderson at the South African National Biodiversity Institute in Pretoria. Their mas-
sive collections, more than twenty-seven thousand specimens, come from almost
seventy different localities that stretch in an ellipse from Little Switzerland and Golden
Gate in the north of the Karoo Basin, to Askeaten, Aasvoëlberg, and Bamboesberg in

89
 90
origin and prehistory

the south. The fossils are preserved as beautiful impressions in dark to yellowish gray
clays and silts. The quality of their preservation is much less impressive than the much
older Rhynie Chert, but what these fossils lack in details of structure they make up for
in the numbers of specimens available and the care with which huge collections have
been assembled.
Collecting the fossil plants of the Molteno Formation was the work of a lifetime for
John and Heidi Anderson. Both grew up in South Africa and despaired of the apartheid
system, but in their professional lives, through decades of near scientific isolation, they
spent many summers and countless weekends befriending farmers and ranchers and
exploring their lands for new localities. The specimens were brought back to their labo-
ratory in Pretoria, where they were carefully catalogued, described, and photographed.
Their first major report focused on the most common fossils in the Molteno Forma-
tion. The second placed the Molteno in the context of all other fossil plant occurrences
from South Africa. Most recently, they published two magnificently illustrated cata-
logues of the rich variety of extinct Molteno seed plants and ferns that lived in the lush
forests and floodplains of what is now the dry southern tip of the African continent.2
Fossil ginkgo leaves are present at about one of every five localities in the Mol-
teno Formation. The Andersons recognize six different species, which they cautiously
assign not to Ginkgo itself but to a very similar genus, Ginkgoites, that paleobotanists
often use for ginkgolike leaf fossils. Some of these leaves, such as Ginkgoites koningen-
sis and Ginkgoites matatiensis, are similar to Schweitzer and Kirchner’s Ginkgo cordi-
lobata from Afghanistan. Others, such as Ginkgoites muriselmata, have leaf lobes that
are more sharply pointed. Ginkgoites telemachus is a variation on the same theme, but
with leaf lobes that are irregularly toothed. All these species, at least in terms of their
leaves, are unmistakably related to living ginkgo, and they occur repeatedly with seed-
bearing structures, which John and Heidi named Avatia, and probable pollen catkins,
which they called Eoasteria. As far as we can tell these seeds and catkins are not very
different from those of living ginkgo, and their relationship to the living tree seems
pretty secure.
The Molteno fossils date from about 220 million years ago; they extend the fossil
record of ginkgo back about 30 million years before Ginkgo cordilobata. Fossil ginkgo
leaves of roughly similar age are known from many other parts of the world, including
Arctic Canada, eastern North America, the southwestern United States and northwest-
ern Mexico. Ginkgolike wood has also been described from just outside the Petrified
 91
ancest ry

Forest National Park in Arizona. However, farther back, obvious ginkgo fossils quickly
disappear. According to the Andersons’ compilations, the oldest occurrence of ginkgo
leaves in South Africa is in the early part of the Middle Triassic, about 240 million years
ago. In the Southern Hemisphere as a whole early ginkgo is widespread and the earliest
record is from the Sydney Basin of Australia, dating from near the end of the Early Tri-
assic at about 245 million years ago. There are no older records in the Northern Hemi-
sphere, and before this the obvious fossil history of ginkgo peters out.3
Tracing the ancestry of living ginkgo still farther back is difficult. The fossils become
less like the living tree, and their relationships become less secure. Rudolph Florin, one
of the great paleobotanists of the twentieth century, a member of the Royal Swedish
Academy and director of its Bergianska Garden, was among the first who were bold
enough to make a strong link between ginkgo and fossil plants from the Permian, the
geologic era before the Triassic. Florin focused on a kind of fossil plant that had first
been described more than seventy years earlier by the brilliant French paleobotanist
Gaston de Saporta.
Rudolph Florin is best known for his contributions to understanding the history
and evolutionary position of living conifers, but his most important work on ginkgo
was published in 1949, based on an examination of three fossils similar to those that
Saporta had named Trichopitys heteromorpha. All of the original material of Tricho-
pitys, including that examined by Florin, is from old coal mines in the region around
Lodève, southern France, which are Early Permian in age and produce fossils that date
from about 275 million to 290 million years ago. Saporta had already suggested that
Trichopitys might be related to ginkgo based on the similarity of the ribbonlike leaves
to some deeply divided but unmistakably ginkgolike leaves from much younger rocks.
Florin supported this idea and added the point that some of the leaves seem to have a
small branch attached near their base, and that some of these branches have seeds at
their tips. He compared these branches with the seed-bearing shoots of living ginkgo,
and also pointed out that living ginkgo sometimes produces abnormal, branched seed-
bearing shoots that may bear as many as ten small seeds.4
On a warm summer day in 1986 I had the chance to examine the original material
of Trichopitys described by Saporta from Lodève in the collections of the Natural His-
tory Museum in Paris. With me was Sergei Meyen, one of the foremost Russian experts
on plant paleontology and a specialist in Permian plants. We were both traveling back
from a conference in Montpellier; Sergei asked whether I would like to see the speci-
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origin and prehistory

mens, and I followed along. They are preserved alongside ancient conifers and other
plants as black fossils on a hard gray slate. The preservation is not very good; it is not
possible to make out microscopic details, and the fossils are not easy to understand.
We puzzled over them together for some time. Eventually, though, with deference to
Florin’s many other paleobotanical accomplishments, I had to agree with Sergei that
the structure of these fossils may not have been quite as Florin described and that the
link to ginkgo was not completely convincing.
One problem is that without more detailed information, the relationship between
deeply divided ginkgolike leaves from the Triassic and those of Trichopitys from the
Permian is problematic, especially given the presence of similar leaves in other groups
of extinct plants. Second, as Sergei pointed out, what Florin interpreted as a branched
seed-bearing axis attached at the base of a highly divided leaf might equally have been,
at least in the specimens that we examined, just another deeply divided but somewhat
flattened leaf segment. Florin’s linking of Trichopitys to ginkgo depends on the seed-
bearing structures’ being borne at the base of a leaf, just like those of ginkgo, which is
far from certain.5
Given these unresolved issues, Trichopitys, long held up as a likely ginkgo ancestor,
the link between unequivocal ginkgo from the Triassic and earlier seed plants, for the
moment is best treated as of uncertain relationship. Indeed, as more and more fossils
have come to light from Permian rocks, Trichopitys increasingly seems to be part of a
much more complicated picture, just one among many interesting but enigmatic fossil
seed plants from around that time. In some general way, these enigmatic Permian fos-
sils seem to provide a bridge between the ancient seed plants of the Paleozoic and the
more modern seed plants of the Mesozoic, but exactly how they relate to one another
and to older and younger plants is still uncertain.6
Sergei Meyen struggled with these kinds of questions, using his unrivaled knowl-
edge of Permian fossil plants from Russia and elsewhere. Toward the end of his life he
tried hard to develop a comprehensive overview of seed plant evolution that would
account not just for living plants but also for the vast diversity of Permian and other
extinct forms. He saw ginkgo as of particular importance and placed it at the center of
what he called Class Ginkgoopsida, which he assembled by “congregational analysis
taking into account as much information as possible.”7
Sergei put a huge amount of care and thought into the development of his ideas,
 93
ancestry

but his concept of Ginkgoopsida is extraordinarily broad and includes plants that dis-
play huge differences in many of their features. Many of the key fossil plants are also
not very well understood. Most are known only from leaves or leaves with associated
seeds. Frustratingly few can be compared point for point with living ginkgo, and while
this is a normal problem with fossil plants, it is a huge impediment to understanding
how they might relate to one another and to living plants.
In the fossil record of the past 350 million years it is rare that anything approaching
a complete plant is preserved. In the early phases of land plant evolution, at the time
of the Rhynie Chert, all of the plants were small and relatively simple, and often they
were preserved more or less intact. As a result they are relatively easy to understand.
However, as plants became larger and more complicated, the fossil record increasingly
becomes one of isolated fragments: a leaf here, a seed there, with stems, pollen cones,
and pollen grains somewhere else. Imagine the fallen leaves, twigs, fruits, and seeds
from a rich patch of forest being washed out into an ancient lake, settling to the mud
on the bottom and then being chipped out of the rock a few million years later. How
would you know which piece goes with which?
Needless to say, this problem greatly complicates efforts to use fossil plants to
understand plant evolution. For example, between about 270 million and 300 million
years ago, around the time of Trichopitys, several other fossils might be important for
a deeper understanding of the ginkgo life story. The Russian paleobotanist Serge Nau-
golnykh has described seed clusters from the Ural Mountains similar to those linked
with ginkgolike leaves in the Triassic. From other sites in the same area he has also col-
lected intriguing fossil leaves called Kerpia, which look a lot like those of ginkgo. The
problem is that in both cases they are just isolated pieces. It is hard to make sense of
them, still less to understand their significance, without more information about the
rest of these ancient plants.8
Paleobotanists with an interest in plant evolution therefore spend a good deal of
time trying to reconstruct something approaching a whole fossil plant from its sepa-
rated pieces. A colleague once called it the Humpty Dumpty game: the aim is to put
the pieces back together again. When the conditions of preservation are favorable, or
where we are lucky enough to have fossils that are attached to each other, then we may
be able to understand which fossil leaf goes with which seed or which pollen cone.
In these fortunate situations it is sometimes possible to link the stems, leaves, seeds,
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origin and prehistory

pollen-producing structures, and pollen that were all produced by the same extinct
species. Such reconstructions provide the key anchor points for understanding the
evolutionary history of plants in what can otherwise be an extraordinarily confusing
imbroglio of isolated bits and pieces of ancient plants.
One group of Permian plants that is relatively well understood and that may be
relevant to the evolution of ginkgo are the so-called glossopterids. Sergei Meyen also
placed glossopterids within his Ginkgoopsida. Glossopterid leaves, generally assigned
to the genus Glossopteris, are relatively simple, with many fine veins that are all of about
the same thickness. In some ways they are comparable to the leaves of ginkgo, but they
differ in one important respect: the leaf veins form extensive elongated reticulations.9
Glossopterid leaves have been well known to paleontologists for more than a cen-
tury and were among the specimens found with the bodies of Captain Robert Falcon
Scott and his party, who perished as they returned from the South Pole in 1912. The
occurrence of glossopterid leaves in all the Southern Hemisphere continents was im-
portant early evidence for the now widely accepted theory of continental drift.10
Botanically, glossopterids are much better understood than Trichopitys. Their seed-
bearing and pollen-producing organs are borne very clearly at the base of the leaf, and
in many cases they are directly fused to it. We also know that glossopterids were large
trees with woody trunks, and sometimes had distinct long and short shoots not dra-
matically different from those of living ginkgo. There are also beautifully preserved
specimens from Australia that provide a hint that glossopterids produced sperm cells
like those observed by Hirase in ginkgo.11
A further contender for an early ginkgo relative is one of the plants described by
John and Heidi Anderson from the Molteno flora of South Africa. The leaves, which
they called Kannaskoppifolia, are much more ginkgolike than those of Glossopteris. They
are wedge-shaped and deeply lobed, and while all the specimens that the Andersons
assign to this genus have some reticulations among the leaf veins, this varies among the
ten species that they recognize; in some, as in living ginkgo, reticulations are relatively
uncommon. There are also many specimens showing that the seed-bearing structures,
which they named Kannaskoppia, were borne at the base of the leaves. However, in this
case the reproductive structures look quite different from those of living ginkgo. The
seed-bearing structures are recurved and cuplike, and borne in large numbers on com-
plex branches. Each recurved structure presumably contained one or more seeds, but
 95
ancest ry

Reconstruction of the possible ancient

ginkgo relative Kannaskoppia, with its
partly divided leaves (Kannaskoppifolia)
and branches bearing numerous
small, recurved seedlike structures
(Kannaskoppia) from the Late Triassic
of South Africa, about 220 million
years before the present.

no details are known. The possible pollen-producing organs, which were almost cer-
tainly produced by the same plant, were given the name Kannaskoppianthus. They also
look strange. What seem to be pollen sacs are borne on a peculiar curled-over structure
that resembles fingers pressed onto the palm of a hand.12
The main practical problem in evaluating the relationships of plants like glossopter-
ids and Kannaskoppifolia-Kannaskoppia to each other and to living ginkgo is that even
in these relatively well-known cases, just as with Trichopitys, we are missing important
information. For example, other than in glossopterids, there are no details of the inter-
nal structure of the seeds of these plants, no information about pollen, and no knowl-
edge of the internal tissues of the stem. Similarly, in cases like Kannaskoppianthus, we
are also uncertain about how the basic structure of the fossils should be interpreted.
With so much information missing and so much uncertainty about key features, it is
hard to compare these plants with each other, still less with a living plant like ginkgo.
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origin and prehistory

However, there is also a theoretical problem: along with the similarities, there are many
differences. For example, in glossopterids the details of the pollen sacs and pollen
grains are quite different from those of living ginkgo. So in trying to track down the
origin of ginkgo, we need not only more information on the relevant fossil plants but
also a method that helps us know what to make of the various patterns of similarity
and difference.
 13
Relationships

Maternity is a matter of fact, paternity is a matter of opinion.

—Proverb

There are many fossil plants, especially from the Permian and Triassic, which may
be important for understanding the origin of living ginkgo, but, as in the case of gloss-
opterids, Kannaskoppifolia-Kannaskoppia, and Trichopitys, it is hard to decide which
are the most important. In some cases this is because the fossils are not known in suffi-
cient detail to allow useful comparison, but there is also the problem of how to account
for and understand the similarities and differences we observe. For example, should we
focus more on the leaves, or is it similarities of the seeds that should be given greater
weight? And how should we choose among different competing ideas that might link
ginkgo to glossopterids on the one hand, or Kannaskoppifolia-Kannaskoppia on the
other? This uncertainty suggests that we might be in need of a different way to think
about this problem, rather than simply trying to trace the ancestry of ginkgo backward
in the fossil record.
Instead of looking for ancestors, an alternative approach is to ask instead: to which
group of living or fossil plants is ginkgo most closely related? For example, is ginkgo
more closely related to cycads than it is to conifers, or vice versa? Or is ginkgo more
closely related to glossopterids or Kannaskoppifolia-Kannaskoppia than it is to any

97
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origin and prehistory

living plant? Framing the question in this way forces evaluation of the evidence for
alternative ideas about the relationships among the different groups of seed plants.
What we know and what we don’t know needs to be clearly spelled out, and in such
a way that we can choose among different competing ideas. This approach, based on
relative degrees of relatedness, is the primary way that these kinds of “origins” ques-
tions are addressed today across the whole of biology, whether the focus is the origin
of mammals or the origin of the HIV virus.
The modern methods used to assess the evolutionary interrelationships of organ-
isms developed from the work of the German entomologist Willie Hennig. Hennig, a
specialist on living and fossil flies, began his research in the 1930s, but his most influ-
ential work, Grundzüge einer Theorie der phylogenetischen Systematik, was begun at the
end of the Second World War during his time as a prisoner of the British. The book
appeared in 1950, and a translation, undertaken at the Field Museum in Chicago, was
published in 1966.
Subsequently, during the 1970s and 1980s, Hennig’s ideas were developed and elabo-
rated through lively, and often acrimonious, debates mainly centered around key par-
ticipants at the American Museum of Natural History in New York and the Natural
History Museum in London. For a young scientist working and teaching in this area on
both sides of the Atlantic in the late 1970s and early 1980s, it was fascinating, although
not always comfortable, to observe this scientific revolution at close quarters. It was
a true paradigm shift in modern evolutionary biology, and even on the fringes of the
intellectual action it was heady stuff. The outcome was a new, theoretical basis through
which questions about evolutionary relationships could be approached and answered.1
Hennig’s breakthrough was to recognize there is a hierarchy among the different
features that organisms share; and that this hierarchy makes sense in terms of a simple
model of the evolutionary process. The features might be characteristic structures, or
they might be components of DNA sequences, but Hennig’s point was that they can be
used to define successively less and less inclusive groups nested one inside the other
like a matryoshka doll. Membership within those groups defines successive levels of
relationship: successive degrees of relatedness.
What this means in practice is that relationships are defined in relative terms. For
example, ginkgo and conifers are regarded as more closely related to each other than
either is to a moss, because both belong to a group of plants, vascular plants, defined
by the presence of water-conducting cells with specially reinforced walls. In the same
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r e l at i o n s h i p s

The twelve major groups of living plants, classified into three larger groups, above, that are nested
like matryoshka dolls of different sizes. These nested relationships can also be expressed in a treelike
diagram, below, that approximates evolutionary relationships.

way, ginkgo and conifers are regarded as more closely related to each other than either
is to a fern because both belong to a group of plants defined by the ability to produce
the massive woody tissues seen in the trunk of a normal tree. No living fern produces
wood or becomes a tree in the same way. At the next level in the hierarchy, ginkgo and
conifers also belong to a more inclusive group, seed plants, that reproduce by seeds
rather than by spores, and so on.2
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origin and prehistory

Obviously, in many cases the hierarchy of features is not quite so clear- cut, making
it more difficult to understand relative degrees of relatedness. Sometimes, different
features suggest different hierarchies, and therefore imply conflicting patterns of re-
lationship, and when fossils are included for which key information is lacking, the
situation gets still more complicated. Often there are different alternative hierarchies of
relationships that could potentially explain the same data, and the number of potential
alternative patterns of relationship increases sharply as the number of different groups
under consideration increases. For four different groups there are only fifteen possible
arrangements, but to assess the potential relationships of ginkgo to nine other plants
the number of possible alternative arrangements among the ten groups under con-
sideration is a staggering 34,459,425.3
In such cases the method used to choose among competing patterns of relation-
ship is to invoke the standard scientific principle that the simplest explanation is pre-
ferred. What this means from an evolutionary point of view is that, for a particular set
of plants and a given set of features, the preferred explanation minimizes the num-
ber of separate evolutionary events needed to explain the origin of these features. For
example, it makes more sense to support an explanation in which the production of
woody tissues and seeds each evolved only once rather than an alternative explanation
that requires multiple origins of each feature.
Obviously, with many plants and many different features, and therefore a very large
number of possible explanations, it may be hard to find the one that really is the sim-
plest. There may be multiple patterns that are equally, or almost equally, simple. For
this reason the application of Hennig’s ideas drew huge new impetus from the devel-
opment of computers and appropriate software through which the large datasets could
be analyzed. Even with the most sophisticated approaches, finding the simplest expla-
nation requires either complex calculations that approximate an answer or exhaustive
mapping of many different features onto an almost overwhelming number of different
potential patterns of relationships, or “trees,” as they are sometimes called. For any
complex analysis involving more than a few plants and more than a few features it is
impossible to do this by hand.4
A further obvious problem is that it is not easy to compare organisms that differ
greatly in their structure and biology. For example, how do you compare ginkgo with
a moss, or a sea urchin with a shark? Point-by-point comparisons are often hard to
make. For this reason the study of evolutionary relationships received a further boost
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r e l at i o n s h i p s

from the recognition that relatively short DNA sequences extracted from very different
kinds of organisms could be obtained rather easily and, with the right analytical soft-
ware, could be compared very straightforwardly. In plants, the first large-scale appli-
cations that used DNA data and computer analyses to apply Hennig’s ideas were done
in the 1990s, and since then there has been rapid progress with understanding how
different groups of plants are interrelated. We now have strong evidence, for example,
that magnolias are more closely related to bay laurels than to water lilies, and that
the sacred lotus is more closely related to poppies and plane trees than to water lilies,
grasses, or palms. For the more than 350,000 species of flowering plants, there is now
a well-corroborated framework of relationships in which other aspects of plant evolu-
tion can be studied and against which the fossil record can be compared.5
At first glance, especially given the very rapid progress that has been made in other
areas where a much larger number of plants is involved and where broad consensus on
the pattern of relationships has been achieved, figuring out where ginkgo fits among
living seed plants ought not to be too difficult. After all, beyond ginkgo itself, there
are only four other living groups: conifers, cycads, flowering plants, and a peculiar
and rather obscure group known as Gnetales. We also have a huge amount of infor-
mation about the similarities and differences among their DNA sequences. The cost of
obtaining longer and longer DNA sequences, or even sequencing the entire DNA of an
organism, also continues to get cheaper and cheaper all the time. It ought to be easy to
determine which one of the 105 potential patterns of relationships for the five groups
is most strongly supported by all the available evidence.6
The question of how the five groups of living seed plants are interrelated has re-
ceived an enormous amount of attention, beginning with papers published thirty years
ago, long before the application of DNA techniques to these kinds of problems. The
focus has mainly been on trying to understand the relationships of flowering plants,
and hence to learn something about their origin. However, there is still no consensus
on which of the many different patterns of relationship that have been revealed by
slightly different kinds of analyses, on slightly different kinds of data, more accurately
reflect the actual pattern of evolution. When the many extinct groups of seed plants
are also included—and the data are thus restricted to the limited information that can
be gleaned from fossils—a still different set of possible answers emerges. In many ana-
lyses based on DNA sequences, and others that include fossils, ginkgo seems to be more
closely related to conifers than to any other group of living seed plants. However, other
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origin and prehistory

One plausible treelike diagram that summarizes the possible relationships among selected
groups of living and extinct seed plants. Dagger (†) indicates extinct groups.
For more on Yimaia and Karkenia see Chapter 15.

analyses come to different conclusions and find instead that conifers are closely related
to Gnetales, with ginkgo equally closely related to both.
Given the effort already invested to try to solve this problem, it may be that a truly
definitive answer to the question of whether ginkgo is more closely related to conifers,
cycads, flowering plants, or Gnetales will never emerge from the brute-force applica-
tion of more and more molecular data. But more important, even if the relationship of
 103
r e l at i o n s h i p s

ginkgo to other living seed plants could be solved, the answer is unlikely to be helpful
with regard to what we really want to know, which is how ginkgo fits into the broader
constellation of living and extinct plant diversity and how its characteristic features
evolved. Solving these problems will require probing more deeply, more thoughtfully,
and in an integrated way, the similarities and differences between ginkgo and other
seed plants, especially those seed plants that are known only as fossils, about which we
still have much to learn. We will also need to see through apparently conflicting signals
from different lines of evidence and reconcile them into a pattern of relationships that
most closely approximates the actual course of evolution.7
At the moment we can only fall back on the lame generalization that ginkgo prob-
ably had its origin among those groups of seed plants with relatively simple leaves and
flattened seeds that diversified during the latter part of the Paleozoic. These groups,
which included fossil plants like Kannaskoppifolia-Kannaskoppia, glossopterids, and
perhaps Trichopitys, represent a second wave of seed plant evolution beginning about
300 million years ago, which succeeded the more ancient seed plants of the Devonian
and Carboniferous. It is all frustratingly vague, but beyond that it is impossible to go
at the moment. Despite our best efforts, exactly how ginkgo fits into the grand scheme
of plant evolution remains elusive.
 14
Recognition

Too much light often blinds gentlemen of this sort.

They cannot see the forest for the trees.
—Martin Wieland, Musarion oder die Philosophie der Grazien

Many of the most famous figures in the history of plant paleontology have writ-
ten about ginkgo, but none has done more to illuminate its long evolutionary history
than the Chinese paleobotanist Zhou Zhiyan. Beginning with a series of influential
studies in the 1980s, Zhou discovered several different kinds of early ginkgolike plants
and came to understand them not just from their leaves but also from their seeds and
other parts. He also placed his new discoveries in the broader context of what earlier
researchers had learned. As a result, the history of the ginkgo lineage is now better
understood, and also much more complicated and interesting, than it was thirty years
ago. In 1994, Zhou’s work was recognized by one of the most prestigious awards given
by the Chinese Academy of Sciences, and the following year Zhou was elected to the
academy, a high honor in a culture where science is held in great esteem. Zhou is the
most respected of all Chinese scholars of the plant fossil record, and his work is central
to a full understanding of living ginkgo.1
Zhou Zhiyan spent the first half of his career in China, but in 1980 he traveled to the
United Kingdom to study with Professor Tom Harris at the University of Reading. It

104
 105
recognition

Tom Harris (1903–1983), the British

paleobotanist who contributed much to
our understanding of ancient ginkgo and
other fossil plants. Photographed at the
University of Reading in summer 1980.

was one of the pivotal experiences of Zhou’s life. His priority was to study a collection
of fossils that he brought with him from China, but had he known that ginkgo would
figure so prominently in his later career, he might have spent his time differently. Har-
ris had also made important contributions to knowledge of ginkgo fossils. They would
have had much to speak about.2
Harris arrived at Reading in 1934 as one of the first professors to be appointed after
the university was founded, and he worked there until his death in 1983. By the early
1980s Harris was in his late seventies, but he was still a force to be reckoned with: every
day he came to the Department of Geology to work on the latest plant fossils to com-
mand his attention. Tall and gangly, with a strong personality, Harris had a steel-trap
mind that saw the world in black and white.
Harris was generally warm to those around him, and he was nothing but kind
and generous to me in my years at Reading, but he did not suffer fools. He had little
patience with those who saw the world in shades of gray. He built his reputation in
the middle of the twentieth century based on clear thinking combined with energetic
efforts to collect new fossils. The resulting stream of published research spanned al-
most six decades. His magnum opus, at least in the second half of his career, was his
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origin and prehistory

comprehensive five-volume treatment of The Yorkshire Jurassic Flora. He became the

dominant force among British paleobotanists in the decades after the Second World
War, and the cumulative impact of his work was enormous.3
Harris went to Cambridge intending to become a doctor, but he was something of a
child prodigy. At the age of eighteen, traveling every day from his home in Leicester to
the University of Nottingham, he had already received his bachelor of science degree.
It was through the influence of H. S. Holden, whom Harris met at Nottingham, that he
first became interested in fossil plants. Holden had been trained at the University of
Manchester, which had long been a stronghold of scientific paleobotany in Britain, and
his enthusiasm for such work evidently rubbed off. Harris quickly discarded medicine
in favor of botany.4
At Cambridge, Harris became part of the circle around A. C. Seward, probably the
foremost paleobotanist of his time. Twenty years after Charles Darwin’s death, Seward
had edited a volume of his letters with Darwin’s son Francis, but his main interest was
fossil plants, and he wrote prolifically on all aspects of the subject. Seward was also a
skilled administrator, and by the 1920s, when Harris arrived, Seward was well estab-
lished, well respected, and well connected.5
Seward made important contributions in many different areas of paleobotany and
worked on a range of fossils from many different parts of the world, but he was particu-
larly intrigued by fossils of flowering plants. It was Seward who was largely responsible
for popularizing a phase used by Darwin, who once remarked that the origin of flower-
ing plants was an “abominable mystery.” With a gentle push from Seward, that phrase
has been quoted more than Darwin could ever have imagined, and perhaps more than
he ever would have wished.6
In the 1920s Seward was interested in fossil flowering plants from about 100 million
years ago and especially in material that was then becoming available from the work of
Danish geologists in West Greenland. He traveled there himself in the summer of 1921
and, soon after, was sent a collection of fifteen packing cases of fossil plants from much
older rocks collected on the other side of the icecap, from the fjords about halfway up
Greenland’s east coast. As Harris told it, the crates were sent to Seward by mistake. The
material had been collected in 1900, on an early geological expedition by the Danish
geologist Nikolaj Hartz. Harris seized the opportunity, made the collection the focus
of his research at Cambridge, and never looked back.7
In 1925 Harris visited Stockholm and Professor Thor Halle, Nathorst’s successor at
 107
recognition

the Swedish Museum of Natural History, to learn the techniques needed to study the
Greenland material. Harris used those same techniques, with little modification, for
the rest of his career. The following year, over dinner at Seward’s home in Cambridge,
he met the Danish geologist Lauge Koch. In Harris’s words, Koch was a “huge, rather
fierce looking man in charge of the Greenland Geological Survey. He asked me—could
I come with him on an expedition to East Greenland for a year, starting next month.
It appeared to me instantly that it was one of those situations where thought does not
lead to a wiser decision, so I said ‘yes.’”8
In the summer of 1926, after a brief stop in Copenhagen and a three-week passage
to the east coast of Greenland, the small party lead by Koch landed at Scoresby Sound.
Along with Koch and Harris, the group consisted of Alfred Rosenkrantz, a Danish ge-
ologist and engineer, two Eskimo hunters, and about fifty sledge dogs. Their aim was to
help understand the geology of that part of East Greenland by making extensive collec-
tions of fossils, and especially to expand the collections of fossil plants made by Hartz.
The expedition had enough supplies to last the winter, but their lives were not easy
in their isolation. The climate was harsh, the terrain vertiginous, and the slopes slip-
pery on top of the permafrost. Living conditions were basic, but in the year they spent
there, much of it during the long winter and using primitive equipment, Harris and
his colleagues amassed a large collection of fossil plants from a geologic section about
three hundred feet thick. Harris remarked, “Each bed had one or a dozen species often
all different from those of a neighboring bed,” and in some beds there were abundant
fossil leaves unquestionably similar to those of modern ginkgo. The party was picked
up again the following summer. Tons of rocks with plant fossils, including many an-
cient ginkgo leaves, were shipped back for study.
In all, Harris worked on the fossils from East Greenland for about a decade, and
these were the most paleobotanically productive years of his life. He described a rich
variety of ferns, horsetails, and clubmosses, as well as conifers, cycads, and a wide
range of peculiar extinct seed plants. He established the East Greenland flora as one
of the best-known fossil floras in the world, and it quickly secured his reputation; he
moved to Reading as the first professor of botany at the age of thirty-one.
Harris carefully described the ginkgolike leaves from East Greenland and recog-
nized that they were more deeply and more regularly divided than the leaves of mod-
ern Ginkgo biloba. He called them Ginkgoites taeniata and noted that they were more
like the leaves produced by ginkgo seedlings, or the early leaves produced from regen-
 108
origin and prehistory

erating shoots, than leaves from a mature tree. Also, using the classic but rather dras-
tic paleobotanical technique that he had learned from Halle, and later taught to Zhou,
Harris employed strong acids, followed by a strong alkali, to dissolve away the coal-like
material of the fossil leaves and leave just the resistant, waxy outer cuticular covering
of the upper and lower leaf surfaces. He then used similarities in their cellular details
to show that although variable in shape and size, all the ginkgolike leaves were most
probably from a single species. Harris also suggested that some of the seeds he found
associated with Ginkgoites taeniata leaves were probably produced by the same trees.
He could not prove it, but he was convinced that they were part of the same plant based
on their consistent association at different localities and similarities in their cuticular
coverings.9
When the Greenland work was completed, Harris, by that time at Reading, needed
to decide what to work on next. Rather than continuing with the Greenland material,
he turned his attention to something new and embarked on a comprehensive revision
of the beautifully preserved Jurassic plants from Yorkshire in the United Kingdom.
These fossils had been collected and studied since the earliest days of scientific pale-
ontology, and by the early 1930s the Yorkshire material had already been worked on
extensively, including by Nathorst in Stockholm and Hugh Hamshaw-Thomas, one of
Harris’s senior colleagues at Cambridge. Thomas especially had done extensive field-
work and made major new collections. Harris, however, was undeterred; in his view
there was plenty still to be done, and Yorkshire was easier to get to than East Green-
land.
Harris never worked again on the material from Greenland, although many years
later he did allow that perhaps this was a pity. Harris once said to me that he would
have made more exciting discoveries had he focused on the Greenland fossils; but he
then added, with his characteristic grin and bob of the head, that he also would never
have enjoyed so many holidays in Yorkshire. Many of those holidays, with his family
in tow, were apparently spent collecting at the classic localities, and walking the North
Yorkshire Moors looking for new sites that might give new information or produce
new kinds of fossil plants.
Through The Yorkshire Jurassic Flora, Tom Harris made major contributions to
understanding the plants that grew in the estuaries, back swamps, and floodplains of
the ancient coastline that existed about 150 million years ago in what is now northeast-
ern England. That coastline, which today is often swept by biting winds off the North
 109
recognition

Fossil leaves of an ancient ginkgo, Ginkgoites huttoni, from Scalby Ness, Yorkshire,
collected from the sands of an ancient meandering river that flowed out toward
what is now the North Sea about 170 million years ago.

Sea, was then home to diverse luxuriant, more or less tropical vegetation of conifers,
cycads, ferns, and many kinds of plants now extinct. It was entirely devoid of mod-
ern kinds of mammals, as well as birds, butterflies, bees, and many other animals that
today we take for granted. Instead, it was home to dinosaurs and pterosaurs; beetles
and early flies would have been among the common insects. Harris devoted the bulk
of his career to bringing the plants of these ancient ecosystems to life. Building hand-
somely on what had gone before, his work established the fossil flora from Yorkshire
as the benchmark against which all other fossil floras of this age are compared.10
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origin and prehistory

Ginkgo leaves occur at several of the classic Yorkshire localities but are especially
common in the Scalby Ness plant bed, which is exposed on the coast just north of Scar-
borough. I collected them there for the first time in the autumn of 1974 with Harris,
his former student Joan Watson, and her students from the University of Manchester.
I have returned many times since and have never been disappointed.
The cliffs at Scalby Ness offer a crude cross section through the sands of an ancient
river, now hardened by time, that once meandered out to sea. The fossil plants occur
tumbled together exactly where they settled among the river sands millions of years
ago. By far the most common fossil plants are the deeply divided leaves of Ginkgo hut-
toni. Over the course of his career, Harris also found seeds very like those of modern
ginkgo at Scalby Ness. They were similar to those that he had collected in Greenland,
and Harris thought that they were probably produced by the plant that bore Ginkgo
huttoni leaves. He also described a single pollen-producing structure similar to that of
modern ginkgo.11
The species of ginkgo that Tom Harris described at different times in his career, first
from East Greenland and then Yorkshire, are unquestionably part of the lineage lead-
ing to modern Ginkgo biloba; they are certainly more closely related to ginkgo than any
other living plant. However, our knowledge of these plants is incomplete in several cru-
cial ways, especially in showing how the seeds were borne. Harris’s painstaking work
did much to clarify the exact similarities between the leaves of fossil and living ginkgo,
but he more than anyone else recognized that not much more could be done until some
fossil ginkgolike plants were known from more complete information. Harris always
emphasized that when the opportunity arose, assembling whole fossil plants should be
the top priority. Without such reconstructions Harris knew that it would be impossible
to make more useful comparisons between the living and fossil species. It was a central
theme in his life’s work and a philosophy he passed on to all of his students, including
Zhou Zhiyan.
 15
Proliferation

Be fruitful, and multiply, and replenish the earth.

—Genesis 1:28

Zhou Zhiyan arrived at the University of Reading in September of 1980, and for
almost a year we worked side by side in the same laboratory. I was in my mid-twenties
and coming to the end of my first job in the Department of Botany. Zhou was in his
late forties and a senior scientist at the Institute of Geology and Palaeontology of the
Chinese Academy of Sciences in Nanjing. He was on his first trip outside China. At that
time, the competition for such opportunities was fierce, and Zhou was among the first
Chinese scientists allowed to travel overseas in the years following the death of Mao.
He brought with him interesting fossil plants to study, but his broader objective was
simple. He intended to learn all he could and make up for lost time: the years spent
on what he calls “unprofessional activities”; the political meetings, manual labor, and
other activities unrelated to science that he endured during the Cultural Revolution.1
Zhou was born in Shanghai in 1933. In the early 1950s he studied at Nanjing Univer-
sity and gained a position at what was then the Nanjing Institute of Geology and Palae-
ontology. Like Tom Harris, he specialized in fossil plants of the Mesozoic, but while
Harris had been a botanist almost from the beginning, Zhou’s training was in geology.
Most of his early work used fossil plants to support geological exploration; his fossils

111
 112
origin and prehistory

Zhou Zhiyan, the Chinese paleobotanist

whose work revolutionized our
understanding of ancient ginkgo based
on studies of fossils from China and
elsewhere. Photographed in early 2009
outside the Museum at the Nanjing
Institute of Geology and Palaeontology
of the Chinese Academy of Sciences.

were useful indicators of geologic age, and his broader goal was to better understand
the commercially important coals that would be crucial for China’s economic devel-
opment. Only later, and especially after his stay in Britain, did Zhou turn his attention
to botanical questions, including the area in which he is now preeminent: elucidating
the history of ginkgo and its relatives. More than anyone else, in his own gentle and
understated way, Zhou Zhiyan has helped make sense of the fossil ginkgolike leaves
from between about 60 million and 225 million years ago and has illuminated what
they tell us about the evolution of the single living species.
Unless they devote themselves entirely to theory, or to working on collections made
by others, all paleontologists need a bit of luck. One way or another they need to have
good specimens that are sufficiently well preserved to give new information and useful
new insights. However, in paleontology as in other areas, fortune favors the prepared
mind. Good paleontologists are always on the lookout for interesting new material,
and when serendipity brings it their way, they know what to do with it. Zhou Zhiyan’s
 113
p r o l i f e r at i o n

studies of early ginkgo fit this profile: the right material came along at just the right
time, he recognized its significance, and he did what needed to be done.
In the mid-1980s, not long after returning from Britain, Zhou was contacted by
Zhang Bole, a mining engineer working in the Yima region of Henan Province, in
northern China. Zhang was a professional geologist, and he knew a good deal about
the coal mines in that area, but he was also a dedicated fossil hunter with a particular
interest in plant fossils. He and his family devoted much of their spare time to collect-
ing fossil plants from the spoil heaps of the large open-cast coal mine at Yima, and the
hours that they had invested had yielded spectacular fossil ginkgo leaves, many of them
beautifully preserved in a soft gray siltstone.
Zhou recognized the potential of Zhang’s material immediately. It was of Juras-
sic age, dating from about 170 million years before the present, and was much better
preserved than the specimens he had studied earlier in his career, including the ones
that he had taken with him to Reading. Most Jurassic plant fossils from elsewhere in
China—for example, from the coal mines around Beijing—occur in rocks that have
been subject to great temperatures and pressures after they were formed; as a result,
the fossil plant material is badly squashed, chemically altered, and almost hopeless
for providing detailed information. In contrast, the Henan material was preserved in
softer rocks that had not been deeply buried. The fossils looked more like some of the
well-preserved material from the Jurassic of Yorkshire that Zhou had seen in Britain.
He quickly realized that the waxy cuticular coverings of the leaves and other plant parts
were still preserved and that, with the right techniques, these fossils would yield useful
information.
An immediate priority was to visit the site and collect more fossils, so in 1986 Zhou
traveled to Henan and spent several days at the Yima coal mine collecting specimens
with his student Xuanli Yao. He also pored over the large collections that Zhang and
his family had made and recognized that among the beautifully preserved ginkgolike
leaves were not just seeds but also the structures on which the seeds were borne. The
leaves, seeds, and seed-bearing structures were so common that they were almost cer-
tainly produced by the same plants.
The first report that Zhou Zhiyan and Zhang Bole published on fossil plants from
the Yima mine was a preliminary announcement of two new types of ginkgolike seed-
bearing structure. They noted that “as a detailed study will take quite a period of time,
we are disposed to announce the important features and to present briefly a prelimi-
nary assessment here for reference to colleagues who are interested in the past history
 114
origin and prehistory

of Ginkgo.” They recognized that both these seed-bearing structures were entirely new
discoveries, and they drew the obvious conclusion; there was more than one kind of
ginkgolike plant preserved at the Yima locality.2
The first of the two fossil plants from the Yima coal mine to be worked out in detail
by Zhou and Zhang was a very ginkgolike fossil plant known from leaves and seed-
bearing structures. They named the leaves Ginkgo yimaensis. Zhou and Zhang strength-
ened their earlier argument that the two different organs were produced by the same
species, based not only on their consistent association and abundance at one particular
level in the Yima coal mine but also on the structure of their waxy outer covering, which
was similar, but not identical, to that in the modern tree. Zhou and Zhang also noted
that the leaves were “more deeply lobed than those of G. biloba” and that the seed-
bearing stalk was branched with each of the five or six seeds borne on a long stalk.3
Zhou and Zhang then turned their attention to the second plant that they had recog-
nized in their preliminary account of the Yima fossils. They named the seed-bearing
structures Yimaia recurva. Each had a simple stalk with a cluster of about eight to
nine seeds at the tip. They were associated with quite different, and much more finely
divided, leaves than had been described by H. C. Sze, a distinguished Chinese paleo-
botanist of the previous generation. Sze had named these leaves Baiera hallei after Thor
Halle, the Swedish paleobotanist who had helped Tom Harris and done much early
work on fossil plants from China. The leaves of Baiera hallei were so highly divided
that the individual leaf segments were almost grasslike.
Zhou and Zhang named the reproductive structures associated with Baiera hallei
after the locality at which they were collected, Yimaia. In this case the fossil was known
not just from leaves and isolated seeds but also from distinctive twigs with obvious
long and short shoots. The different plant parts occurred massed together in the Yima
coal mine a little below the level from which Ginkgo yimaensis was described. At the
time they wrote the paper, Zhou and Zhang had about fifty specimens of Yimaia re-
curva and hundreds of Baiera hallei leaves.4
After the original description of Ginkgo yimaensis and Yimaia recurva from the
Middle Jurassic of Henan, Zhou went on to discover similar fossil plants of about the
same age from elsewhere in China. In the space of eighteen years Zhou and others
have described several different species of Yimaia and clearly established it as one of
the best-known ancient early relatives of living ginkgo. The picture that emerges is that
Yimaia-like plants were reasonably common in the vegetation of the Northern Hemi-
sphere all the way from Europe to China during the middle of the Mesozoic, between
 115
p r o l i f e r at i o n

Leaf of an ancient ginkgo relative, Ginkgo

australis, from the Early Cretaceous
Koonwarra fish beds, Victoria, Australia, about
130 million years before the present.

160 million and 200 million years ago, and that, at the same time, these same ancient
landscapes were also home to other plants, such as Ginkgo yimaensis, that were even
more like modern ginkgo.5
In their initial studies of fossil plants from the Yima coal mine in the late 1980s, Zhou
and Zhang recognized two different ginkgolike plants, but surprisingly, almost fifteen
years later, they were also able to recognize a third. Zhou assigned this third ginkgo-
like plant to the genus Karkenia, a group of fossils first described in the mid-1960s
from the Early Cretaceous of Tico, Santa Cruz Province, Argentina, by the pioneering
Argentinean paleobotanist Sergio Archangelsky. Archangelsky had also worked with
Tom Harris earlier in his career, and he too had been thoroughly inculcated with the
importance of reconstructing ancient plants from their different, dispersed parts.6
The Tico flora is beautifully preserved and dates from about 130 million years ago.
It contains common ginkgolike leaves that Archangelsky named Ginkgoites tigrensis.
Associated with them are seed-bearing organs for which Archangelsky created a new
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origin and prehistory

name, Karkenia incurva. Karkenia presents an interesting puzzle. On the one hand
the leaves are very like those of a deeply divided modern ginkgo leaf; it is easy to see
the relationship. However, the associated seed-bearing structures are very different,
so different that when I first read about these fossils in the mid-1980s, I was skeptical
whether they had anything whatever to do with living ginkgo. The seeds are tightly
packed in masses of more than a hundred, almost making a cone. They are quite dif-
ferent from the simple seed stalks of living ginkgo that bear just one or two seeds. In
addition, in Karkenia, each seed has its own stalk and is curved back on itself, so the tip
of the seed faces inward, back toward the cone axis. However, there is no disputing the
ginkgolike leaves, and the evidence for linking the isolated fossil leaves from Argen-
tina, China, and elsewhere with the same kind of seed-bearing structures is strong.
Sergio Archangelsky’s initial deduction of the link between the leaves and seeds has
been completely borne out by later work and in hindsight is now seen as a key step in
expanding our knowledge of the biography of living ginkgo.
The Karkenia that Zhou and his colleagues recognized from China is much less
common in the fossil flora from Yima than either Ginkgo yimaensis or Yimaia recurva.
It comes from a level in the coal mine a little below that from which Yimaia was de-
scribed. Only about five seed-bearing structures are known, but these discoveries add
to evidence of diverse ginkgolike plants during the Mesozoic. In a single coal mine in
northeastern China, Zhou had now discovered three quite different fossil plants that
were all more closely related to ginkgo than to any other living plant.7
Given the presence of Karkenia in both the Northern and Southern Hemispheres, an
important question is whether plants more like ginkgo or extinct Yimaia ever existed
in the Southern Hemisphere. Undeniably ginkgolike leaves have been recorded from
many places in Africa, Australia, and South America, for example, and also from India,
which at that time was also part of the mass of continents aggregated together in the
Southern Hemisphere. However, in no case have these leaves been associated with typi-
cal ginkgo or Yimaia reproductive structures. They may all be Karkenia-like plants, or
perhaps further kinds of ancient ginkgo that we do not yet understand.8
Zhou Zhiyan made his key discoveries of fossil ginkgolike plants decades ago, but
he has continued to add meticulous descriptions of new fossil material from other
localities. These new fossils tended to confirm, rather than expand, the picture already
developed from studies at the Yima coal mine, but in 2003 Zhou made another break-
through in understanding the fossil history of ginkgo.
 117
p r o l i f e r at i o n

The relative position of the southern continents around 100 million years before the present
as they separated from Antarctica, showing the presence of similar ginkgolike leaves
in South America, India, and Australia.

During the 1990s some of the most sensational paleontological discoveries of all
time began to emerge from a remarkable fossil deposit in northeast China. This so-
called Jehol Biota, collected from the Yixian Formation in Liaoning Province, is of
Early Cretaceous age and dates from about 120 million to 125 million years ago. It has
continued to make headline after headline as it has yielded one spectacular animal fos-
sil after another. Especially prominent has been a remarkable series of early birds and
related dinosaurs, some of which show clear evidence of feathers or the downy feather-
like structures that were probable feather precursors. There are also early mammals
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origin and prehistory

and amphibians, a great variety of insects, and a wide range of fossil plants, including
ferns and even mosses. The seed plants include conifers and some of the most infor-
mative and earliest fossils of flowering plants.9
It is surprising, given the widespread occurrence of ginkgolike plants through the
Jurassic and Cretaceous, that few fossils relevant to ginkgo’s life story are found in the
Yixian Formation. However, in 2003, Zhou Zhiyan and his colleague Shaolin Zheng
described unequivocal ginkgo fossils from these beds for the first time, and these fos-
sils help fill the gap between the truly ancient Ginkgo yimaensis and essentially mod-
ern ginkgo. These new ginkgo fossils were not from the classic Jianshangou Bed of the
Yixian Formation near Jinzhou, which has yielded the most spectacular animal fos-
sils, but from the Zhuanchengzi Bed, which outcrops to the southeast on the southern
slope of Yinwoshan Mountain near Toudaohezi Village. The two beds are thought to be
roughly the same age, but they contain a slightly different assortment of fossil plants.
Fossil collecting at Yinwoshan Mountain has turned up about a dozen indisputable
ginkgo leaves, along with seven seed-bearing structures that seem to have been pre-
served at slightly different stages of maturity. The leaves are unusually small compared
with the leaves of modern ginkgo, none of them much more than about an inch long,
and they are deeply divided. The seed-bearing structures are also small, and in all cases
they were more or less unbranched. At their tips they bore up to about three to six tiny
seeds. Among the specimens were a few seed stalks with immature seeds still attached,
but there were also some with a large seed attached and large scars from which others
had been shed. Zhou and Zheng named the leaves and the associated seed-bearing
structures Ginkgo apodes.
The importance of Ginkgo apodes is that it fits nicely in the age gap between Ginkgo
yimaensis, at about 170 million years, and fossils that are essentially the same as mod-
ern ginkgo, at about 65 million years. It is also conveniently intermediate in structure.
Ginkgo yimaensis has a seed-bearing stalk that divided into three or four branches,
each bearing a single seed at its tip. However, in Ginkgo apodes the branches of the seed
stalks are almost nonexistent and the seeds are attached close together at its tip. This is
much more like modern ginkgo, although in the living species there are normally just
two seeds on each seed-bearing structure rather than three to six. Zhou Zhiyan con-
cluded that the pervasive trend in the evolution of ginkgo, at least in the seed-bearing
structures, has been one of reduction from about six seeds on separate stalks in Ginkgo
yimaensis to about two seeds on an unbranched seed stalk in the living plant.10
 16
Winnowing

People from a planet without flowers would think we must be mad

with joy the whole time to have such things about us
—Iris Murdoch, A Fairly Honourable Defeat

Since Zhou Zhiyan began his work in the Yima coal mine a quarter of a century ago,
what we know about the fossil record of ginkgo and its relatives has expanded dramati-
cally. New information and new discoveries continue to accumulate and have revealed
an astonishing variety of ancient ginkgolike plants. This unexpected diversity changes
the way we think about the evolution of the single living species. Studies of fossil leaves
had hinted at the existence of such diversity, but until more was known about their cor-
responding seeds and other parts, the real plants lurking behind the isolated leaf fossils
remained enigmatic and difficult to compare with the living tree. Work by Zhou Zhiyan
and his colleagues has changed all that.1
Given what we know now, it is possible to begin to consider the major patterns in
the evolution of living ginkgo and its extinct relatives. Again, it is Zhou Zhiyan who has
led the way based on some simple analyses made possible by his unrivaled knowledge
of the relevant fossils.
As an initial attempt to gain a sense of the changing diversity and abundance of
ginkgolike plants through time, Zhou made a graph showing the number of different

119
 120
origin and prehistory

kinds of ginkgolike leaves that have been distinguished as different genera through
the long fossil history of the group. Setting aside Trichopitys and other Permian plants,
which may or may not be related to the ginkgo lineage, Zhou showed that from four
different kinds of ginkgolike leaves in the Early Triassic, the number increases over
about 50 million years to six in the Middle Triassic, and twelve in the Upper Triassic.
The number remains high through the Jurassic and into the Early Cretaceous, vary-
ing from seven to eleven between about 100 million and 200 million years ago, before
declining to four in the Late Cretaceous and just one and two in the Paleogene and
Neogene. There are fewer ginkgo-related reproductive structures, but the pattern is the
same: numbers for the Late Triassic through Early Cretaceous range from five to three,
whereas only a single kind of reproductive structure is known after about 100 million
years ago. In both cases the patterns are crude but nonetheless revealing.2
Because of potential confusion about which fossil leaves can be securely related to
the ginkgo lineage, Zhou also took a more conservative approach to see whether the
pattern would still hold up. He focused on the fossils he knew best and traced through
time the number of different species of the three main leaf types that he had worked on
from the Yima coal mine: Baiera, the leaf of Yimaia; Ginkgoites, the leaf of ginkgo; and
Sphenobaiera, the leaf of Karkenia. Again, the bulk of the diversity is right in the middle
of the Mesozoic, from the Upper Triassic to Early Cretaceous, between about 100 mil-
lion and 225 million years ago, but drops off rapidly thereafter. In the Early Cretaceous
of China, Zhou recognized twenty-two different kinds of ginkgo leaves, ten different
kinds of Baiera leaf, and ten different kinds of Sphenobaiera leaf, but in the Late Creta-
ceous all had disappeared except for a single kind of Ginkgoites leaf. However you look
at these simple statistics they show an astonishing and consistent pattern of decline in
the number of ginkgolike plants. From some point in the middle of the Cretaceous,
around 100 million years ago, the world of ginkgo began to change.3
In parallel with the decline in the number of different kinds of ginkgolike plants,
the importance of ginkgo and its relatives in ancient vegetation seems likely to have
declined substantially as well. We might expect that these plants steadily became less
common on Mesozoic landscapes. This is harder to assess from what has been pub-
lished on the fossil record, but Zhou was able to gain a rough idea of how widespread
these different kinds of ginkgolike plants were by looking at the number of counties
in China from which they had been recorded. In the Early Cretaceous the number of
counties with records of ginkgo was eight, compared with thirty-one for Ginkgoites,
 121
winnowing

Key dates in the history

of ginkgo and its relatives
over the past 300
million years.

eighteen for Baiera, and eleven for Sphenobaiera. Thereafter, from the Late Cretaceous
onward, there are no records of Baiera or Sphenobaiera at all, just one for Ginkgoites,
and five for ginkgo itself. Again, the pattern is crude but the conclusion is clear. For
ginkgo and its relatives, the middle of the Cretaceous period was a time of transition.
It is possible only to speculate about what might be behind the apparently perva-
sive decline of ginkgolike plants about 100 million years ago, but one obvious potential
cause is competition with flowering plants, a new group of highly successful plants that
rose rapidly to dominance during the middle part of the Cretaceous. Of course, there
were many other environmental changes during the mid-Cretaceous; for example, ac-
celerated rates of continental drift created new configurations of land and sea, making
possible new kinds of currents in the oceans and atmosphere, which must have resulted
in new kinds of climates. However, it is also hard to believe that the explosive increase
in the diversity and abundance of flowering plants in the Cretaceous did not have some
impact on the ginkgolike plants that had previously been so prominent. By the Late
Cretaceous Yimaia, Karkenia, and similar plants seem to have disappeared. The ginkgo
group as a whole was a shadow of its former self, and the survivors were those ginkgo-
like plants most like the single living species.4
The period between about 65 million and 100 million years ago, immediately after
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origin and prehistory

the initial rise of flowering plants, was an interesting time in the history of life on land:
a time when familiar plants grew alongside unfamiliar animals. It was a moment when
magnolias were food for Triceratops, and hadrosaurs made their nests among groves of
ancient plane trees. During this time, about three-quarters of the known mammals be-
longed to an extinct group called the multituberculates, small marsupial-like animals
that are often compared with rodents. It has been speculated that these small mammals
may have fed upon and dispersed ginkgo nuts.5
One of the best insights into the strange ecosystems during this last gasp of the age
of dinosaurs comes from fossils preserved in the Horseshoe Canyon Formation of cen-
tral and southern Alberta. These rocks have provided us with some of the best known
of all dinosaurs—carnivores, such as Tyrannosaurus rex and Albertosaurus, as well as
herbivores, such as Triceratops and Stegosaurus. Fossil plants from these deposits have
received less attention but are important because they were the ultimate source of
energy on which the dinosaurs and other animals living on these ancient landscapes
depended. The plants of the Horseshoe Canyon Formation are also important because
of the ways in which they are preserved. Sometimes they are squashed and flattened,
but other times they are petrified by various minerals including calcite, phosphate, and
silica.
A detailed study by Kevin Aulenback shows that the plants of the Horseshoe Canyon
Formation included mosses, clubmosses, horsetails, and various ferns, as well as sev-
eral different kinds of seed plants. Flowering plants included aroids and gingers, as
well as trees similar to living swamp gum, dove tree, and hornbeam. Conifers similar
to living dawn redwood, cypress, and China fir are also present alongside leaves and
seeds that are very like those of living ginkgo. Many of the living relatives of these coni-
fers are still found in eastern and southwestern China, not far from those areas that are
home to living populations of ginkgo.6
Farther south and east of Alberta, in Montana and North Dakota, ginkgo is also
widespread in the sands and silts laid down on the ancient floodplains of the Hell Creek
Formation. Fossils from the Hell Creek Formation provide some of the last known oc-
currences of dinosaurs before they disappeared during the mass extinction at the end
of the Cretaceous. From the standpoint of animal evolution, the impact of the end-
Cretaceous mass extinction was devastating. The losses were selective but widespread,
and many groups of animals, both in the ocean and on land, were eliminated. Their
extinction left an indelible mark on the trajectory of animal evolution.
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winnowing

Interestingly, the long-term effects of the end-Cretaceous extinction on the history

of plant life were seemingly much less profound. Over the short term, many of the pre-
viously dominant plants were lost, and a careful study by Peter Wilf and Kirk Johnson
in southwestern North Dakota showed that almost two of every three plant species
that were present below the boundary became extinct. However, while there is strong
evidence of regional extinction, it is less clear that major groups of plants were lost at
a global scale.
In southeastern North Dakota only a few of the plants known from below the
boundary reappear above it: one of them is ginkgo. Ginkgo leaves are present at many
fossil localities in the Hell Creek Formation and have been collected just a few feet
below the boundary, but they are back again, not long after the apparent cataclysm, in
the overlying Paleocene rocks of the Fort Union Formation. This pattern speaks well to
the capacity of ginkgo for survival, but it makes the great ginkgo decline in the middle
part of the Cretaceous, which does not coincide with a massive extinction in the world
of animals, still more striking. It underlines the point that in some ways plant and ani-
mal evolution is decoupled. Somehow, it seems, plants and animals dance to a differ-
ent evolutionary beat.7
Taken as a whole, the large-scale patterns in the evolution of the ginkgo group ex-
pand the way that we should think about the evolution of the living tree. We might
have thought that we could trace ginkgo back through a series of ancestors to ancient
plants, perhaps like Trichopitys, that lived in Permian or older times. Today, however,
as a result of the work of Zhou Zhiyan and others, we have a very different picture, a
picture that is complicated by a great deal of previously unrecognized diversity. We
now know that many species of ginkgolike plants lived at the same time in the past and
perhaps even alongside one another in the same kinds of plant communities. This may
have been the case with Ginkgo yimaensis, Yimaia recurva, and Karkenia incurva, which
grew in similar kinds of habitats 170 million years ago in the coal swamps of northern
China, but it was probably also true of ginkgo and its relatives in many other ancient
plant communities between about 100 million and 225 million years ago.
When we delve deeply into the fossil record of many groups of plants or animals,
the situation that has come to light for ginkgo turns out to be quite common. For ex-
ample, in the case of the modern horse, we have not just one ancient lineage leading in
ladderlike progression to the single living species; rather, we have fossils of many kinds
of extinct horses, some of which browsed together in the same ancient landscapes. The
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origin and prehistory

pattern in the fossil record of horses is a bush of many closely related species that was
winnowed by extinction over time to a single lucky winner.
Based on Zhou Zhiyan’s work on different kinds of ancient ginkgolike fossils,
and his painstaking analyses of the ginkgo fossil record worldwide, a similar picture
emerges of once-great diversity among ginkgo and its relatives, followed over time
by the gradual emergence of a single winner and many losers. From the Jurassic on-
ward, and perhaps from the latest Triassic, there is strong evidence of plants similar to
the living species. However, at first, this lineage was just one among many, just as the
lineage leading to ourselves was one among many hominids on the savannah of East
Africa between a half-million and a million years ago. There were once other lineages
of ginkgolike plants, and each of those also had multiple species. Whether it is the evo-
lutionary history of horses, of ginkgo, or of ourselves, the pattern is the same: sudden
appearance and proliferation, followed inevitably and inexorably by many losses, in
which most species fall by the wayside. In the cases of ginkgo, horses, and ourselves,
only one species survived.8
 17
Persistence

The “tendency to persevere,” . . . it is this that in all

things distinguishes the strong soul from the weak.
—Thomas Carlyle, Chartism: Past and Present

My own small contribution to what we know about the prehistory of ginkgo came
from work on fossils much younger than those from the Triassic and Jurassic studied
by Tom Harris and Zhou Zhiyan. It began in the late summer of 1982 as I was driving
back to Indiana University after a long trip collecting fossils with my colleague David
Dilcher. I was just wrapping up a year of work in David’s laboratory before moving to
the Field Museum in Chicago. The trip had started with fossil hunting in the drylands
of eastern Oregon, followed by a long swing through Idaho, Wyoming, and Colorado.
Now in North Dakota, with the end of summer approaching, I was getting ready to
move on. Ginkgo could not have been farther from my mind. We had seen no trace of
it at any of the fossil sites that we had visited that summer.1
As the last stop on our trip, we were trying to track down the source of an interest-
ing collection of fossil plants that David had been given by Rudi Turner, a colleague at
Indiana. Rudi, an enthusiastic fossil collector in his spare time, had come across these
fossils at various rock and fossil shows organized by amateur and commercial collec-

125
 126
origin and prehistory

The leaf of an ancient ginkgo, Ginkgo cranei, from the Late Paleocene of
Almont, North Dakota, about 57 million years before the present.

tors. He had traced the material to a newly discovered site a little west of Bismarck,
North Dakota.
As we pulled the car off I-94, the main east-west highway, the signs were not encour-
aging. The road headed north through flat agricultural land. There were no road cuts or
badlands to explore, mostly just fields of sunflowers, until eventually, at the designated
spot, we came to a single figure bent over by the side of the road among low piles of
sharp, yellow-brown shale. As we pulled up, he greeted us warmly, and the first fossil he
showed us was an exquisitely preserved ginkgo leaf, laid out complete and absolutely
perfectly on the hard shale.
Ginkgo leaves were the main target for our new friend and other local fossil col-
lectors who were working at the site. The fossils were attractive, reasonably common,
and easily recognized, often with all the fine veins nicely picked out in red and brown
against the ochre background. It was easy to see why these distinctive fossils, dating
from about 57 million years ago, were a prized curiosity in the now nearly treeless land-
scape of this part of North Dakota.
On this first visit to what we later called the Almont locality, the focus of local collec-
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persistence

tors on ginkgo leaves was a boon to David and me. We had broader interests, and the
heaps of discarded material were packed with other fascinating plant fossils, many of
them beautifully preserved. Especially common were the flying saucer–shaped fruits of
the Asian wheel wingnut. Circular in outline, and about an inch across, these were the
fruits that had first attracted David’s interest. Also prominent in the Almont collections
were the leaves, pollen catkins, and characteristic spiny fruits of an early extinct rela-
tive of hazels and hornbeams, which turned out to be similar to fossils I had described
a year or two earlier from a locality of about the same age in southern England. The
collecting that afternoon was some of the most exhilarating I have ever experienced;
there was something new on almost every piece of shale. It quickly became obvious
that there were many interesting fossils at Almont, especially a rich assortment of fossil
fruits and seeds, which would be worthy of detailed study.2
Along with the ginkgo leaves, it was also easy to recognize ginkgo seeds among the
fossils in the Almont shale. With the leaves so common, and with so many other kinds
of fruits and seeds present, it would have been strange if they had not been there as
well. Often the leaves and seeds occurred together on the same piece of shale. Usually,
the only part of the seeds preserved was the hard inner shell, about half an inch long,
with a distinctive raised longitudinal seam running round the outside and a small point
at the top. However, in a few cases the hard layer was enclosed in a shiny covering
almost certainly formed from the remains of a silvery, waxy coat like that of modern
ginkgo seeds.
It was also striking that both the leaves and the seeds had the same characteris-
tic red-brown appearance and shiny, leathery texture, probably a result of their thick
waterproof covering and the resin in the original tissues. It was obvious that they “be-

The seed of an ancient ginkgo, Ginkgo cranei, from the

Late Paleocene of Almont, North Dakota, approximately
57 million years before the present.
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origin and prehistory

longed together” and had been shed from the same kind of tree. Somehow they had
been washed into a small pond on the ancient floodplain, buried in mud, and pre-
served together until we collected them on that August afternoon 57 million years later.
After several hours sifting through piles of shale discarded by previous collec-
tors, and with the sun going down, we wrapped the last fossil and loaded the last box.
Almont was a paleobotanical treasure trove that was hard to leave. The fossils were
superbly preserved and partly impregnated with silica. It was clear that Almont would
provide more detailed information on ancient plants than any of the other fossil locali-
ties that we had seen that summer. The quality of preservation was better than that of
any fossil locality of similar age anywhere in the world. We could not resist coming
back the next morning for a few more hours, and when we eventually eased back onto
the interstate heading for Chicago and Indiana, David’s aging Oldsmobile was well
down on its springs.
Over the next few years I returned to Almont several times, and on two or three
occasions I sent a small team from the Field Museum on the long drive from Chicago to
collect there. Other paleobotanists found their way there, too. The collecting was easy,
the landowner was more than tolerant, and the fossils turned out to be just as infor-
mative as we had anticipated back in the early 1980s. Eventually, David Dilcher, Steve
Manchester, and I published an overview of the Almont fossil flora, detailing not only
the ginkgo leaves and seeds but also—importantly for ginkgo’s biography—some new
information on how the mature seeds were borne on this ancient fossil plant.3
On that first and many subsequent visits to Almont, we collected many plant frag-
ments that were puzzling. We knew that there was plenty of work to be done, and just
because we didn’t immediately understand what a particular fossil was, that didn’t
mean that we would not collect it. We brought all the oddities back and puzzled over
them in the laboratory. Among them were some especially enigmatic fossils that looked
as if they ought to be somehow connected to the ginkgo leaves and seeds. They had the
same texture and color, and were superficially like the stalk of a ginkgo leaf, but they
didn’t look quite right. For one thing they never seemed to be attached to a leaf blade,
and for another they had a peculiar knobbly structure at one end.
Back in Chicago, under a microscope these strange fossils provided a bit more infor-
mation, and eventually I decided to rummage around under a few living ginkgo trees
to see what was left in the leaf litter beneath them. This unconventional approach has
served me well over the years. Textbooks often provide a beautiful portrayal of a plant
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persistence

in all its structural details, but for a paleontologist such images are excessively sani-
tized. When you are trying to interpret fossils, what you really want to know is what a
tree looks like when it falls apart: which pieces normally become detached, and what
do they look like as they start to decay? This is the best way to find out what pieces of
a plant have a chance of ending up as a fossil.
A couple of minutes under an old female ginkgo tree were enough to get me what
I needed. Among the rotten fruits were exactly the same kinds of knobbly stalks that
we had collected from the Almont fossil site, and among the other ginkgo debris it was
easy enough to figure out what they were. The unusual pieces of the Almont Ginkgo
were the shed stalks on which the mature seeds had been borne. In living ginkgo the
seeds and the stalks fall from the tree at about the same time, becoming separated from
each other in the process. In the fossil ginkgo from Almont, as in the living species,
there was usually only one large seed scar on each stalk, indicating that only one of the
two seeds on each seed stalk had matured. However, at the tip of both the fossil seed
stalks and those from living ginkgo, the remains of the other young seed that had failed
to develop were often easy to make out. It was obvious that the fossil, like the living
species, had two seeds on each seed stalk but that normally only one of those seeds de-
veloped.
Apart from this minor triumph, which solved one of the small problems raised by
the Almont fossils, the recognition that the ginkgo growing in North Dakota around 57
million years ago bore its seeds in exactly the same way as the living species was useful
new information. It showed that the similarities between this ancient ginkgo and its
living relative extended beyond the form of the leaf and the shape and size of the seeds
to other aspects of the plant. This was one more piece of evidence to justify using the
Latin name Ginkgo for these leaves, with the important implication that the rest of this
ancient plant, if we understood it in all of its details, would not differ significantly from
the living tree.4
One of the interests of the late Tom Schopf, a former colleague at the University of
Chicago, was “living fossils,” and as one of his last projects Tom wrote a commentary
about the supposed living-fossil status of a wide variety of plants and animals. Classic
examples from the world of animals include the clamlike creature Lingula, which still
lives in the cold shallow waters off northwestern Europe and appears to have survived
more or less unmodified from the Silurian period, more than 450 million years ago.
Among animals with backbones, the coelacanth first occurs in rocks about 350 million
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origin and prehistory

years old but survives in deep ocean waters off the Comoros in the Indian Ocean, and
Malaysia in the South China Sea. Other well-known living fossils include the horseshoe
crab, the paddlefish, and the platypus.5
From this compilation Tom pondered some of the questions that living fossils raise
for our ideas of plant and animal evolution, especially whether the apparent absence of
change over vast periods of geologic time reflects reality or illusion. He wanted to know
whether the absence of information misleads us into thinking that living and fossil
organisms are the same, when we would clearly recognize them as different from each
other if we knew more about them. Tom raised the possibility that significant changes
could be occurring in these organisms through time but were going undetected be-
cause they are invisible in those parts of organisms that are preserved as fossils.
Dianne Edwards, a paleobotanical colleague at the University of Wales, uses the
analogy of the Mini, a car first introduced in the United Kingdom in 1959 and now
manufactured by BMW. From the outside the 1960s version and the twenty-first-
century version are unmistakably similar, but inside they could scarcely be more dif-
ferent. From the computer chips in the engine to the liquid crystal displays on the
dashboard, the technology of the modern Mini is rather different from that of its pro-
genitor. But if you had only a photograph of the outside, how would you know?
Tom asked, just because the shells of Lingula from the Silurian and today look the
same, does that mean that the animals inside were identical? If we could go back in a
time machine and take our microscopes and laboratory with us, would we find that the
ancient Almont Ginkgo of 57 million years ago was more or less the same as or quite
different from that of today? I suspect that we would find the former. There is much else
in the landscape of 57 million years ago that we would find strange, but the evidence
we have so far suggests that we would find the ancient ginkgo comfortingly familiar.
Even though the paleontological lens through which we view these ancient plants is
obviously imperfect, it seems that the more we find out about the Almont Ginkgo,
the more it looks like its modern counterpart. The similarity of the fossil seed-bearing
stalks from Almont to those of living ginkgo was one more piece of evidence pointing
in that same direction.
Of course, it is possible that future research will discover significant differences—for
example, in the structure of the pollen cones, pollen, or wood—but I suspect that the
reverse is much more likely to be true: as we find out more about the Almont Ginkgo, it
will come to look more and more like the living tree. It will be interesting to see whether
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persistence

the next generation of paleobotanists to turn their attention to Almont will take up this
question. For example, among the small catkins that Steve Manchester, David Dilcher,
and I described is one that we perhaps passed over too quickly; now, with the benefit
of hindsight, it looks very like a shed ginkgo pollen catkin. The fossil pollen that we
extracted from a similar specimen doesn’t look right, but fossils like these certainly de-
serve a more careful look. The prediction is that if it is a ginkgo pollen catkin, its struc-
ture and its pollen grains will turn out to be more or less identical to that of the living
species.6
Just as the structure of ginkgo seems to have remained virtually unchanged for tens
of millions of years, it seems likely that aspects of its ecology have also remained con-
stant. Dana Royer from Wesleyan University, Leo Hickey from Yale, and Scott Wing
from the U.S. National Museum of Natural History analyzed forty- eight separate fos-
sil occurrences of ginkgo leaves from western North America, ranging from about 55
million to 65 million years ago, all roughly similar in age to the Almont fossils. In all
but two cases the geological situation in which the fossils occurred suggested that the
leaves were preserved in ancient muds and sands and deposited in, or close to, river
channels. Occurrences of ginkgo in lake deposits, in deltas, or in other situations were
either nonexistent or very rare. Dana Royer and his colleagues concluded that these an-
cient ginkgos consistently favored streamside habitats, and they pointed out that these
are also the kinds of environments where ginkgo seems to flourish today.
In general, modern ginkgo dislikes being deeply shaded by other plants. It grows
best on sites that are partly or fully exposed. Living ginkgo trees in China show that
they favor steep rocky slopes, cliff edges, and—most significant for comparison with
the fossils—stream banks. They like to be partly in the open and partly in the shade.
In addition, as many of the great ginkgo trees of Asia, such as the giant Yongmunsa
Ginkgo, show us, ginkgo does especially well where its roots have good access to water.
The same was probably true for ancient ginkgo that grew 60 million years ago in the
American West. They thrived in partly open habitats and especially along rivers. By a
happy accident, this also happens to be the right place for their leaves, seeds, and other
parts to be incorporated into the river mud and sand accumulating nearby. Long-term
constancy in its ecology, combined with its tough and easily recognized leaves, is one
of the reasons why ginkgo has such an excellent fossil record.7
 18
Prosperity

Everything in the world may be endured except continued prosperity.

—Johann Wolfgang von Goethe

John Starkie Gardner is not among the truly great paleontologists of the later nine-
teenth century, but he was nevertheless energetic and talented. Between about 1879
and 1887 he produced several publications on fossil plants, including the two-volume
British Eocene Flora, before abruptly bringing all his paleontological work to a close
and devoting himself instead to a second career as an expert on decorative ironworks.
He created Victoria Gate in London’s Hyde Park and the iron gates and screens at
Edinburgh’s Holyrood Palace. As such a radical and sudden change of direction might
suggest, Gardner was a forceful character who held strong views.1
In the 1880s Gardner clashed spectacularly in the pages of the scientific journal
Nature with one of his German contemporaries, Baron Constantin von Ettingshausen,
over the scientific reliability of some of the latter’s work. The two had previously col-
laborated on a study of ancient ferns, conifers, and ginkgo from southern England, but
they parted company on how to deal scientifically with the massive collections of fossil
leaves that Gardner had accumulated from his excavations along the cliffs at Bourne-
mouth in Hampshire. These leaves, from localities now long obliterated or covered by

132
 133
prosperity

development in this popular seaside town, date from the Middle Eocene, about 40 mil-
lion to 50 million years before the present.2
What bothered Gardner was Ettingshausen’s tendency to rush into print with new
identifications and formal Latin names for fossils that had not yet been studied thor-
oughly. He disapproved of Ettingshausen’s cavalier attitude toward linking these leaves
with specific living plants. It was true that Ettingshausen had studied the patterns of
veins in the leaves of different kinds of broadleaved plants more carefully than anyone
else, and that he had also published a beautiful atlas of the leaves of living plants, which
he had carefully “skeletonized” to reveal their delicate vein patterns. The illustrations
in his book were striking white- on-black silhouettes. However, Gardner was right:
Ettingshausen’s way of working was to match fossil and modern leaves without care-
ful comparison. If they looked broadly similar in shape and vein pattern, he was quick
to say the fossil and living plants were the same. Gardner was exasperated. He knew
that the numbers of plants to which the fossils had to be compared was overwhelming,
and he knew also that the leaves of even distantly related plants often look confusingly
similar. He was acutely aware of the possibility of making massive, misleading mis-
takes. Gardner could not sign on to an approach that he regarded as patently unscien-
tific.
Fortunately, identifying ginkgo in the fossil record is much less problematic than
identifying the often rather nondescript leaves of flowering plants that make up most
fossil floras from the middle of the Cretaceous onward. One of the wonderful things
about ginkgo, from a paleobotanical point of view, is that the leaves are so distinctive:
they are unlikely to be overlooked or confused with anything else. So given how wide-
spread ginkgo had been during the Jurassic and Cretaceous, it is perhaps significant
that in the whole of Gardner’s massive collections of fossil leaves from Bournemouth,
there is no example of anything like a ginkgo leaf. In their early work together, before
their falling out, Gardner and Ettingshausen had described fossil ginkgo leaves from
Scotland, but there is not a single ginkgo leaf among the many hundreds of specimens
collected from Bournemouth, which today occupy many tens of drawers in the Natural
History Museum in London.
Ginkgo is also strangely absent from another massive and important collection of
fossil plants from southern England, which also dates from the Eocene. Fossils col-
lected from the beaches of the Isle of Sheppey in the Thames Estuary, just forty miles
 134
origin and prehistory

east of central London, provide an unusually detailed window into the plant life of
between about 45 million and 55 million years ago. This so-called London Clay flora
has been collected since the earliest days of scientific paleontology. Largely through
the efforts of Eleanor Reid and Marjorie Chandler, two pioneering women paleon-
tologists, what has been learned from the London Clay fossils exceeds that from any
other collection of fossil plants of similar age. Reid and Chandler’s classic work, The
London Clay Flora, published by the Natural History Museum in 1933, is a hefty tome.
It described and illustrated more than four hundred species and set new standards of
accuracy in the comparison of living and fossil plants. Its importance rests in part on
its being based on studies of beautifully preserved fossil fruits and seeds rather than of
fossil leaves.3
Fossils from the London Clay are preserved in three dimensions in iron pyrite, a
mineral sometimes called fool’s gold. They weather out from the soft, chocolate-brown
cliffs along the northern shore of the Isle of Sheppey, and are concentrated on the
muddy foreshore by the tides and the waves. Collections from Sheppey are especially
extensive, but similar fossils have been collected from other places around the coast of
southeastern England, particularly from Herne Bay in Kent, Bognor Regis in West Sus-
sex, and Walton-on-the-Naze in Essex. However, at all localities the fossils are unusual
because they occur in clays that were clearly deposited in the sea. Fruits and seeds are
found alongside the teeth of sharks and rays, the shells of marine snails, and the fos-
silized carcasses of crabs and shrimps. This is a little strange; it is much more common
for plant fossils to be preserved in muds and sands laid down in ancient freshwater
lakes, ponds, and river systems on the land. In the case of the London Clay, the fruits
and seeds were washed out to sea, eventually sank, and were then literally petrified in
the fetid mud at the bottom of the ocean.
These peculiarities of the London Clay flora bring with them several advantages
over fossil plants collected from ancient floodplains. First, because the fossil fruits and
seeds had drifted out to sea, they provide an unusually broad sample of the plants that
grew along the ancient shoreline of the London Clay Sea and the banks of rivers that
emptied into it. Second, the preservation in iron pyrite means that the tissues of the
fruits and seeds are well preserved, not just on the outside but also on the inside. Often
exquisite details of internal structure are visible, enabling these ancient fruits and seeds
to be compared carefully with their living counterparts.
The London Clay flora also has a third advantage; it has been collected intensively
 135
prosperity

A nineteenth-century botanical
print of the living stemless nypa
palm, which grows today in tropical
mangrove habitats around the coast
of southeast Asia. Abundant fossil
fruits of nypa have been recovered
from the London Clay, on the Isle
of Sheppey in the Thames Estuary.
The presence of nypa indicates the
warmth of the climate about
50 million years ago.

for more than 150 years. Hundreds of people have spent untold hours on the beach at
Sheppey and elsewhere looking for these fossils. Many thousands of specimens have
been collected. The London Clay flora is by far the richest and most informative Eo-
cene flora currently known. As at Bournemouth, though, ginkgo has so far failed to put
in an appearance in the London Clay. The hard inner shell of a ginkgo seed is distinc-
tive, would be readily preserved, and would quickly have been recognized by Eleanor
Reid, Marjorie Chandler, and the many other specialists who have studied the Sheppey
fossils over the years.4
An important clue as to why ginkgo might be missing from Gardner’s Bournemouth
collections, and also from the London Clay, comes from the most conspicuous plant
fossils collected from the beach at Sheppey; the distinctive fruits of the stemless palm
 136
origin and prehistory

nypa. Today nypa is native to tropical southeast Asia and entirely restricted to brack-
ish, often partly tidal, mangrove habitats that are protected from full exposure to the
ocean. Not only does good representation of such a coastal plant in the London Clay
make ecological sense, but extrapolation from its living relative strongly suggests that
the vegetation of London Clay times grew in a tropical or near tropical climate.5
Other plants that are common in the London Clay flora, such as palms and mem-
bers of the sweetsop family, support this idea. They point to dramatically warmer tem-
peratures than those in the Thames Estuary of today. For example, there are abundant
climbers, from the grape and moonseed families, as well as true mangroves. In the flora
as a whole, plants that are now characteristic of tropical southeast Asia are especially
well represented. There are also bay trees, magnolias, and a variety of other plants,
which while not truly tropical, prefer warmer climates. The strong overall impression
is of an ancient coastal plain clothed with lush subtropical-to-tropical vegetation. The
London Clay flora, together with other fossil floras from elsewhere in Europe, tell us
that climates during the Eocene, even at midlatitudes, were very warm.
To the north, however, in slightly older rocks that seem to have deposited in slightly
cooler climates, the situation is different. The columnar basalts exposed along the rug-
gedly beautiful southwest coast of the Isle of Mull in Scotland rival those at the Giant’s
Causeway on the other side of the Irish Sea in County Antrim, Northern Ireland. They
date from around the Early Paleocene, about 60 million to 65 million years before the
present, and were formed by successive outpourings of molten basalt from a deep geo-
logical rupture in the Earth’s crust that ultimately helped create the North Atlantic. On
the Giant’s Causeway, on Mull, and also on the Isle of Staffa, where they create Fingal’s
Cave, the hot basalts cooled into layers of tall hexagonal columns.6
Ardtun Head, near the southwestern tip of Mull, is formed by several layers of mas-
sive black columnar basalt, but as early as 1851 the landowner, the Duke of Argyll, who
also had a keen interest in the emerging science of geology, described the details of the
rock section exposed in the cliffs. He was also among the first to draw attention to the
softer rocks between the basalts and the fossil plants that they contained. The main fea-
tures of the geological section that the Duke of Argyll described are still clearly visible
on Ardtun Head today. They show three main layers of basalt that range from ten to
forty-eight feet thick, between which are siltstones laid down not by volcanic activity
but by water. As successive layers of basalt cooled, they were colonized by vegeta-
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prosperity

tion, just as happens today, for example, on Iceland or on the Big Island of Hawai’i. In
the muds created by the streams, ponds, and lakes on this new landscape, leaves and
other plant parts became entombed as fossils before they were eventually buried more
deeply, and for the long term, by a new outpouring of lava.7
It was to these fossils, preserved in the soft layers between the basalts on the Isle
of Mull, that John Starkie Gardner brought the same enthusiasm for collecting that
he brought to the cliffs at Bournemouth. When challenged by the massive hard black
basalts on the Isle of Mull, he resorted to dynamite, and with great effect. Spectacular
plant fossils from Mull are now housed along with those from Bournemouth in the col-
lections of the Royal Scottish Museum in Edinburgh, the Hunterian Museum in Glas-
gow, the Natural History Museum in London, and elsewhere. Together with ginkgo,
the fossils that Gardner collected include the leaves of ancient hazel, oak, laurel, and
katsura trees. Taken together, they point to cooler climates than those of the London
Clay.8
This pattern seen in Britain, where ginkgo appears to be missing from more tropical
floras but is present in fossil floras from cooler climates, seems to hold across Europe
from about 40 million to about 65 million years ago. For example, ginkgo is absent
from the remarkable fossil assemblage collected in an old oil shale mine at Messel not
far from Frankfurt, Germany. The truly exceptional preservation of ancient animal life
at Messel includes mammals with the remains of skin, hair, and gut contents, as well as
a hummingbird with feathers. The abundance of bats and crocodiles is one of several
indications of tropical conditions, which is also confirmed by the plants, which include
palms, ferns, and aroids. In the past few years, a superb study by the young paleobota-
nist Selena Smith showed that one especially puzzling plant fossil, which had remained
enigmatic for many years, is actually the fruiting stalk of the Panama hat palm, one of
a group of plants that today grows only in tropical Latin America.9
In North America, the picture is the same. The Eocene fossil floras from Kentucky
and Tennessee have been collected extensively for more than a hundred years, espe-
cially in the early twentieth century by E. W. Berry and those who worked with him.
More recently, during the 1960s, 1970s, and 1980s, enormous collections were made
by David Dilcher and his students. There are tens of thousands of specimens from the
Eocene fossil floras of Kentucky and Tennessee in the collections of many museums
across the United States. The collections of the National Museum of Natural History in
 138
origin and prehistory

Washington, D.C., and the Field Museum in Chicago, as well as the Dilcher collections
now at the Florida Museum of Natural History, are especially rich. Among all this ma-
terial, however, there is not a single ginkgo leaf.
Ginkgo is also missing among the rich collections of Eocene fossil leaves made from
Castle Rock near Denver by Kirk Johnson and his team from the Denver Museum of
Natural History. This locality came to light in the early 1990s, when excavations were
made to broaden the interstate highway that climbs up from Denver through the Front
Range of the Rocky Mountains. The Castle Rock plants have a distinctly tropical look
to them, and many species have the large leaves characteristic of tropical plants, but
again there is no ginkgo. Ginkgo has also never been found among the large collec-
tions of fossil plants made from the Green River Fossil Basin of Colorado, Utah, and
Wyoming. Palms and many other warmth-loving plants are well represented, and there
are crocodiles and turtles among the animals of “Fossil Lake,” but among the collec-
tions of fossil leaves from these deposits there is no evidence of ginkgo.10
Ginkgo is present, however, in one of the most intensively studied and most infor-
mative Eocene fossil floras from North America: that from the ancient mudflow of
Clarno Formation, not far from the town of Fossil in eastern Oregon. The fossil plants
at Clarno are preserved as jumbled fruits, seeds, and leaves in an ancient mudflow
created by nearby volcanic activity. Like the fruits and seeds preserved in the London
Clay, the fossils have been literally turned to stone, in this case not by iron pyrite at
the bottom of a stagnant ocean but by silica dissolved in the warm waters produced by
nearby volcanoes.
Fossil fruits and seeds from the Clarno fossil assemblage have been painstakingly
collected over several decades, especially by the late Tom Bones, with the help of gen-
erations of high school students through programs run by the Oregon Museum of
Science and Industry. Steve Manchester, one of the graduates of that program and
now a professor at the Florida Museum of Natural History, has provided the defini-
tive description of the Clarno flora. Thousands of fossil fruits and seeds have been col-
lected, as well as hundreds of specimens of leaves, but so far the only clear evidence of
ginkgo is a piece of petrified ginkgo wood and a single unmistakable ginkgo leaf. Again
the flora is strongly indicative of warm conditions. There are fruits of palms and the
sweetsop family, and as in the London Clay there is an abundance of climbers. But in
this case ginkgo was present, although apparently rather rare.11
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prosperity

Similarly, not too far away from the Clarno Formation, and in rocks of about the
same age, ginkgo puts in an appearance in the rich fossil floras preserved in ancient
lakes during early evolution of the northern Cascade Mountain Range. These lakes,
which were gradually filled with ash and other fine volcanic debris, preserved fish,
insects, and occasional mammals, as well as the leaves and other parts of plants that
grew nearby. Several of these fossil-producing ancient lakes straddle the U.S.-Canadian
border in the Okanagan Highlands of northern Washington State and adjacent British
Columbia, but by far the most thoroughly studied is that around the small town of Re-
public in northeastern Washington, not far from where the Columbia River passes into
Canada.
Like the fossil plants at Clarno, those from Republic have been extensively collected,
not only by teams of professionals but also by many schoolchildren led by the late Wes
Wehr, working from a base at the Burke Museum of the University of Washington in
Seattle. The dedication of this multifaceted and truly lovely man resulted in the col-
lection of tens of thousands of specimens from the Republic site, among which were
many spectacular fossils. The flora looks much more temperate than that from Clarno
and the Green River; in place of palms and tropical climbers there are currants and
witch hazels, along with ancient representatives of the birch, elm, oak, rose, and wal-
nut families. Ginkgo is also present, but extremely rare; it is represented by just one or
two fragmentary but unmistakable leaves. Floras of similar age from China that include
ginkgo are also known from Liaoning Province.12
One explanation for the presence of ginkgo in the Republic flora and perhaps at
Clarno, in contrast to its absence among the fossil plants from the Green River Forma-
tion and from Kentucky and Tennessee, may be slightly increased elevation, as well as
higher latitude and hence slightly cooler temperatures. At the time that the leaves were
washed into the Republic Lake, they may have been about 2,300–3,000 feet above sea
level. Ginkgo seems to prefer these cooler temperatures, and this preference is dramati-
cally confirmed by the fact that at roughly the same time ginkgo was growing at very
high latitudes within the Arctic Circle, not too far from the North Pole.13
On Ellesmere Island, Canada, high in the Canadian Arctic, Jim Basinger and the
late Elizabeth McIver of the University of Saskatchewan described ginkgo among the
ancient plants growing in and around high-latitude peat bogs. The flora, which is not
rich compared to those farther south, includes a few different kinds of broadleaved
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origin and prehistory

trees alongside early firs, pines, and the unmistakable leaves of the dawn redwood. The
presence of forests with ginkgo in the Arctic around 55 million years ago is a vivid re-
minder that the familiar configuration of our planet, with permanent ice at the poles
and year-round warmth only in the tropics, is the exception rather than the norm in
the past 200 million years of life on Earth.14
 part iv

Decline and Survival

OVERLEAF A Japanese family crest of three stylized ginkgo leaves.
 19
Constraint

What does not kill me, strengthens me.

—Friedrich Nietzsche, Nietzsche contra Wagner

Today, ginkgo can be grown easily in many parts of the world, including over most
of Europe and the United States, as well as much of eastern Asia, but it is confined
mostly to what might loosely be called the temperate regions. For example, in Europe,
ginkgo flourishes from Paris in northern France to Montpellier in the south, and the
connection to Goethe has made ginkgo popular throughout Germany. It is not a tree
that survives across most of Finland, though, or that thrives at the other end of Europe’s
climatic spectrum—for example, in Sicily or the Greek Isles, where both high tempera-
tures and scarce water are problematic. Similarly, in Australia, ginkgo grows happily
outdoors in Melbourne, and also in Sydney, but you will not find it on the streets of
Cairns or Darwin. There seems to be a reasonably clear band of latitude within which
ginkgo flourishes but outside of which it struggles and eventually dies. There are con-
straints of some kind, probably a combination of constraints, which limit the survival
of ginkgo both toward the poles and toward the tropics.1
The different aspects of climate that govern the distribution of ginkgo and other tree
species are complicated, and made more so by the fact that local conditions, microcli-
mates, and the availability of nutrients and water in the soil can vary dramatically over

143
 144
d e c l i n e a n d s u r v i va l

short distances depending on aspect, elevation, or proximity to fresh and salt water. All
gardeners know that whether a plant flourishes, struggles, or dies in a particular place
depends on many different things. However, a key constraint on the growth of ginkgo
toward the poles and at high elevations is obviously cold temperature. In the plant
hardiness zones recognized by the United States Department of Agriculture, which are
widely used by gardeners in North America as a guide to the likely tolerances of par-
ticular plants, the northern limit of ginkgo is zone 5, defined by average annual mini-
mum temperatures down to minus 20 degrees Fahrenheit.
In China, ginkgo grows well in climates with an average annual temperature in the
fairly broad range of 50–65 degrees Fahrenheit and an annual rainfall in the range of
twenty-four to forty inches a year, but from the North American hardiness zones it is
also clear that ginkgo’s tolerances are broad. The tree can tolerate very low winter tem-
peratures and very high summer temperatures, at least in short bursts. This makes it
possible for it to thrive in places like Chicago, where winter temperatures can go down
to minus 27 degrees Fahrenheit and summer temperatures can top out at 108. Simi-
larly, ginkgo can get by in Minneapolis–Saint Paul, where winter temperatures are even
more extreme and the winters even longer. Nevertheless, there are obviously limits.
Ginkgo will not survive in North American hardiness zones 1 and 2, or colder places
where winter temperatures may plummet to minus 45 or 50 degrees or lower.2
Ginkgo can survive, although not always happily, in all fifty U.S. states, including
Alaska. Along the East Coast of North America, ginkgo grows well from Charleston,
South Carolina, to Montreal in Quebec, but there are no ginkgo trees in Labrador or
the southernmost parts of Florida. In Hawai’i ginkgo grows well only in the mountains.
Along the West Coast, ginkgo grows from San Diego to Vancouver, but farther north
in Alaska, or farther south in Baja, it is a different story. Nursery owners in Juneau,
southern Alaska, report that the tree can hang on for a number of years but it does not
thrive, and it will not survive farther north in Fairbanks.
The situation is similar in Europe. For example, in Copenhagen, Denmark, or even
Lund, southern Sweden, ginkgo trees grow outside without special protection, but a
little farther north—for example, around Stockholm or Uppsala in central Sweden—
ginkgo can normally be grown only inside. Ginkgo trees in the garden of the Komarov
Botanical Institute in Saint Petersburg, at nearly 60 degrees north latitude, are among
the most northerly in Europe. However, ginkgo cannot be grown outside in Moscow,
away from the ameliorating influence of the Baltic.
Part of ginkgo’s success in tolerating extreme cold is its deciduousness. As in most
 145
constraint

In early spring clusters of tiny ginkgo leaves emerge from the buds in which they have
been protected all winter. The scars of last year’s leaves show the traces of the two strands
of conducting tissue that supplied the leaves with water.

temperate broadleaved trees, shedding the leaves is one way to avoid the damage that
would be caused by ice crystals forming in the living tissues or by water loss through
the leaves, during the time when the groundwater is frozen. In effect, deciduousness
shifts the problem of winter survival to the easier task of protecting the young leaves
inside the overwintering buds. In ginkgo the buds are small, rounded, and well pro-
tected by a mass of tightly packed, overlapping, light brown, papery bud scales. Some
temperate trees, and perhaps also ginkgo, also have an ingenious mechanism by which
the buds are actively supercooled so that chances of ice crystal formation are further
minimized. However, this mechanism appears to fail at temperatures of about minus
40 degrees Fahrenheit: at colder temperatures ice crystals form spontaneously and
damage the living tissues. It seems likely that poleward limit of where ginkgo can grow
outside in part reflects the extent to which it can protect the next season’s leaves inside
their buds.3
While absolute winter temperatures are one factor that helps set a poleward limit for
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d e c l i n e a n d s u r v i va l

the growth of ginkgo, the length of the growing season is probably just as important.
If the leaves can’t harvest enough energy in a short summer to cover what we might
think of as “the costs of maintenance,” then growing in those places is probably not a
viable proposition. In most cases where there is a tree line, a clear observable limit to
tree growth, either with increasing latitude or with increasing altitude, these seem to
reflect not winter temperatures that are too cold for trees to survive but rather some
critical limit in the length and warmth of the growing season. This supposition also fits
with the observation that in many tree species photosynthesis still works at low tem-
peratures, but if levels of activity are insufficient, then the energy is stored and saved
for later rather than used directly for growth.4
The susceptibility of ginkgo and other trees to extreme cold also varies considerably
depending on the stage of life at which a particular tree encounters frigid temperatures.
Young plants are generally much more sensitive than plants that are well established,
so the regularity and timing of extremely cold temperatures in relation to the plant’s
development is important. If a large tree loses a few leaves in a late frost, it will prob-
ably have enough stored reserves of energy to grow new ones. For a seedling with just
a few leaves and limited food reserves, however, it is quite a different story. At Kew, ex-
perience with some frost-sensitive trees, such as certain eucalypts, showed that if their
time as seedlings happened to coincide with consecutive mild winters, which allowed
them to become well established, then their chances of surviving when the next hard
winter came along were greatly increased. However, the reverse is also true. In south-
ern England the winter of early 2010 was one of the harder ones of recent decades. A
colleague at Kew who had ginkgo seedlings growing outside lost almost half of them.5
In ginkgo a further complication is that the length of the growing season affects the
development of the embryo in the maturing seed and therefore the opportunities for
the young plant to get established in its first season. A study by Peter Del Tredici shows
how important such effects can be. Using careful observations and some simple ex-
periments, Peter studied how reproduction in ginkgo is affected by temperature based
on comparisons of plants growing at the Arnold Arboretum of Harvard University, at
about 42 degrees north latitude, and in Guizhou Province, China, at about 25 degrees
north latitude.6
At the Arnold Arboretum ginkgo pollen cones generally emerge from the winter
buds around mid-May, and pollination takes place soon after. Fertilization happens
about four and a half months after that, and the resulting seeds, which are generally
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shed from the tree in late October, germinate the following spring. On the face of it
there is no problem with the reproductive cycle, and under normal conditions it seems
as though ginkgo ought to be able to reproduce itself by seed in the climate of present-
day Massachusetts. However, by comparing the timing of these various stages in Mas-
sachusetts with the timing in Guizhou, Peter showed that the situation in the two places
is actually quite different, and that those differences could be significant for plants try-
ing to establish themselves in the wild.
By carefully following seed development in trees growing outside, and comparing
them with seeds kept in a warm greenhouse over the winter, Peter was able to show
that while the tempo of the annual reproductive cycle is remarkably similar in both the
Chinese and North American trees, the way in which the seeds develop is very differ-
ent. Seemingly irrespective of temperature and latitude, fertilization typically occurs
about 130–140 days after pollination. The time from pollination to germination was
also remarkably constant at around 233–234 days for the Guizhou plants and the seeds
kept in the greenhouse in Massachusetts. Maturation of the embryo from fertilization
to germination was also similar, taking roughly 100 days in both cases. The embryo de-
velops continuously without a pause, and there is no natural period of dormancy.7
However, seeds developing outside, without the protection of a greenhouse, experi-
ence very different conditions in Massachusetts compared with Guizhou. In both cases
embryo development slows or stops completely during the winter. This helps prevent
the seeds from germinating too early, which increases the seedlings’ chances of sur-
viving the cold part of the year. It does, however, increase the time to germination. In
the relatively short growing seasons and long winters of the northeastern United States
this means that germination is significantly delayed compared with southern China,
where the winters are shorter and less severe, and the growing season is correspond-
ingly longer.
In the northeastern United States, pollination takes place around mid-May and fer-
tilization takes places around the end of September or the beginning of October. A
month or two later, the seeds are shed. The embryo, whose maturity at the time of seed
drop depends largely on local conditions, continues to develop on the ground. This
means that the embryonic plant has only a month or so to develop before growth slows
or shuts down completely with the onset of cold weather. As a result, development is
carried over into the following season, and because of the relatively late spring and the
additional time needed for the embryo to undergo its development, germination does
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not take place until mid- to late June. The whole process takes thirteen months from
pollination to germination, and the young seedling has only about five months to be-
come established before the next winter, with its potentially fatal cold temperatures.8
The contrast with the life cycle of ginkgos growing in Guizhou is stark. Here, with a
much earlier start to the growing season, pollination takes place in mid-March to early
April, as much as two months earlier than in Massachusetts. Seeds are shed in mid-
September, and germination occurs in mid-March of the following year. The whole
process goes a little more quickly, just twelve versus thirteen months, but more im-
portant is the earlier completion of embryo development, which allows germination a
full three months earlier than in Massachusetts. In Guizhou, the seedlings have about
eight months, rather than five, to put on good growth and become securely established
before facing their first winter. And winter, in any case, is relatively mild compared to
that in the northeastern United States. All these considerations show that while tem-
perature is important in determining where ginkgo will grow, its effects are complex,
and made more so by interactions with other factors, particularly soil conditions and
the availability of water.9
One factor that does not seem to matter for the growth of ginkgo, however, is the
annual distribution of light. High-latitude areas experience a strange light regime, with
several months of almost complete darkness in the winter and several months of near
complete light in the summer. Unless something drastic has happened to the angle at
which the Earth’s axis of rotation is currently tilted, which seems unlikely, this light
regime at high latitudes would have been the same 55 million years ago as it is now.
Fossils from the high Arctic—such as the north slope of Alaska, at nearly 70 degrees
north, or Ellesmere Island or Spitsbergen at nearly 80 degrees north latitude—show
that ginkgo and other trees were indifferent to spending part of the year in nearly
round-the-clock sunlight and a corresponding amount in round-the-clock darkness
during the winter. Peter Del Tredici’s experiments also imply that the Eocene winters
in these high-latitude areas were also rather mild, and certainly not cold enough to cut
off a ginkgo seedling in its prime.10
While the control on the northward limit of ginkgo today is broadly related to dif-
ferent aspects of temperature, especially shorter growing seasons, frigid absolute tem-
peratures in the winter, and the period for which those harsh conditions need to be
tolerated, exactly how ginkgo is limited at the other end of the climatic spectrum is
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harder to understand. A quick survey in North America indicates that while ginkgo
thrives in many places, it is not a characteristic part of the landscape in New Orleans
or in the year-round warmth of southern Florida. Ginkgo is obviously an appropriate
tree to plant at Dinosaur World in Disney’s Animal Kingdom theme park in central
Florida. It was certainly part of the world of dinosaurs, but along with other trees from
more distinctly temperate climates, it does not look especially happy there. According
to Jeff Courtney, a horticulturalist at Disney’s Animal Kingdom, the trick to keeping
the park’s handful of ginkgos alive is to find a good spot with the right microclimates,
out of the direct sun and in places where there is plenty of water.
In Mexico, ginkgo grows well at the relatively high altitude of Mexico City, where
winters can be chilly and average low temperatures in January are around 43 degrees
Fahrenheit, but ginkgo is not a plant that you will find growing in the lowlands of
Chiapas or Oaxaca. In Brazil, ginkgo grows in the southern provinces of São Paulo,
Minas Gerais, and Rio Grande do Sul, planted in outdoor plazas, and also in the tra-
ditional Japanese gardens that reflect the cultural influence of a million and a half
Japanese-Brazilians. However, it does not thrive in Amazonas, or in the hot, dry tropi-
cal climates of the interior of Bahia. Similarly, in Asia ginkgo nuts are common in
the Chinese cuisines of Hong Kong and Singapore, but they are all imported. Ginkgo
grows happily in the warm temperate southern provinces of Guizhou and Yunnan, but
not in the extreme south of China—for example, at Xishuangbanna, near the border
with Laos and Myanmar. Tough as it is, ginkgo cannot tolerate the climates of the true
tropics.11
As with its poleward limit, the factors that exclude ginkgo from the tropics, defining
its southern limit in the Northern Hemisphere and its northern limit in the South-
ern Hemisphere, are likely to be complex. Absolute temperatures and the availability
of water undoubtedly play roles, but also important is a situation well known to the
world’s winemakers. We tend to associate vineyards with hot summer days when we
would be more comfortable taking our glass of wine in the shade rather than out in
the full sun, but we should not forget the winter. Much as grapes flourish in hot sum-
mers, they do not grow well in places with year-round warmth. They need a distinct
cold period as part of their normal annual cycle of growth. The same is true of many
fruit trees, which also need a cold winter, a period of what gardeners call vernalization,
to flower and fruit the following year. In many cases, deciduous trees from temperate
A ginkgo flourishing in Weimar, Germany, one of the historic cultural centers
of Europe and once home to Goethe. Ginkgo easily withstands the cold of
continental Europe and seems to thrive in its warm summers.
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climates do not do well when planted in botanical gardens in the tropics. They leaf out
in a seemingly uncontrolled way and eventually die. In most cases, plants from the
temperate zone don’t survive long in frost-free climates.12
The reasons behind this need for a cold period are not well understood, but they
seem to be connected to the internal biological mechanisms that result in rapid and
coordinated bursting of the buds, as well as flowering, in the spring. In many decidu-
ous trees from the temperate zone, unless there is a period of about one to two months
where the mean minimum temperature is about 41 degrees Fahrenheit or below, bud
burst and subsequent leafing out do not seem to proceed normally.13
Water is also crucial, and again its effects may be subtle. In particular, it is also not
just the absolute amount of rainfall that is important but also how rainfall is distrib-
uted across the year and how water is stored in the soil. Water is quickly lost from
thin, sandy soils, and in these circumstances it can quickly become scarce, which can
in turn affect the growth of trees. At Kew, for example, where the soil is generally well
drained, trees often responded to long dry summers by shedding their leaves early, or
even shedding large branches. In other circumstances—for example, where there are
deep loamy soils—soil water is retained for longer periods, which helps to smooth out
short-lived deficits of rainfall. Ginkgo is sensitive to extreme waterlogging; its roots
cannot tolerate permanent drowning, but water is indispensable for it to really flourish.
Taken together, these aspects of temperature, seasonality, and water availability
seem to be crucial factors that limit the distribution of ginkgo today, both toward the
tropics and toward the poles. Similar factors potentially influenced the changing distri-
bution of ginkgo in the past. However, while the ginkgos planted all over the world tell
us something about where the tree manages to grow, this is not the same as saying that
ginkgo would be able to survive in the wild under those conditions. That is altogether a
taller order, requiring not just an ability to tough out the climate and set seed but also
an ability to be successful in an ecosystem, alongside a varied mix of plants and ani-
mals, as well as microbes, pests, and diseases. The ability of ginkgo to survive in culti-
vation in many different places is only a partial test of its ability to survive in the wild.14
 20
Retreat

It isn’t so much that hard times are coming;

the change is mostly soft times going.
—Groucho Marx

Between about thirty-five million and sixty-five million years ago ginkgo was wide-
spread across the Northern Hemisphere, but that period of great prosperity eventually
came to an end as the climate began to cool. In the Southern Hemisphere, the ginkgo-
like plants that had persisted from the Cretaceous were still part of southern land-
scapes, but they too soon disappeared. The last evidence of ginkgolike plants in the
Southern Hemisphere is in Tasmania about forty million to sixty-five million years ago.
After that, even though there are many younger fossil floras from Australia and South
America, ginkgolike plants seem to have been lost from the Southern Hemisphere until
they were reintroduced, tens of millions of years later, by people.1
Around thirty-five million years ago the global climate not only became markedly
cooler but in many places also became drier. In North America the continuing uplift of
the Sierra Nevada, the Cascades, and the Rocky Mountains sucked the moisture from
winds that had passed over the Pacific and intensified the rain shadow over much of
the continent. This created opportunities for expansion of a new kind of landscape,
the prairie. In Asia too, as the climate changed, forest was replaced with steppe. In

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Fossil localities with ginkgo from about 40 million to 100 million years ago compared with localities
with ginkgo from about 2 million to 40 million years ago. In earlier times ginkgo was present at very
high latitudes in the Northern Hemisphere, but later it was restricted to middle-latitude regions
(maps show the present-day distributions of continents).

these new and more open habitats, new groups of plants, such as grasses, sunflowers
and their relatives, and herbaceous species of many kinds flourished. There was an ex-
plosion in the evolution of flowering plants, which created much of the plant diversity
that we enjoy today. At the same time, forest habitats suitable for the growth of ginkgo
became steadily more restricted.
In Europe, ginkgo was certainly present around sixty million years ago, for example
at Mull in Scotland, on the eastern margin of the opening Atlantic Ocean, but ginkgo
leaves are absent from the many Eocene floras known from Britain, France, Germany,
and surrounding countries. This was almost certainly because the climates were too
warm. More puzzling is that leaves of ginkgo are also absent from some of the classic
Oligocene floras from Europe, as well as from most of the rich and intensively studied
Miocene and Pliocene fossil floras associated with the massive deposits of brown coal
from the Rhine Basin in western Germany.2
There is, however, a truly remarkable occurrence of ginkgo in the Selárdalur flora
from Iceland. Following up on earlier paleobotanical studies, extensive new collect-
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ing and comprehensive studies by Thomas Denk and his colleagues have shown that
around fifteen million years before the present, ginkgo was growing in a forest of an-
cient oak, swamp cypress, coast redwood, magnolia, and grapevines on the volcanic
landscapes of Iceland’s west fjords, places that were perhaps not much different from
those that ginkgo inhabited on Mull almost forty million years earlier.
A hotspot for the occurrence of ginkgo in Europe in the Miocene and Pliocene is in
fossil floras from eastern and southeastern Europe. In this area, ginkgo appears to have
been absent in the Paleocene and Eocene, but its characteristic leaves occur widely in
Middle Miocene, Late Miocene, and Pliocene fossil floras, ranging in age from about
five million to sixteen million years ago across a broad area from Munich in the west
to Ukraine and Russia in the east. In the extreme southeast, ginkgo is known even
from northwestern Greece. In this area the leaves are most common in fossil floras
that reflect the vegetation growing along the banks of ancient rivers. The characteristic
ecology of ginkgo recognized by Dana Royer and his colleagues in the Late Cretaceous
to Early Miocene of western North America also applied in Europe and seems to have
persisted.3
The later appearance of ginkgo in fossil floras from eastern and southeastern
Europe, combined with its presence in fossil floras from the Paleocene and Eocene in
places such as Scotland and Spitsbergen but absence over much of northwest Europe,
suggests that ginkgo migrated into this area as it was forced south by changing climatic
conditions, perhaps a combination of cooling and drying. Obviously—important in
light of what happened later—this implies that its dispersal system was still working.
In North America the pattern is similar to that in Europe. Ginkgo is absent from
some of the most thoroughly collected and best studied Oligocene floras, such as the
Bridge Creek flora from the John Day Basin, and it is very rare in the Oligocene fos-
sil flora from Florissant, Colorado. However, it is present in a smaller number of fossil
floras, many of which are mainly, but not exclusively, western. For example, unmistak-
able fossil ginkgo leaves occur in the Oligocene Lyons fossil flora in western Washing-
ton, which is about the same age as that from John Day Basin. Ginkgo is also present in
other Oligocene fossil floras from the Pacific Northwest, including from the Willamette
flora near Eugene, Oregon. These occurrences might suggest that ginkgo was most
common in places not far from the Pacific Coast. As it does today, that part of western
North America may have had higher rainfall than areas inland.4
Similar complex patterns persist into the Miocene and Pliocene. Ginkgo does not
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occur in the Clarkia flora of northern Idaho, which is one of the most remarkable fossil
floras anywhere in the world for the quality of its preservation. Rich collections from
the Clarkia assemblage have been assembled over many years by the late Jack Smiley
and Bill Rember of Idaho State University, but the unmistakable leaves of ginkgo have
never been seen, even though other characteristic Chinese plants like China fir are
well represented. Today Clarkia is about 370 miles from the Pacific Coast, and the area
experiences only about half the annual rainfall of places like Portland and Eugene.
Perhaps inland the climates were already becoming too dry or too cold for ginkgo to
flourish.5
A classic occurrence of ginkgo from the Miocene is at Ginkgo Petrified Forest State
Park, not far from Vantage, Washington. Here it is known not from fossil leaves but
from its unmistakable wood. The stumps of approximately 15.5 million–year–old
ginkgos occur among those of other forest trees growing on the slopes of an ancient,
active, Cascades volcano, petrified by silica from the ash and preserved where they
grew. Other species that were growing nearby include plants like bald cypress, sweet
gum, oaks, and sycamores. This is one of the last known occurrences of ginkgo in
North America until its eventual return, with the help of people, millions of years later.6
 21
Extinction

Many rivers to cross

But I can’t seem to find my way over.
—Jimmy Cliff, “Many Rivers to Cross”

Given its long fossil history, the presence of ancient ginkgo across much of the
Northern Hemisphere for most of the past 65 million years is not so surprising. Ginkgo
and its extinct relatives were seemingly nearly everywhere on the planet for eons, and
despite their clear decline about 100 million years ago, ginkgo managed to persist in
many places. However, looking back from today, the fact that ginkgo was growing wild
in Bulgaria and Greece just 5 million years ago nonetheless seems strange. It reminds
us that not so long ago the world was a very different place. In the grand sweep of geo-
logic time the distribution of animals and plants on our planet has changed rather
quickly; where they live and grow now bears a strong imprint of history.1
Fossil floras from the Late Miocene and Pliocene provide irrefutable evidence that
in addition to ginkgo, there were many other plants in western North America and
Europe between about five million and fifteen million years ago that no longer grow
there. In terms of the trees the vegetation in these areas was much richer then than
now. For example, fossils from the fill of an ancient sinkhole at Willershausen near

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extinction

Göttingen, Germany, show a mix of broadleaved and coniferous forests. On richer

soils broadleaved forest included species of maple, birch, hickory, beech, ash, oak, and
elm among about thirty-four tree species. Conifer forest included many trees that no
longer occur today in Europe but can be found growing in the warm temperate forests
of eastern Asia: the umbrella pine, for example, as well as the Chinese swamp cypress,
the katsura, the dawn redwood, and the hardy rubber tree. Like ginkgo, in Europe, they
all disappeared relatively recently.
In North America, fossil evidence from Clarkia, Idaho, shows exactly the same pat-
tern. Again, the Chinese swamp cypress and the katsura are both present, along with
the dawn redwood and the China fir. All of these plants are today restricted to east-
ern Asia. At both Clarkia and Willershausen there was also the Cathay silver fir, a rare
conifer discovered as a living plant only in 1955. Today it has a scattered and restricted
range in southwestern China. After about five million to fifteen million years ago, these
plants were never seen in Europe and North America again, but somehow they man-
aged to persist in the East.2
It is hard to understand exactly when and how these species were eliminated from
Europe and North America because in most cases the fossil record is not sufficiently
complete to provide a detailed picture of how their distribution gradually changed
from being widespread in the past to being much more restricted today. We can, how-
ever, get some idea of how they may have fared by tracing the fate of a few of their
associates that have especially distinctive pollen grains. Pollen grains are produced and
preserved in the fossil record in vast numbers, and when they are sufficiently diagnos-
tic of a particular tree, and readily recognized in fossil assemblages, they can be used
to get a fine-grained look at how that plant fared as global climates deteriorated.
Particularly instructive is the history of the Caucasian wingnut, a tree in the walnut
family that has especially distinctive pollen grains. These pollen grains disappear and
reappear through successive glacial and interglacial phases in southern Britain. After
each of the first few glacial advances up to about 500 thousand years ago, pollen grains
of the Caucasian wingnut reappear in the intervening warm interglacials. These plants
seem to have been forced south by successive glacial advances, but they evidently mi-
grated back again into Britain, presumably from the south and east, as the climate
warmed. However, these distinctive pollen grains are last seen in Britain during the
Hoxnian interglacial between about 374 thousand and 424 thousand years ago. For
some reason, in the two most recent interglacials, the Eemian, which lasted from about
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114 thousand to 130 thousand years ago, and the present Holocene, which began about
10 thousand years ago, the Caucasian wingnut never made it back.3
It would be wonderful if we could follow the history of ginkgo in a similarly detailed
way, but unfortunately its pollen grains are too easily confused with those of other
plants. However, the example of the Caucasian wingnut does raise a potentially impor-
tant but unanswered question concerning the ecology of ancient ginkgo: having been
displaced from particular places by changing climates, did it have the ability to recolo-
nize? Colder and drier climates may have progressively restricted ginkgo’s geographic
range, but why did it not bounce back? Surely it should have been able to recolonize
those places where it obviously grows so well today.
In most plants, the ability to colonize an area depends on the effectiveness with
which seeds are dispersed. Seed dispersal provides plants with the ability to emulate
an animal and move from one place to another, albeit much more slowly, generation
by generation. The fruits and seeds of many plants show specializations to increase the
effectiveness of dispersal, from the parachute-like fruits of dandelions that are blown
along by the wind to the seeds of blackberries that are gobbled up along with the fleshy
fruits in which they develop and are dispersed in the droppings of birds. A key ques-
tion in the case of ginkgo is whether one of the factors responsible for its decline over
the past few million years was a poor system for dispersing its seeds.
In 1982 the tropical ecologist Dan Janzen and the paleontologist Paul Martin pub-
lished a provocative article with an arresting title: “Neotropical Anachronisms: The
Fruits the Gomphotheres Ate.” Their central idea flowed from the observation that
many of the common plants in Guanacaste National Park in Costa Rica, where Janzen
had worked for many years, appeared to have no natural means of dispersing their
seeds. They noted that this was particularly the case for some of the plants in which the
fruits and seeds were relatively large, such as guanacaste itself and another legume tree,
divi-divi. Today, the fruits and seeds of these trees are eaten by horses and cattle, but
these animals have been introduced by people from elsewhere only relatively recently.
There are no indigenous animals that appear capable of dispersing them. Janzen and
Martin argued that this mismatch arose because these plants had been dispersed in the
past by animals that are now extinct. The plants had survived, but the animals capable
of dispersing their seeds had not.
Janzen and Martin suggested that such plants used to be dispersed by the large
mammals that once inhabited South and Central America but disappeared relatively
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extinction

suddenly, perhaps as a result of hunting by humans, climate change, or both factors

acting together, about ten thousand years ago. These now-extinct animals would have
included the gomphotheres, massive extinct relatives of modern elephants, that were
obviously plant eaters and flourished in Central America for most of the past five mil-
lion years. Living alongside them were other fruit eaters like ground sloths, glypto-
donts, extinct horses, extinct bears, giant armadillos, flat-headed peccaries, and others.
Janzen and Martin’s point was that the gomphotheres, along with other extinct large
mammals, probably played an important role in the ecosystems of Central America
over the past few hundred thousand years and that their relatively recent extinction
has left us trying to understand an ecosystem that is missing some of its most impor-
tant parts.4
What was most important about Janzen and Martin’s idea was its focus on the im-
portance of history for interpreting the world around us. The survival of the plants,
after the extinction of the gomphotheres and other animals that may have dispersed
them, was an accident of history. In effect the evolutionary histories of the plants and
their associated animals were now out of phase. With some slight rhetorical license,
Janzen and Martin called those plants that had lost their dispersal agents the “living
dead.” The implication was that without the dispersers with which they had evolved,
their days were numbered.5
Janzen and Martin’s ideas proved hugely influential, and in 1984 my paleobotanical
colleague Bruce Tiffney of the University of California, Santa Barbara, suggested that
something similar might have happened in the history of ginkgo. Bruce argued that
ginkgo, like Janzen and Martin’s tropical trees, was also one of the “living dead,” a plant
that had lost its dispersers. He speculated that the strange and strong-smelling ginkgo
seed might have been a specialization for attracting dinosaurs, or perhaps early kinds
of mammals that are now extinct.
Of course, an idea like this is hard to prove, but it does begin to hint at another rea-
son, other than local extinction due to climate, as to why living ginkgo very nearly went
extinct. The apparent migration around fifteen million to twenty-five million years
ago of ginkgo into eastern and southeastern Europe, areas where it was not previously
present, seems to suggest that dispersal was still possible long after the demise of dino-
saurs and ancient extinct mammals. However, Bruce’s point was nevertheless a good
one. A lack of effectiveness in the dispersal of ginkgo seeds may have played a part in
its progressive restriction, and the fact that this may reflect more recent extinctions,
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rather than ancient extinctions at the time of the dinosaurs, is in some ways beside the
point.
Unfortunately, even though its smelly seeds are one of its most well-known and dis-
tinctive features, we know very little about how seed dispersal works in living ginkgo.
However, germination does improve after the fleshy seed coat has been removed—for
example, by passing through the gut of an animal. In one of the potentially wild ginkgo
populations in China it is also documented that the seeds are eaten by a wild cat, and in
Japan they are eaten by badgers. Dogs are sometimes attracted to them too. A friend re-
calls his dog feasting on ginkgo seeds one autumn on the University of Minnesota cam-
pus. It would be helpful to have more information on the kinds of animals attracted to
ginkgo seeds today, but even if various mammals are known to collect and eat ginkgo
seeds, this is not quite the same as knowing that ginkgo has a reliable seed disperser.6
If Bruce is broadly correct, and sometime toward the end of the Mesozoic, or more
likely during the Cenozoic, ginkgo lost the animals on which it depended for disper-
sal, then the effects of climatic restriction would have been greatly amplified. It would
have meant that ginkgo, unlike the Caucasian wingnut, for example, was not able to
easily recolonize areas from which it had been displaced. It would have continually lost
ground, and its populations would have become smaller, moving it ever closer to what
conservationists sometimes call the extinction vortex. Colder or perhaps drier climates
would have eaten away at ginkgo’s once widespread geographic range, and limited
powers of dispersal would have reduced ginkgo’s ability to recolonize. The effect would
have worked like a ratchet; once ginkgo lost ground it was unable to take it back. In
North America and Europe the impact over the past few million years may have been
especially pronounced if, as seems likely from the fossil evidence, the geographic ex-
tent of ginkgo in those areas had already been reduced by climatic drying and other
vegetational changes. The mountains and valleys of southern and western China may
have provided a greater variety of potential refuges.
Whatever the reason, the pattern of regional extinction could not be clearer. Ginkgo
has a more or less continuous record in Asia beginning with the early fossils described
by Zhou Zhiyan and his colleagues more than 200 million years ago. It continues
through the Jurassic and Cretaceous, to the presence of ginkgo in fossil floras from
the Pliocene of Japan. However, in Europe and North America the pattern is different.
Here the fossil record of ginkgo is also deep but it is abruptly truncated relatively re-
cently.7
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extinction

These insights provide a clear example of the importance of fossils to fully under-
stand how our modern world came to be. The natural world is full of patterns, some of
them completely unexpected, that can be explained only by reference to history, and
as I never tire of saying to my students, if you want to understand the way anything is
today, whether it is a plant, a person, an ecosystem, an organization, or a country, then
you need to understand its history. It is a mantra that is hardly original, but one that is
easily forgotten in our modern preoccupation with the here and now. In biology, these
kinds of historical complications are the reason why we ignore evolution, and the di-
rect historical evidence that comes from paleontology, at our peril.
In particular, the fossil record of ginkgo and similar plants helps make sense of a
somewhat enigmatic observation made by botanists since the time of Linnaeus: that
there are surprising similarities between the plants of eastern North America and east-
ern Asia. Highlighted at the end of the eighteenth century by the Italian botanist Luigi
Castiglioni, and then later by the American Thomas Nuttall, the full extent of these
similarities did not become clear until the work of the great nineteenth-century Ameri-
can botanist Asa Gray.8
Gray and his contemporaries were at a loss to explain how the pattern had come
about. For Darwin, writing to Gray at Harvard in 1856, this was one of the “many utterly
inexplicable problems” of botanical geography. Darwin was completely puzzled about
why there should be stronger similarities between the flora of eastern North America
and eastern Asia than between the floras of eastern and western North America. The
fossil record shows beyond doubt, just as Gray later inferred, that these seemingly
strange and widely separated occurrences are the result of regional extinction, espe-
cially in Europe and western North America, of plants that were once much more
widespread. In the case of ginkgo regional extinction went even farther; the species
was completely eliminated from Europe, from eastern and western North America,
and also from Japan. Even in China its extinction was very nearly total.9
 22
Endurance

Here on this rugged and woody hillside has grown an apple-tree,

not planted by man, no relic of a former orchard, but a natural growth,
like the pines and the oaks.
—Henry David Thoreau, “Wild Apples”

The oldest and largest ginkgo trees on the planet occur today in eastern Asia, and it
is only in China that we find the combination of really ancient trees of large size grow-
ing alongside seedlings that are regenerating naturally in what appears to be a more or
less wild situation. However, whether truly wild populations of ginkgo trees still sur-
vive in China is an open question. One problem is that China is a vast country; even
today it is not completely explored from a botanical point of view. Many new species
of plants continue to be described from China, and in just the past decade, new and
potentially wild populations of ginkgo have been discovered and have not yet been
fully studied. Another problem is that it can be hard to tell whether a particular popu-
lation is really wild or not.1
The possibility of identifying truly wild populations of ginkgo is an almost irresist-
ible attraction to botanists interested in plant evolution. If we could identify living
populations of wild ginkgo trees, we might be able to learn how their lives interconnect

162
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endurance

Ginkgo seedlings regenerating

naturally along a path on the campus
of EWHA University, Seoul, South Korea.
The seedlings flourish in partly shaded,
partly open areas of the woodland
understory.

with those of other plants and animals that live in the same ecosystem. We could see
whether ginkgo has natural enemies, such as insects that can feed on its leaves, and we
could observe whether there are mammals that collect or eat its seeds. We could also
study the bacteria and fungi that live on, or in, its roots and stems. It would be interest-
ing to know whether some of the plants that occur at Almont and in other fossil floras
still live alongside ginkgo in modern forests.
A key problem in trying to recognize natural populations of ginkgo is the difficulty
of distinguishing truly wild trees from those with a history of cultivation. China has a
long history of human occupation, and most of the potentially wild ginkgo populations
occur in areas where people have lived for millennia. It is hard to know whether a par-
ticular tree grew from a seed that came there naturally or from one that was planted,
and because ginkgo will regenerate readily from seed under the right kind of condi-
tions, the distinction is a fine one to make. Even a wild-sown seed may have the hand
of people somewhere in its background. On the campus of EWHA University in Seoul,
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d e c l i n e a n d s u r v i va l

I have seen many ginkgo seedlings growing on a steep slope in the understory of a small
piece of seminatural woodland. All of them had germinated from seeds that were natu-
rally sown, but from seeds produced by female ginkgo trees planted along a road a bit
farther up the slope. Given that ginkgo has probably been planted as a nut tree in China
for centuries, the difficulties are obvious.
Western botanists first encountered ginkgo in Japan at the end of the seventeenth
century. In the eighteenth century they also became aware of ginkgo and the use of
ginkgo nuts from European traders in China, Japan, and Korea. However, early bota-
nists visiting eastern Asia were mainly confined to the coasts. It was not until the mid-
nineteenth century, as a result of the forced opening of trade with China, Japan, and
Korea by Western mercantilist powers, that European plant explorers were able to
travel more widely in Japan and Korea and to penetrate the interior of China. Large
ginkgo trees were among the spectacular plants that they encountered, many of them
entirely new to Western science. The accounts of the adventures of early Western plant
hunters in China make fascinating reading. As a result of their explorations, and the
preserved and living plant specimens that began to flow back to Europe, it quickly be-
came clear not only that the native flora of China was exceedingly rich, but that there
were also plants of commercial interest to which the European powers wished to have
access.2
The most important of the early botanist-explorers to journey in China was Robert
Fortune, an intrepid Scotsman from Berwickshire. Fortune began his career as a gar-
dener at the Royal Botanic Garden in Edinburgh, before moving south in 1840 to
work in the garden of the Royal Horticultural Society, then based in west London. Just
months after the signing of the treaty of Nanjing, which brought to an end the First
Opium War of the late 1830s and early 1840s, Fortune was dispatched to China by the
Royal Horticultural Society and arrived in Hong Kong in July 1843. It was the first of
four incident-filled visits, and Fortune was also one of the first Western botanists to
travel in Japan.3
Fortune’s adventures are vividly recounted in his several books. His botanical focus
was on the collection of living plants, and his second expedition on behalf of the British
East India Company had a direct economic purpose: the introduction of tea to India
against the wishes of the Chinese government. By mastering the Mandarin dialect,
adopting local dress, shaving his head except for a queue, the brusque Scottish botanist
was able to pass himself off as a native Chinese from a distant province and gain entry
 165
endurance

into otherwise forbidden areas. Over the course of his expeditions in Asia he shipped
more than twenty thousand seeds and seedlings from China and Japan, ultimately
introducing more than a hundred plant species into European horticulture.
On his first journey to China, Fortune writes of ginkgo, using its then accepted
botanical name: “The only tree which I met with of very large size in [Shanghai] is the
Salisburia adiantifolia, commonly called the Maiden-hair tree, from the resemblance
its leaves bear to a fern of that name. This is one of the plants which the Chinese are
fond of dwarfing, and it is, consequently, often seen in that state in their gardens. Its
fruit is sold in the markets in all Chinese towns by the name of ‘Pa-Kwo,’ and is not
unlike dried almonds, only whiter, fuller, and more round. The natives seem very fond
of it, although it is rarely eaten by Europeans.”4
After Fortune, a second wave of plant explorers gained access to China as a result
of changing political circumstances, this time the Second Opium War. The resulting
Convention of Beijing in 1860 greatly benefited both the French and the British. From
Britain, Augustine Henry was a key link between Robert Fortune and collectors who
came later, such as Joseph Rock, Ernest Henry Wilson, George Forrest, and Frank
Kingdon-Ward. Also influential as great plant explorers were three French missionar-
ies who were dispatched to China in the 1860s, of whom the first and most important
was the Basque Lazarist monk Père Jean Pierre Armand David.5
Augustine Henry went to China as an employee of the British Imperial Maritime
Customs Service and collected extensively in central China between 1882 and 1889.
He sent more than fifteen thousand dried specimens and seed samples to Britain.
Most were described by botanists at Kew, such as William Botting Hemsley and Daniel
Oliver. Henry brought to light a wealth of exciting new species from China, many
of which remain important in European horticulture and bear his name, such as the
Henry Honeysuckle and the Henry Lily. He came across ginkgo on several of his expe-
ditions. In his “Notes on Economic Botany of China,” published in 1893, his comments
are brief: “Pai- kuo, Ginkgo biloba L. Seeds eaten.” However, in his seven-volume work
The Trees of Great Britain and Ireland, published in 1906, seven pages of text are de-
voted to ginkgo, along with three photographs that include an ancient ginkgo beside a
temple in central China.6
Henry is also important because of his influence on a young British plant collector
who followed him, Ernest Henry “Chinese” Wilson. Through his roughly two thousand
plant introductions, Wilson had a major impact on the cultivation of Asian plants in
 166
d e c l i n e a n d s u r v i va l

A large old ginkgo photographed by

Ernest Henry Wilson at a temple in
Nara Prefecture, Japan, in April 1914.
In this ancient ginkgo the trunk has
split in two and another large tree
is growing in the cleft.

the gardens of the West and became one of the best-known plant collectors of all time.
In the process he also made substantial contributions to knowledge about the plants
of China.7
Wilson came across ginkgo many times during his Chinese explorations. In his
book A Naturalist in Western China with Vasculum, Camera, and Gun, Wilson gives his
impression of what he had seen of ginkgo during his Chinese travels: “This strikingly
beautiful tree is associated with temples, shrines, courtyards of palaces and mansions
of the wealthy throughout the length and breadth of China, and also in parts of Japan.
But nowhere is it truly wild.”8
Nevertheless, with further botanical exploration it has become clear that there are
many large ginkgo trees in China that are not associated with temples, and Chinese
botanists have raised the possibility that wild ginkgo continues to grow in several
places in the rich forests along the Yangtze River and especially in Zhejiang Province
in the area around Tianmu Mountain.
 167
endurance

For someone wishing to get a taste of the true richness of the forests of eastern China
within easy striking distance of Shanghai and Huangzhou, there is no better place to
visit than Tianmu Mountain, which reaches almost five thousand feet and is one of
the highest mountains in Zhejiang Province. Its varied terrain produces equally varied
vegetation ranging from subtropical evergreen forests to temperate deciduous forests.
The area with ginkgo and the especially rich flora is located on the south-facing slope
of the west peak. The vegetation is luxuriant and includes subtropical evergreens more
typical of forest farther south, mixed in with more typical temperate deciduous broad-
leaves and a variety of more temperate conifers. The flora is exceptionally rich, with
about fifteen hundred species of vascular plants, a number of species comparable to the
entire native flora of the United Kingdom in an area of little more than sixteen square
miles.9
The ginkgo population at Tianmu Mountain has provided important information
about ginkgo and how it behaves under natural or near natural conditions. For ex-
ample, fieldwork by Peter Del Tredici located 167 ginkgo trees, more than a third of
which had two or more substantial trunks. It seems likely that the extra trunks were
produced by the activation of the basal chi-chi. As in other plants that produce basal
lignotubers that are capable of sending out shoots and roots following a disturbance
to the tree, they may help to stabilize the plant growing on steep slopes.
There is, however, a nagging worry that ginkgo, like several other trees growing on
Tianmu Mountain, may have been introduced from elsewhere, perhaps a rather long
time ago. It may never be possible to fully understand the origin of the ginkgo popula-
tion on Tianmu Mountain, but over the past few decades the discovery of other poten-
tially wild populations in China, together with the introduction of new techniques
from molecular biology, has provided further opportunities to probe whether ginkgo
survives in the wild in some parts of that vast country.
 23
Relic

By their fruits ye shall know them.

—Matthew 7:15

With recent rapid developments in plant molecular biology, new tools have become
available to help understand the history of ginkgo in eastern Asia over the past few
hundreds of thousands of years. All ginkgo trees look more or less the same from the
outside, but we can now look into their DNA to see just how similar or different they
really are. We can sample different ginkgo trees growing in different places to see how
they might be related to one another and also how much variability there is in their ge-
netic makeup. We can use evidence from DNA to assess whether the individual trees in
a population are genetically very different from each other or whether they are all more
or less the same. Armed with that information we can also develop ideas about which
populations could potentially have given rise to which, based on the assumption that
those populations with the most genetic diversity might be living plants not too differ-
ent from those from which genetically less diverse populations in other places might
have been derived.
These kinds of arguments have been widely used to infer the origins of different
crop plants. The different parts of the world where particular crops were originally do-
mesticated from wild plants, as demonstrated by archaeological, botanical, and other

168
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relic

The famous Five Generations Ginkgo growing in the Tianmu Mountain Reserve,
Zhejiang Province, China.

evidence, often show relatively high levels of genetic diversity, whereas other parts of
the world into which these plants have been introduced often show relatively lower
levels. This is exactly what would be expected because it is unlikely that the full genetic
diversity would be represented in a subgroup of plants that had been moved to other
places. For example, at a global scale, potatoes and tomatoes have their greatest ge-
netic variety in the northern Andes, where wild species that still grow in those regions
were most likely domesticated in ancient times. Using arguments of this kind, although
without the benefit of modern genetics, the great Russian plant scientist Nikolai Vavi-
lov identified the likely wild relatives and probable source areas of many of our most
important crop plants.1
In trying to track down wild populations of ginkgo, there has been particular inter-
est in studying the genetic diversity among the Tianmu Mountain ginkgo trees and
 170
d e c l i n e a n d s u r v i va l

comparing that with the genetic diversity seen in other potentially wild populations
elsewhere in China. Early studies tended to suggest that the Tianmu Mountain popu-
lation had relatively low genetic variability, but how this compared with other popula-
tions of ginkgo elsewhere in China had not been tested until a series of studies over the
past decade or so by Chinese scientists at East China Normal University in Shanghai
and Zhejiang University in Huangzhou.2
In one initial study the Chinese team sampled nine populations of possible ancient
ginkgo trees from different parts of China: Guizhou in southwestern China, Henan and
Hubei in central China, and Jiangxi and Zhejiang in eastern China, which included the
Tianmu Mountain population. For each population they extracted DNA from between
ten and thirty trees, and for each tree they used two approaches to assess the genetic
variation.3
Their first approach used the so-called RAPD technique: an acronym for the much
less user-friendly technical designation, random amplified polymorphic DNA. This
technique is relatively crude compared with the level of sophistication of modern mo-
lecular biology, which could potentially “decode” the entire DNA sequence of a ginkgo
tree in a few weeks, but RAPDs are nevertheless relatively quick and effective, and were
once widely used. For good reason, in the modern slang of molecular biology this ap-
proach is usually known as “RAPIDS.”
After extracting the DNA from individual ginkgo trees the Chinese group created
large quantities of short sections of DNA, each based on a “starter” section, a so-called
primer, with a precise sequence of the genetic code. The basic idea was to see how
many fragments each primer would produce from the DNA of each tree and how big
those fragments would be. Then the researchers characterized the resulting fragments
in terms of their size by placing each sample from each tree on a gelatin-like slab with
an electric current passing through it. Drawn along by the current, each sample moves
along a “lane” in gel, much like a sprinter in a hundred-meter race; smaller fragments
of DNA move more quickly and travel farther, while larger fragments of DNA move
more slowly and do not travel as far. In total, from all 164 trees sampled, the research-
ers recovered forty-seven DNA fragments of different sizes, and by comparing how
many of the fragments were represented in each ginkgo tree and each group of trees,
they were able to come to a rough estimate of genetic diversity for the different ginkgo
populations growing in different places in China.4
The results showed that from a regional standpoint, and in terms of the number of
 171
relic

different DNA fragments recognized, the ginkgo populations from southwest China
were the most diverse. Three of the four populations from Guizhou showed especially
high levels of diversity. The ginkgo trees of Tianmu Mountain from Zhejiang were
more or less intermediate in diversity along with those from Henan and Hubei. Com-
parison of the RAPD results from each population with every other population high-
lighted the ginkgo population from Jinfo Mountain in Chongqing Municipality, as the
most distinct. The population from Tianmu Mountain was also relatively distinct but,
on the whole, more similar to the other populations studied.
The Chinese team then followed up with a second approach to assessing the genetic
diversity of the different populations of ginkgo trees. In this case, they made many
duplicates of specific pieces of the DNA from the leaves of 158 trees collected from four
places in China. Having obtained a sufficient quantity of each piece of DNA from all of
the trees, they used three enzymes, each of which had the potential to cut the DNA in
a specific place. When they analyzed the resulting pieces, they found that among the
158 trees there were eight patterns. The population from Jinfo Mountain was again the
most diverse: of the eight patterns the Jinfo population showed six.5
So what can we conclude from these initial insights into the DNA of Chinese ginkgo
populations? First, these kinds of studies provide only a crude estimate of the true ge-
netic situation. In both cases the technique looks at only a tiny portion of the overall
genetic composition of these trees, and whether that portion is really representative is
an open question. There is also a slight worry, inherent in the use of both techniques,
that the fragments produced, while treated the same for purposes of comparison, may
actually be slightly different.6
Nevertheless, more recent studies by Chinese scientists at Zhejiang University and
their collaborators, using more sophisticated techniques, support these earlier conclu-
sions. Again they highlight that the populations from southwestern China are geneti-
cally more diverse than those in eastern China, but on the whole they tend to support
the idea that ginkgo may have survived the Pleistocene glaciations in two refuges: one
on Tianmu Mountain not too far from the Chinese coast and the other farther inland
in the protected valleys along the southern margin of the Sichuan Basin.7
If we take the results at face value, and also recognize that they provide only a guide,
rather than a definitive picture of the history of different populations, it is certainly true
that a grove of ginkgo trees with low or only moderate genetic variation, like those that
occur elsewhere in China, would be unlikely to be the source of a genetically more di-
 172
d e c l i n e a n d s u r v i va l

Location of two potentially native populations of ginkgo in China at Jinfo Mountain,

Chongqing Municipality, and Tianmu Mountain, Zhejiang Province.

verse population, like that at Jinfo Mountain or even at Tianmu Mountain. However, it
does not necessarily follow that the reverse is true. Less diverse populations may have
been derived from the more diverse ones, but especially in a case like ginkgo it is pru-
dent to be cautious. There has obviously been widespread extinction, including per-
haps even in historical times, through forest clearance. Both the genetically diverse and
genetically depauperate populations could be different kinds of relics from formerly
widespread and still more diverse populations that are now extinct.
Against this background, the results from the DNA analyses do not completely settle
one way or the other whether the ginkgo trees on Tianmu Mountain are relics of a natu-
 173
relic

ral population, or whether they were introduced by Buddhist monks, as some have
suggested. They do, however, point the spotlight at Jinfo Mountain and perhaps other
places in southwest China as of exceptional interest and worthy of much more detailed
study.8
Jinfo Mountain in Chongqing Municipality lies near the border of Guizhou and
Sichuan Provinces. It is only a little farther south than Tianmu Mountain, but rather
than being relatively close to the coast it is deep within the interior of China. And in
addition to massive ginkgo trees there are other trees in the same region, such as the
Cathay silver fir and Chinese fir, that have a long fossil history and that were once much
more widely distributed. In this part of China there are many small, scattered popu-
lations of ginkgo that seem to be regenerating naturally from seed. Of all the ginkgo
trees in China these are the ones that seem most likely to be the tiny remnants from a
time when the geographic range of ginkgo was once much more continuous.9
This page intentionally left blank
 part v

History
OVERLEAF A plate decorated with ginkgo shoots and leaves, produced between
about 1700 and 1730 in the Okawachi Kiln in Hizen, Kyushu, Japan.
 24
Antiquity

How cunningly nature hides every wrinkle of her inconceivable

antiquity under roses and violets and morning dew!
—Ralph Waldo Emerson, “Works”

Ginkgo now grows all around the world, but almost everywhere it has been brought
there by people; for most of us ginkgo is a plant of parks, gardens, or city streets, all
human-created habitats. These trees are also of modest size; nowhere outside eastern
Asia are there ginkgo trees of truly massive proportions. Even the Old Lion at Kew, one
of the oldest trees in Europe, has a trunk only a little more than five feet in diameter. In
China, Japan, and Korea, the situation is different; here there are some true giants, and
in a few places in China, there are huge ginkgo trees that seem to be growing wild. At
Jinfo Mountain in Chongqing Municipality on the border of Guizhou and Sichuan, a
tree recorded in the 1950s had a trunk more than twelve feet across. Another recorded
in 1999 was clearly much older than the nearby town, with a trunk more than eleven
and a half feet in diameter. In all, seventy trees at Jinfo Mountain were found to have
trunk diameters of more than forty inches, and eight had trunks more than six and
a half feet across. There are many trees in that region with trunks the size of the Kew
Ginkgo, and they are growing alongside saplings and seedlings, which show that natu-
ral regeneration is occurring.1

177
 178
history

The big trees found on the forested slopes of Jinfo Mountain and Tianmu Mountain
are unusual; most of the really large ginkgo trees in China, and also in Japan and Korea,
are not forest dwellers. Some are associated with temples or shrines, but they are often
found alone in the countryside, growing by themselves. A review of large ginkgo trees
in China turned up 138 with trunks more than about six and a half feet across; many
are singletons, and it is obvious that they are not growing in natural forests.2
Among the most impressive of these trees is the Grand Ginkgo King, which grows
near the small hamlet of Li Jiawan in Guizhou, southern China, towering over the agri-
cultural bottomland of a small valley. Perhaps it started life as part of an ancient ginkgo
orchard, but equally probable, given the seemingly wild populations of ginkgo not too
far away at Jinfo Mountain, it could be the last remnant of an ancient ginkgo grove
that somehow survived when the forest was cleared for farming. Like the ginkgos that
survived at Hiroshima, perhaps it was resistant to fire, the key tool that early farmers
used to clear the land.3
The Li Jiawan Grand Ginkgo King is a big tree, nearly a hundred feet tall with
a trunk about nineteen feet across at ground level; more massive than that of any
other ginkgo so far recorded. However, even from a distance it is obvious that this tree
has had a complicated history. There are four main upward thrusting leaders, each of
which is more than sixty feet tall. Studies by Chinese colleagues suggest that it may
have sprouted from the base at least four times in the course of its long life. Its ring of
partly separate trunks surrounds a hollow center so large that it once provided shelter
for a local farmer and his cattle. The hollow stem rules out a full count of the annual
rings and makes the age of the Li Jiawan ginkgo hard to assess, but by one estimate it
might be as old as forty-five hundred years. Such great antiquity seems improbable,
but from its size alone the Grand Ginkgo King must surely be among the most ancient
of all living ginkgo trees.4
Great ages have also been suggested for other ginkgo trees in China. One in Dong-
kou Xian County, Hunan Province, is thought to be more than thirty-five hundred
years old. Another at Dinlinsi Temple in Juxian County, Shandong Province, is re-
ported to be more than three thousand years old. In Zhouzhi County, Shaanxi Prov-
ince, and also in Tancheng County, Shandong Province, are trees that are thought to
be more than two thousand years old, while another very large tree in Fuquan City,
Guizhou Province, is likely to be of similar age. Altogether there are about a hundred
ginkgo trees in China that are thought to be a thousand years old or more.5
 179
antiquity

The estimate of forty-five hundred years for the Li Jiawan ginkgo, if anywhere near
correct, would make it among the oldest of all living trees, but this seems unlikely.
Individual bristlecone pines in the White Mountains of California are dated reliably as
more than forty-seven hundred years old, and direct evidence from annual rings shows
that several other species, especially conifers, live for more than two thousand years.
Some coast redwoods are into their third millennium, while a few giant sequoias are
into their fourth. But these and other methuselahs are exceptions; most trees have life
spans measured in a few centuries rather than in multiple millennia.6
It is often hard to be confident about the great ages attributed to giant trees. The
massive baobabs of Africa, some of which have trunks that exceed twenty or thirty feet
in diameter, provide a classic case. The great German explorer Alexander von Hum-
boldt never saw a baobab, but nevertheless declared it to be one of the “oldest inhabi-
tants of our globe.” The French botanist Michel Adanson, from whom the baobab takes
its scientific name, had more direct experience. He came across two trees on the Îles de
la Madeleine off the coast of Senegal in 1749 on which earlier sailors had carved their
names and dates. Estimating the amount of wood added since that time, and taking
into account the diameter of the trunk, he deduced the age of the two trees to be more
than 5,000 years. Adanson concluded that they must have been alive before the Great
Flood. It was a clever and perhaps slightly mischievous extrapolation that allowed him
to goad the religious establishment, but recent research indicates that most massive
baobabs are much younger, between 500 and 800 years old. So far, the oldest radiocar-
bon date from a baobab is 1,275 years, from a tree in Namibia that had a trunk about
one hundred feet across.7
Figuring out the age of baobabs is particularly difficult because they do not have
well-defined annual rings, but even for trees with rings that are easily counted, accurate
age estimates are often not easy to come by. One problem, as in the case of the Li Jia-
wan Grand Ginkgo King, is that the oldest parts of many truly ancient trees have been
lost through decay. Another difficulty is that even though foresters have developed a
special tool for taking a small core from the trunk of living trees, this becomes harder
to do as the trunk gets bigger. It also is not always easy to find the center of a living tree
with a complicated trunk.
A further problem is that even if a reliable age can be obtained for one tree of a par-
ticular species, the relationship between girth and age is not sufficiently consistent to
extrapolate reliably to other specimens. Rates of growth can vary drastically in the life
 180
history

of a single tree—for example, when the sapling is growing in deep shade, compared
with when it has an opportunity to grow quickly into the canopy. Growth rates also
vary depending on where the tree is growing. Cross sections of two stumps of Japanese
cedars on display in the forest museum on the island of Yakushima, Southern Japan,
nicely illustrate the difficulty.
The massive Japanese cedars on Yakushima are among the great botanical wonders
of the world. Jōmon-sugi, the largest of these great trees, was discovered in the mid-
1960s in the dripping wet forests about three thousand feet above sea level. It is only
about eighty feet tall, but its fifty-three-foot girth is truly impressive. Age estimates for
the massive tree range widely, from 2,170 to 7,200 years old. The younger suggestion
may be plausible, but seven millennia seems much too old. However, part of the dif-
ficulty in narrowing the age range further is in knowing how to extrapolate from the
ages of cedars felled in other parts of the island.8
The two stumps of young Japanese cedars in the Yakushima Forest Museum are
both about two and a half feet across, and the age of each is known precisely by count-
ing the annual rings. One, which grew at a higher elevation, is 225 years old; the other,
which grew at a lower elevation, is a mere 64 years old. In the older of the two the an-
nual rings are tightly spaced, and each is perhaps no more than about one twentieth of
an inch thick. In the other, the annual rings are much thicker. Trees with good growing
conditions and a longer, warmer, growing season, with access to plenty of water and
light, can grow fast, and the annual rings will be broad. Trees growing in less ideal con-
ditions will grow more slowly; their annual rings will be narrow and tightly packed.9
A further way to try to try understand the age of massive Asian ginkgos is to look
for historical or cultural clues. But this also has its problems, because the clues most
often available are legends, which are rarely reliable. A good example is the massive
and much-loved ginkgo at the Tsurugaoka Hachiman-gū Shrine in Kamakura, not far
from Tokyo.
When I visited the Tsurugaoka Big Ginkgo on a warm spring Saturday several years
ago, the shrine was bustling with families. The grand old tree had still not leafed out,
and its nakedness revealed how much its crown had been sculpted by generations of
tree surgeons. Nevertheless, it was wonderfully imposing. Standing next to the steep
steps leading to the main building of the shrine, and with a traditional rope made of
rice straw wrapped around its massive trunk, it was resplendent in the early spring
sunshine.
The Tsurugaoka Big Ginkgo at the Tsurugaoka Hachiman-gū Shrine in Kamakura, Japan,
photographed in spring 2006. The great tree fell after a storm in March 2010.
 182
history

The Tsurugaoka Hachiman-gū Shrine was built in Kamakura in 1180 by the first Sho-
gun, Minamoto no Yoritomo, and in 1219 was the site of the notorious assassination of
the third Shogun, Minamoto no Sanetomo, on the same stone stairs that visitors still
ascend on their way to the main shrine. According to legend, the assassin hid behind
the ginkgo tree as he waited to make his attack; by that measure, the Tsurugaoka Big
Ginkgo would today be nearly a thousand years old.10
Shihomi and Terumitsu Hori, the two foremost scholars of the cultural history of
ginkgo in Japan, carefully reviewed the evidence to determine whether the legend and
such a great age for the tree might be true. The Azuma Kagami, an official record of
events that occurred under the Kamakura bakufu between 1180 and 1266, reports the
assassination in detail, with the date, time, weather, and even the words of the assassin
as he attacked. The Eukan-sho, written in 1220, just a year after the assassination, by the
priest and poet Jien, gives still more detail, even describing the assassin’s clothes. How-
ever, in neither of these near-contemporary accounts is there any mention of the tree.
Suspiciously, the ginkgo appears only in accounts written much later; the first is in the
Kamakura Monogatari, written around 1659, more than four hundred years after the
fact. It seems very likely that the ginkgo is a later addition to the legend. Rather than
a thousand years, a more modest age of perhaps five or six hundred years seems more
likely for the Tsurugaoka Big Ginkgo.11
These kinds of problems apply to the great ages ascribed to ancient trees of all kinds,
including ancient ginkgo trees in China. For example, one legend associated with the Li
Jiawan Grand Ginkgo King links its origin to a scholar named Bai who lived in the Tang
Dynasty (618–907), while another makes a connection to a somewhat similar legend
from the Ming Dynasty (1368–1644). Similarly, it is said that the massive ginkgo trees
at the ruins of the ancient Huiji Temple, not far from Nanjing, were planted by Zhao-
ming, a famous prince of the Liang Kingdom from the early sixth century. This would
make them about fifteen hundred years old, but again there is no written evidence, and
an age of about half that seems much more likely.12
It is hard for anyone, still less a Westerner, to get to the bottom of these kinds of
legends, but Joseph Needham’s classic Science and Civilization in China is nevertheless
a useful guide. Needham devoted his life to understanding the early history of science
in China, and in 1986 he reviewed the history of Chinese botany. A little later, Nicholas
Menzies, one of Needham’s colleagues, provided a similar account of Chinese forestry.
Together, these two works review some of the most important early literature on the
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antiquity

history of ginkgo in China. Especially illuminating is the example cited by Menzies

of an inscription beside an old tree at the Fu-Yen Ssu monastery “on the slopes of the
sacred peak of Hēng Shan in Hu-Nan.” According to the inscription it was planted by
“the venerable abbot Hui Ssu, in the second year of Kuang-Ta of the Chhēn dynasty.”
This would mean that the tree was planted in 568. However, Menzies cites work by for-
esters in the Nan-Yüeh Forest who had counted its annual rings and found it to have
an age of six hundred years. This was a long-lived and no doubt majestic tree, but it
was less than half the age that generally had been ascribed to it.13
Examples like these suggest that the great ages assigned to many ancient ginkgo
trees are not reliable. Legends tend to be embellished over time, and it is not easy to
obtain reliable estimates of the age of a living tree. For the largest and most spectacu-
lar specimens this has rarely been done. Given these complications, it seems unlikely
that even the most ancient ginkgo trees approach the three thousand years of a giant
sequoia or come close to the four and a half millennia of the bristlecone pine. A more
reasonable view is that the age of some might approach fifteen hundred years, but that
most have ages that are best measured in centuries rather than millennia. This is also
in line with data from written sources, which points toward younger, rather than older,
ages for the world’s largest and most impressive ginkgo trees.
 25
Reprieve

A burning stick, though turned to the ground, has its flame drawn upward.
—Saskya Pandit, “Elegant Sayings”

It is not known exactly when, or why, ginkgo first became associated with people,
but in Science and Civilization in China, Nicholas Menzies finds the two earliest refer-
ences to ginkgo unconvincing. The dating to the sixth century of the old tree at Fu-Yen
Ssu Monastery is known to be unreliable. The other, the mention of a fruit referred to
as phing chung in the poem “Rhapsody on the Capital of Wu” by Tso Ssu, which dates
from the Jin Dynasty in the third century, has nothing to link it to ginkgo except men-
tion of its silvery color. Similarly, while ginkgo is sometimes identified in carvings and
paintings from the fourth to the eighth centuries, these show few botanical details and
are also probably cases of mistaken identity. As far as I know, there is also no firm evi-
dence to support the sometimes-cited use of ginkgo nuts during the Han Dynasty in
the third century.1
The earliest reliable written reference to ginkgo is from 980 in the Ko Wu Tshu Than,
or Simple Discourses on the Investigation of Things, written by Tsan-Ning, a “learned
monk.” A little later, during the Song Dynasty in the eleventh century, the most widely
cited and earliest undisputed historical reference to ginkgo is a famous exchange of
poems between the early Chinese historian Ouyang Hsiu and the poet Mei Yao-Chēn.

184
 185
reprieve

Both refer to ginkgo by the name duck’s foot, ya chio, but they also use the name yin
hsing, silver apricot.
Ouyang Hsiu begins the dialogue by presenting his friend with ginkgo nuts from
a tree planted in Kaifeng, one of the Seven Ancient Capitals of China. Mei Yao-Chēn
responds with his thanks and reminisces about ginkgo in his native Xuancheng. The
exchange concludes with a poem from Ouyang Hsiu that sets out a delightful brief his-
tory of ginkgo cultivation.2

Ya chio (duck’s foot) grows in Chiangnan with a name that is not appropriate.
At first it came in silk bags as a tribute, and as yin hsing (silver apricot) it be-
came cherished in the middle provinces. The curiosity and effort of the Noble
Prince [Li] brought roots from afar to bear fruit in the capital. When the trees
first fruited they bore only three or four nuts. These were presented to the
throne in a golden bowl. The nobility and high ministry did not recognize them
and the emperor bestowed a hundred ounces of gold. Now, after a few years,
the trees bear more fruits.

These poems and the other early record from the late tenth century both point to
a first association of ginkgo with people about a thousand years ago. There are many
earlier Chinese works that deal with plants, and with cultivated plants in particular, but
none mentions ginkgo. Ginkgo is also unlike many other trees, which are often first
noted as growing wild and are only later mentioned for their uses. This might suggest
that before it was cultivated ginkgo was a rather rare tree. From the beginning, ginkgo
is treated as a cultivated plant, a tree grown for its nuts. Compared with other plants,
like rice and soy, which have a history of cultivation in China that goes back several
millennia, the cultural roots of ginkgo seem relatively shallow.3
After the exchange of poems between Ouyang Hsiu and Mei Yao-Chēn, ginkgo ap-
pears commonly in Chinese literature, and by the Yuan Dynasty, established in the late
thirteenth century by the ethnic Mongols under Kublai Khan, there are many refer-
ences to ginkgo being grown for its nuts. According to Menzies, in some areas the nuts
became an important commodity. He cites the Nung Sang Chi Yao from 1273 as the
earliest work to give details of how ginkgo should be cultivated, and notes that these
horticultural instructions were repeated verbatim in later manuals such as the Chung
Shu Shu, the Pien Min Thu Tsuan, and the Nung Chēng Chhüan Shu.4
From the beginning, it was also recognized that both male and female trees are
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history

needed to produce nuts, but then, as now, there is no easy way to tell whether a nut
would produce a male or female tree. The Nung Sang Chi Yao offers the following ad-
vice: “The ginkgo has male and female trees. The male [seed] has three ridges, the
female has two. They must be planted together. When they are planted next to a pond,
they face their own reflection and so can bear fruit.” There is no evidence that three-
ribbed seeds, which in any case are rare compared with the two-ribbed forms, grow
into male trees. However, the idea of planting female trees near water may be helpful.
Ginkgo thrives when it has good access to water, and trees stressed by drought are un-
likely to produce a good crop of seeds. Later horticultural treatments also recommend
grafting a male branch onto a female tree, the reverse of what was done by Jacquin in
his experiments. It is an easy and sensible way around a potential twenty-year wait for
seeds if there are no male trees nearby.
Based on the comments of Mei Yao-Chēn reminiscing about ginkgo near his home,
and mentioning that he had collected nuts from the wild, Chinese authors have sug-
gested that the ginkgo is native to the area around Xuancheng in Anhui Province. More
recently, attention has also focused on Jinfo Mountain in Chongqing Municipality, as
well as Tianmu Mountain in Zhejiang. It is still not completely certain whether ginkgo
is native to these areas, but it does seem likely that the cultivation of ginkgo began in
the south and spread to the north. This northward movement of ginkgo is also con-
sistent with the comment in Shihuazonggui by Ruan Ye: “In the capital—now Kaifong
City, Henan Province—there was no ginkgo. Since the coming of Mr. Li Wienhe, the
emperor’s son-in-law, from the south, he introduced it and planted it in his private
house. Then it yielded and was propagated and developed there.”5
The fact that none of the largest ginkgo trees in China are from the three northern
Chinese provinces of Liaoning, Jilin, and Heilongjiang, on the landward side of the
Korean Peninsula, also fits this pattern. The northerly expansion of ginkgo cultivation
to Korea was probably along the coastal trade routes of China and then across the Yel-
low Sea to the southern tip of the peninsula.
Ginkgo would certainly have been cultivated on the Korean Peninsula well before
Japanese troops under Toyotomi Hideyoshi invaded at the end of the sixteenth century,
and in the seventeenth century Hendrick Hamel and his shipwrecked companions are
reputed to have dreamed of their escape under the massive ginkgo at Gangjin-gun in
the far south of the peninsula. A list of large ginkgo trees in Korea includes twenty-
one with ages estimated to range from four hundred to a thousand or more years. The
 187
reprieve

age of the great Yongmunsa Ginkgo is said to be about eleven hundred years, and two
others on the Korean Peninsula are reputed to be even older.6
In North Korea, at the Anbulsa Temple there is a huge female ginkgo, 140 feet tall
and 19 feet in diameter. It was protected on orders of President Kim Il Sung during
the Korean War and in 2003 was visited by his son Kim Jong Il. It is said to be 2,120
years old and to produce more than 650 pounds of seeds each year. In South Korea
the ginkgo in Yeongwol, with a trunk diameter of around 15 feet, is said to be 1,000–
1,200 years old. It once stood in front of the demolished Daejeongsa Temple, but the
60-foot-tall tree still gives shade to the people in the village that has now grown up
around it.7
In Japan there are fossils of ginkgo from before the ice ages and massive living trees
that are evidently of great age, but there is no evidence that ginkgo is native to the Japa-
nese Islands. Like many other useful plants, ginkgo was introduced from mainland
Asia during historic times. As in China, many of the oldest and most venerable ginkgos
in Japan are on the grounds of temples, but large trees are also found in agricultural
landscapes, where, like the Li Jiawan Grand Ginkgo King, they may mark the sites of
former ginkgo orchards rather than former temples that have disappeared completely.
Shihomi and Terumitsu Hori have reviewed the early cultural history of ginkgo in
Japan and identified eight great trees that range from seven hundred to fifteen hun-
dred years in age according to their associated legends. The Jyonichiji Ginkgo, a large
female tree on the grounds of the Jyonichiji Temple in Toyama Prefecture, is consid-
ered the oldest. According to local legend, this was already a big tree when the temple
was founded in 682. The Nigatake Ginkgo, sometimes called the Chichi Ginkgo or Uba
Ginkgo, at the Ubagami Shrine in Miyagi Prefecture, is thought to be almost as old. The
Hōryō Ginkgo in Aomori Prefecture, which is thought to be eight hundred to twelve
hundred years old, is said to have been planted to commemorate the founding of the
now lost Zenshoji Temple in the Heian period.8
It is hard to probe the veracity of these legends from Korea and Japan, but as in
the case of the Tsurugaoka Big Ginkgo, such extreme ages are probably not correct.
In Japan unambiguous written evidence for ginkgo appears even later than in China,
and ginkgo is notably absent from classic early literature, where it might be expected
to have appeared. Traditional Japanese thirty-one-syllable poems called waka have a
history that goes back about thirteen hundred years. The Man’yōshū, the oldest an-
thology of such poems, was compiled at the end of the Nara period, during the late
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history

eighth century, but while there are many references to yellow leaves in these poems,
these could belong to many other plants. Similarly, the Horis conclude that the word
chi-chi mentioned in the poems of Otomo-no-Yakamochi, which date from the middle
of the eighth century, more likely refers to the widely grown Japanese fig.9
There are no unequivocal references to ginkgo in the anthologies of waka collected
by Imperial Command in the tenth and thirteenth centuries, either. Nor does ginkgo
feature in The Tale of Genji by Murasaki Shikibu, one of the oldest novels in Japan, or
in the Makura no Soshi, a collection of ancient essays written by Sei Shonagon in the
late tenth to early eleventh centuries. Considering how prominently nature is featured
in this literature, and how often ginkgo features in later works, its absence may be sig-
nificant.
The first indisputable reference to ginkgo comes in two Japanese dictionaries and a
textbook from the mid-fifteenth century. The dictionary Ainosho from 1446 by Gyoyo
is published in a characteristic style, also used in early Chinese literature, which poses
direct, specific questions and then uses previous references to answer them. To the
question “What is ginkgo?” the answer comes back “There is no name in Wamyo,”
which is believed to be an earlier dictionary, the Wamyo-sho, published about 934. The
Ainosho seems to set a lower and upper limit for the introduction of ginkgo into Japan
between 934 and 1446.10
In the Ainosho and also in the textbook Sekiso Orai, written by Ichijo in the fifteenth
century, the Japanese name for ginkgo is written in kanji characters as “silver apricot,”
exactly as in earlier Chinese sources. But it is also accompanied by the phonetic pro-
nunciation in katakana script that gives the pronunciation as icho. In the Kagaku-shu, a
dictionary from 1444, ginkgo is listed in the same way, but “duck’s foot,” another older
Chinese name, is also mentioned, along with an explanation of why that term became
the name of a tree. It is a reference to the similarity between the shape and diverging
veins of the ginkgo leaf and the webbed feet and diverging toes of the mandarin duck,
a lone bird revered for its symbolism. In Japanese literature the earliest reference to
ginkgo appears in the travel diary of the poet Socho in 1530, who writes of giving a gift
of beautiful yellow autumn ginkgo leaves.
Taken together, evidence from historical documents indicates that ginkgo was in
cultivation in China by the eleventh century, or perhaps the late tenth century. It per-
haps spread through the gifting of seeds and cuttings, which grew increasingly com-
mon in China from the tenth century onward. It clearly came to Korea and Japan some-
 189
reprieve

what later. In Japan there is no reliable evidence for ginkgo before the early part of the
fifteenth century, three or four hundred years after ginkgo is well-established in China,
probably with an association with Buddhism. The implication is that any ginkgo tree in
Japan is unlikely to be older than about seven hundred or eight hundred years; none is
likely to approach an age of a thousand years.11
The link between ginkgo and Buddhism may have been part of the motivation for
its transfer to Japan, but ginkgo was perhaps even more important for food and medi-
cine. And these motivations may not have been mutually exclusive; as in Europe, the
early development of medicine in eastern Asia is closely linked to the development of
religious beliefs. Buddhist monks would have been among the most important early
practitioners of Chinese traditional medicine in medieval Japan.12
As to the likely place of ginkgo’s introduction to Japan, it seems most plausible to
look to the west and the south. In particular, the spotlight settles on Kyushu, the south-
ernmost and westernmost of the four main Japanese islands, and a place with espe-
cially strong historical connections with mainland Asia. For centuries through medi-
eval times, there was a thriving trade through the Kyushu area, in which ceramics
played an especially important role. Beginning toward the end of the Heian period in
the twelfth century, and continuing through the Kamakura and Muromachi periods
in the thirteenth through sixteenth centuries, ceramics were greatly valued by the elite
and were imported into Japan in large quantities. Colonies of immigrant Chinese be-
came established in Japan and helped fuel trade in everything from precious metals to
inkstones and water droppers for calligraphy. They were also importers of medicinal
and other useful plants from the mainland; ginkgo may well have been among them.13
 26
Voyages

He goes a great Voyage, that goes to the Bottom of the Sea.

—Thomas Fuller, Gnomologia

In May 1975 a fishing boat working in an area of high tides and strong currents off
the northwest coast of Jeungdo Island, off the southwestern Korean Peninsula, dredged
up in its nets six pieces of light green Chinese celadon ceramics and white porcelain:
the first hint of the Shinan Ship, a discovery that ranks alongside the Vasa of Sweden
and the Mary Rose of Britain among the most remarkable in underwater archaeology.
At first the Korean authorities were uncertain how to proceed, but when the site at-
tracted looters in the autumn of 1976, a high-profile salvage project was launched by
the Korean Cultural Heritage Administration with the support of the Korean Navy.
This massive nine-year effort opened an extraordinary time capsule from the four-
teenth century and a treasure trove of exquisite objects that provided new perspectives
on trade between China, Korea, and Japan in medieval times.1
The circumstances under which the Shinan Ship sank are not known. It was prob-
ably blown over in a storm, and as it went down, with the crew aboard, it crashed on
underwater rocks before settling on the muddy seabed. Over time, in about sixty-five
feet of water, the ship was partly buried by mud and sand. The parts left exposed were
destroyed or washed away, but those parts of the cargo and hull protected from decom-

190
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v o ya g e s

position and shipworms remained in excellent condition. In 1990 the Marine Antiques
Preservation Center was established at Mokpo on the mainland to help deal with the
huge amount of material recovered and to oversee the ship’s preservation.2
Excavating the wreck of the Shinan Ship was a formidable task. The Korean authori-
ties had never undertaken an underwater excavation, and the navy divers found them-
selves working in darkness in muddy, fast-flowing waters. Initially, operations were
aimed mainly at protecting the site, preventing looting, and assessing whether further
work was warranted. However, the nearly two thousand pieces of ceramics and more
than six thousand coins recovered by those early efforts made it clear that a careful,
large-scale excavation would be well rewarded.
Salvage started in earnest in July 1977 with the support of two naval vessels and
about sixty deep-sea divers. The initial work recovered tens of thousands of objects,
many still preserved in the wooden crates in which they were originally packed. There
was no doubt that the Shinan Ship was a trading vessel, and at ninety feet long, twenty-
five feet wide, and about 260 tons, it was a relatively large one for its time, packed to
the brim with merchandise of all kinds.3
Over the following summers until September 1984, the excavations recovered a fur-
ther vast collection of objects, as well as the remains of some of the crew. An area of
more than half a mile around the wreck was also checked with dragnets to scour up
any last evidence about the ship, what it was carrying, and where it was going. The
ship’s timbers were also removed from the seabed for study, restoration, and exhibit,
and many small objects were scooped or siphoned from the mud of the seabed in those
final stages of the excavation. Among the plants recovered there was a single but un-
mistakable seed of ginkgo.4
Altogether, more than twenty thousand objects were brought to the surface after
six and a half centuries on the seabed. The bulk was pottery of various kinds. There
were about five thousand pieces of white porcelain and about three thousand pieces
with varied glazes, but the majority, more than twelve thousand pieces, was distinctive
green celadon. There were figurines, water droppers, incense burners, teapots, cups,
cup stands, mortars and pestles, and even a ceramic pillow, as well as a variety of beau-
tiful dishes, bowls, and vases. The preponderance of Chinese celadon makes it obvi-
ous that the Shinan Ship was on its way from China when it sank. Only seven pieces of
Goryeo celadon were from Korean kilns.5
Some of the plant materials recovered from the Shinan vessel were part of the cargo,
 192
h istory

Reconstruction of the
Shinan Ship, a trading
vessel that sank in June
1323 on its way from China
to Japan. A seed of ginkgo
and other exotic plants
were among its cargo,
which consisted mainly
of fine ceramics from
Chinese kilns.

and had probably come from southeastern Asia, perhaps the Malay Peninsula, or even
farther afield. There were more than a thousand pieces of sandalwood, most cut into
pieces about six feet long. This exotic aromatic hardwood, which was used to make
high-quality furniture or burned to produce fragrant smoke, could have come from
southern China or southeast Asia, but it may have been brought from as far away as
India. Also from the tropics was a large amount of black pepper in a rectangular stor-
age box, as well as massive quantities of croton fruits, which are used medicinally as a
purgative.
Other plant materials were found in much smaller quantities, but of sixteen species
identified, fourteen were used in traditional Chinese medicine. They included two dif-
ferent kinds of ginger root as well as cinnamon sticks. There was also charcoal powder,
made from dense logwood, which is used to help stop bleeding and treat dysentery.
Along with the ginkgo seed, there were pieces of lychee, betel nut, apricot, peach, wal-
nut, and hazelnut. All may have been part of the medicine chest on board the ship, a
conclusion also supported by other objects discovered in the wreck, such as millstones
for grinding medicines, as well as spoons and scales suitable for measuring and weigh-
ing herbal ingredients.
The celadon pottery showed that the Shinan Ship was carrying a cargo from China,
and that the bulk of these ceramics were from the Longquan kilns. Other bluish white
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v o ya g e s

and white porcelain came from the Jingdezhen kilns. Both are in the hinterland of the
important port area of Qing Yuan Lu, which is now the area around Ningbo. Further
evidence that Ningbo was the port of departure comes from one of the scale weights,
which is inscribed Qing Yuan Lu, as well as some of the curious 364 wooden tags re-
covered from the wreck. Like some of the other fragile objects from the Shinan Ship,
these were recovered mainly during the final phases of the excavation from the mud in
and around the wreck site.
Each tag is a short flat piece of wood up to about six to eight inches long with a hole
or two notches at one end that were evidently used to tie the tag to the cargo. Remark-
ably, some of these tags identified the owner of that piece of the cargo, and included
the owner’s signature, the number of units or weight of the cargo, and its type. The tags
not only confirm the port of departure as Qing Yuan Lu but also tell us when the ship
left on its ill-fated voyage. More than a hundred tags record the shipping date. One is
labeled April 20, six are labeled April 23, thirty-seven are labeled May 11, one is labeled
June 1st, nine are labeled June 2, and fifty- eight are labeled June 3. And on the eight
tags where the year is mentioned it is 1323. The ship probably left port in early June 1323
and may have been at sea for several weeks before it went down in a summer storm or
perhaps an early typhoon.
The tags also provide direct evidence of who owned the cargo and where it was to be
delivered. A hundred and one tags were labeled Gangsa, the title of a Buddhist monk
responsible for aspects of temple administration. Forty- one tags were labeled for the
Tōfukuji Temple in Kyoto, while others included the Hakozaki Shrine and Chojuan,
one of the smaller branch temples belonging to the Jotenji Temple, both in Fukuoka.
Historical records for this time, during the thirteenth and fourteenth centuries, show
that some of the larger temples in Japan dispatched licensed trade ships to China, espe-
cially from Fukuoka on Kyushu, where there was a naturalized Chinese population.
Among the tags designating the names of private cargo owners, about twelve seem
to belong to Buddhist monks; a further twelve are clearly Japanese. The Chinese or
Korean nationality of the others is uncertain.6
There would have been many possibilities for the movement of ginkgo from China
to Korea and Japan through well-established trade routes plied by craft like the Shinan
Ship. The ginkgo nut from the Shinan Ship, which dates from the early fourteenth cen-
tury, is also consistent with the documentary evidence for the presence of ginkgo in
Japan by the 1440s. The trees probably would have needed to have been in Japan and
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history

yielding nuts for a few decades before they attracted sufficient interest to be included in
a dictionary. This suggests that ginkgo may have been in Japan by the 1300s.7
By the time of the first Western contact with Japan in 1543, ginkgo had probably been
assimilated into Japanese culture for a century or two. By then there is also abundant
evidence for an early association between ginkgo and Buddhist temples. The Okazari
no sho from 1523 records the furniture, tools, and utensils that the Shogun Ashikaga
Yoshimasa kept in the great Ginkaku-ji Temple, the Temple of the Silver Pavilion, that
he founded as a private villa in Kyoto in 1460. Among them is mention of icho-guchi,
a “ginkgo-mouthed flower vase.” Also, then as now, the distinctive leaves of ginkgo
proved an irresistible motif. From the late fifteenth century or early sixteenth century
there is a long box (nagafbako) used for the storage and transportation of scrolls that
is decorated with a crest (mon) of five ginkgo leaves. The Kikigaki Shokamon, a book of
heraldry written in the fifteenth century, includes some 260 family crests, one of them
composed of three ginkgo leaves. From the Azuchi-Momoyama period at the end of
the sixteenth and beginning of the seventeenth century, there are kimonos decorated
with ginkgo leaves. One of the national treasures in the National Museum of Japan in
Tokyo is a man’s silk hip-length jacket (kodofuku) decorated with embroidered ginkgo
leaves. By late medieval time the Japanese elite had already recognized ginkgo as some-
thing special.8
 27
Renewal

The only real voyage of discovery consists not in

seeking new landscapes but in having new eyes.
—Marcel Proust, À la recherche du temps perdu

After millennia of decline, the reprieve that ginkgo found through its association
with the cultures of eastern Asia was followed in the eighteenth century by an even
more marked turnaround in its fortunes. This renewal began on September 25, 1690,
with the arrival of the German-born physician-botanist Engelbert Kaempfer in what
is now the heart of the Japanese city of Nagasaki. Kaempfer stayed just two years, but
what he learned, and what he later wrote, distinguish him in the West as “the first in-
terpreter of Japan.” He was also the first to introduce ginkgo to Western science.1
Located on the western shore of Kyushu, the farthest east and south of the four main
Japanese islands, Nagasaki stands at the head of a fine natural harbor where the ancient
trade routes that connect mainland Asia and Japan come together. To the north, there
is a short crossing to the Korean Peninsula via the ancient port of Hirado and the island
of Tsushima. To the south, the western shore of Kyushu leads to the gentle arc of the
Ryuku Islands and on to Okinawa and Taiwan. From there, Canton, Hong Kong, and
Macau are not far to the east. To the south, across the Luzon Strait, the Philippines are
the gateway to the islands of southeast Asia.

195
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history

From Nagasaki the most direct route to China is west to the Goto-retto Archipelago,
on to Jeju Island, and then southwest, out across the East China Sea to the Zhoushan
Archipelago just off the Chinese coast, and from there to the ancient Chinese cities of
Ningbo and Huangzhou. This was the lucrative trade route that the Shinan Ship was
taking and that connected the rich ancient cultures of China to Korea and Japan. By
the time Kaempfer arrived in Japan at the close of the seventeenth century, ginkgo
had been assimilated into Japanese culture for perhaps two hundred or three hundred
years, and by the 1690s Kaempfer records that it was grown “almost everywhere in
Japan.” This makes it likely that he was not the first Westerner to see it.2
In discussing trees from China, which were then very poorly known, John Evelyn,
who published his great Silva: a Discourse of Forest Trees in 1664, mentions two great
trees; one “a certain Tree called Ciennich (or the Tree of a thousand years) in the Prov-
ince of Suchu near the City Kien, which is so prodigiously large, as to shrowd two
hundred sheep under one only branch of it without being so much as perceived by
those who approach it,” the second “a greater wonder . . . in the Province of Chekiang,
whose amplitude is so stupendiously vast, as four score persons can hardly embrace.”
There are several possibilities for the identity of these trees, including figs, which are
often striking and achieve a great size, but there is just a chance that they could also be
ginkgo.3
The first contact between Japan and the West had come even earlier with the ar-
rival of the Portuguese in 1543, nearly 150 years before Kaempfer landed at Nagasaki.
The Portuguese came from the south, bringing goods from Europe, as well as ceramics
and other valuables from China, especially after establishing a trading base in Macau
in 1557. European guns, first introduced by the Portuguese in the sixteenth century,
later proved decisive in internal power struggles within Japan. The Portuguese also
brought bread and tobacco, most likely chilies too, as well as new words that found
their way into the Japanese language. The Japanese word tempura comes from the Por-
tuguese word tempero, “seasoning.” However, the most important Portuguese intro-
duction into Japan was Christianity, which, intertwined with trade, was instrumental
in the establishment of Nagasaki as a major port. Just a few years after first contact
with the Portuguese, Jesuit missionaries began to arrive, and widespread conversion to
Christianity followed. By the late 1580s the Catholic influence in southern Japan was
widespread and growing. Steadily and predictably, it became an increasing cause of
friction.4
 197
r e n e wa l

The fan-shaped man-made island of Deshima in Nagasaki Harbor where Engelbert Kaempfer,
Carl Peter Thunberg, and Philipp Franz von Siebold were based while working for the
Dutch East India Company.

In 1587, concerned with the growing influence of the Jesuits in local politics, the feu-
dal ruler Toyotomi Hideyoshi, the great unifier of Japan in the late sixteenth century,
ordered the expulsion of the missionaries. Nine years later, on suspicion of a potential
invasion, he ordered the crucifixion of twenty-six Catholics in Nagasaki, but the ten-
sion continued. In 1614 the Shogun Tokugawa Ieyasu finally banned Catholicism and
followed through on the expulsion of the missionaries.
In 1636, in a further attempt to curb the influence of the Portuguese, Tokugawa
Ieyasu confined them to the man-made island of Deshima constructed in Nagasaki
Harbor. From Deshima, their activities could be monitored more readily and the lucra-
 198
history

tive trading activities could be more easily controlled. When asked in what shape the
island should be made, Tokugawa Ieyasu is reputed to have simply spread out his fan.
Deshima was thus created by cutting a canal through a small promontory and fashion-
ing the resulting island as an arc linked to the mainland by a single bridge. This tiny
island served as the main point of contact between Japan and Europe for more than
two hundred years. It was the cultural and commercial conduit through which ideas
and materials flowed in both directions.
While Deshima was founded as a base for the Portuguese, it soon became home
to the Dutch. In April 1600, the Dutch ship Liefde arrived off the coast of present-
day Usuki after a catastrophic voyage through the Straits of Magellan and across the
Pacific. On board, one of the few lucky survivors was the Englishman William Adams.
He became a key figure in early Western contact with Japan and nearly four hundred
years later was the model for John Blackthorne, the fictional Englishman at the center
of James Clavell’s novel Shōgun.5
Adams flourished in Japan through the patronage of Tokugawa Ieyasu. He became
the Shogun’s personal adviser, was established as a samurai, and is best known for as-
sisting the Japanese in the development of Western-style sailing ships and new trading
ventures. Adams helped strengthen faltering early trade between Britain and Japan,
but more important, Adams assisted the Dutch East India Company in breaking the
near monopoly enjoyed by the Portuguese. With his help the Dutch began trading from
Hirado, also on Kyushu, in 1609. With the expulsion of the Portuguese after the Shima-
bara Rebellion of 1637 and 1638, the Dutch became the new occupants of Deshima; this
relationship became the most important European influence on Japan until the mid-
nineteenth century.6
In the seventeenth century Deshima was a far-flung outpost of the Dutch trade net-
work, but the writings of several of those who were stationed there became the main
source in the West for early information on Japan. In Japan it also became known as
a center of learning about certain branches of science and technology. It was a mag-
net for scholars of Rangaku, or “Dutch studies,” and through the inquisitiveness of a
succession of three physician-botanists, Kaempfer, Thunberg, and Siebold, all in the
employ of the Dutch, Deshima also became the route through which Western science
first learned about the plants of East Asia. Almost certainly it was also one of the ports
from which ginkgo first embarked on its journey from eastern Asia to Europe.
Engelbert Kaempfer, the first of the Dutch physician-botanists to be based at De-
 199
r e n e wa l

shima, grew up in Lemgo, in northern Germany, about midway between Amsterdam

and Berlin. As the second son of the pastor of Saint Nicholas church, he came from
an educated family, and from 1674 to 1676 he studied languages, history, and medicine
at the University of Cracow. This was followed by four years at Königsberg in Prussia
studying natural sciences and medicine, and in 1681, Kaempfer moved to Sweden and
the University of Uppsala.
The journey that took Kaempfer to Japan had an unlikely beginning in Stockholm at
the court of Charles XI. It then followed an even more unlikely route, overland via Rus-
sia to Iran when Kaempfer was appointed secretary to the Swedish Legation to the Per-
sian Court. The legation traveled to Isfahan in central Iran by way of Moscow—where
Kaempfer had an audience with Peter the Great and his half-brother Ivan V—and it
eventually arrived at the court of the sultan in March 1684. Ultimately, the Swedish
mission was not a success, but Kaempfer then took employment with the Dutch East
India Company, working for a time as a physician at the Dutch trading base at Bandar
Abbas on the Persian Gulf. While there, he studied the local plants and also made care-
ful observations of the date harvest, about which he published a detailed account on
his return to Europe.
In June 1688, in the employ of the Dutch, Kaempfer left Persia on a voyage that took
him to Muscat, the coast of India, Sri Lanka, and then on to the Dutch East India Com-
pany headquarters in Batavia, close to present- day Jakarta in Indonesia. He arrived
there in September 1689, and in early May of the following year he left for Japan. In his
posthumous work The History of Japan, Kaempfer recounts his voyage from Indonesia
to southern Japan, including a month spent in Siam, modern-day Thailand.
When Kaempfer was at Deshima, the Dutch were virtual prisoners; their activities
and especially interactions between them and the Japanese were strictly controlled.
Over the winter Deshima was home to only a few dozen Dutch residents supported
by Japanese who worked as everything from interpreters to cooks, but when the ships
arrived, the port would have swarmed with both Dutch and Japanese as the cargo was
unloaded, trades were made, orders were taken, and the new cargo was stowed for the
long journey back to Europe.
As the Dutch contingent posted on Deshima came and went, the year followed a
regular cycle. In the late summer and early autumn the boats would arrive from Bata-
via loaded with goods from Europe and also from the tropics. During the early autumn
they would unload and be reprovisioned in time for departure before the weather de-
 200
h istory

teriorated and winter set in. In the spring, the head of the Dutch mission, treated like a
Japanese feudal lord (daimyo), would lead a delegation to Edo, modern Tokyo, to pay
tribute and present his respects and gifts at the court of the Shogun. This was the same
journey that Siebold took more than a century later. It took about a month in each di-
rection, and was the best opportunity for the Dutch to see and learn more about Japan.7
In his two years in Japan, Kaempfer made the journey to Edo twice. In The History
of Japan he gives a vivid account of “the Author’s two journies to the Emperor’s Court
at Jedo, the City of his Residence,” describing the preparations for the journey and the
buildings and other structures that he saw. He describes how he traveled, the accom-
modations in which he stayed, the people that he encountered, and what took place
along the way. He also describes in detail his stay in “the city of Jedo, its Castle and
Palace, with an account of what happened during our stay there; our Audience and
Departure.” He writes: “I went to the Emperor’s court twice myself to my very great
satisfaction: the first time, in the year 1691 with Henry von Butenheim, a gentleman of
great candour, affability and generosity . . . the second time, in 1692, with Cornelius van
Outhoorn, brother to the Governor General at Batavia, a gentleman of great learning,
excellent sense, and well skilled in several languages . . .”
Kaempfer continues:

In the first place we set out from Nagasaki, to go by land across the Island
Kiusju, to the town of Kokura, where we arrive in five days time. From Kokura
we pass the streights in small boats going over to Simonoseki, which is about
two leagues off where we find our abovemention’d barge riding at anchor and
waiting our arrival, this harbour being very convenient and secure. . . . At Simo-
noseki we go on board our barge to proceed from thence to Osacca, where we
arrive in eight days, more or less, according as the wind proves favourable or
contrary.

He goes on:

Osacca, is a city very famous for the extent of its commerce, and the wealth
of its Inhabitants. It lies about thirteen Japanese water leagues from Fiogo,
which we make in small boats, leaving our large barge at that place to await
our return. From Osacca we go again by land, over the continent of the great
Island Nipon, so far as Jedo, the Emperor’s residence, where we arrive in about
fourteen days or more. The way from Osacca to Jedo is by the Japanese call’d
 201
r e n e wa l

Tookaido, that is, the Sea, or coast-way. We stay at Jedo about twenty days, or
upwards, and having had our audience of his Imperial Majesty, and paid our
respects to some of his chief ministers and great favourites, we return to Naga-
saki the same way, compleating our whole journey in about three months time.8

Kaempfer made good use of his journeys to Edo and his time in Japan. When he
boarded his ship on the last day of October 1692 to return to Batavia and then to
Europe, he took with him a rich collection of specimens, books, and cultural objects:
the first effort by any European to take samples reflecting the environment and culture
of Japan. By May of the following year Kaempfer was in the Cape, and on October 6,
1693, he arrived in Amsterdam.
Kaempfer had intended to devote his time to writing an account of his travels, but
his book Amoenitatum Exoticarum was not published until 1712, twenty years after he
had left Japan. It is an extraordinary work, full of original observations and the story of
his journey overland to Persia and then to southeast Asia. He also provides accounts of
cultural and natural items that caught his interest, such as Japanese tea, and describes
the plants that he had encountered, many of which were completely unknown to West-
ern science. Included are the first Western descriptions of many Japanese plants, in-
cluding many, like ginkgo, that had been introduced from China.9
On page 811 of the Amoenitatum Exoticarum, in the section dealing with fruit and
nut trees, Kaempfer gives an illustration of a twig of ginkgo with its distinctive leaves
inscribed “ᱻ䨢 Ginkgo. vel Gín an, vulgò Itsjò. Arbor nucifera folia Adiantino.” This is
the first Western illustration of ginkgo and probably was drawn by Kaempfer himself.
The original sketch is among the Kaempfer papers in the British Library.
Decades later, in 1775, the position held by Kaempfer was taken by Carl Peter Thun-
berg, a talented and inquisitive student of Linnaeus. Building on what Kaempfer had
accomplished, Thunberg’s Flora Japonica was the first detailed account of the plants of
Japan. A few years later, in 1779, Isaac Titsiugh arrived as Opperhoofd of the Dutch trad-
ing station. As a surgeon, scholar, and trader, he became a key figure in late-eighteenth-
and early-nineteenth-century European relations with Japan and China.10
Throughout its history the export of ceramics was a lucrative part of trade passing
through Deshima, and some of these early- eighteenth-century ceramics give direct
evidence that the aesthetic possibilities of ginkgo had already caught the eye of artisans
who were creating high- quality porcelain in southwestern Japan. Ginkgo leaves ap-
 202
history

The drawing of ginkgo published by Engelbert Kaempfer in his Amoenitates, 1712:

the first illustration of ginkgo by a Western botanist.

pear occasionally among the many plant motifs that decorate the extraordinary bowls,
dishes, and many other objects that were created as special gifts for the shogunate
from the kilns of the Arita region. In the Kyushu Ceramic Museum one such white
bowl shows three ginkgo leaves arranged in a circle. Still more striking is a beautiful
large shallow dish with two stout upright ginkgo branches picked out in a cobalt-blue
 203
r e n e wa l

underglaze and covered with ginkgo leaves. Strong horizontal elements, a little above
the midpoint, represent mist passing through the branches. It was produced in the
Okawachi Kiln in Hizen between about 1700 and 1730, not long after Kaempfer was in
nearby Nagasaki. Several large ginkgo trees still growing in that part of Japan would
have been alive during the time of Kaempfer, Thunberg, and Siebold.11
 28
Naming

If you do not know the names of things, the knowledge of them is lost too.
—Carl Linnaeus, Philosophia Botanica

Perhaps the most curious thing about Kaempfer’s introduction of ginkgo to the
West is the word itself, and much has been written about how Kaempfer came up with
this seemingly strange name. To understand how and why Kaempfer fastened upon
the name ginkgo it is important to understand how the plant was referred to in Japan
around the time he was there in the late seventeenth century.1
When ginkgo was introduced into Japan, perhaps in the thirteenth or fourteenth
century, the names most commonly attributed to it in China probably came with it.
Among them was the name still used there today, silver apricot. In Chinese charac-
ters this is written 䞬ᱻ, and this is also how ginkgo is written in the Chinese character
form (kanji) of Japanese. The characters must have seemed strange to Kaempfer, but he
carefully copied them into his book. He wrote “silver apricot” as ᱻ䞬 rather than 䞬ᱻ
because at that time in Japan the characters were written from right to left. Kaempfer
then followed those characters with the transliteration of the two words that his Japa-
nese translators were using to refer to the tree: Gín an and Itsjò.
When ginkgo first appears in Chinese historical texts it has two names: 䞬ᱻ, silver
apricot, and 剔㚟, duck’s foot. Later, beginning in the Yuan Dynasty, other names also

204
 205
naming

began to be used: ⮩᳈, white fruit; ‫ژ‬႗ὥ, grandfather-grandchild tree; and ⮩ⱨ,
white eye. Among these, pei kuo or bai guo, meaning “white fruit,” is the most widely
used colloquial name in China, although 䞬ᱻ, silver apricot, is used most often in writ-
ing. In modern Japan, however, as in Kaempfer’s time, when ginkgo is written in Chi-
nese characters, 䞬ᱻ, it can be pronounced in two ways, icho and gin’nan. Shihomi and
Terumitsu Hori reviewed how these pronunciations arose, and they suggest that they
are corruptions of ways that the two original Chinese names of ginkgo, duck’s foot and
silver apricot, were pronounced. In standard Chinese these characters are pronounced
yajiao or yinxin, but in the dialect of southern Jiangsu and northern Zhejiang, the part
of China from which the Shinan Ship came and with which Japan had strong links
through trade, 剔㚟 is pronounced ai cho and 䞬ᱻ is pronounced nin an.2
The links between what Kaempfer wrote as “Itsjò” and the modern Japanese “icho,”
and between Kaempfer’s “Gín an” and “gin’nan,” are clear enough, but where did
“ginkgo” come from? Shihomi and Terumitsu Hori investigated this carefully with the
help of a distinctive feature of the Japanese language. Frequently, when a Chinese char-
acter is used, it may be accompanied by a phonetic spelling, a so-called kana.
By examining the kana associated with the Chinese characters 䞬ᱻ, silver apri-
cot, in various Japanese dictionaries and books published between the fifteenth and
eighteenth centuries, the Horis made an extraordinary discovery. In almost all the
works they looked at, the pronunciation of 䞬ᱻ is generally given either as “icho” or
“gin’nan,” or a clear variant of one of these two possibilities. However, in two Japanese
pictorial dictionaries from the seventeenth century that were in use around the time
Kaempfer was in Japan, the pronunciation is given as “ginkyo.” In the Kagaku-shu,
a dictionary published between 1617 and 1619, the pronunciation of 䞬ᱻ is given as
“icho” and “ginkyo,” while in the Kinmo Zui, published in 1666, the pronunciation is
given as “gin’nan” and “ginkyo.”
In the 1940s A. C. Moule carefully examined Kaempfer’s manuscripts, now pre-
served in the British Library, and found about ten mentions of the plant to which
Kaempfer gave the name ginkgo, and noted in particular that “Gin’nan 32” is referred
to as Catalogus Plantarum Iaponicarum in Kin mo chjju i. The Horis point out that this
almost certainly refers to the Kinmo Zui, and they make a further important connec-
tion by realizing that Kaempfer obtained copies of this book while in Japan. There can
be little doubt that Kaempfer referred to the Kinmo Zui both when he was in Japan and
when he was writing the Amoenitatum Exoticarum, and that this is where the name
 206
h istory

ginkgo came from. With its later endorsement from Linnaeus it has become perhaps
the most internationally recognized of all seventeenth-century Japanese words.3
There is one final puzzle, and perhaps the most difficult one: why did Kaempfer
write “ginkgo” rather than “ginkyo”? This is particularly intriguing because in the
preface of his book Kaempfer explains that he went to great lengths to transcribe the
Japanese language as accurately as possible. He listened carefully to how words and
sounds were pronounced, and developed a system of rules to commit them to paper.
Because Kaempfer was so meticulous, Shihomi and Terumitsu Hori point out, the
plant names that he transcribed into the Roman alphabet in Amoenitatum Exoticarum
vividly capture how those names were pronounced by the Japanese people at the end
of the seventeenth century. So if Kaempfer heard “ginkyo,” why did he transcribe it as
“ginkgo”?
Some have thought that the spelling of ginkgo was a simple mistake, a misspelling
by Kaempfer or an error by the typesetter, but the Horis think that ginkgo is the spell-
ing Kaempfer intended. They suggest that the mysterious second g in his spelling of
ginkgo is a legacy of his own roots in northern Germany. In northern German dialect,
the sound of j is often written as g. For example, Kaempfer might have said “jut” but
would have written it “gut.” The Horis suggest that in the context of his own back-
ground, Kaempfer transcribed “ginkyo” just as he heard it.4
Whatever the origin of the second g in ginkgo, the broader significance of Kaemp-
fer’s journey to Japan was widely recognized after the publication of the Amoenitatum
Exoticarum, and after his death on November 2, 1716, his collections were put up for
sale. They were quickly snapped up by Sir Hans Sloane, a wealthy and acquisitive
patron of the sciences during the Enlightenment. It was Sloane, more than anyone
else, who ensured that Engelbert Kaempfer and his work were not forgotten, and it was
through Sloane that significant parts of Kaempfer’s unique collections, which docu-
ment one of the earliest Western encounters with ginkgo and with Japan, were saved
for posterity and came to be in London.5
Sloane was an inveterate collector with a particular interest in plants. He collected
plants in southern England and also in the south of France, but he will be forever asso-
ciated with the plants that he collected during his fifteen months as physician to the
governor of Jamaica, the Duke of Albemarle. Sloane came back from the West Indies
in 1689 with a collection of eight hundred dried plant specimens and enough original
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naming

observations for a large two-volume book; he also found that while he was away, the
Catholic King James II had been replaced by the Protestant William of Orange.6
Returning to London, Sloane established a lucrative medical practice, based in
Bloomsbury, which catered to some of the most influential and wealthy figures of the
day. As a result Sloane too became well known, wealthy, and increasingly influential.
He was elected to the Royal Society in 1685, became its secretary in 1693, and in 1696
published his Catalogus Plantarum, a list of all the plants that he had encountered
while in Jamaica. His better-known two-volume work Natural History of Jamaica ap-
peared in 1707 and 1725. Volume 1, which deals mainly with the plants, contains the
first record in English of the use of chocolate. In 1719 Sloane became president of the
Royal College of Physicians, a post he held until 1735. He succeeded Sir Isaac Newton
as president of the Royal Society in 1727.7
One of Sloane’s great scientific passions was acquiring important collections of
plants being gathered during the European exploration of different parts of the world
at that time, and it was this that preserved the legacy of Engelbert Kaempfer. Sloane’s
extraordinary collection of collections is the most extensive and historically important
accumulation of early botanical specimens anywhere and vividly documents the rapid
expansion of scientific knowledge about the diversity of plants during the seventeenth
and early eighteenth centuries. All told, the Sloane collection consists of 337 separate
collections preserved in 265 huge luxuriously bound books with green leather covers
that are today housed in the former Department of Botany, now part of the Depart-
ment of Life Sciences, in the Natural History Museum in London.8
Sloane collection 211 is labeled “H.S. 211, ff Volumen Plantarum in Japonia collec-
tarum ab Engelberto Kempfero M.D. annis 1691 et 1692. Additae sub finem Plantae
aliquot ab eodem in Persia et Insula Ceylan repertae.” It is one of the most consulted
herbaria in the Sloane collection. No other herbarium contains specimens of such an
early date collected from Japan, and it is therefore a key resource for the scientific study
of Japanese plants. For this reason, Sloane’s Kaempfer herbarium has been studied by
some of the greatest figures in the history of botany, including Daniel Solander, who
accompanied Sir Joseph Banks on Cook’s first voyage to Australia, and Robert Brown,
the great botanist who first observed “Brownian motion.” It was also consulted by Carl
Peter Thunberg and Franz von Siebold, Kaempfer’s successors on Deshima.
The Kaempfer collection was also well known to J. E. Smith, who, in 1786, in one of
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history

the more curious episodes in the history of scientific study of ginkgo, took it on him-
self to rename ginkgo Salisburia adiantifolia, “in honour of Richard Anthony Salis-
bury, Esq., F.R.S. and F.L.S., of whose acuteness and indefatigable zeal in the services
of botany no testimony is necessary in this [the Linnean] society, or in any places
which his writings have reached.” Salisburia is used in favor of Ginkgo in some of the
nineteenth-century botanical literature, but it never really caught on, and under the
modern international rules of botanical nomenclature the name Ginkgo takes prece-
dence. Salisburia is considered an invalid synonym.9
In addition to obtaining Kaempfer’s collection of plants, Sloane also obtained
Kaempfer’s papers, and Sloane manuscript 2914, which is now in the British Library,
is titled in Kaempfer’s own hand “Delineato Plantarum Japonicarum manu Engelberti
Kempferi.” It is a volume of 217 folio-size drawings of Japanese plants, accompanied
by an alphabetical list of names in Japanese with references to his Amoenitatum Exoti-
cum and other works.
Realizing the significance of what Kaempfer had written, and also how much Kaemp-
fer had not covered in his Amoenitatum Exoticarum, Sloane gave Kaempfer’s papers to
John Gaspar Scheuchzer, his personal librarian, to be translated from Kaempfer’s High
German dialect. The work that Kaempfer was preparing for publication, The History of
Japan, was completed and published by Scheuchzer in 1727.10
Ginkgo is mentioned specifically in Kaempfer’s chapter 9, which is titled “Of the
Fertility of the Country as to Plants,” and along with short accounts of “Mulberry
trees,” “Paper-tree,” “Varnish-tree,” “Bay-tree,” “Tea-shrub,” and many other kinds
of trees and useful plants, Kaempfer comes to a nut called “ginau”: “Another sort of
Nuts, call’d Ginau, as big as large Pistaches grown very plentifully almost everywhere in
Japan, on a fine tall tree, the leaves of which are not unlike the large leaves of an Adian-
thum. The Japanese call it Itsionoki. The nuts afford plenty of Oyl, which is also much
commended for several uses. As to a more accurate description if this Tree, I refer the
Reader to the Amoenitates Exotic.”11
The History of Japan proved to be an enormously influential work. According to
some, Jonathan Swift’s Gulliver is something of a composite of Kaempfer and the sailor
William Adams. Some of the incidents in Gulliver’s journey seem to connect directly
to similar descriptions and incidents in the Scheuchzer translation of the The History
of Japan. Perhaps Sloane circulated Scheuchzer’s translation among his friends at the
Royal Society, in which Jonathan Swift was also a fellow.12
 29
Resurgence

At your return visit our house; let our old acquaintance be renewed.
—William Shakespeare, Henry IV, Part 2

Through his Amoenitatum Exoticarum, Kaempfer was the first to bring ginkgo to the
attention of Western science, but it is unlikely that he was the first to introduce living
seeds or plants to Europe. This probably did not happen until later, perhaps not until
three or four decades into the eighteenth century. The evidence for this time lag between
Kaempfer’s recognition of ginkgo and its cultivation in the West is mainly negative, but
in this case it has at its center the most reputable source of information on all the plants
known to science at that time: the Swedish physician and botanist Carl Linnaeus.1
Linnaeus, one of the greatest of all cataloguers and describers of new plants, spent
the years 1735 to 1738 in Holland. Then a young man, he was employed by the wealthy
Anglo-Dutch banker and great patron of the sciences George Clifford. Clifford was
one of the directors of the Dutch East India Company, which at that time had been in
existence for more than a hundred years. A few decades earlier Clifford’s predecessors
had been Kaempfer’s employers. Clifford, Linnaeus, and their scientific circle were well
connected and would have been on the lookout for interesting exotics from the East,
but there is no mention of ginkgo in Linnaeus’s catalogue of the plants growing in Clif-
ford’s garden, which was published in 1738. There is also no specimen of ginkgo in the

209
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history

Clifford collections, which were acquired by Sir Joseph Banks and are now preserved
in the Natural History Museum in London.2
Kaempfer’s work was published in 1712, and Linnaeus was certainly aware of its
importance. He owned a copy of Kaempfer’s book and commemorated Kaempfer by
naming a genus of tropical ginger after him. However, even by 1753 Linnaeus was still
apparently unaware of the existence of ginkgo or, more likely, unwilling to commit it to
print until he had seen a specimen with his own eyes. In his monumental work Species
Plantarum, a comprehensive compilation of all the world’s plants known to him at the
time, there is no mention of ginkgo.3
A key figure in the circle of people in correspondence with Linnaeus at that time,
and also someone with an interest in procuring plants from all over the world, was
John Ellis, a leading scientist of his day and a prominent member of the Royal Society.
In 1767 Ellis received the Copley Medal, the society’s most prestigious award, for his
work showing for the first time that corals are animals rather than plants. Ellis was also
an excellent networker who used his connections to advance both science and Britain’s
commercial interests. In 1770 he published an account of how seeds and living plants
should be transported from overseas. It was Ellis who played a key role in bringing
ginkgo to the attention of Linnaeus.4
In a letter dated April 25, 1758, Ellis writes to Linnaeus:

If you want a correspondent here that is a curious gardener, I shall recommend

you to Mr. James Gordon, gardener at Mile End London. This man was bred
under Lord Petre and Dr. Sherard, and knows systematically all the plants that
he cultivates. He has more knowledge on gardening than all the gardeners and
writers on gardening in England put together; but he is too modest to publish
anything. If you send him anything rare, he will make you a proper return. We
have got a rare double jessamine (Gardènia flórida) from the Cape, that is not
described: this man has raised it from cuttings, when all the other gardeners
have failed in the attempt. I have lately got him a curious collection of seeds
from the East Indies, many of which are growing, but are quite new to us. He
has got the ginkgo (Salisbùria), which thrives well, and when he has increased
it, he will dispose of it.5

The anticipated arrival of ginkgo took another nine years. In a letter dated July 3,
1767, Ellis wrote to Linnaeus, “James Gordon intends you a plant of the Ginkgo of
Kaempfer. I shall send your specimen of the Siren lacertina at the same time.”6
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resurgence

James Gordon owned the then famous Mile End nursery in east London, and this
was the conduit for many new plants into horticulture. According to Smith, he was al-
ready growing ginkgo in his nursery at Mile End in 1754. Further evidence of the early
cultivation of ginkgo by James Gordon comes from the notes of Dr. John Hope, king’s
botanist to Scotland and professor of botany at Edinburgh. His trip to London in the
summer of 1766 is an important source of information about the plants growing there
at that time.7
Gordon was not only among the first to introduce ginkgo into British horticulture
but he was also the first to provide Linnaeus with a living plant, probably as a cutting
rather than as seed. It was probably the first ginkgo specimen that Linnaeus ever saw.
Gordon wrote to Linnaeus in 1769, just a few years before Linnaeus’s death:

London, Mile End

October 26, 1769
My Lord
Being many years much beholden to your Lordship’s most ingenious and
learned Labours, please to accept the underwritten plants. If there is any thing
particular, that is acceptable, which falls in my way it will be the greatest plea-
sure and honour done me, to furnish your Lordship with it
I am my Lord,
Your Lordship’s obliged servant
James Gordon

1 Ginkgo kaempferi
1 Magnolia acuminata
1 Andromeda mariana
These stand in the natural ground, and bears the severest frost of our winters,
without damage.

Linnaeus wrote back thanking Gordon for the ginkgo plant that he had sent earlier
that year. Since the plant was not yet mature, Linnaeus could not assign to it a definite
systematic position. A specimen, probably preserved from the plant that Gordon sent
Linnaeus, taken after the plant had been established at Uppsala, is now in the Linnaean
herbarium of the Linnean Society of London. This is the specimen on which Linnaeus
based the scientific name Ginkgo biloba. It is a typical long shoot with deeply bilobed
leaves of the kind seen especially clearly on leaves near the tip of the shoot. If the speci-
men had come from a more mature plant, perhaps Linnaeus would not have chosen the
 212
h istory

The ginkgo specimen in the

collection of Carl Linnaeus on
which the name Ginkgo biloba is
formally based. Linnaeus almost
certainly made this specimen near
the end of his life from a living
plant sent to him in 1769
by the British nurseryman
James Gordon.

specific name biloba. Linnaeus first published the name Ginkgo biloba in 1771, almost
at the end of his life, in his Mantissa Plantarum Altera. Linnaeus, who was by then in
poor health, published little after that; he died in Uppsala in 1778.8
Gordon was clearly a key figure in propagating ginkgo from limited early sources,
but as far as I know, Gordon’s records no longer exist, so it is not certain how he ob-
tained the plants, or where they came from. One possibility is that through the connec-
tions of his wealthy friends and patrons he was able to obtain cuttings from the very
few other trees that were in cultivation in Europe by then. It seems less likely that he
was able to obtain seeds from Japan. However, there is also the strong possibility that
he obtained material from China, and again it may have been John Ellis who provided
the key connection.9
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resurgence

As well as Linnaeus’s specimens, the collections preserved in the Linnean Society of

London also include his extensive personal library. Pasted into the cover of Linnaeus’s
copy of Kaempfer’s Amoenitates is a single handwritten sheet of paper headed: “Re-
marks on Kaempfer’s Amonitat. Exotic by John Bradby Blake Esq. Canton 1770.”
John Bradby Blake, a Londoner from a seafaring family, was born in November 1745
and was the son of Captain John Blake, who worked for the British East India Company
and later ran a successful business supplying fresh fish to markets in London. Blake fol-
lowed his father in working for the East India Company and first sailed to China when
he was twenty-one, in 1766 as a supercargo. On one of his visits home he brought with
him the Chinese boy later painted by Sir Joshua Reynolds. Eventually the company
based him more permanently in China.10
While in Canton, Blake pursued a wide range of scientific interests. He sent back
specimens of the kaolin and petuntse used in the making of Chinese ceramics. There
are references to the “Chinese porcelain materials sent from China by Mr. Blake” in the
experiment books of Josiah Wedgwood. He also sent back seeds of all the useful plants
that he came across. According to Henry Laurens, writing in 1773: “Mr. Blake has an
excellent opportunity by the aid of his Son who resides SuperCargo at Canton in China
of introducing many things from thence which may become peculiarly beneficial to
Carolina. He Seem’d most benevolently disposed to do all the good he can in that way,
& only wants the assistance of minds a kin to his own, on our Side of the Water. . . .
Severaral [sic] of the articles from Mr. Ellis were derived from Mr. Blake.”
From Ellis or his associates some of these plants found their way to the Americas.
For example, “the fine Cochin China rice” was distributed to growers in Jamaica, South
Carolina, and the present-day Dominican Republic. It seems likely that Blake trans-
mitted living ginkgo seeds too.11
The note written by Blake in 1770, and now in Linnaeus’s copy of the Amoenitates,
lists information on eight plants, referring to them by their numbers in Kaempfer’s
book.

811
Ginkgo vel Gin-nan pronounced Maulon ging:hang
Pao=Zuo
from the Northward—grows in gardens
but does not flourish at Canton the nuts are eat in
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history

various ways: and are brought down for sale dryed

by fire in great quantities to Canton. Procuring seeds
this hint must be particularly alluded to as also in
many other seed.

It seems unlikely that the original ginkgo plants grown by James Gordon in his Mile
End nursery came from John Bradby Blake. Blake was not sent to Canton until 1766,
eight years after Ellis’s letter to Linnaeus proves that Gordon was already growing it,
but it is not impossible that Gordon obtained living ginkgo in the 1750s from one of
Ellis’s other correspondents, perhaps even Blake’s father. A few years later, Gordon,
and perhaps others in England, may well have received ginkgo collected in China by
Blake via John Ellis. After all, we know that Blake had been sent out for the express
purpose of collecting living plants, and from the note in Linnaeus’s copy of Kaemp-
fer we have direct evidence that he was focused on making sure his seeds were alive.
Confirmation that Ellis was being sent seed also comes from an aside in the account
of ginkgo written by Loudon in his Arboretum et Fruticetum Britannicum; or, The Trees
and Shrubs of Britain: “The nut, when examined by Sir J. E. Smith, from specimens in
his possession, which were sent from China to Mr. Ellis, was found to be larger than
that of the pistachia . . .”12
From this, it seems likely that the enigmatic piece of paper from John Bradby Blake,
pasted into Linnaeus’s copy of Kaempfer, also came from the Ellis correspondence
which he gave to the Linnean Society of London. The note from Blake also makes it
likely that the early ginkgo plants grown in Europe were the result of several separate
introductions from diverse sources, from both China and Japan. The introduction of
ginkgo into Europe almost certainly happened more than once and over a long period
of time. Whatever the ultimate source of the early ginkgo trees growing in Europe, the
evidence suggests that living ginkgo was rare there at least until the late 1750s, but only
a decade or two later it was widely distributed.
It is also clear that James Gordon played a key role in its propagation and spread.
He was probably the source of the ginkgo planted in the Royal Estate at Kew, now the
precious Old Lion. He was growing ginkgo at the Mile End nursery already in 1754.
There is also a tree at Blaise Castle, Henbury near Bristol, which is reputed to have been
planted in 1762, and there is another at Whitfield, Warmbridge in Herefordshire, which
is reputed to have been planted in 1776. Gordon’s nursery may have been the source
of all of them.
part vi

Use
OVERLEAF A large ginkgo tree, probably a male, towers over an orchard
of smaller female trees in early spring near Sobue, Honshu, Japan.
 30
Gardens

The garden suggests that there might be a place where we can meet nature halfway.
—Michael Pollan, Second Nature

Ginkgo owes its resurgence in historical times not just to its utilitarian value but also
to some kind of irresistible biological charisma that has taken hold in both Eastern and
Western cultures. In the East, ginkgo may have made the transition from wild forests
to gardens almost by accident. Buddhist and Taoist priests have long nurtured tracts
of forest with ancient trees around their temples. Chinese and Japanese Buddhism in
particular believe that achieving nirvana, a state of spiritual liberation, is available to
all life, including trees. With their unusual leaves and bizarre chi-chis, ancient ginkgos
perhaps embodied an element of Buddha nature. Adopted also by Confucianism and
Shintoism, ginkgo became widely revered as a symbol of vitality, longevity, and resil-
ience.1
In the eighteenth century, ginkgo expanded from the forests, temples, and orchards
of the East to the botanical gardens and grand estates of the West. Only a few decades
after Kaempfer’s first description, living ginkgo, probably as seeds but perhaps as seed-
lings or cuttings, began to arrive in Europe. Exactly when this took place is uncertain,
but that it was no earlier than about 1700 and no later than about 1750 fits the evidence

217
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from many different sources. This means that there is no living ginkgo outside China,
Japan, and Korea that is more than about three hundred years old.
Given the strong Dutch connections with Japan from the early seventeenth century
onward, the earliest introductions of ginkgo into Europe may have been into the Low
Countries, and two trees, little more than a hundred miles apart, vie for the title of the
oldest living ginkgo outside of Asia. The largest is an old female that stands next to the
church in Geetbets, near Hasselt in Belgium. With a trunk now about five feet across,
it was reportedly planted around 1730. It is said to have been brought to Belgium by
missionaries returning from China. Unfortunately, there is no corroborating evidence,
but from its size alone this tree must be an early introduction, certainly from the eigh-
teenth century. Whether it dates from as early as 1730 is less certain, and that age would
be unusual for a female tree. Most early ginkgos in Europe are males, probably propa-
gated as cuttings from one or a few original sources; the first record of a ginkgo pro-
ducing seeds in Europe is not until 1814. A female tree planted in 1730 ought to have
been mature in the 1760s or 1770s.2
In competition with the Geetbets tree for the distinction of being the oldest in the
West is the large old tree in De Oude Hortus, the old Botanic Garden, close to the cen-
ter of Utrecht. This tree is a male with a trunk about four feet across, roughly a foot
less than the Geetbets tree, but still large. It is unquestionably among the oldest ginkgo
trees in Europe, but again its exact age is uncertain. It is not mentioned by Linnaeus,
who was in Holland between 1735 and 1738, would have visited the Utrecht garden, and
had an eye for unusual plants. It is also not mentioned in the catalogue of the Utrecht
garden from 1747 or in the list of plants growing at Utrecht sent to André Thouin at
the Jardin de Roi in Paris in 1780. However, by the end of the eighteenth century the
Utrecht ginkgo was flourishing. In 1787 the Swiss naturalist Frederick Ehrhart visited
the garden and mentions that the tree was thirteen feet tall.3
In 1838 John Loudon, who covered ginkgo in his comprehensive account of the trees
and shrubs grown in the British Isles, quotes the director of the Utrecht Botanic Gar-
den in the 1830s: “Professor Kops informs us in a letter dated December 7, 1835, that it is
a branchy tree and still continues to grow vigorously. He adds that, when he succeeded
to the directorship of the garden in 1816, it was then calculated to be between 70 and
80 years of age; and hence, it must now (1837) be between 90 and 100 years old; and, if
so, it must have been planted at Utrecht before the tree was introduced into England.”4
A literal reading places this ginkgo as having been planted somewhere between
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gardens

about the mid-1730s and mid-1740s, but its absence from the 1747 catalogue may sug-
gest that it dates from the 1750s or 1760s, around the time when we know that James
Gordon already had ginkgo growing in his London nursery. Perhaps this ginkgo came
to Utrecht together with fine ceramics on one of the annual Dutch shipments from De-
shima, but it is also possible that it was sent by Gordon from London. Whatever its true
pedigree, the Utrecht tree is a living monument to early Dutch contact with Japan and
reminder of the key role of both countries in the introduction of plants into the world’s
gardens.5
In Britain it is part of garden folklore that the oldest ginkgo at Kew was among a
select group of five specimens, the so-called Old Lions, introduced from the estate of
the third Duke of Argyll, who had extensive nurseries, not far along the Thames at
Whitton, near Twickenham. According to the story, when the duke died in 1761, his
nephew John Stuart, the third Earl of Bute, moved some of the best young trees to the
Royal Estate at Kew, bringing them down the Thames by barge.6
The reality is less romantic. Among the Old Lions, only a black locust and a plane
have documented links with the Argyll Estate. The big Japanese elm, planted in 1760, is
unlikely to have been acquired by Princess Augusta because it sits beyond the bound-
aries of the original arboretum. One of the others, the Pagoda Tree, is thought to have
come to Kew in 1760 from the nursery of James Gordon, who introduced the tree into
Britain in 1753. Given that Gordon was also an early source of ginkgo in Europe, it
seems likely that he was also the source of the Old Lion ginkgo at Kew. It might be that
both the Pagoda Tree and ginkgo were part of the same batch of seeds that Gordon
somehow received, probably via John Ellis, from China or Japan.7
Early in the second half of the eighteenth century in Europe, ginkgo was already
being propagated from cuttings and passed from garden to garden. In 1785 a ginkgo
was planted in the Hortus Botanicus at Leiden, the oldest botanical garden in the
Netherlands. In 1787 Giorgio Santi, director of the Botanical Garden of the University
of Pisa, planted a ginkgo in the new arboretum, which still stands today just a five-
minute walk from the Leaning Tower. Conrad Loddiges, who ran a prominent English
nursery specializing in exotics, supplied a plant to the royal estate of Schönbrunn just
outside Vienna in 1781, and this may also have been the source of the old male tree in
the botanical garden in Vienna.8
Ginkgo first found its way to France in the late 1770s. Loudon cites the planting of a
specimen in Rouen in 1776. Translating the account given by the French botanist André
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Thouin, he also gives an account of the eccentric introduction of ginkgo into the gar-
dens of Paris, which also explains the origin of the unusual name of the tree in French,
l’arbre aux quarante écus:

In 1780, a Parisian amateur, named Pétigny, made a voyage to London, in order

to see the principal gardens; and among the number of those that he visited
was that of a commercial gardener, who possessed five young plants of Ginkgo
biloba, which was still rare in England, and which the gardener pretended that
he then alone possessed. These five plants were raised from nuts that he had re-
ceived from Japan; and he set a high price on them. However, after an abundant
déjeuné, and plenty of wine, he sold to M. Pétigny these young trees of Ginkgo,
all growing in the same pot, for 25 guineas, which the Parisian amateur paid
immediately, and lost no time in taking away his valuable acquisition. Next
morning, the effects of the wine being dissipated, the English gardener sought
out his customer, and offered him 25 guineas for one plant of the five that he
had sold the day before. This however was refused by M. Pétigny, who carried
the plants to France; and as each of the five had cost him about 120 francs, or
40 crowns (quarante écus) this was the origin of the name applied to this tree
in France, of “arbre aux quarante écus”; and not because it was originally sold
for 120 francs a plant.9

According to André Thouin, almost all of the early ginkgo trees in France were
propagated from these five plants, one of which was given to the Jardin de Plantes in
Paris. There are, however, other well-documented ginkgo introductions, for example,
the ginkgo sent by Sir Joseph Banks to Pierre Marie Auguste Broussonet in 1788, which
probably came from Gordon’s stock. Broussonet gave the plant to Antoine Goüan, who
planted it at the Jardin des Plantes in Montpellier.10
Around the turn of the eighteenth century the first generation of ginkgos planted
in Europe began to mature; in horticultural vernacular they began to “flower,” even
though ginkgo has nothing resembling a flower in the normal sense of that word. The
first is thought to have been the Old Lion at Kew, which produced pollen cones in 1795.
Male trees also “flowered” in Pisa in 1807, and at Montpellier and Rouen in 1812. When
a female ginkgo was eventually recognized in Europe in 1814, perhaps sixty years after
the first introductions, it created a stir.11
The botanist Augustin Pyramus de Candolle from the Geneva Botanical Garden was
the first to notice the female reproductive shoots on the ginkgo at the Bourdigny Estate
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gardens

just outside Geneva, but in the absence of males the seeds did not mature. However,
cuttings from the female tree were circulated to botanical gardens across Europe and
grafted onto many male trees, including in Strasbourg and Kew. The first viable seeds
produced in Europe were borne on a male tree in the Montpellier Botanic Garden in
1835 that had been “covered . . . with grafts.”12
Ginkgo made the leap across the Atlantic to the New World with the help of William
Hamilton, a wealthy late-eighteenth-century botanist, plant collector, and landscape
designer. Inspired by a year spent in England, Hamilton wrote to Thomas Parke on
September 24, 1785: “The verdure of England is its greatest beauty and my endeavours
shall not be wanting to give the Woodlands some semblance of it. . . . Having observed
with attention the nature, variety and extent of the plantations of shrubs, trees and
fruits and consequently admired them, I shall . . . endeavour to make it smile in the
same useful and beautiful manner.”13
His efforts resulted in the Hamilton Woodlands, a country estate on the Schuylkill
River south of Philadelphia, one of the first American landscape gardens designed in
the “natural” style of English estates of the time. Groves of native and exotic trees were
set among pathways, meandering streams, and carefully maintained lawns. We can
thank Hamilton for several familiar trees that he introduced into North America; the
elegant Lombardy poplar, the tough Norway maple, and, less happily, the aggressively
invasive tree of heaven. In 1784 Hamilton also cultivated the first ginkgo trees to grow
on North American soil in several million years.
As the winter of 1785 approached, Hamilton wrote to his private secretary somewhat
anxiously on November 2: “Secure the tender plants from the severe weather, otherwise
all my pains will have been to no purpose. The Cistus’s, the Heaths, eleagnus, Ginkgo,
Laurus’s, Tamarisks, Yucca glorioso, the Carolina mahogany, Zantoxylon sempervirens
&c, should be secured by skreens of Dry straw or some other means, but by [no] means
let dung be put to their roots for it will inevitably kill them.”14
Hamilton’s ginkgo was one of several acquired from England, perhaps through a
connection to Peter Collinson, a British scientist, gardener, and fellow of the Royal
Society. Through Hamilton’s social connections, young ginkgo plants found their way
to the gardens of some of the most prominent early American botanists. He gave one
to his cousin and neighbor the prominent early American naturalist John Bartram,
also a correspondent and friend of Collinson’s. This male tree, which still survives in
Bartram’s Garden just a few miles from the former Woodlands estate, is now the oldest
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living ginkgo in North America. Another of Hamilton’s cuttings went to the botanist
and physician David Hosack. This is probably the ultimate source of the large ginkgo
on the Vanderbilt Mansion Estate at Hyde Park that Hosack acquired in 1828.15
Within a few years of the arrival of ginkgo in the northeastern United States, André
Michaux, a diplomat and botanist to the King of France, also introduced the tree into
the southern United States. Having first attempted to establish a botanical garden in
New Jersey, where the climate proved too harsh, Michaux moved to Charleston, South
Carolina, in 1787 and developed what became known as the French Botanical Garden.
Charleston’s agreeable climate allowed him to cultivate and tend his plants year round,
and it was here that he introduced many Old World plants to America, including the
mimosa or silk tree, crepe myrtle, and camellia, and planted many others, including
ginkgo. With the interruption of the French Revolution the garden was abandoned, but
Michaux’s son François, returning in 1802, recalled: “I found in this garden a superb
collection of trees and plants that had survived almost total neglect . . . some of which
were in the most flourishing state. I principally remarked two Ginkgo biloba, that had
been planted about seven years, and which were then upward of thirty feet in height.”16
Besides their interest in gardens and exotic plants, pioneering ginkgo growers in
the United States shared another attribute: political prestige. André Michaux moved
in the circles of Thomas Jefferson, Benjamin Franklin, and George Washington. John
Bartram had cofounded the American Philosophical Society with Franklin in 1743.
Peter Collinson had frequently exchanged correspondence with Franklin about elec-
tricity and other matters, and supported the American Philosophical Society with gifts
of books. David Hosack was the founder and first president of the New York Horti-
cultural Society, the first horticultural organization in America, in which John Adams,
Thomas Jefferson, and James Madison were honorary members. On the Fourth of July
in 1788, the official acceptance of the United States Constitution was celebrated with a
parade that ended with seventeen thousand of the young country’s citizens picnicking
at the Woodlands.17
Given the company kept by some of the early ginkgo growers in the United States,
it is perhaps not surprising that another prominent politician, Henry Clay, probably
played an important role in the early cultivation of ginkgo through the American
South. Several stately ginkgos that grow today in Kentucky are rumored to have ar-
rived in Washington from Japan as seedlings and were subsequently sent by Clay to
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gardens

Kentucky. Attempts to document this connection have so far failed, but a group of
large and undoubtedly early ginkgos survive on Clay’s former estate, namesakes for
the estate’s Ginkgo Tree Cafe. Two such trees also grow on the grounds of the former
Kentucky Military Institute. Magnificent large ginkgos at Cave Hill Cemetery in Louis-
ville, the former Cave Hill Farm, seem also to be of similar age, and they too may have
connections to Henry Clay.18
Whether or not the ginkgos planted at the Kentucky Military Institute came from
Henry Clay, they are significant as the first to bear seeds in the United States. Nurseries
had distributed ginkgo as cuttings since 1810, but the availability of abundant seeds al-
lowed ginkgo to be cultivated more widely. The Arnold Arboretum at Harvard received
seeds from Kentucky on January 7, 1878. In 1890 William R. Smith, the curator of the
United States Botanic Gardens in Washington, reported that the Kentucky female was
the garden’s “chief source of supply” of ginkgo seed: “Up to a recent date the Ginkgo
was a very rare tree, nurserymen asking £1 apiece for them. The first Japanese Embassy
brought over some seeds and presented them to the Botanic Garden and the trees
raised from those seeds are now bearing seed.”19
By the late nineteenth century, ginkgo was well established in the preeminent bo-
tanical gardens across the United States. In 1859 the Missouri Botanical Garden was
founded by Henry Shaw, a British merchant from Sheffield; he planted several ginkgos
that are now large trees in his magnificent garden. The New York Botanical Garden,
founded in 1891, in the Bronx in New York, boasts massive old ginkgos growing near
the Botanical Garden station, which was once part of the garden’s estate. The largest,
planted in 1898, is seventy-seven feet tall, with a crown spreading to fifty-seven feet.
The garden’s records indicate that by the 1890s ginkgo was readily acquired as small
trees from nurseries.
Further enthusiasm for planting ginkgo came with the new excitement about its
evolutionary significance following Hirase’s discovery in 1896. As an icon of turn-of-
the-century botanical science, it was planted on many university campuses. At the
University of California, Berkeley, the elegant old ginkgo that stands next to Giannini
Hall dates from around this time and is one of the most beloved trees on campus. Its
rare display of autumn color among the otherwise dominant California evergreens has
inspired poetry from professors with offices that face it. At the City College campus of
the City University of New York, one of the most diverse colleges in America, students
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A carefully pruned espaliered ginkgo growing on the wall of the Department

of Plant Sciences at Cambridge University.

complain about the smelly seeds of several old ginkgos in more than ninety languages.
At Cambridge University, a carefully tended and unusual espaliered ginkgo cloaks the
south side of the Plant Science Building on the Downing Street Site.20
The large female ginkgo that stands beside Botany Pond at the University of Chicago
was planted almost at the founding of the university and is embedded in the memory
of thousands of University of Chicago students and staff. A recent lighthearted com-
petition among University of Chicago alumni to find the best haiku resulted in many
entries about Botany Pond and its towering ginkgo. This was one of the best:

Rustling gold ginkgo,

Languid koi circling below
In Botany Pond.21
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gardens

As the meaning of ginkgo has changed in gardens of the world, so too has the plant
itself. In the wild, new variants arising by mutation or unusual genetic combinations
are quickly eliminated by natural selection. However, such novelties—for example,
with an unusual growth habit or strange leaves—are easily perpetuated in gardens
using cuttings or grafts. In this way, astonishing diversity has emerged from the gene
pool of those few native ginkgo populations that survived into historical times in
China. Today more than 220 different horticultural cultivars have been documented,
and at least 28 cultivars are distinguished based on the details of their nuts.22
A landscape gardener recommended Princeton Sentry for the limited space in our
own front garden in Oak Park. Tall and slender varieties like Mayfield and Tremonia
can be also squeezed into narrow spaces. The varieties Golden Globe and Globosa have
crowns as round and full as an apple tree’s. Fastigiata, with its pyramidal form, looks
a bit like a Christmas tree, while varieties like Pendula, Umbrella, and Horizontalis
sprawl and weep like willows and Japanese maples. Mariken and Troll are dense, squat,
and shrublike, barely resembling trees at all. Also much sought after are cultivars with
unusual leaves. Saratoga, named for the Saratoga Horticultural Research Foundation
in California, where the variety was developed, has drooping, triangular leaves with
frayed edges like a fishtail palm. Variegata has striking variegated leaves with white
streaks. Tubifolia has leaves that are almost entirely fused into trumpetlike funnels.
Ginkgo bonsai are also in high demand. They have the same dramatic fall color and
abrupt leaf fall seen in full-size ginkgo trees, and are especially prized for these reasons.
Even more coveted are chi-chi ginkgo bonsai, which begin life as cuttings taken from
the descending woody chi-chi of old ginkgo trees. Planted upside down, chi-chi bonsai
have an unusual conical form: the stalactite becomes a stalagmite, but with roots and
protruding branches. Since there are few ginkgos outside of Asia that are old enough
to produce chi-chi, these especially strange bonsai are available in the West only as im-
ports and at great expense.23
 31
Nuts

Sweetest nut hath sourest rind.

—William Shakespeare, As You Like It

Long before it became popular in gardens, or as a memory-enhancing supplement,

ginkgo was valued for its edible nuts. With a plump, soft, partly creamy, partly waxy
white “meat” not much bigger than a peanut, the ginkgo nut has a taste that has been
variously described as like “mild Swiss cheese,” “pine nuts,” “potatoes crossed with
sweet chestnuts,” “green pea crossed with Limburger cheese,” or just “fishy.” Today, de-
spite their enigmatic flavor, ginkgo nuts are common in the cuisines of China, Japan,
and Korea, and they can be bought almost everywhere that people from those parts of
the world have settled.1
It is possible that people have collected ginkgo nuts from wild trees for thousands of
years. However, the first written accounts of ginkgo and its value as a nut tree appear
much later. During the Song Dynasty, in the early eleventh century, when the poet and
historian Ouyang Hsiu presented his friend Mei Yao-Chēn with ginkgo nuts, he was
prophetic about how the association between ginkgo and people was likely to grow:

In the past Chang Chien (second century B.C.) introduced grape and pome-
granate (from central Asia). We can imagine that when these first came, they

226
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nuts

must have been similarly highly valued as these nuts. But now these plants are
common all over China, growing along fences and walls. The very things are
still the same, but human nature changes in time. Someone should record the
beginning so that future generations can know its origin. This is thus not only
continuing your verse, but also contributing to history.2

Chun Chu Chi Wen, another early source, describes four large trees in Kaifeng, the
ancient capital of China in eastern Henan Province, which produced several bushels
of ginkgo nuts each year. A big female ginkgo can produce seeds in huge quantities,
and such bounty may have been what first attracted the attention of people. Peter
Del Tredici mentions a tree a hundred feet tall in Yang Tang village, Zhejiang Prov-
ince, China, which produced more than eight hundred pounds of cleaned nuts in a
single season. Today, production of ginkgo nuts is focused in ginkgo orchards of much
smaller trees. In China production extends to more than twenty provinces and cities.
By one estimate, there may be as many as 800,000 trees in China that produce an aver-
age of up to seven thousand tons of dried nuts each year.3
Collecting nuts for food is something that we inherited from our nonhuman ances-
tors. Nuts provide proteins, carbohydrates, vitamins, and minerals, and in the hands of
an intelligent primate the hard shell makes them easy to keep and transport. Some of
the earliest human tools may have been used to free the meats of nuts from their hard
shells. At the same time, both consciously and unwittingly, people have spread nut-
producing plants everywhere they have gone. The Old Testament tells us that Joseph’s
brothers carried pistachios down to Egypt. In northern Spain, the expansion of beech-
dominated forests after the last glaciation seems to have followed human settlement
and the use of beech nuts in animal husbandry. Native Americans may have used fire
to maintain and increase populations of important nut trees like chestnut, hickory,
and oak, and after the first European encounters with the New World, peanuts quickly
found their way to China, now the world’s largest peanut producer. Ginkgo is just one
success story among many of the nut-people symbioses.4
In strict botanical parlance, a nut is a hard-walled fruit with a single seed inside,
a definition that does not apply to ginkgo. Ginkgo is different; its hard shell is part
of the seed rather than the fruit wall. However, the function of the “meat” in ginkgo
seeds and the true nuts of other plants is the same: to provide food for the embry-
onic plant. Packed with carbohydrates, fats, and proteins, the “meat” helps sustain the
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young seedling as it develops until it can become self-sufficient. Ginkgo nuts are rich
in starch and protein but low in fats. Compared to pine nuts, for example, ginkgo nuts
contain about one-twentieth of the fat and a third of the calories. They also contain
about 6 percent sucrose, which may account for the faint sweetness in their flavor.5
One of the Chinese names for ginkgo is kung sun shu, meaning “grandfather-
grandchild tree.” The name alludes to the patience needed to grow ginkgo from seed:
the tree you plant today yields nuts in your grandchildren’s generation. However, the
time from seed to seed is generally much shorter, usually about twenty to thirty years.
Even so, commercial growers are not prepared to wait. Grafting gets the trees into pro-
duction earlier, often after only about ten years. It also brings the advantage that grafts
can be chosen from trees known to produce large nuts.
Most of the ginkgo nuts consumed in China, Japan, and Korea are grown in com-
mercial orchards often operated by small holders using methods that have changed
relatively little over hundreds of years. In Japan, the area around Sobue, west of Na-
goya in central Honshu, is a traditional region for ginkgo nut production. I visited on
a bright, cold, early spring day in 2007. Around Sobue the land is flat, low lying, and
often wet underfoot, with scattered houses, rice fields, and small farms; everywhere
there are ginkgo trees.
The life of a ginkgo tree in the nut orchards around Sobue begins with a three- or
four-year-old rootstock grown from seed, onto which several branches from female
trees are grafted, usually in March and early April, just before the leaves appear. A
useful by-product of grafting is that when normal branches from “mother trees” are
grafted to the rootstock, they somehow “remember” how they were growing and con-
tinue to grow in a more or less horizontal fashion. They seem to have lost their ambi-
tion to produce the strong upward growth of a normal ginkgo tree.
This phenomenon, a so-called topophytic effect, is well known to horticulturalists
and has been exploited in some species of conifer to produce the low-growing pros-
trate forms that are useful for gardeners. In ginkgo, the effect is to accentuate the nor-
mally spiky form of a young tree and produce the characteristic shape seen in ginkgo
orchards: small trees with low, spreading branches. Such trees would be very inconve-
nient if planted along narrow streets where there is little space, but in a ginkgo orchard
keeping the trees small and the branches low makes for easier harvesting.
Around Sobue even the largest ginkgo orchards consist of only a few hundred trees,
and most of the seeds are produced by small-scale farmers who tend a few dozen trees
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on small plots of land close to their homes. Trees in orchards are pruned more regu-
larly and kept in a uniform shape. Most are no more than about twenty or thirty feet
tall, with a crown of spiky branches from the top of the trunk six to ten feet above the
ground. Each tree is carefully tended and is often perched on a low mound of fertilizer
and soil that is covered by a mat of straw to keep the fertilizer in, the weeds down, and
the roots moist.
Whether the production of ginkgo nuts is on a small or large scale, the objective is
the same: to produce good yields of the large nuts that command the best prices. In
both China and Japan female trees that produce larger than normal seeds are much
prized and often allowed to grow to reach their massive natural size. Over several cen-
turies selection for larger nuts has produced a modest increase in size. Nuts from the
Chinese cultivar King of Donting Mountain are typically a little more than an inch
long, while those from a normal female ginkgo are two-thirds that size. Many cultivars
are distinguished by the size and shape of the nuts.6
Nuts start to ripen on the trees in August and can then be collected by hand. Usually,
though, they are left longer to fill out, to give the shell more time to harden and to help
ensure that the nuts are not damaged during harvesting. Timing is critical; the nuts
cannot be left too long: they need to be gathered before the young embryo inside grows
too much.
Traditionally, after collection, the seeds are buried to encourage decay of the smelly
flesh, then dug up, washed, dried in the sun, and made ready for sale. Alternatively the
seeds are collected in a bucket of water, where the pulp is allowed to partially rot until
it can be rubbed off in changes of water until all of the flesh is gone. However, growers
wanting to get paid more quickly are even more direct. In the countryside around
Sobue, seeds are loaded into small vats that are stirred by large screwlike blades that
help break up and remove the fleshy seed coat. The putrid flesh is then washed off as
a gut-wrenchingly malodorous slurry. Active depulping needs to be done carefully, to
avoid damaging the shells and squirting the noxious juices.
When the flesh is gone and the shell is exposed, the nuts are sorted by size, dried
quickly in the sun, and sent on the way to the packers and distributors the next day. The
nuts cannot be kept. Often the shell is removed and the “meats” are vacuum-packed
before they find their way into the shops. More traditionally, ginkgo nuts are air-dried
for one or two weeks, stored in a cool place, and then shipped to market, sometimes
after having been roasted.7
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The annual ginkgo harvest is also carried on every autumn by Asian families in
the West, in streets and parks from Montreal to San Francisco. In New York City, a
Harlem homeowner speaks of a new and unwelcome tradition: waking in early Novem-
ber to find people climbing the gangly ginkgo in front of her home, knocking down
seeds with long poles. Other New Yorkers encounter the seed collectors under the old
ginkgos that accent the northern boundary of Central Park. These foraged urban seeds
mainly find their way onto local dinner tables rather than to the markets of China-
town.8
Recently, a new group of ginkgo gatherers has appeared at the autumn harvests.
Motivated by an ethos of eating local, seasonal food, intrigued urban “locavores” are
on the lookout for new experiences, and recount their exploits in magazines, on food
blogs, and over dinner parties. Sara Crosby, a writer for Gourmet magazine, went door
to door among Chinatown restaurants toting a reeking bag of uncleaned ginkgo seeds,
seeking advice on their preparation. After attempts at boiling, salting, and roasting all
produced a snack she found too bitter to be palatable, her southern instincts came to
the fore and she deep-fried them: ginkgo seeds, like most things, taste good that way.
For curious cooks unwilling to delve into putrid pulp for the full experience, there is
also the option of visiting the local Chinatown. A pound of plump, bleached, cream-
colored bai guo sold in bulk, or a vacuum-sealed package of cooked and shelled “semen
ginkgo,” can be had for under two dollars.9
Interest in ginkgo nuts has mainly centered on their value as a food, but they have
also been used in ritual and medicine. In sixteenth-century China, the nuts were often
used at weddings and feasts, sometimes dyed red and substituted for lotus seeds. In
traditional Chinese medicine raw ginkgo nuts are used to “send down adversely rising
chi, remove toxic substances, and destroy parasites.” Generally, however, it is not ad-
visable to eat the nuts raw. They contain toxins that are partly broken down by heat, so
ginkgo nuts are generally eaten only after cooking. Even then, they should not be eaten
in quantity like almonds or peanuts. Cooked inside the shell, they emerge slightly yel-
low in color, with a waxy texture, and a slightly bitter taste.10
As Engelbert Kaempfer noted in the History of Japan, ginkgo nuts have also been
used in Japan to produce oil. They may also have been used in this way in China. In an-
cient times plant oils provided an important alternative to animal fats for lighting and
cooking, and this may have encouraged widespread ginkgo cultivation. The modern
use of mineral oils and the introduction of gas and electricity for lighting and cooking
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Ginkgo nuts baked in their shells at a banquet in Nanjing, China.

have pushed this use of ginkgo aside. Nevertheless, a long list of edible oils from nuts
such as almond, cashew, hazel, peanut, pine nut, and walnut are still relatively easily
obtained for various culinary, manufacturing, and medicinal uses. The fact that ginkgo
oil is not among them may suggest that in the time of Kaempfer it was used mainly for
nonculinary purposes.11
Culinary uses of ginkgo nuts in Asian cuisines vary by region and occasion; the nuts
may appear in savory dishes and desserts, in formal banquets, and in day-to-day fare.
In China, they may be enjoyed in many traditional dishes, boiled into soups, fried with
celery and lily bulbs, or served in sweet dessert soups along with Chinese dates. At a
dinner in Singapore not long ago, the dessert menu offered the options of “Teochow
yam paste with gingko nut” and “Chilled snow fungus with lotus seed and gingko nut.”
The spelling was not quite right, but the taste for these nuts has evidently followed the
Chinese diaspora to places well outside where ginkgo trees can be grown. Most often I
have come across ginkgo nuts still in their shell, just baked on a tray or roasted in the
oven in tinfoil. This is the easiest way to prepare them, but thorough cleaning of the
seed is vital: as with poison ivy, sensitive people can have an allergic reaction if they are
caught in the smoke.
In Japan, as in China, the culinary uses of ginkgo nuts were recorded soon after the
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Vendor selling roasted ginkgo nuts at the Tsurugaoka

Hachiman-gū Shrine, Kamakura, Japan.

first written records of the plant itself. Ginkgo nuts are listed as a fruit or as tea cakes in
Shinsen Ruiju Orai, a text written sometime between 1492 and 1521, and ginkgo nuts are
also mentioned in tea ceremony records written between 1533 and 1596. In the Miyoshi-
tei Onariki from 1561 ginkgo nuts are listed as sweets and a dessert on the menu of a
meal served to Shogun Ashikaga Yoshiteru and his followers.12
Kaempfer, Thunberg, and Siebold may all have eaten ginkgo nuts during their time
at Deshima, perhaps in traditional dishes little different from those of today. Edward
Morse, a Harvard archaeologist and naturalist who was among the first Westerners to
teach at Tokyo University in the late 1870s, considered himself an enthusiast of tradi-
tional Japanese food, but he remarked after a dinner at a Japanese tea house: “There
were many things I tasted for the first time. The bulb or root of the lily was an excellent
substitute for the potato; there were a number of water plants similar to the watercress;
a preparation of fish, like macaroni; the nut of the gingko tree, which I did not like, and
a preparation of tea, which I did.”13
In Japan, as early as 1785, a book on traditional Japanese cuisine entitled Kaiseki-
ryori-cho mentions ginkgo nuts as a side dish for sake drinkers, a tradition that con-
tinues. It is sometimes claimed that ginkgo seeds help prevent drunkenness and are an
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effective hangover cure. Research provides just a hint that such optimism is not entirely
wishful thinking. An enzyme in the seeds seems to speed the breakdown of alcohol.
In one study, laboratory animals were given enough alcohol to get them nicely inebri-
ated: those fed ginkgo nuts beforehand were able to better clear the alcohol from their
blood.14
Perhaps the best way to eat ginkgo nuts is out beneath the trees themselves; roasted
ginkgo nuts are often sold by street vendors, like traditional European or North Ameri-
can chestnuts. To me, they taste a little like chestnuts, too. I saw them for sale one sunny
spring weekend at the Tsurugaoka Hachiman-gū shrine in Kamakura not far from
Tokyo. The brightly colored open-air stall advertised them with a big sign in Japanese,
“delicious.” The vendor was roasting them in a large wok over a gas burner, selling
them in small bags and calling out in Japanese to the visitors to the shrine, “Freshly
roasted ginkgo nuts, easy to eat, ready to eat.” However, extracting the meat from the
shells of roasted seeds can be a trial for heat-sensitive fingers. At home, my Japanese
friends crack open the shells with pliers.15
On my visit to Sobue, we had lunch at a restaurant specializing in the use of ginkgo
in Japanese cuisine. Our guides were enthusiastic for us to sample the local produce.
There were ginkgo nuts in just about every dish of our traditional Japanese lunch.
They appeared as a garnish with sashimi, for example, on a plate of finely sliced octo-
pus. In nabe-ryori they were boiled with vegetables, fish, and meat in a hot miso-like
soup. They were also deep fried in the tempura batter around shrimp. However, most
memorable was the chawan mushi, a savory, pale yellow, custardlike dish with one or
two ginkgo seeds at the bottom. This steamed, lightly set egg soup usually containing
pieces of chicken, fish, or vegetables is one of the most common places to encounter
ginkgo nuts. They are similar in color to the soup itself, and their flavor, like the flavor
of the whole dish, is delicate and subtle: not quite sweet, but not fully savory either.
This typical Japanese dish, like many others from Asia, extends the normal experience
of the Western palate. Like the ginkgo tree itself, the versatile yet mysterious nuts con-
tinue to serve up more questions than answers.
 32
Streets

I had to compete. In the concrete, in the jungle.

—Tower of Power, “Back on the Streets Again”

Ginkgo is among the most widely planted street trees in the world. With most of the
world’s people now living in cities, it is seen by millions of people every day. Along with
other street trees, ginkgo has a role to play in sustaining connections between people
and the natural world. In the United States alone, the trees growing in backyards,
streets, parks, and the urban reserves comprise about 74.4 billion trees that account
for about 8 percent of the total national tree canopy. These trees are important in the
lives of those three-quarters of Americans who work and live in metropolitan areas.1
Interest in planting trees in cities has never been higher. The United Nations En-
vironment Programme’s Plant for the Planet: The Billion Tree Campaign in only five
months secured pledges for a billion new trees globally. Two years after the program
was launched in 2006, more than 1.8 billion trees had already been planted. In Lon-
don the British charity Trees for Cities organizes its work around five themes: Trees
for Food, Trees for Learning, Trees for Play, Trees for Streets, and Trees for Volunteer-
ing. It has planted nearly a quarter of a million trees around the world in the past few
years. Initiatives to plant millions of new trees have taken off in New York, Los Angeles,

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Memphis, Miami, Denver, and Philadelphia. Delhi has a target to increase its tree cover
by 13 percent. These large-scale efforts have been aided by the simple fact that planting
trees can be a good investment. They make people feel good about their environment,
and within a few years economic and other returns more than make up for the costs of
their planting and maintenance.2
Trees in cities contribute to floodwater reduction, temperature moderation, pollu-
tion abatement, energy savings, and improved property values. The crown of a mature
tree can intercept as much as 100,000 gallons of rainfall each year, allowing much of it
to evaporate before it flows into overloaded storm drains. Trees also take up substan-
tial amounts of water through their roots and moderate temperature by breaking up
and shading concrete and asphalt surfaces that absorb and intensify heat. Urban heat
islands may have temperatures up to 10 degrees higher than places nearby. A well-
placed tree that provides shade in the summer and shelter in the winter can reduce
household energy costs by a third. In front of a home, a single tree can increase prop-
erty values by about 6 percent. Studies also show that we are more likely to linger, and
open our wallets, in business districts where the hard landscape and hard selling have
been softened by trees.3
At the USDA Forest Service Center for Urban Forest Research, Greg McPherson and
his team quantify the benefits of urban trees with their computerized valuation pro-
gram i-Tree Streets. Using this tool, property owners, urban forestry groups, and city
governments can quantify the benefits and cost savings that trees provide. The pro-
gram uses data on the different kinds of trees and their sizes, together with local and
regional information on climate, species growth rates, property values, energy prices,
water prices, air pollutant emissions, stormwater costs, and costs of tree maintenance.4
Results from i-Tree show that the economic performance of ginkgo as a street tree
varies substantially according to location. In Minneapolis, the average annual value of
each of the city’s 5,002 ginkgo trees at the time of inventory was $11.52. In San Fran-
cisco the savings were double that, at $23, reflecting differences in property values and
costs of energy and water. With 16,184 ginkgo trees at the time of the i-Tree survey, not
only are the ginkgos in New York City more numerous than anywhere else in the coun-
try, but they are also more valuable. Even so, $82 a tree is low compared to the New
York City street tree average of $209. Ginkgo’s lower value reflects in part a “penalty”
for its relatively modest leaf surface area compared with other trees of similar height
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and spread. On the positive side, ginkgo scores high for its aesthetic value; people like
it. Greg McPherson, the developer of i-Tree Streets, was quick to confess, “Ginkgo is
my favorite city tree.”5
Street trees also help us in ways that do not translate easily into monetary values.
Neighborhood trees increase the time that children spend playing outdoors and the
amount of supervision they receive. Parents and children spending more time out-
doors helps to strengthen community bonds and reduce crime through improved vigi-
lance. Tree-lined streets also help decrease road rage and improve the attention of
drivers. By creating the illusion of a narrower street, trees prompt drivers to drive more
slowly. Trees and nature generally also affect us psychologically in ways we can’t always
describe or explain. In one study, patients whose hospital room looked out on to a view
of trees and water had shorter stays and more positive evaluations and needed fewer
painkillers than a group of similar patients with windows facing a brick wall.6
In New Haven, Connecticut, the Urban Resources Institute, a nonprofit university-
community partnership based in the Yale School of Forestry and Environmental Studies
has worked for more than a decade to foster community-based land stewardship, en-
vironmental education, and urban forestry. Its work has touched every neighborhood
in New Haven and has engaged thousands of residents, including unemployed ex-
convicts and proud homeowners, most of whom might previously never have given
trees a second thought.7
The scale of current campaigns for urban forestry is unprecedented, but the idea
that trees are important for healthy cities has a long history. More than three thousand
years ago in Egypt, Ramses III had trees planted along the streets for promenading
and recreation, and in ancient Greece trees were planted to beautify cities and shade
the pathways leading to market. In the sixteenth century, as the influence of the Italian
Renaissance spread across Europe, the idea of a garden allée was widely embraced and
soon moved into the city. In 1615 Amsterdam became the first city to formally incor-
porate buildings, transportation, and trees in its Plan of Three Canals, and King Louis
XIV, in 1670, ordered that the walls around Paris be destroyed and replaced with new
tree-lined boulevards for pedestrians and carriages.8
Aside from their aesthetic and recreational roles, urban trees were also thought to
cleanse the city air and prevent the “miasmas” that were believed to cause illnesses and
disease. The first public greening campaign in the United States was in Philadelphia,
which passed an ordinance in 1700 that every owner of a house “should plant one or
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more trees before the door that the town may be well shaded from the violence of the
sun . . . and thereby be rendered more healthy.” In 1732 the Assembly decreed that
“walks may be laid out and trees planted to render [the town] more beautiful and com-
modious.” In 1792 citizens of Philadelphia petitioned to have trees planted in public
areas because “It is an established fact that trees and vegetation . . . contribute to . . .
the increased salubrity of the air.” In 1872 in New York, the city commissioner of health
declared that street trees should be planted to mitigate heat and reduce the death rate
among infants.9
By 1773 Savannah already had a blueprint for avenues lined with live oaks, and in
1791, when George Washington appointed Major Pierre L’Enfant to design the capi-
tal city that would bear the first president’s name, tree-lined avenues were planted
throughout the city. In the early 1870s ninety ginkgo trees were planted as an avenue
on the grounds leading to the Department of Agriculture, one of the first major uses of
ginkgo as a street tree. A 1929 article gave the D.C. ginkgo trees a hearty endorsement
that foreshadowed their broad acceptance in cities elsewhere:

Visitors to Washington, D.C., are always much impressed with the beauty of the
avenues of Ginkgo trees that line the approaches to the Department of Agricul-
ture and that ornament the city in many other places. There is no good reason
why Washington should be the only city in the country especially favored with
this famous tree, sacred to the Chinese and Japanese and grown for centuries
in their temple courts. It does very well in all parts of the United States where
the winters are not too severe and can at least survive as far northwest as cen-
tral Iowa. . . . A great virtue of the Ginkgo is the almost complete freedom from
the fungus disease and insect pests that bedevil practically all of our other orna-
mental trees.10

One problem with the use of ginkgo as a street tree in Europe and North America
is the disagreeable smell of the seeds. I have heard of several urban homeowners who
have taken matters into their own hands and tried to kill unwanted female ginkgos on
streets outside their homes. Every year arborists in Washington, D.C., spray large quan-
tities of the herbicide chlorpropham on female trees to prevent them from seeding.11
Today, reputable nurseries circumvent the problem of smelly female ginkgos by
selling only male trees propagated as cuttings. In New York City, the Department of
Parks and Recreation, as a formal policy, hasn’t planted a female ginkgo tree in twenty
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Three young ginkgo trees among

the many thousands growing on the
streets of Seoul, South Korea.

years. Ordering trees from nurseries that propagate only cloned males is one way to
ensure that the gender ban is enforced, but older trees that were planted before such
regulations and that now are grandfathered into city tree heritage laws complicate the
removal of healthy old female trees. Moreover, females often sneak back, sometimes
through the planting efforts of less cautious residents and sometimes with approved
permits.12
However the issue of female trees is handled, ginkgos remain common in cities
from Beijing to Berlin. Often they are a defining feature of the streetscape. In London
ginkgos are among the plantings around the Tower of London; on the other side of the
city they accent the streetscape near Imperial College and outside the nearby Natural
History Museum. In Manhattan, ginkgo accounts for 10 percent of the urban forest and
is the third-most-common of all street trees. Whether on the streets of Chelsea, on Fifth
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streets

A young ginkgo being nursed back

to health after having been
transplanted during the renovation
of the streetscape in Sejongo, Seoul,
South Korea.

Avenue, along the northern border of Central Park, or through Harlem and Washing-
ton Heights, the spiky forms and distinctive leaves of ginkgos are ubiquitous.13
In Japan, ginkgo accounts for around 11 percent of all street trees and is the most
widely planted tree in the country. Its use was nurtured there in the late nineteenth and
early twentieth century as the country underwent rapid modernization. In Tokyo in
particular, a 1907 plan focused on ten fast-growing and resilient trees, of which ginkgo
was one. After the fires from the Great Kanto Earthquake in 1923 and the bombing of
the Second World War, which destroyed close to half of the 270,000 street trees planted
along the roads of Japan, city planners again turned to ginkgo to enliven and soften
their streets. Today, more than half a million ginkgo trees are planted along Japanese
roadsides.14
The same attributes that have allowed ginkgo to survive for thousands of millennia
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may have also contributed to its success as a hardy and resilient street tree. The lives
of street trees are “nasty, brutish and short”: on average, they survive a mere seven to
thirteen years, compared with the sixty years the same species might expect to enjoy
in a park and the hundreds of years it might live in its native forest. Deborah Gang-
loff, executive director of San Francisco’s nonprofit group American Forests, explained
some of the reasons why: “They’re stuck in a concrete box, get bikes chained to them,
with dogs relieving themselves and cars hitting them. . . . It’s a hard life.” From the salt
that attacks their roots in winter to the ozone that assaults their leaves in summer, the
constant barrage of chemicals faced by street trees is well beyond the level of abuse that
evolution designed them to endure.15
Street trees also face challenges belowground. Urban soils are often made of land-
fill rubble, building material, and other contaminants. Such soils are highly variable
in nutrient content, low in organic matter and fertility, and generally lacking in soil
microorganisms needed for healthy plant growth. The ginkgo trees wedged into the
pavement of my old neighborhood in the South Loop area of Chicago were pushing
their roots through the remains of the old railway tracks that once entered Dearborn
Station, but they did well nevertheless.16
When you next see a large street tree with an elegant crown, try to imagine what it
looks like underground. An average tree has only about a fifth or sixth as much bio-
mass belowground as above, but for a decent-sized tree that is still a huge amount of
roots. With impermeable pavement running almost to the base of the trunk, those
roots receive only a tiny fraction of the oxygen and rainwater that they would receive
in the wild. Typically the roots of successful street trees are able to make do with little
water and oxygen. Many of the hardiest and most widely planted, such as London
plane, sweet gum, swamp cypress, and red maple, are floodplain plants with roots that
are used to getting by with little access to oxygen. The long association of ginkgo with
rivers may give them an edge as well.
The extensive networks formed by tree roots below sidewalks, driveways, and roads
often torment homeowners and local officials. A former colleague in the United King-
dom made a successful living as an expert in the identification of tree roots. His ser-
vices were much in demand from individuals and insurance companies wishing to find
out whose tree was undermining the foundations of whose house. The problems flow
the other way too. Most of the fibrous roots, which are the most important for absorb-
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ing water and nutrients, occur in the first foot of soil. When they are crushed by con-
crete, vehicles, and pedestrians, the aboveground parts of the tree begin to die.
Ginkgo tolerates root compaction better than most trees, but sometimes prevailing
in a battle of wills with the sidewalk triggers other consequences. In the spring of 2007
in Everett, Pennsylvania, a pedestrian tripped on a pavement upended by the roots of
a sixty-five-foot-tall ginkgo in front of the local library. Removal of the tree was rec-
ommended to the library board. However, it didn’t take long for citizen advocates
to launch a “Save the Ginkgo” campaign, with a petition circulated by the local high
school, a benefit concert, and T-shirts, as well as bowls and wine stoppers made from
the ginkgo’s trimmed branches. Their efforts not only saved their prize ginkgo but also
raised the $15,000 needed for pavement repairs.17
Along the way, renewed interest in the tree revealed that it was truly a survivor. It
was the last of three ginkgos planted in 1861 to honor the three sons of the founder of
the city, then called Blood Run, as they left to fight for the Union in the Civil War. The
tree also turned up in old photographs of the Fourth of July parade on Main Street
during in the 1920s, and could also be seen rising above the waters of the infamous
flood that swamped the town on Saint Patrick’s Day in 1936. It also turned out that the
Everett ginkgo had also been saved once before, in 1985, from the city’s attempts to
widen its street. A group of elderly ladies had threatened to chain themselves to it. Re-
silience may be only part of the story of ginkgo’s success, with charisma making up the
rest.
 33
Pharmacy

In all natural things there is something wonderful.

—Aristotle, “On the Parts of Animals”

One Friday afternoon at Kew, at the end of a harrowing week, my long-suffering

secretary confided that she was among the estimated ten million Europeans regularly
taking ginkgo leaf extract. “Oh yes,” she said, “I’ve been taking ginkgo for my mem-
ory for quite a while—when I remember.” Ginkgo is now a common herbal medicine
in the West. In the East, interest in the health-giving properties of ginkgo goes back
much farther. For almost as long as ginkgo has been prized for its nuts, it has also been
valued in medicine.1
According to some sources, the medicinal use of ginkgo dates back to 2800 B.C.,
to the first pharmacopeia of traditional Chinese medicine, attributed to the perhaps
mythical figure Shen Nung. However, the first undisputed written records of ginkgo
come much later, and no original copies of Shen Nung’s work survive. Ginkgo first ap-
pears in copies of the Shen Nung pharmacopeia around the eleventh and twelfth cen-
turies, about the same time as it begins to appear in other written records. It is certain
that ginkgo has been used in medicine for nearly a thousand years, but probably not
for three or four millennia.
In the legends and folklore of China, Korea, and Japan, ginkgo is often associated

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pharmacy

Stalactite-like zhōng-rǔ, or chi-chi,

bound with prayer ribbons on an
ancient ginkgo at the Huiji Temple
in Tang-Quan County, not far from
Nanjing, China.

with health and longevity. For example, in South Korea, the legend of the two ginkgo
trees in Myeoncheon makes a link to Bok Ji-gyeom, one of the founders of the Goryeo
Dynasty. It portrays him suffering from an incurable disease that has led his daugh-
ter to climb Ami Mountain and pray for one hundred days. There she meets a Taoist
hermit, who tells her, “Brew Dugyeonju [wine from azalea flowers] . . . and drink the
wine.” Then after another hundred days, he tells her, “plant two ginkgos, dedicate your
entire mind to praying, and it will cure your father.” His daughter having dutifully fol-
lowed the instructions, Bok Ji-gyeom recovers.2
Ginkgo is also often linked with fertility, and here the chi- chi, an eastern mani-
festation of herbalists’ “doctrine of signatures,” are especially important. Sometimes,
chi-chi in the shape of a phallus become a particular focus for the red prayer ribbons
that are often tied to the parts of old ginkgo trees. More often, the link is made to the
nursing of infants. In Japan, the word chi-chi refers directly to breasts. Legend has it
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that the massive Nigatake Ginkgo in the city of Sendai was planted as the dying wish of
Byakkonni, a wet nurse of the Emperor Shoumu. Women with problems in producing
their own milk still worship at the tree.3
In traditional Chinese medicine, the seeds of the ginkgo are used most often; the
best are said to be “large, dry, white, full and heavy.” Seeds from Guangxi Province in
southwest China are considered to be of especially high quality. They are used in the
treatment of lung and respiratory ailments, sometimes combined with other plants
such as the dried stems of ephedra, licorice root, and mulberry bark. They have also
been used to treat a broad range of conditions from nocturnal seminal emissions and
vaginal discharge to cavities, ringworm, scabies, and sores. Today ginkgo seeds con-
tinue to be used as an antitussive, expectorant, and antiasthmatic, as well as in the
treatment of bladder infections.
Different medicinal uses call for different modes of preparation: dried unprepared
seeds, generally known as yin hsing (silver apricot), are used to clear phlegm and kill
parasites, whereas dry, fried, or baked seeds known as cha bái guo (charred white fruit)
are pulverized and used to treat wheezing and vaginal discharge. To treat sores, fresh
ginkgo seeds are cut in half or a poultice is made from the powdered seed. A colleague
from Singapore also tells me that ginkgo seeds are good for the complexion.4
The chemistry of ginkgo has been studied for nearly two hundred years, and more
than 170 different chemicals have been extracted and described from the seed and
leaves. Some of these underpin the use of ginkgo in medicine, while others are respon-
sible for some of its less desirable attributes. In 1927 the Japanese scientist Kawamura
separated three novel, allergenic chemicals, ginkgolic acid, ginkgol, and bilobol, from
the fleshy seed coat, and found them chemically similar to the compounds responsible
for the allergic reactions produced by poison oak and poison ivy.5
Also problematic is butyric acid in the fleshy seed coat. This is what has given ginkgo
the nickname “ginkgo stinko” among some American urbanites and led to the outlaw-
ing of female trees in many Western cities. Descriptions likening the smell of fallen
ginkgo seeds to vomit and rancid butter are completely accurate; butyric acid is the
main volatile compound in all three sources. Nevertheless, in spite of the smell, the
fleshy coat from ginkgo seeds has sometimes found a use. In ancient China, it was
mixed with lye to make soap, and it has also been used by fishermen who apply it to
their bait to catch carp. In February 2007 the marine conservation group Sea Shep-
herd protested against a Japanese whaling expedition by spraying butyric acid at the
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whalers. Extracts from the smelly flesh show activity against disease- causing fungi,
drug-resistant strains of bacteria, and even the snail hosts of the parasitic schistosoma
fluke. The true value of the unwelcome waste product from ginkgo seeds remains to be
seen.6
Notwithstanding their culinary and medicinal uses, an unfortunate truth is that
ginkgo nuts are potentially toxic. In adults, ginkgo poisoning is rare, because generally
only eating a large quantity would trigger a reaction. However, there is no authorized
safe dose, and deaths have been reported from consumption of as few as fifteen or as
many as 574 nuts. It is especially recommended that children under the age of six limit
their consumption. However, the risk of ginkgo poisoning is very small, and I have en-
joyed ginkgo nuts prepared in many ways; they are an essential part of the experience
of being in eastern and southeastern Asia.
The effects of ginkgo seed poisoning have been known since at least 1709, when a
case was first described in Yamato-Honzo, an old Japanese scripture. Symptoms range
from irritability and vomiting to convulsions and loss of consciousness, which may
begin one to twelve hours after ingesting. The toxin interferes with the body’s ability
to absorb Vitamin B6, which is crucial to maintaining functional nervous and immune
systems, as well as many other vital processes. During food shortages in Japan between
1930 and 1960, reports of ginkgo poisoning increased significantly.
Despite a long history of selection and cultivation by people, the toxic compound
ginkgotoxin, also known as MPN, has not been reduced or eliminated. Nevertheless,
the toxin is water soluble and can be reduced by soaking. Levels of ginkgotoxin are
also reduced by cooking; concentrations can be over forty times greater in raw seeds
than in their cooked equivalents. In Yunnan Province, the Naxi ethnic group first soak
ginkgo nuts, then sauté them with onion, garlic, apple cider vinegar, soy sauce, sesame
oil, chili pepper, black pepper, and salt.7
As the medicinal use of ginkgo has spread from the East to the West, it has taken
a surprisingly different trajectory. In the East the seeds have been used most widely,
but in the West attention has focused almost exclusively on an extract from the leaves,
which has been promoted mainly as a memory enhancer. There are early references
from China to medicinal uses of the leaves. Dian Nan Ben Cao, written by Lan Mao in
1436, recommends the use of the leaves to treat freckles as well as skin and head sores.
Slightly later, the medical text Ben Cao Pin Hue Jing Yaor recommends ginkgo leaves
for internal use. In this and other Chinese materia medica, the leaves are described as
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used for treating dysentery, asthma, and cardiovascular problems. However, these uses
have not been widely adopted in the classic texts on traditional Chinese medicine.8
The active compounds in ginkgo leaves come from an impressive battery of chemi-
cals produced as a normal part of the plant’s growth and include two main classes of
chemicals, terpenoids and flavonoids. Flavonoids are the diverse chemicals responsible
for the colorful pigments of certain flowers, absorption of potentially harmful ultravio-
let radiation, protection against pathogens, and many other functions. Flavonoids are
often suggested to have health benefits because they are an important source of anti-
oxidants in food.9
Ginkgo leaves contain more than forty different flavonoids. Generally the leaves are
harvested just as they begin to change color in the autumn, when the flavonoid content
is highest. Green and yellow ginkgo leaves harvested in the autumn hold three times
the amount of one key flavonoid than spring and summer leaves. On the other hand,
the green spring and summer ginkgo leaves have a more potent content of other flavo-
noids and terpenes than the yellow ones, and also command a greater price in China
as a tea.10
In Europe the main conditions for which ginkgo leaf extract is prescribed are periph-
eral vascular diseases, or the narrowing of arteries surrounding the heart and brain,
resulting in reduced blood supply. In particular, ginkgo leaf extract has been widely
applied to the symptoms of “cerebral insufficiency” often seen in elderly people, in-
cluding difficulties of concentration and memory, absentmindedness, confusion, lack
of energy, and other symptoms.
Standardized ginkgo leaf extracts developed by Schwabe Pharmaceuticals in Ger-
many first appeared in 1964. Standardization is especially important for products from
a plant like ginkgo in which the content and concentration of flavonoids and other
chemicals may vary significantly through the season or depending on where the plant
is growing. The extraction process consists of twenty or more steps to enrich the active
and desirable substances in the leaves, while eliminating or greatly reducing the in-
active or potentially harmful substances. The first commercialized ginkgo product was
made available by Beaufour Laboratories in France in 1973, and was introduced to the
market in 1975 by the subsidiary group IPSEN under the registered name of Tanakan.
Soon after, INTERSAN and Schwabe laboratories released the ginkgo products Rökan
and Tebonin forte on the German market.11
By 1988 doctors in Germany were writing more prescriptions for drugs containing
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The Ginkgo Museum, a haven for ginkgo lovers in Weimar, Germany.

ginkgo extract than for any other plant-derived drug. Such use for symptomatic treat-
ment of deficits in memory, concentration, and certain kinds of depression is approved
by the public health insurance system. Currently ginkgo leaf extracts are among the
leading prescription medicines in both Germany and France, accounting for 1 and 1.5
percent of total prescription sales, respectively.12
Annual global sales of crude ginkgo leaf extract were around $1 billion in the late
1990s, mainly from sales in Germany, but also from elsewhere in Europe, as well as in
the United States and Asia. It is estimated that two billion daily doses have been used
over the past twenty years. Recently, ginkgo has become a top-selling herbal medicine
in the United States, despite the lack of the U.S. Food and Drug Administration’s ap-
proval for the standardized purified extract. There are currently dozens of products
based on ginkgo extracts, which can be administered intravenously, ingested in liquid
form, or taken as tablets.13
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According to some studies, the clinical effects of ginkgo, taken in the form of the
standardized purified extract, include improved memory and learning capacity, in-
creased brain tolerance to low oxygen, and improved circulation and microcircula-
tion. Negative side effects appear to be few, and some of those occasionally reported,
such as skin reactions and stomach upset, may be related to residual ginkgolic acids.
Headaches may be a product of increased blood flow. Studies that reported significant
improvements on one or more of the outcomes measured generally used dosages be-
tween 120 and 300 mg administered daily for between three and twelve weeks. When
the extract was being taken to affect physiologic functions, such as memory or mood,
four to six weeks were needed before positive results were noted. A review of 188 dif-
ferent studies conducted with humans and animals and in the laboratory found ginkgo
extract to be promising in demonstrating a range of neurological and physiological im-
provements, including some that can take effect in a matter of hours.14
In spite of such positive indications and its widespread use, the effectiveness of
ginkgo leaf extract remains a controversial subject. As with all herbal medicines, the
use of ginkgo leaf extract raises eyebrows among researchers from a more analyti-
cal scientific tradition. They want evidence, based on extensive trials, and at the same
stringent levels required of synthetic pharmaceuticals. In the United States, because
ginkgo leaf extract is classed as an herbal treatment, manufacturers are not required
to test the drug’s safety and effectiveness, as they would be if ginkgo leaf extract were
regulated by the FDA. The headline from a 2003 article in Scientific American, “The
Lowdown on Ginkgo biloba,” set a fairly typical tone from the orthodox end of the
medical practitioner spectrum: “This popular herbal supplement may slightly improve
your memory, but you can get the same effect by eating a candy bar.”
However, the assessment of the efficacy of ginkgo leaf extract in the article that fol-
lowed was balanced, fair, and based on a published review in a reputable scientific
journal by three respected neuroscientists. They found “evidence that Ginkgo enhances
cognitive functions, albeit rather weakly, under some conditions.” Overall, though,
they felt that more information was needed “to state conclusively whether Ginkgo
does or does not improve cognition.” They pointed out that too few experiments of the
right kind had been conducted, and generally at too small a scale to provide definitive
results. They summed up their feelings on the current state of research into the effec-
tiveness of ginkgo leaf extract as follows: “there are enough positive findings to sustain
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our interest in conducting further research on Ginkgo,” but they added an important
caveat. “Many years of experience with investigations of new drugs have demonstrated
that the initial positive results from studies involving a small number of subjects tend
to disappear when the drugs are tested in larger numbers of subjects from diverse
populations, so the true test of Ginkgo’s efficacy lies ahead.”15
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 part vii

Future
OVERLEAF Three statues carved from ginkgo wood by the Buddhist priest Mokujiki Shonin
in the early nineteenth century at the Ojiya Temple, central Honshu, Japan.
 34
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We see in many cases . . . that rarity precedes extinction; and we know that
this has been the progress of events with those animals which have been
exterminated, either locally or wholly, through man’s agency.
—Charles Darwin, On the Origin of Species by Means of Natural Selection

Perhaps more than any others, Dave Raup and Jack Sepkoski, working at the Uni-
versity of Chicago in the 1980s and early 1990s, initiated the modern quantitative study
of extinction in the fossil record. Dave led science at the Field Museum when I arrived
there in 1982, but his move to the University of Chicago later that year gave new impe-
tus to the development of that university’s distinctive tradition of paleontology, which
still continues. Along with Steve Stanley, at Johns Hopkins University, Dave was the
driving force behind the development of a fresh and analytical way to study the fossil
record of animals. As it developed in the late 1980s and early 1990s the approach that
Dave, Jack, and others fostered came to be known, sometimes with affection, and occa-
sionally dismissively, as the Chicago School of Paleontology.1
Jack was also a creative and brilliant thinker, and much of his best work emerged
from his analysis of the massive database that he developed over more than two de-
cades. In effect Jack built a giant table summarizing the history of occurrence of nearly

253
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every kind of animal through its entire fossil record based on specimens described
from anywhere in the world. From this mind-numbing compilation he was able to
chart and analyze the appearance and disappearance of different kinds of creatures
over about the past 550 million years. The main focus was on the history of life in an-
cient oceans, and his database eventually summarized the fate of about 3,500 different
families of marine animals. It was a herculean undertaking, but Jack’s ambition went
even farther. He developed a second database summarizing the comings and goings of
11,800 different genera of animals across the same vast expanse of geologic time.2
The work of assembling such a huge mass of data was tedious in the extreme, but
also required careful attention to detail. Jack spent untold hours trawling through
libraries to pull together thousands of fragments of information, often from obscure
publications written in many different languages, but in the process he had to assemble
the data in a consistent way, as far as possible standardizing different treatments of ge-
ology written by different authors at different times in different parts of the world. He
also did what he could to untangle the complexities that come from different authors
using different names for the same kinds of fossils.
Out of Jack’s synthesis came strong evidence, assembled with new rigor and ana-
lyzed with new insight, of large-scale patterns in the history of life, and with Dave
Raup also contributing his expertise, what emerged was the first well-founded attempt
to quantify changing levels of extinction and origination through geologic time. One
result, beguilingly simple but underpinned by a vast amount of data and huge effort,
was a bar chart showing the first and last appearances of families of animal fossils at
different times in the past. It revealed many ups and downs in the history of life, but
five intervals stood out as seemingly cataclysmic moments in which an especially large
number of families were lost never to appear again.
One such spasm of extinction took place around the boundary between the Ordovi-
cian and Silurian periods about 444 million years ago; a second occurred about 75 mil-
lion years later in the Late Devonian, and this was followed by a third, more dramatic
than any other, around 251 million years ago at the transition between the Permian and
Triassic periods. The fourth great extinction occurred at the end of the Triassic, about
200 million years ago, while the fifth, the so-called K-T extinction that extinguished
the dinosaurs and about which so much has been written, occurred at the boundary
between the Cretaceous and Cenozoic, about 65 million years ago. Jack and Dave dis-
tinguished these five extinction episodes from the background extinction that is an in-
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evitable part of the evolutionary process, and they also showed that while there have
been twelve significant extinction events over the past 250 million years, the “big five”
were of an entirely different order.3
The ancestors of living ginkgo, perhaps plants like glossopterids, evidently survived
the mass extinction at the end of the Permian. This extinction, the so- called Great
Dying, which wiped out perhaps 96 percent of all the different kinds of animals living
in the oceans, may be the closest that life has come to going completely extinct. Plants
very similar to living ginkgo also survived the extinction at the Triassic-Jurassic bound-
ary, as well as the mass extinction at the end of the Cretaceous. Nevertheless, despite
its resilience, ginkgo very nearly succumbed to the massive ecological reorganization
that has occurred over the past few million years and especially during the Quaternary
ice ages. Ginkgo came within a whisker of becoming one of those plants, like so many
others, that we know only from the fossils they left behind.
There is direct evidence from fossils that the climatic trauma of the past few million
years resulted in the loss of ginkgo from both western North America and Europe; a
similar loss probably also occurred in eastern North America, although we don’t have
the fossils to prove it. The cause was probably complex. Drier climates may have been
one problem. Repeated assaults by colder conditions, harder winters and shorter sum-
mers, and eventually the advance of glaciers from the north certainly constituted an-
other.
Young ginkgo seedlings were probably the first to succumb, cut off before they could
become established and start producing seeds of their own. Mature ginkgo trees may
also have been killed as they struggled to find water or were eventually ground down
by relentless frigid winters, shorter growing seasons, successive late frosts, and perhaps
also by temperatures that occasionally went so low that the young leaves were killed
inside the buds. If this increased mortality was coupled with limited capabilities for
dispersal, as seems likely, the overall effect would have been to reduce the geographic
range over which ginkgo could grow, and steadily deplete the number of living trees.
As our world emerged from the last ice age, the remaining living populations of ginkgo
in China may have been very small indeed.4
When speaking about extinction, Dave Raup would often use a memorable rhetori-
cal flourish. He would ask, “Did they go extinct because of bad genes or bad luck?” It
was a gentle rebuttal to the common assumption that animals or plants become extinct
because they are somehow poor performers: that species lose out to other species that
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are superior ecological competitors. Dave’s point was that such groups need not nec-
essarily have been competitively inferior, they could just have been unlucky: perhaps a
small population in the wrong place at the wrong time, or a large population that had
an unfortunate encounter with an asteroid. In the case of ginkgo, losing the animals
that dispersed its seeds might also be thought of as “bad luck.”5
At the same time, Dave’s favorite poser also had another subtext; it emphasized the
importance of what are sometimes thought of as “random” events in evolution. At one
end of this spectrum are certain mass extinctions, a form of extreme “bad luck”; it
could hardly be the fault of dinosaurs or ammonites that they fell afoul of an asteroid
from outer space. But in the same way, the cumulative impact of many small chance
events, building contingently and relentlessly one upon the other, might also be sig-
nificant. As Jimmy Stewart’s character finds out in It’s a Wonderful Life, Frank Capra’s
classic film, the present is the result of many small, seemingly insignificant, and some-
times chance episodes that over a lifetime add up and make a difference.
Both ideas speak to the significant role of chance and contingency in evolution.
Stephen Jay Gould explored these ideas in his book that played off the title of Capra’s
film. Gould’s particular focus was the early evolution of animals and how that early
diversification influenced what came later; but his broader point was that chance and
contingency have influenced much of the history of life. With regard to our own place
in biological evolution, he posed the provocative question: “Would we appear at all if
we could rewind the tape . . . and let it run again?”6
As Darwin made clear, as Dave Raup emphasized, and as modern-day conservation-
ists also recognize, the risk of extinction by simple “bad luck” is greatly increased when
species are reduced to just a few individuals. The risk is increased still further when
those last remaining holdouts live in just one place or a few places where they could
be wiped out by a single catastrophic event. The International Union for the Conser-
vation of Nature (IUCN) formalizes such thinking in the different categories of threat
that it recognizes in its so-called Red List. To qualify under the IUCN scheme as criti-
cally endangered, which is defined as facing “a very high risk of extinction in the wild,”
a species must meet several formal criteria, but basically they boil down to whether or
not the species is known from just a small number of individuals or from only one or
a few restricted locations.7
In the current IUCN Red List the most complete data are for the 5,488 known species
of mammals, all of which have been carefully assessed based on strict criteria; 188 are
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critically endangered. More than one in ten of all mammal species are either critically
endangered or endangered; seventy-six are thought to have gone extinct since 1500.
The scimitar oryx and Père David’s deer are extinct in the wild and survive only in cap-
tivity. The situation is similar for all other vertebrate animals and also, as far as we can
tell, for plants. A recent assessment of all 800 different kinds of conifers and cycads in
the world suggests that more than a third are threatened with extinction.8
Another important consideration in Red List assessments is whether there is already
evidence that the population size of a species is declining. Measuring such declines is
central to the Living Planet Index, a different approach to assessing the health of bio-
logical diversity. Developed by the World Wide Fund for Nature, the index is based on
a compilation of data on the changing size of populations of 1,313 animal species, in-
cluding fish, amphibians, reptiles, birds, and mammals. Since 1970 the Living Planet
Index has fallen by about 30 percent, mirroring the gloomy picture from the IUCN
Red List. Whichever way one looks at the current state of biodiversity, the news is not
good; much diversity has already been lost, and the trends are consistently downward.
The situation is urgent. It raises important questions about conservation priorities and
what the most effective approaches should be.9
 35
Insurance

They took all the trees and put them in a tree museum,
And they charged all the people a dollar and a half just to see ’em,
Don’t it always seem to go that you don’t know what you got till it’s gone?
—Joni Mitchell, “Big Yellow Taxi”

In a famous phrase from A Sand County Almanac, Aldo Leopold, the best-known
graduate of the Yale School of Forestry and Environmental Studies, argued: “If the
biota, in the course of aeons, has built something we like but do not understand, then
who but a fool would discard seemingly useless parts? To keep every cog and wheel is
the first precaution of intelligent tinkering.” The message for modern conservation is
clear, but too often we are forced to make choices, and exactly how conservation pri-
orities should be determined is a complicated issue on which there are many differ-
ent views. Nevertheless, focusing conservation attention on those species at greatest
risk is just common sense, and this means devoting particular effort to species that
are known only from small and threatened populations. Doing what we can to protect
those species where they occur, reducing the rate at which they are dying, and encour-
aging their reproduction are all critical steps toward the same basic goal: stabilizing or
increasing the number of living individuals of that species in the wild.1

258
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Zoos recognized long ago that the threats to some animal species in the wild were so
severe, and the chances of their succumbing to simple “bad luck” or to the poacher’s
gun were so great, that the best hope was to rear them in captivity. In some cases it
has proved possible to introduce these animals back into the wild. In the same way,
for plants, conservation through cultivation is an important part of the toolkit needed
to preserve the variety of plant life for the future. Everything possible should be done
to conserve plants where they grow, as part of the broader ecosystem in which they
evolved and the broader ecological processes in which they play a part; this should
always be our main objective. However, as in other spheres of life where the risks of ir-
reversible loss are great, it also makes sense to take out insurance. In the world of plants
the actions taken to assure the long-term survival of the Wollemi pine are instructive.
Like ginkgo, it is one of those species for which an encounter with people has helped
make it more, not less, secure.2
The initial discovery of the Wollemi pine in 1994 in the Blue Mountains, just west
of Sydney, was dramatic. David Noble, a park ranger working for the New South
Wales National Parks and Wildlife Service, rappelled down into an otherwise inacces-
sible gorge and came across a peculiar species of tree with strange-looking leaves and
“bubbly” bark. Specialists at the Royal Botanic Gardens, Sydney, quickly saw that this
was a new species, but almost equally quickly they also saw that it was vulnerable to
what Dave Raup would have called “bad luck.” The total population comprised only
about 110 individuals, all restricted to three localities that were very close together. The
“bad luck” of a bush fire in the wrong place or the introduction of a stray pathogen on
the boot of a visiting hiker could be enough to extinguish it. To the authorities in New
South Wales the potentially catastrophic consequences of such chance events added up
to a compelling case for swift conservation action.
The first step was to protect the area itself. The exact location of the Wollemi pine
populations is kept secret, and access, even for the most legitimate reasons, is highly
restricted. It was also decided not to intervene in a pristine habitat that was seemingly
untouched by direct human influence. However, right from the start, a major effort
was mounted to bring the Wollemi pine into cultivation and to distribute it to other
gardens. Some of the first plants outside Australia were grown at Kew and Wakehurst
Place. Tens of thousands of Wollemi pines, all ultimately clones from the hundred or
so wild trees, were distributed by the National Geographic Society for $99.95 each.
With seedlings now available from many sources, this unique and distinctive tree, un-
 Seed cones, seeds, and shoots of the dawn redwood, with its distinctive opposite leaves
and branching. The dawn redwood is a rare plant that was recognized as a fossil before being
discovered in 1946 living in the forests of Sichuan Province, China. Like ginkgo and the
Wollemi pine, it is now widely grown all over the world.
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known less than twenty years ago, is increasingly common in the world’s gardens. In
the coming decades and centuries, even if the few remaining trees in the Blue Moun-
tains should succumb to fire, disease, or climate change, the species is now safe. Its
long-term survival has been secured by a combination of effective vegetative propaga-
tion, clever marketing, and high-profile publicity.3
At the end of the last ice age the number of ginkgo trees living in China may not
have been quite as small as the number of Wollemi pines in the Blue Mountains, but
its situation may have been almost as precarious. However, ginkgo has also been for-
tunate; through its association with people, the population of ginkgo trees growing
around the world has been vastly increased. Even though many of the ginkgo trees
in cultivation may all be genetically rather similar and could be more or less equally
susceptible to a new pest or disease, ginkgo is now so widespread that there is little
chance it will go extinct simply through “bad luck.” By bringing the tree into cultiva-
tion in large numbers and in many different places around the world, we have greatly
increased its chances of long-term survival, and we are all among the beneficiaries.
This kind of ex situ approach to plant conservation is needed because everywhere
the world’s trees are under threat. According to the Global Trees Campaign, led by Sara
Oldfield at Botanic Gardens Conservation International, more than eight thousand
species, approximately 10 percent of all known trees, are threatened with extinction.
More than seventy are thought to be extinct, about eighteen are now known to exist
only in cultivation, and these depressing figures are almost certainly an underestimate.
Like ginkgo, each of these trees has its own story to tell; each helps fill in part of the
grand evolutionary puzzle.4
The drautabua is a small tree restricted to steep narrow ridge tops high in the moun-
tains of Viti Levu, Fiji. Like the Wollemi pine, it is one of the world’s most endangered
conifers; fewer than ten small and widely dispersed populations are known, and one
has already been lost. Fossils similar to the drautabua occur in Australia and Antarc-
tica, but the only other living species grows on New Caledonia. The drautabua is on the
brink of extinction. It is under threat from a nearby copper mine as well as from the
impacts of climate change in the special mountain habitats where it lives. It is not easy
to grow, but the few plants in botanic gardens provide invaluable insurance against its
loss in the wild.5
The Mulanji cypress, the national tree of Malawi, faces a similar crisis. It grows only
on Mount Mulanji and has been decimated by overcollecting for its highly prized,
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decay-resistant, timber. It is no longer reproducing in the wild. Its close relatives, the
Clanwilliam cedar, one of the most majestic trees of the Western Cape of South Africa,
and the Willowmore cedar, restricted to the Eastern Cape, have similar problems.
Three of the four species in the genus are in trouble. The Millennium Seed Bank at
Kew and its partners in Malawi and South Africa are collecting seeds, working to im-
prove the survival of seedlings, and bringing the plant into cultivation as a prelude to
its reintroduction into the wild.6
There are many similar examples. Franklinia, named for Benjamin Franklin and
discovered in what was then Britain’s American colony of Georgia by John and William
Bartram in 1765, is another tree that is widely cultivated but no longer known in the
wild. The Hawaiian cotton tree is known from only four trees growing in their original
habitat; many more are conserved in botanic gardens. It is a small tree with spectacu-
lar red hibiscus-like flowers; pure good luck brought its ancestor to Hawai’i, perhaps
three million years ago; a run of bad luck could easily wipe it out. The Robinson Crusoe
cabbage tree, known only from the Juan Fernández Islands, was down to just three
surviving plants in the 1980s; the Café Marron, known only from Rodrigues, survives
as a single wild plant; the Saint Helena ebony is known from just two individuals; the
Toromiro tree, from Easter Island, is now extinct in the wild. All have a safe haven in
the living plant collections at Kew and other gardens around the world.7
These examples provide instances of welcome insurance against extinction provided
by cultivation, sometimes combined with the protection of the species in a seed bank.
Ex situ conservation, by itself, is assuredly not enough. It cannot preserve the processes
that maintain species in their natural habitat, nor can it sustain the ecological services
provided by the community of which the species is part. However, ex situ conserva-
tion is a key tool to preserve plant diversity for the long term. Both in situ and ex situ
conservation are needed, and must be integrated, to ensure the long-term survival of
species that might otherwise be lost.8
Some contend that efforts to encourage ex situ conservation undermine efforts to
conserve plants in their native habitats or draw funds away from in situ conservation.
Such concerns are understandable, but fail to account for the realities of our current
predicament or different opportunities for conservation funding; in conservation, as
in many other areas, we should be careful that the perfect does not become the enemy
of the good.9
Ex situ conservation, as zoos have long recognized, is worth doing and brings many
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other benefits. It draws attention to the plight of species that are threatened, it creates
opportunities to assess the state of populations in the wild, and it helps build collec-
tive capacity around the world to preserve biological diversity. Ginkgo and the Wollemi
pine are excellent examples from the world of plants. Both are known from the paleon-
tological record, but surely everyone must agree that we are fortunate to know them as
living trees rather than only as fossils. As living plants we can continue to enjoy them,
study them, and better understand their secrets.10
Ginkgo and plants like it illuminate World History on the grandest scale, what some
have called Big History. Just as history taught solely from a Western perspective is in-
complete, history limited to a narrow time horizon only partly captures the contingen-
cies through which our modern world came to be. This is not just about our delight or
enjoyment; studying species and where they came from is one of the most important
windows through which we can understand the world we live in and our place in it.
It is sometimes said that the reasons to preserve species differ little from the reasons
to save a great work of art. It is a helpful analogy; it reminds us of what we stand to lose
when species go extinct. It underlines the cultural, emotional, and ethical dimensions
of animal and plant conservation. But it is also incomplete. Conserving species is not
just an indulgence for a few “tree huggers” who like “wild nature.”
For me, there is a fundamental difference between losing something created by
human genius over days, weeks, or years and losing something created by nature over
millennia. Either would be a tragic loss, but comparing the loss of species with the loss
of human achievements, equating human creativity with the creativity of nature, some-
how fails to capture the enormity of the situation we face. I prefer another analogy that
one also hears from time to time: letting species go extinct when we have the power to
intervene is like letting the library burn just as we are learning how to read the books.
It is a waste of information and a loss of knowledge about our world. Losing species
is a wasted opportunity to better understand the past, and understanding the past will
be necessary for managing the future. Preserving ginkgo and other species preserves
information on our own origins, our own history, and the history of the biological and
geological systems of which we are part. Extinction destroys the evidence of how our
world, and everything in it, came to be.11
 36
Gift

In communities drawn together by gift exchange, “status,” “prestige,”

or “esteem” take the place of cash remuneration.
—Lewis Hyde, The Gift

In June 1992 the nations of the world gathered in Rio de Janeiro at the first Earth
Summit. Attended by 172 countries, and more than a hundred heads of state, as well
as 2,400 representatives of nongovernmental organizations, it was one of the largest
United Nations conferences ever convened. The aim was to address increasing concern
about deterioration of the global environment. Greater awareness of pollution and
declining environmental quality, as well as their impacts on human health, had been
building since the 1950s and 1960s. Rio was a culmination of that process and a key
moment in the growth of the global environmental movement.1
One focus at Rio was desertification, the loss of vegetation in the drylands of the
world, and the threat that poses to poor people in developing countries. Another was
climate change, and it was in Rio that the United Nations Framework Convention on
Climate Change was introduced to the world; it led to the Kyoto Protocol and much
later to the climate change conferences in Copenhagen, Cancun, Durban, and else-
where. Also opened for signature in Rio was the Convention on Biological Diversity,
and in December 1993, ninety days after ratification by the thirtieth country, the con-

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vention entered into force. All the countries of the world, except Andorra, the Holy See,
South Sudan, and the United States of America, are now parties to the convention.2
The Convention on Biological Diversity (CBD) has as its aim the conservation
and sustainable use of biological diversity. It was a response to growing misgivings
about the fate of the animals and plants with which we share our planet. From Rachel
Carson’s warning about the devastating impacts of widespread synthetic pesticide use,
to the attention brought by Chico Mendes to the loss of the rain forest and abuse of its
peoples, to the coining of the term biodiversity itself in the mid-1980s, the CBD was the
result of a multidecade process through which the nations of the world sought a com-
mon approach to reduce the destruction and degradation of biodiversity. The Conven-
tion on International Trade in Endangered Species had already come into force in 1975,
in an effort to reduce threats to species from overexploitation, but there was a strong
feeling that more international action was needed.3
Early in the negotiations leading to the CBD it became obvious that biological di-
versity was not evenly distributed across the world’s surface. The richest places on the
planet for plant species are the tropical regions of Central and South America, espe-
cially in the foothills of the Andes in both Colombia and Ecuador, as well as the rain
forests of central Africa and southeast Asia, especially Malesia. Important outliers of
high species diversity also occur in a few other so-called hot spots, including the south-
ern tip of Africa and the extreme southwest of Australia. The comparison with other
parts of the world is stark. In the Atlantic rain forest of eastern Brazil, only 5 percent
of the original forest remains, yet it contains approximately twenty thousand plant
species, of which nearly half occur nowhere else in the world. The native flora of the
island of Madagascar totals about twelve thousand plant species, about 90 percent of
which occur only there. The native flora of the United Kingdom is meager by com-
parison. It comprises only about fourteen hundred native plant species, and almost all
occur elsewhere in Europe.4
The striking pattern is that many countries that are relatively rich in biodiversity
are relatively poor economically and struggle with massive day-to-day problems to
improve the lives of their people. As a result, concerns about the fate of animals and
plants, emanating especially from the wealthier countries of the world, often seem to
collide with more immediate concerns about the well-being of people in poorer coun-
tries. One outcome of this tension was that the process of negotiating the CBD became
highly charged and deeply politicized.
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A specific manifestation of this political reality was a perceived conflict between

the conservation of biological diversity and its use by people. While there are concerns
over the long-term fate of species of animals and plants, poverty, malnutrition, infant
mortality, and the many other pressing problems highlighted, for example, in the Mil-
lennium Development Goals are critical issues of basic human justice that must be ad-
dressed. Nevertheless, while it is easy to portray this tension as a simple dichotomy,
nature versus people, the reality is much more complicated. The fates of people and the
fates of their environment are inextricably interlinked. The real question is: how can
the Millennium Development Goals be met while also preserving biological richness
and the ecological goods and services that ensure healthy and sustainable supplies of
energy, food, medicines, and water, as well as a good quality of life?
It was not the role of the CBD to provide a definitive answer to such a complex ques-
tion, but it tried to find middle ground by establishing the principle that the conserva-
tion and use of biological diversity are two sides of the same coin; conserving biologi-
cal diversity enables us to continue to use it in sustainable ways. Implicit was the idea
that positioning conservation and use as complementary activities strengthens the case
for conserving biological diversity by emphasizing its value. It is a position that pivots
around a simple point; biological diversity should be conserved because it is useful.
In broad terms this makes sense: we keep what we use, and we are more likely to
conserve what we value. This key principle of the CBD takes an important step with the
emphasis that utilization should be sustainable. By any sensible biological definition
this means that the total number of individual living plants or animals should be main-
tained, not depleted. Populations of a species need to be allowed to reproduce and re-
place themselves at a rate equal to or greater than the rate at which they are being lost.
However, on top of these principles the political context of the CBD negotiations was
muddied by the arguments by overenthusiastic conservationists that a key reason to
protect natural habitats, most iconically the rain forest, was that the species living in
those places were an Eldorado of riches waiting to be exploited. It was a well-meaning
argument that went further down the utilitarian route, but it had unforeseen conse-
quences. Because it held out the promise of substantial future profits, it fueled nation-
alistic and protectionist tendencies in what one might have hoped would have been
more principled negotiations about the future of life on Earth.
One outcome of these complexities was that while the text of the convention ac-
knowledged that the conservation of biological diversity “is a common concern of
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human kind,” it nevertheless enshrined into international law for the first time the
principle that biological resources within the borders of individual countries are a na-
tional patrimony. At one level, this simply reaffirmed practices already implemented
in many countries, where the use of animals and plants living within their borders was
subject to national laws, but when the issue was raised in international negotiations
leading to the CBD, the question of under what terms biological resources should be
shared internationally was given new prominence. Inevitably, it inspired new nation-
alistic sensitivities and further complicated the political landscape.5
The struggle with these complexities resulted in another key provision of the CBD:
“the fair and equitable sharing of the benefits arising out of the use of the genetic re-
sources inherent in living organisms.” Again, a seemingly reasonable argument was
made that countries that are engaged in the conservation and sustainable use of bio-
logical diversity within their borders ought to share in any benefits that might arise
from its utilization. Heightened expectations of financial reward, concerns about pos-
sible “biopiracy,” and a general lack of trust took the convention still farther down this
road, to the point of linking the sharing of benefits not just to the commercial utiliza-
tion of biodiversity but also to access to it. Legally binding “access and benefit sharing
agreements” were one result.6
In these ways the convention sought to place the onus on countries to conserve the
biological diversity within their borders, while also seeking to provide economic incen-
tives by leaving open the option for sustainable utilization and introducing the prin-
ciple of benefit sharing. It was a seemingly sensible approach, but it had at its heart
one fundamental consequence: no longer were the plants and animals bequeathed to
all of us by 4.6 billion years of planetary evolution part of our common human patri-
mony; instead, they became the proprietary interests of nations. Trees, birds, flowers,
and all other kinds of organisms from insects to bacteria, and the genetic material they
contained, were taken as property by the people living inside the borders of individual
countries.
The ramifications of this fundamental shift are still being played out in never-ending
negotiations around the implementation of the CBD, but one important practical effect
is that many countries are now very sensitive about sharing their perceived biological
wealth with others. Their overriding concern is not to give away what they regard to be
valuable natural assets. One unintended consequence is that by limiting access through
complex permitting regulations the CBD actually helped stifle commercially oriented
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work that could potentially generate revenue. Another is that many countries greatly
restrict scientific access to their native plants and animals, even for noncommercial
work in collaboration with in-country scientists that helps support animal and plant
conservation. A third consequence is that the CBD unwittingly created a serious issue
regarding how the genetic diversity of the world’s most important crops, which was
once freely exchanged among countries, should be treated.
In essence, the International Treaty on Plant Genetic Resources for Food and Agri-
culture, which was adopted by the FAO conference in November 2001 and came into
force as of June 29, 2004, is an international agreement that excludes sixty-four of
the world’s most important crop and forage plants from some of the more unfortu-
nate provisions of the CBD. Ironically, for our most important plants this international
treaty seeks to restore the preexisting situation by attempting to create a multilateral
system through which crop genetic resources may be shared for purposes of breeding
new varieties. Despite broad agreement about the potential shared benefits it would
bring, in the atmosphere created by the CBD, the international treaty took seven years
to negotiate, failed to reach agreement on several key crops, remains highly conten-
tious, and is being implemented halfheartedly if at all by certain countries, even those
who are signatories. Terms can be revisited only by consensus among all 127 of its con-
tracting parties. It is symptomatic of where we have come to in seeking to manage
biological diversity in the twenty-first century: something is wrong in the way that we
think about the natural world and the common interests of humanity.7
The reality, two decades after Rio, is that the CBD has struggled to find its place
among national and international priorities. It has also not been well funded by those
governments that have ratified it. It would not be hard to argue that despite huge
investments of time and effort, and vast amounts of money spent mainly on the asso-
ciated meetings, the CBD has so far produced little revenue for countries that are rich
in biodiversity; nor has it resulted in a real advance toward its overriding goal of the
conservation and sustainable use of biological diversity.8
For the future of ginkgo, the Convention on Biological Diversity has been neither
a help nor a hindrance. The convention does not apply retroactively, and ginkgo was
already ubiquitous before we embarked on our clumsy global approach to conserv-
ing the variety of plant and animal life. Nevertheless, an interesting question is, if a
“new ginkgo” were discovered today in some far-off land, would the CBD make it more
secure or more vulnerable?
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Saplings of Wollemi pine for sale at

Wakehurst Place, the country garden
of the Royal Botanic Gardens, Kew
near Ardingly, West Sussex.

The answer would depend entirely on the attitude of the country in which this new
plant was discovered. Under the current regime of the CBD and the atmosphere that
it has created in many countries around the world, a “new ginkgo” would be jealously
guarded. Unfortunately, that is not the same as saying that it would be well protected
or that its long-term future would be secure. Other countries, however, would perhaps
take a more enlightened approach, and the Wollemi pine provides an encouraging ex-
ample through which to explore how such a thought experiment might turn out.
Australia, or more strictly the government of New South Wales, to which implemen-
tation of the CBD is delegated under Australia’s federal system, took a sensibly prag-
matic attitude to ensure the long-term future of the Wollemi pine. In effect the gov-
ernment sought to gain some short-term financial reward to assist conservation of the
species while also trying replicate what occurred with ginkgo through its interaction
with people. The authorities in New South Wales decided that in addition to protect-
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ing the tree in the wild, they needed to get it into cultivation—and, crucially, not only
in Australia. They did not hold out for some illusory long-term economic advantage.
First, they entered into a contract with a company to handle the propagation and
distribution of the Wollemi pine. The company raised thousands of young plants that
were ultimately distributed for sale around the world. The government then mounted a
successful and well-coordinated publicity campaign, which included selling off some of
the first plants to be propagated at a high-profile international auction at Sotheby’s in
New York. The funds raised from the auction and sales flowed back through the com-
pany into conservation of the remaining small population in the natural habitats.
The Wollemi pine provides an excellent example of an integrated approach to con-
servation of a rare plant. Protection of the wild population has been enhanced and this
unique species has been insured against “bad luck” by being brought into cultivation.
The Wollemi pine is now growing successfully in gardens all over the world, and in
the coming decades and centuries it is inevitable that the plant will be propagated by
others. Step by step the Wollemi pine, as it is grown in gardens far from its home in the
wild, will join many other wonderful gifts of Australia to the world.
To the extent that the CBD is an impediment to such effective ex situ conservation,
is perceived as such, or is used as a reason to prevent such exchanges of plant material,
it can do more harm than good. In the garden world there is a well-worn adage: “If
you have a rare plant; give it away.” The idea is simple: increase the number of plants
being grown and eliminate the risk that something valuable might be lost just through
“bad luck.” Unfortunately, the CBD steers us in the opposite direction, away from that
simple principle of common sense.
It is also an interesting question, for a plant like ginkgo, to think about what the
CBD really means. It would be hard to argue with the idea that ginkgo in some sense
belongs to China, but in an equally valid sense ginkgo belongs to us all. As the last of a
line of plants that was once much more diverse and grew on every continent, it is part
of the shared natural heritage that binds all people together. Ginkgo is both a gift of the
world to China, and a gift of China to the world. When we see a ginkgo on the street in
London, New York, or Tokyo, we should realize that it is a gift for which we all should
be grateful. It is a gift that enhances the esteem in which China is held by others; even
more so because it is a gift that has been freely given.
This line of thinking asks us to look beyond science and economics for guidance
about how we should view the natural systems of which we are part, and how we should
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gift

seek to manage them. It takes us into a realm where ethics, moral values, spiritual re-
lations, and a broader sense of our place in the world, and indeed the universe, come
into play. In many parts of the world, religions are central to the way that people think
about these issues and play important roles in constructing the moral frameworks
through which we interact with other people and the environment.9
In some religions, however, ideas about the natural world are peripheral rather than
central. For example, the cultural historian Thomas Berry offers the view that Chris-
tianity, with its emphasis on the individual, and that individual’s relationship with his
or her own God, in many ways directs us inward. It predisposes us to a worldview that
is fundamentally centered on the relation of humans to God and that too often turns its
back on the natural world. At the same time, Berry argues that a purely secular, mecha-
nistic, and scientific worldview has narrowed our perspective, and blunted our sense
of wonder. Fortunately, for many people around the world, and not just botanists, gar-
deners, or conservationists, trees still have the power to inspire, to evoke awe, and to
lift our spirits. They embody the intrinsic value of nature. Thomas Berry would have it
that in some sense we are part of them and they are part of us. Trees meet a basic bio-
philic need embedded deep within us: a need that we have inherited from our ances-
tors and that is increasingly starved by our highly urbanized and dominantly indoor
existence.10
The international approach that resulted in the CBD, with its parochial emphasis on
benefits and commoditization, brings into sharp focus fundamental questions about
our currently unsatisfactory relationship with the natural world. Does it really make
sense to try to manage the global environment on a country-by-country basis? Are we
comfortable with a view that so clearly asserts that nature is simply there for human
benefit? Is it morally or ethically right for the demands of people always to trump long-
term survival of species of plants and animals? And is it really in our long-term interest
to further extend our hegemony over nature? The ways in which such questions are an-
swered will be important for the future of all of humanity. If we take a broader view of
the history of our planet, and recognize that we have evolved over millennia as part of
complex global system of which we still have only limited knowledge, placing humans
so explicitly at the center seems arrogant, shortsighted. It might also be risky. To bor-
row a phrase from my friend Paul Falkowski, “Our destiny lies in understanding that
humility leads to enlightenment and that hubris leads to extinction.”11
 37
Legacy

In the empires of usury the sentimentality of the man with the soft heart
calls to us because it speaks of what has been lost.
—Lewis Hyde, The Gift

Tony Kirkham, who runs the arboretum at Kew, has become something of a celeb-
rity. He featured prominently in the BBC programs A Year at Kew, which appeared on
British television between 2004 and 2006. He then graduated to a television series of
his own: The Trees That Made Britain. With his colleague John Hammerton he explored
the contribution of trees to British history, landscape, and culture, from the preser-
vation of ancient yews in rural churchyards to the use of their wood in the medieval
longbow. He sampled cider made from the British apples in Somerset, went sailing in
ash-frame coracles, and investigated the sunken timbers of Britain’s sixteenth-century
warship the Mary Rose.1
However, Tony’s “day job” is to care for the nearly fifteen thousand trees on the Kew
estate. He makes sure that they are healthy, that they grow well, and that information
relating to their planting, growth, and management is up to date in the massive data-
base used to keep track of Kew’s collections of living plants. It is also Tony’s responsi-
bility to make sure that none of his trees is a danger to Kew visitors. Every tree is moni-
tored carefully every year. Tony is also an expert in the potentially dangerous business

272
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of climbing trees, and he recently revised the classic book on pruning trees and shrubs
written by one of his predecessors in the Kew Arboretum. These days, his team does
most of the ropework needed to remove dead limbs from high in the canopy.2
Tony also has responsibility for adding to Kew’s collections of living trees. With
his colleagues, especially Mark Flanagan, who is keeper of the Savill Gardens and the
Valley Garden Arboretum in Windsor Great Park, he has collected seed of different
species of trees in Japan, Taiwan, South Korea, and China. Back in Britain this seed
is the source for the new saplings planted out every year at Kew and in other gardens
and arboreta around the country. Tony has seen many of Asia’s ancient ginkgos on his
travels. In 2006 he visited Lengji, near Luding in western Sichuan, to check on the great
ginkgo photographed by Ernest Henry Wilson on August 1, 1908. It is still there, with
a small shrine at its base, crowded between the houses in the middle of the village, in
the valley of the Dadu River.3
Tony is a tree person through and through. Outside his family he has no greater pas-
sion. Like a doctor he does his best to avoid emotional involvement with his patients,
for from time to time he needs to perform major surgery, or even remove a tree com-
pletely, but he has not lost the sense of awe and wonder that drew him to trees in the
first place. He also knows that he must plant many more trees than he takes down to
allow for losses to disease and storms as his saplings grow to maturity. In 2009 on the
250th anniversary of the founding of the Royal Botanic Gardens, Kew, he led the plant-
ing of 250 new trees on the Kew estate. In May 2009 the last two trees in Tony’s yearlong
effort were a Wollemi pine planted by the Duke of Edinburgh and a ginkgo planted by
Her Majesty the Queen. Tony takes the long view; he understands that he is planting
for those who will come to Kew a hundred, or perhaps two hundred, years or more in
the future.
The Old Lion is one of more than fifty ginkgo trees under Tony Kirkham’s care. He
keeps a close eye on all of them, but the Old Lion is special. In the past few years he has
cleared the shrubs that once grew around its base; he saw no need to make it compete
with plants of lesser importance. He has stripped away the nearby grass and mulched
a broad circle around the base of the trunk. He has removed part of the tarmac path
that ran beneath the tree to allow more air and water to the roots; and he has used com-
pressed nitrogen to break up the compacted soil around them.
The care that Tony lavishes on his trees, and the concern that he feels for the Old
Lion and the other long-lived charges under his care, is paralleled not just in Britain
The Old Lion ginkgo at the Royal Botanic Gardens, Kew: one of the oldest and most celebrated ginkgo
trees in the United Kingdom, planted around 1761. It is a living link to the eighteenth century, when
King George III often spent time with his family on his Kew estate, but it also connects us to the
glorious and powerful deep history of plant and animal life on our planet.
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but everywhere around the world. In Oak Park, just outside Chicago, when the ginkgo
outside the Frank Lloyd Wright Home and Studio showed signs of stress, the tree sur-
geons were called in, dead branches were removed, the tree was carefully pruned, and it
emerged rejuvenated. Concerned that the Yongmunsa Ginkgo in Korea might be struck
by lightning, the temple authorities built a steel lightning tower close by to ensure
that this would never happen. When the great ginkgo at the Tsurugaoka Hachiman-gū
Shrine in Kamakura, Japan, fell after heavy rains in March 2010, work began immedi-
ately to propagate it so that the tree could live on at the same spot. When the Mizufuki
Ginkgo at the Nishi Honganji Temple in Kyoto, Japan, showed signs of decline, care-
ful pruning, soil renewal, and protection from the feet of thousands of temple visitors
brought it back to health.4
In the mid-nineteenth century, when the ancient Huiji Temple in Tang-Quan
County, not far from Nanjing, China, fell into disrepair and was lost, it was the two
great ginkgos that grew close to where it once stood that kept its memory alive. They
also came to preside over the small ginkgo orchard planted beneath them. When I
visited in August 2008, the temple was being rebuilt and both ginkgos were wrapped
in red ribbons; some had wishes of all kinds written on them. Other visitors had ex-
pressed their hopes more directly in neat vertical columns of Chinese characters writ-
ten on small bare patches of wood where the bark had been worn away. These trees are
important in their community; local people still go there to pray. Late in the afternoon,
as we left to return to Nanjing, a young woman arrived. She looked nervously behind
her to be sure that we were leaving, then made her way to one of the two great ginkgos
and stood alone in a few moments of quiet contemplation. Many others, from previous
generations, had stood there before her.5
In rural Aomori Prefecture in northern Honshu, the Hōryō Ginkgo sits on just a
sliver of uncultivated land sandwiched between commercial forests and agricultural
fields. It is approached with reverence down an aisle of closely spaced, moss-covered
stepping-stones. Local people visit it regularly. They lovingly clear away the twigs and
branches that fall in the winter snows, they care for the rice straw rope wound around
the trunk, they explain the tree’s legends to local schoolchildren, and they work to
spread word of its importance. This tree was a friend to their grandparents; it will prob-
ably also be a friend to their grandchildren.
These scenes are repeated many times over, in many ways, in many parts of the
world. People feel connected to trees; old trees in particular are objects of affection
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future

that have earned and deserve our love and respect. Despite the comings and goings of
nations, times of conflict and times of peace, years of austerity and years of plenty, the
great ginkgos of Asia, and other ancient trees around the world, provide continuity
in communities with the passing of generations. It is not only “plant people” who are
moved by the grandeur of a coast redwood, or humbled by the antiquity of bristlecone
pines. These feelings resonate among anyone who takes the time to reflect on the true
meaning of ancient and long-lived plants. In cultures around the world trees place our
own ephemeral existence in proper perspective.
When I was growing up in the Midlands of England, the countryside around my
home was dominated by majestic avenues, each composed of hundreds of elms. They
were all planted more than two centuries before by a wealthy local landowner, the sec-
ond Duke of Montagu, sometimes known as John the Planter. It is said that his plan
was to plant an avenue of elms and lindens all the way to London, but thwarted by the
difficulties of persuading other landowners to join in the project, he satisfied himself
with planting his great avenues around and across his own estate. Altogether the ave-
nues he created were more than seventy miles long. Then, sometime in the late 1960s,
I remember hearing for the first time about the scourge of Dutch elm disease; the dis-
ease quickly took its toll. In just a few years the elms of my childhood, the elms that
are such a feature of Constable’s paintings, the elms across the whole of Britain, were
all gone. A magnificent tree that had been part of the English landscape for thousands
of years was almost obliterated. A bald spot appeared in the Kew arboretum.6
One morning in early October 2006, soon after I arrived back in Chicago from our
years at Kew, I drove down to the University of Chicago with my daughter, who was
attending school on campus. There had been a big storm overnight. Where we live in
Oak Park, to the west of the city, there was just lots of water. As we approached the
South Side, the scene was different; everywhere there were trees with big limbs on the
ground, and others had been completely uprooted. On campus, in the Quadrangle out-
side the Divinity School, a big oak, perhaps older than the university itself, was down
on its side and already being dismantled by staff from the grounds department. More
than nine hundred large mature trees came down that night on the South Side of Chi-
cago; a thousand more were badly damaged. In a matter of hours trees were lost that
had always been there, trees that were there before my daughter’s grandparents were
born. We had taken them for granted, and we hadn’t known what we had until they
were gone. Trees take time to grow, but their loss can be quick and easy. It takes only
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a moment to dynamite an acre of rich Appalachian forest to plunder the coal beneath.
It takes just a few hours to fell giant dipterocarps in the rain forest of Borneo, or old-
growth hemlocks in the Pacific Northwest of the United States. Trees and forests that
have stood firm for centuries in the face of repeated natural assaults have no power to
resist our fleeting, but often devastating, attention.
These timescales—hours, days, or even a few years—contrast with the timescales of
ginkgo’s life story. Hundreds, thousands, millions, tens of millions, hundreds of mil-
lions of years: these spans are not easy for us to grasp, but they are perhaps more rele-
vant to the way we should think about ourselves, and our true place in the world. They
should make us pause. Trees help calibrate the speed of current environmental change:
they provide a context more in tune with the tempo of the Earth. They slow us down,
they teach us the virtue of patience, and they remind us to think about all that has gone
before and what is to come; the legacy we received and the legacy that we will leave. The
modern-day mantra of more, better, faster is all very well; but followed unthinkingly it
is a recipe for disaster. Trees, especially trees like ginkgo, which connect us to the deep
history of our planet, ask us to reflect more often and think more carefully about all we
lose when the short view rules our world and everything in it.
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 Appendix
List of Common Plant Names Used
in the Text and Latin Equivalents

(fossils indicated by †) †Avatia = fossil seed-bearing structure related

African oil palm = Elaeis guineensis to living ginkgo
Alder = Alnus spp. azalea = Rhododendron spp.
†Allicospermum xystum = fossil seed related to †Baiera furcata = fossil leaf related to living
living ginkgo ginkgo
Almond = Prunus dulcis †Baiera gracilis = fossil leaf related to living
American beech = Fagus americana ginkgo
American elm = Ulmus americana †Baiera hallei = fossil leaf related to living
apple = Malus pumila ginkgo
apricot = Prunus armeniaca bald cypress = Taxodium spp.
†Archaefructus = early fossil flowering plant banyan = Ficus benghalensis
†Archaeopteris = fossil “progymnosperm” baobab = Adansonia digitata
related to living and fossil seed plants bay tree = Laurus spp.
argan = Argania spinosa bearded fig = Ficus citrifolius
aroids = several genera in the family Araceae beech = Fagus spp.
aromatic ginger = Kaempferia galanga betel = Piper betle
ash = Fraxinus spp. bigleaf hydrangea = Hydrangea macrophylla
aspen = Populus spp. (syn. Hydrangea otaksa)
†Asteroxylon = fossil plant from Rhynie Chert birch = Betula spp.
related to living fir clubmoss blackberry = Rubus fruticosus
Atlantic cedar = Cedrus atlantica black locust = Robinia pseudoacacia

279
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black tupelo = Nyssa sylvatica currant = Ribes spp.

bright-green cave-moss = Cyclodictyon laete- cycads = about eleven genera in several
virens (syn. Hookeria laete-virens) families of the order Cycadales
bristlecone pine = Pinus longaeva (other †Cyclocarya brownii = fossil fruits of the living
closely related, long-lived species, which are wheel wingnut in the Almont fossil flora
sometimes also called bristlecone pine, are cypress = several genera in the family
Pinus aristata and Pinus balfouriana) Cupressaceae
butterfly bush = Buddleja spp. dawn redwood = Metasequoia glyptostroboides
cabbage palmetto = Sabal palmetto dense logwood = Croton congestus
Café Marron = Ramosmania rodriguesii dipterocarp = several genera in the family
camellia = Camellia spp. Dipterocarpaceae
Carolina mahogany = Persea borbonia divi-divi = Caesalpinia coriaria
castor oil = Ricinus communis dove tree (handkerchief tree) = Davidia
Cathay silver fir [yin shan] = Cathaya involucrata
argyrophylla dragon tree = Dracaena draco
Caucasian wingnut = Pterocarya fraxinifolia drautabua = Acmopyle sahniana
cedar of Lebanon = Cedrus libani elm = Ulmus spp.
celery = Apium graveolens English oak = Quercus robur
cherry = Prunus avium †Eoasteria = fossil, probable pollen-producing
chestnut = Castanea sativa structures related to living ginkgo
chili = Capsicum annuum ephedra [ma huang] = Ephedra spp.
†China fir = Cunninghamia cheneyi fossil eucalypt = Eucalyptus spp.
shoots of living China fir in the Clarkia fig tree = Ficus spp.
fossil flora fir = Abies spp.
China fir = Cunninghamia lanceolata, fir clubmoss = Huperzia selago
Cunninghamia konishii (two closely related fishtail palm = Caryota spp.
forms that may just be a single species) Franklinia = Franklinia alatamaha
Chinese date = Ziziphus jujuba garlic = Allium sativum
Chinese swamp cypress = Glyptostrobus pensilis giant sequoia = Sequoiadendron giganteum
cinnamon = Cinnamomum aromaticum ginger = Zingiber officinalis
Clanwilliam cedar = Widdringtonia ginkgo = Ginkgo biloba
cedarburgensis †Ginkgo adiantoides = fossil leaf related to
clubmoss = several genera in the family living ginkgo
Lycopodiaceae †Ginkgo apodes = fossil leaf related to living
coast redwood = Sequoia sempervirens ginkgo
cork oak = Quercus suber †Ginkgo australis = fossil leaf related to living
crepe myrtle = Lagerstroemia indica ginkgo
crocus = Crocus spp. †Ginkgo cordilobata = fossil leaf related to
croton = Croton tiglium living ginkgo
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†Ginkgo cranei = fossil leaf related to living hydrangea = Hydrangea spp.

ginkgo Japanese cedar = Cryptomeria japonica
†Ginkgo florinii = fossil leaf related to living Japanese elm = Zelkova serrata
ginkgo Japanese fig = Ficus erecta
†Ginkgo huttoni = fossil leaf related to living Japanese maple = Acer japonica
ginkgo †Kannaskoppia = fossil seed-bearing structure
†Ginkgo orientalis = fossil leaf related to living possibly related to living ginkgo
ginkgo †Kannaskoppianthus = fossil pollen-producing
†Ginkgo rajmahalensis = fossil leaf related to structure possibly related to living ginkgo
living ginkgo †Kannaskoppifolia = fossil leaves possibly
†Ginkgo yimaensis = fossil leaf related to living related to living ginkgo
ginkgo †Karkenia asiatica = fossil seed-bearing
†Ginkgoites matatiensis = fossil leaf related to structure related to living ginkgo
living ginkgo †Karkenia incurva = fossil seed-bearing
†Ginkgoites muriselmata = fossil leaf related to structure related to living ginkgo
living ginkgo katsura = Cercidiphyllum japonicum
†Ginkgoites taeniata = fossil leaf related to †Kerpia = fossil leaf possibly related to living
living ginkgo ginkgo
†Ginkgoites telemachus = fossil leaf related to koompassia = Koompassia spp.
living ginkgo laurel = Laurus spp.
†Ginkgoites ticoensis = fossil leaf related to licorice = Glycyrrhiza glabra
living ginkgo lilac = Syringa spp.
†Ginkgoites tigrensis = fossil leaf related to lily bulbs = Lilium spp.
living ginkgo linden (lime) = Tilia spp.
gnetum = Gnetum spp. live oak = Quercus virginiana
grape = Vitus spp. Lombardy poplar = Populus nigra
grape family = Vitaceae London plane = Platanus x acerifolia
guanacaste = Enterlobium cyclocarpum lotus = Nelumbo nucifera
handkerchief tree (dove tree) = Davidia lychee = Litchi chinensis
involucrata magnolia = Magnolia spp.
hardy rubber tree = Eucommia ulmoides mahogany = Swietenia spp.
Hawaiian cotton tree = Kokia drynarioides ma huang (Ephedra) = Ephedra spp.
hazel = Corylus spp. maidenhair fern = Adiantum spp.
heath = Erica spp. maize = Zea mays
hemlock = Tsuga spp. maple = Acer spp.
Henry honeysuckle = Lonicera henryi Mediterranean fig = Ficus carica
Henry lily = Lilium henryi mimosa = Mimosa spp.
hickory = Carya spp. mistletoe = Viscum album
hornbeam = Carpinus spp. monkey puzzle = Araucaria araucana
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Monterey pine = Pinus radiata pumpkin = Cucurbita pepo

moonseed family = Menispermaceae quaking aspen = Populus tremuloides
Mormon tea = Ephedra spp. red maple = Acer rubrum
mountain cedar = Widdringtonia nodiflora rhododendron = Rhododendron spp.
Mulanji cypress = Widdringtonia whytei rice = Oryza sativa
mulberry = Morus spp. Robinson Crusoe cabbage tree = Dendroseris
Norway maple = Acer platanoides litoralis
nypa = Nypa fruticans rockrose = Cistus spp.
oak = Quercus spp. rose = Rosa spp.
oil palm [African] = Elaeis guineensis rubber tree = Hevea brasiliensis
onion = Allium cepa Sago cycad = Cycas revoluta
pagoda tree = Styphnolobium japonicum Saint Helena ebony = Trochetiopsis ebenus
†Palaeocarpinus = fossil fruit in the birch sandalwood = Santalum spp.
family sassafras = Sassafras spp.
palmetto = Sabal palmetto sesame = Sesamum indicum
palm family = Arecaceae sessile oak = Quercus petraea
Panama hat palm = Carludovica palmata she-oak = Casuarina spp.
Pau Brasil = Caesalpinia echinata silk tree = Albizia spp.
pea = Pisum sativum silverberry = Elaeagnus spp.
peach = Prunus persica silver fir = Abies alba
peanut = Arachis hypogaea snowdrop = Galanthus spp.
peepal (pipal) = Ficus religiosa Spanish dagger = Yucca gloriosa
†Petriellaea = fossil seed-bearing structure †Sphenobaiera umaltenis = fossil leaves related
potentially related to fossil Kannaskoppia to living ginkgo
pine = Pinus spp. spruce = Picea spp.
pinyon pine = Pinus subsection Cembroides †Stalagma samara = enigmatic fossil plant
and Nelsonianae perhaps related to living conifers
pistachio = Pistachio vera stonecrops = Chara spp.
plane (sycamore) = Platanus spp. stone pine = Pinus pinea
plum = Prunus spp. strawberry = Fragaria x ananassa
plum yew = Cephalotaxus spp. strawberry tree = Arbutus unedo
poison ivy = Toxicodendron radicans sugar maple = Acer saccharum
poison oak = Toxicodendron diversilobum swamp cypress = Taxodium distichum
pomegranate = Punica granatum swamp gum = Eucalyptus regnans
Portuguese laurel = Prunus lusitanica sweet gum = Liquidambar styraciflua
potato = Solanum tuberosum sweetsop family = Annonaceae
prickly ash = Zanthoxylum spp. sycamore = Platanus spp.
†Psygmophyllum = fossil leaves of uncertain tamarix = Tamarisk spp.
relationship teak = Tectona spp.
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titan arum = Amorphophallus titanum willow = Salix spp.

Toromiro tree = Sophora toromiro willowmore cedar = Widdringtonia schwarzii
tree of heaven = Ailanthus altissima witch hazel = Hamamelis spp.
†Trichopitys heteromorpha = fossil seed plant Wollemi pine = Wollemia nobilis
of uncertain relationship yam = Dioscorea spp.
umbrella pine = Sciadopitys verticillata yevaro = Eperua purpurea
valley oak = Quercus lobata yew = Taxus spp.
Venus flytrap = Dionaea muscipula †Yimaia capituliformis = fossil seed-bearing
walnut = Juglans spp. structure related to living ginkgo
watercress = Nasturtium officinale †Yimaia qinghaiensis = fossil seed-bearing
water lily = several genera of aquatic plants in structure related to living ginkgo
the family Nymphaeaceae †Yimaia recurva = fossil seed-bearing structure
watermelon = Citrullus lanatus related to living ginkgo
welwitschia = Welwitschia mirabilis yin shan (Cathay silver fir) = Cathaya
wheel wingnut = Cyclocarya and Asian argyrophylla
Pterocarya spp.
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 Notes

1. Time
1. Epigraph: Conan Doyle (1912, 63). Note that Conan Doyle’s original spelling transposed the g
and the k, a common error in the spelling of ginkgo.
2. In March 2011, the Old Lion ginkgo at Kew stood about sixty-three feet tall and had a trunk
diameter of a little more than five feet just below the level where the trunk splits into its two main
leaders (Tony Kirkham, Royal Botanic Gardens, Kew, personal communication).
3. Darwin referred to another botanical oddity, Welwitschia, as a platypus for the vegetable king-
dom. He wrote to J. D. Hooker on December 18, 1861: “Your African plant seems to be a vegetable
Ornithorhynchus, and indeed much more than that”; see Darwin and Seward (1903, 281).
4. The new kinds of plants that rose to prominence around 100 million years ago, and that now
dominate almost all terrestrial environments with about 350,000 living species, are the angiosperms
(flowering plants). The geologic timescale used in this book is that of Gradstein et al. (2004).
5. Earlier records of modern humans in China, for example from 100,000 years ago (Liu et al.,
2010), are controversial and could potentially be interpreted as gracile forms of Homo erectus (Den-
nell, 2010).

2. Trees
1. The Marquess of Blandford who planted the arboretum on the Whiteknights estate later be-
came the fifth Duke of Marlborough.
2. Bill Burger, a colleague at the Field Museum in Chicago, first introduced me to Warren Woods.

285
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n o t e s t o pa g e s 9 – 1 1

He describes the significance of this special woodland in Chicago Wilderness magazine; see Trigg
(2008).
3. Nearly eighty thousand people every year visit the coast redwoods at Muir Woods, just north
of San Francisco.
4. The photographs in Meetings with Remarkable Trees and Remarkable Trees of the World, by
Thomas Pakenham (2002, 2003), come closer than most to revealing the true power of the world’s
great trees.
5. Eiseley (1958) argues that the prehensile hands and forward-directed vision that evolved in
humankind’s ancestors developed as adaptations to life in the trees and facilitated movement through
a three-dimensional maze of branches.
6. My climb up the koompassia took place at the Danum Valley Field Centre in Sabah, Malaysia,
probably the leading center for rain forest research in the Old World tropics. The center is adjacent
to the Danum Valley Conservation Area, one of the largest remaining, most important, and best-
protected areas of pristine lowland rain forest in southeastern Asia. Koompassia trees are often left
standing in previously logged areas because their timber is poor and silica deposits in the wood make
them hard to fell.
7. The story of Adam and Eve is from Genesis 2:9; mention of the tree of knowledge is from
Genesis 2:17.
8. The giant peepal at the Mahabodhi Temple in Bodh Gaya, northern India, is known as the Sri
Maha Bodhi, or simply Bodhi tree. The Buddha is said to have achieved Enlightenment under this
tree. Bodhi means “awakening” or “enlightenment” in Sanskrit. The name is also used to refer to other
trees propagated from the Sri Maha Bodhi. Ongoing studies by John Rashford are elucidating the sig-
nificance of fig trees in Candomblé.
9. The significance of the palmetto, sometimes called the cabbage palmetto, to South Carolina
dates to the Revolutionary War, when the British fleet retreated from its attack on Sullivan’s Island
on June 28, 1776, after the palmetto-log fort withstood its cannon fire. In 1950 Mrs. John Raymond
Carson incorporated the story into the pledge to the state flag: “I salute the flag of South Carolina and
pledge to the Palmetto State love, loyalty and faith.” For the significance of the bearded fig in Barba-
dos and its presence on that country’s coat of arms, see Rashford (2007). In China, in 1942, the scien-
tist, poet, and historian Guo Moruo proposed ginkgo as the national tree. This also became the goal
of Jiangsu Congress Deputy Ju Zhangwang, who from 2003 through 2008 proposed such adoption to
the National People’s Congress. In a national poll sponsored by the State Forestry Administration in
2005, ginkgo won more than 1.7 million of the total 1.8 million votes cast. Ginkgo remains to be offi-
cially designated as China’s national tree.
10. The Council Oaks and the Emancipation Oak are live oaks. The Hooker Oak is a valley oak.
Joseph Dalton Hooker (1817–1911) was the second director of the Royal Botanic Gardens, Kew. He
succeeded his father, William Jackson Hooker (1785–1865), in 1865 and retired in 1885. See Allen
(1967) for more on these first two directors of Kew, and Desmond (1999) and Endersby (2008) for
more on Joseph Dalton Hooker.
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n o t e s t o pa g e s 1 2 – 1 5

11. According to some estimates the Yongmunsa, “Dragon Gate,” Temple was first built in 913 and
expanded in 1392, although it has been rebuilt several times. It burned down in 1592, and many of
the old structures were burned in 1907 by the Japanese after a military uprising. The temple was also
badly damaged during the Korean War. Reconstruction of the current temple was completed in the
1980s; see Sky News (2007).
12. The Silla Dynasty began in 57 B.C. and ended in A.D. 935. If the Yongmunsa Ginkgo was
planted at the end of this period, it has survived for almost eleven centuries. The Yongmunsa Ginkgo
reaches an impressive 200 feet. It remains imposing and among the tallest ginkgo trees on the planet;
see Invitation ForestOn, “Story of forest: Old gigantic trees in Korea.”
13. Between 1983 and 1996, the National Forest Seed Centre in Burkina Faso distributed seventeen
tons of seeds of about sixty tree species to help restore degraded woody vegetation and stock local
plantations and nurseries. This project was expanded and enhanced through the work of the Mil-
lennium Seed Bank and also has yielded important data on storing and conserving seeds from the
semiarid tropics. (Moctar Sacande, Royal Botanic Gardens, Kew, personal communication); see also
Sanon et al. (2004).
14. Estimates of U.S. paper consumption are from the World Resource Institute’s 2005 Statistics;
see Kahl (2009) and Nadkarni (2008).
15. Olson et al. (2001) document the “OneTree Project.”
16. Bill Vaughan (1915–1977) wrote a regular feature, “Starbeams,” for the Kansas City Star.
17. The Angel Oak is a massive live oak on John’s Island just outside Charleston, South Carolina.
The “tree sit” on the University of California, Berkeley, campus is recounted by Burress (2008).
18. President Clinton’s 1993 “Forest Plan for a Sustainable Economy and Sustainable Environ-
ment,” known as the Northwest Forest Plan, aimed to integrate management and conservation and
balance different demands on forest resources; see Tuchmann et al. (1996). Recognizing that the cut-
ting of forests contributes perhaps between 6 and 17 percent of annual global carbon dioxide emis-
sions, REDD (Reduced Emissions from Deforestation and Forest Degradation) has been a recurring
issue in the global negotiations about climate change since it was first introduced in the United Na-
tions Framework Convention on Climate Change, Conference of the Parties 13, in Bali in 2007. For
an estimate of CO2 emissions from forest loss, see Van der Werf et al. (2009).
19. The Millennium Ecosystem Assessment, carried out between 2001 and 2005, concluded that
over the past fifty years, humans have modified ecosystems “more rapidly and extensively than in any
comparable period of time in human history,” resulting in “a substantial and largely irreversible loss
in the diversity of life on Earth.” Cultivated systems now cover one-quarter of our planet’s terrestrial
land. “Global forest cover loss” is estimated at around 628,206 square miles between 2000 and 2005;
see Hansen et al. (2010) and Millennium Ecosystem Assessment (2005).
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n o t e s t o pa g e s 1 6 – 2 0

3. Identity
1. Epigraph: inscription on plaque by the ginkgo trees near the LuEsther Mertz Library, New
York Botanical Garden.
2. The descriptions of William Hooker’s character are from William Henry Harvey, an Irish bota-
nist who specialized in algae and named the bright-green cave-moss (Hookeria laete-virens) for his
lifelong friend; see Allen (1967).
3. For the history of the Royal Botanic Gardens, Kew, see Desmond (2007). An outline timeline
of the development of the Royal Botanic Gardens, Kew, can be found at www.kew.org/heritage/index.
html; see also Blomfield (1994, 2000) for a broader history of Kew village and the Kew community.
4. In leading the revitalization of Kew, William Hooker helped to realize the vision of one of his
mentors, Sir Joseph Banks. Banks wished to see Kew present interesting, beautiful, and useful plants
from all over the world to the public, and distribute plants of economic importance throughout the
British Empire; see Desmond (2007) and Allen (1967).
5. See Drayton (2000) for a view of how Kew and other botanic gardens became instruments of
broader colonial objectives in Victorian and earlier times. See Griggs et al. (2000) for an introductory
overview of the Kew Economic Botany collection.
6. See Quin (1882, 199) and also Prendergast et al. (2001) for more on John Quin and his collec-
tion at Kew.
7. The Qin (or Ch’in) Dynasty began in 221 B.C. and ended in 206 B.C.
8. The ginkgo leaf in the Kew collection is of no great value except for the unusual way that it
combines an archetypical Chinese image with one of the most potent natural symbols of the East.
Ancient ginkgo trees occur in Shizilin, the Lion Grove Garden, and also in Liu Yuan, the Lingering
Garden, in Suzhou, China. The ginkgo growing by a small pavilion in the Lingering Garden is 110 feet
tall.
9. Some have speculated that the resemblance of the ginkgo leaf to a fan, a revered symbol, may
have been important in the tree’s adoption into Buddhism and Shintoism.
10. The Nongso, Seonsan, Ginkgo is in the city of Seonsan in the South Chungcheong Province,
South Korea. Information on this and other ginkgo trees designated as natural monuments in South
Korea can be found at http://english.cha.go.kr/.
11. For a more complete list of Japanese names that incorporate icho, “ginkgo,” see Hori and Hori
(1997).
12. Larry Kirkland’s mural can be viewed in the Keck Center Lobby of the National Academy of
Sciences building in Washington, D.C., at Sixth and E Streets.
13. Frank Lloyd Wright may not have removed the ginkgo tree, but he did complain about its
smelly seeds in the autumn, according to my former colleague Laurel Ross (personal communica-
tion). Her father apprenticed with Frank Lloyd Wright in Oak Park and often recounted Wright’s
comments about the female ginkgo. The Ginkgo Dreams Web site features a wide variety of prod-
ucts and art that use the ginkgo leaf as part of the design; see www.ginkgodreams.com. See also, for
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n o t e s t o pa g e s 2 0 – 2 3

example, the ginkgo collection of Michael Aram; www.michaelaram.com and Schmid and Schmoll
(1994) for the influence of ginkgo on diverse decorative arts.
14. For the use of ginkgo in the Art Nouveau architecture of Nancy and Prague see Kwant, “Ginkgo
biloba and Art Nouveau in l’Ecole de Nancy,” and Kwant, “Ginkgo biloba and Art Nouveau in Prague.”
15. Gilbert and George cite their first experience smelling ginkgo in New York City as their inspi-
ration for the exhibit. They first attributed the smell to a dog, then concluded, erroneously, that “the
female ginkgo leaf simply smells like that.” For a review of the Gilbert and George retrospective see
Vogel (2005), Wyman (2008); see also the catalogue (Birnbaum and Bracewell, 2005).
16. The ginkgo tree under which Confucius is reputed to have sat and taught is often depicted as
an apricot, a mistake stemming from use of the Chinese term hsing for “silver apricot.” For more on
Confucianism, ginkgo, and Wofo Si, the Temple of the Reclining Buddha, see Taylor and Choy (2005)
and Porter and Johnson (1993).
17. For a summary of ancient ginkgo trees in Japan see Hori and Hori (1997, 395). For a more com-
prehensive list, with many photographs of individual trees, see Hori and Hori, (2005). Yasukuni Jinja
is a Shinto shrine devoted to the spirits of those who died fighting in the service of the Emperor of
Japan since 1853.
18. See the Ginkgo Pages Web site for lists of many individual ginkgo trees in different parts of the
world, as well as much other fascinating information. See Zukowski, “In Hoboken, Trees for 9/11,”
for more on the Hoboken Memorial. Yoko Ono’s living sculpture in Detroit’s Times Square comprises
a ginkgo tree, a block of granite, and a bronze plaque with the inscription “WISH TREE for Detroit.
Whisper your wish, To the bark of the tree. Yoko Ono 2000 spring.” It is one of a series of Wish Tree
installations; others can be found in Brazil, California, Italy, and Japan. Ono explained that she was
moved by ginkgo trees in the temple gardens of Japan where she grew up, with tiny rolls of paper bear-
ing hopes and prayers adorning the branches like blossoms; see Nawrocki and Clements (2008, 49).
19. Six ginkgos survived the explosion of the atomic bomb at Hiroshima, one less than a mile from
its epicenter; see Kwant, “A-bombed ginkgo trees in Hiroshima, Japan.” On walks during his later
years, Harry Truman often paid tribute to the old ginkgo tree near his home in Independence, Mis-
souri, with a friendly pat and a few words. According to the pastor Thomas Melton, who accompanied
Truman on occasion, he told the tree, “You’re doing a good job” (McCullough, 1992). The ginkgo is
marked by a plaque as a stop on the Truman Historic Walking Tour; see Fischer (2010).
20. The Morton Arboretum is the second-oldest arboretum in the United States, after Harvard’s
Arnold Arboretum. Its seventeen hundred acres include more than forty- one hundred species of
trees and shrubs from around the globe. In addition to its role in conserving plant species and en-
gaging the public, the arboretum is committed to creating greener communities by planting and pro-
tecting trees in urban areas: see Ballowe and Klonowski (2003) and www.mortonarb.org.
21. For more on Goethe and Marianne Willemer see Unseld (2003). Goethe’s book Versuch die
Metamorphose der Pflanzen zu erklären—“An attempt to explain the metamorphosis of plants”
(Goethe, 1790)—is widely regarded as the first scientific study of plant form.
22. “From top to bottom a plant is all leaf ” is a translation from part of a letter that Goethe wrote
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n o t e s t o pa g e s 2 7 – 3 1

to Johann Gottfried van Herder, May 17, 1787. For more on Goethe’s search for a “Bauplan” and a
“fundamental organizational theme” in plant form, see Kaplan (2001).

4. Energy
1. Epigraph: Coolidge (1919, 13). According to Arnott (1959), some rejoining of the veins occurs
in 13.4 percent of ginkgo long-shoot leaves and 8.2 percent of short-shoot leaves, with at most five
junctions on a single leaf. On average, leaf vein unions are found on fewer than one in ten leaves. For
other studies of leaf venation in ginkgo see Florin (1936b) and Arnold (1947).
2. Creating artificial photosynthesis, and improving on its efficiency, which is two to three times
less than the best photovoltaic devices under optimal conditions, has long been a goal in solar energy
research: see Hohmann-Marriott and Blankenship (2011) and Blankenship et al. (2011). Gary Brud-
vig’s research group has created “artificial leaves” that split water into oxygen gas and hydrogen gas,
and that can be used to power a new kind of fuel cell. While solar panels provide electricity only dur-
ing the day, these new fuel cells potentially provide a constant energy supply.
3. It is possible to examine the stomata on well-preserved fossil ginkgo leaves. Measurements
of stomatal density from fossil ginkgo leaves have been used to trace changing levels of atmospheric
carbon dioxide in the distant past. When concentrations of atmospheric carbon dioxide are high, the
density of stomata needed to ensure an adequate supply of carbon dioxide into the leaves is low, and
vice versa; see Retallack (2001), Royer et al. (2001), Beerling and Royer (2002), and Royer (2003).
4. In almost all plants the leaves are the main site of photosynthesis, but in ginkgo the seed also
photosynthesizes. The nutritive tissues in the developing seed contain chlorophyll and provide a sig-
nificant contribution to the energy required for its own growth and ultimately growth of the embryo
that it nourishes. Light penetrates the fleshy coat and hard seed wall at levels equivalent to full shade
on a clear day; see Friedman and Goliber (1986).
5. Among the several kinds of photosynthesizing bacteria, the cyanobacteria (blue-green algae)
first exhibited the kind of photosynthesis seen in plants.
6. A single ginkgo leaf of average size may contain perhaps fifty million cells, based on rough
estimates from scanning electron microscope images of upper and lower ginkgo leaf surfaces and
several leaf cross sections. The so-called thylakoid membranes, in which chlorophyll molecules are
embedded, are approximately five nanometers thick (five millionths of a millimeter).
7. Another important class of light-gathering pigments, the carotenoids, is responsible for the
yellow and orange shades in the autumn colors of many deciduous trees.
8. As they branch and diverge from the leaf base, the veins in a ginkgo leaf gradually become
finer and contain fewer and fewer water-conducting cells (tracheids). A more simple but comparable
system is seen in the needles of many pines: see Zwieniecki et al. (2006).
9. The mucilage-like resin in ginkgo leaves is deposited in a single file in each interveinal area.
Late leaves contain much more resin than leaves produced early in the season; see Critchfield (1970).
This number of ginkgo trees on the streets of Japan reflects an increase of about twenty thousand
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n o t e s t o pa g e s 3 2 – 3 7

from the 1992 statistics cited by Handa et al. (1997); (Toshiyuki Nagata, Hosei University, personal
communication, 2011).
10. For more on factors that cause trees to shed their leaves, see Treshow (1970).
11. For more on the recovery of nutrients from senescing leaves, see Andersson et al. (2004),
Buchanan-Wollaston (1997), and Killingbeck (1996). “The Two Ginkgo Leaves,” by Otto Crusius,
former president of the Bavarian Academy of Sciences, cited in the translation of Unseld by North-
cott (Unseld, 2003): “Ginkgo leaves tired of summer, brilliant as a brimstone butterfly. Flutter down
onto the bench, Flutter, whisper, ‘do you remember?’”
12. The Meiji-Jingu Park in the Kasumigaoka district of Tokyo commemorates the Meiji Emperor
who died 1912. Completed in 1926, it contains sports and cultural facilities the centerpiece of which
is Seitoku Memorial Museum (Meiji Memorial Picture Gallery). The gallery is approached along an
avenue about a quarter of a mile long with ginkgo trees on either side: see Handa et al. (1997, 272).
Similarly, at the Memorial Showa Garden, constructed on the site of a former American military base,
the long central canal that is the centerpiece of the landscape design is flanked by a double row of
ginkgos planted in 1983; see Handa et al. (1997).
13. The ginkgo of the Dorsch Public Library in Monroe is the partial namesake for the Lotus
Ginkgo Show, the self-described longest-running program on Michigan’s Monroe Public Access Cable
Television.
14. “The Consent,” Nemerov (1977, 476). A small group of ginkgo trees that grew outside Neme-
rov’s office on the campus of Washington University in Saint Louis was the inspiration for this poem
(Peter Raven, Missouri Botanical Garden, Saint Louis, personal communication). This poem has a
different tone from “Ginkgoes in Fall,” appearing one page before: “Their fallen yellow fruit mimics
the scentˆ/ˆOf human vomit”; the same leaves described as “fluttering fans of light” are “filtering a
urinary yellow light” in the other; Nemerov (1977, 475).

5. Growth
1. The comment on artistic portrayals of trees is from Hargraves (2010). Ginkgos are easy to spot
from a distance, especially in winter, from their distinctive pattern of branching. When I have occa-
sionally been confused, the other tree has generally turned out to be a conifer, most often a pine.
2. The tradition of learning how to identify trees from their twigs in winter comes from Germany,
and for more than a century, German botany students have benefited from Camillo Karl Schneider’s
classic Dendrologische Winterstudien, which includes photographs and illustrations of bark and buds
from 434 species of trees and shrubs; see Schneider (1903). A friend at Berkeley tells the story of a
colleague who grabbed a naked ginkgo twig hoping to bamboozle a prospective Ph.D. student. He left
chastened when the candidate quickly identified the specimen as ginkgo (Bruce Baldwin, University
of California, Berkeley, personal communication).
3. During the April 2007 cold snap, gardens all across the midwestern United States suffered
as spring flowering trees and shrubs were damaged by wind chills that fell to minus 30° Fahrenheit
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(roughly minus 34° Celsius). Because ginkgo is pollinated at the same time that the leaves emerge (see
Chapter 7), the cold snap also reduced significantly the production of ginkgo seeds that year.
4. For more on the expression and development of short and long shoots in ginkgo, see Gunckel
et al. (1949).
5. See Chapter 3 and Unseld (2003) for more on Goethe and the significance of the poem he sent
to Marianne Willemer. See Chapter 28 for more on Linnaeus and the origin of the name of ginkgo.
6. Studies by Leigh et al. (2010) on the structural and physiological differences between long- and
short-shoot leaves document an interesting difference in vein density. Even though long-shoot leaves
have a lower density of veins than short-shoot leaves, they are significantly more effective at conduct-
ing water. It is not completely clear how this is accomplished, which highlights the fact that there is
still much research to be done before we fully understand how the strange leaves of ginkgo actually
work. One possibility, which needs to be tested by future measurements and experiments, is that the
veins in long-shoot leaves are better connected to the upper and lower leaf epidermis and that the
epidermal cells play an important role in leaking water out of the veins and into the surrounding leaf
tissues.
7. The cells in the outer part of the woody cylinder through which the water passes in most trees
are dead. They form from the cylinder of actively dividing living cells (cells of the cambium), but once
formed they quickly undergo preprogrammed cell death; see Chapter 6.
8. Early in the season in many trees, as the sap begins to rise, sugars stored in the roots and the
living tissues in the lower part of the stem, including the ray cells in the wood, are mobilized and
taken into solution within the cells. This draws in water from the soil, which expands the volume of
fluid in the lower parts of the plant and causes the sap to rise. This is the process that is exploited by
tapping a sugar maple, and in this special circumstance, at this particular time of the year, there is
pressure from below.
9. The volume of water taken up by the yevaro tree (260 gallons a day) was the maximum re-
corded in a review of fifty-two studies conducted over thirty years on sixty-seven species. In this re-
view, 90 percent of trees around sixty-five feet in height take up only between 2 and 44 gallons of
water a day. For more details and a breakdown of water requirements by species, see Wullschleger
et al. (1998).
10. The same minute pores are also present in conifers. For more on the structure of ginkgo wood,
see Dute (1994).

6. Stature
1. Epigraph: from Eknath Easwaran’s translation of the classic Buddhist text (Easwaran, 2007,
126). For a biography of Philipp Franz von Siebold (1796–1866) see Thiede et al. (2000), Kouwen-
hoven and Forrer (2000), and the exhibits and publications of the Siebold Museum in Leiden (www
.sieboldhuis.org) and Nagasaki (www.city.nagasaki.nagasaki.jp/siebold). Glover’s house still stands
above Nagasaki harbor, with statues of Puccini and Cho-Cho San looking out to sea from the garden.
For more on the origins of Madame Butterfly, see Van Rij (2001).
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2. The Siebold Museum in Nagasaki, modeled on Siebold’s former house in Leiden, is located
next to the former site of Siebold’s medical school, Narutaki Juku.
3. In Flora Japonica, Siebold named Hydrangea otaksa after his nickname for Sonogi. The ac-
cepted name for this species now is Hydrangea macrophylla (Thunb.) Ser. Today, bigleaf hydrangea,
as it is commonly known, is the most popular species of hydrangea planted in home gardens (Kou-
wenhoven and Forrer, 2000).
4. The map supplied to Siebold had been completed in 1818 and engraved only in 1823. It was the
result of exhaustive surveying of the coasts and islands of Japan by a team of fourteen and was the
most detailed account for northern Japan then available; see Murdoch (2004, 555–558).
5. An alternate version of the incident from Murdoch’s A History of Japan, based on Siebold’s
diary entry for December 16, 1828, describes his betrayal by Yoshio Tsujiro, an interpreter who was
helping him translate from Japanese books; see Franz (2005, 37); Totman (1993, 510). Siebold tran-
scribed the maps for the archives at Deshima before handing them over.
6. Oine was an inquisitive child and at the age of nineteen was instructed by Ninomiya Keisaku
in obstetrics. She was appointed to the post of imperial obstetrician in 1877.
7. For more on the life of Oine see Kouwenhoven and Forrer (2000, 24).
8. All three works were published in multiple parts over the next quarter century. Nippon (Sie-
bold, 1832–1852) describes the ethnography and geography of Japan, including an account of his
journey to Edo. Fauna Japonica (Siebold et al., 1833–1850) was a series of monographs based on
the collections of Siebold and his Deshima successor, Heinrich Bürger. Flora Japonica built on the
work of Siebold’s predecessors Kaempfer and Thunberg and was undertaken in collaboration with
the German botanist Joseph Gerhard Zuccarini (Siebold and Zuccarini, 1835–1870). It was begun in
1835 and came to a halt after the death of Zuccarini in 1848, but ultimately, after Siebold’s death in
1866, F. A. W. Miquel from the National Herbarium in Leiden published some additional parts. After
thirty-five years, Flora Japonica was completed in 1870. The first volume contained twenty parts and
the second volume ten. Plate 136, a superb illustration of Ginkgo biloba, was published in the second
volume (see Chapter 5).
9. After William Jackson Hooker’s death, his library, a collection of around 4,000 volumes, was
purchased by the British Government for £1,000 in 1866; an additional £1,000 bought his correspon-
dence, manuscripts, portraits, and other miscellanea. The collections of botanical art at Kew comprise
more than 200,000 items, including astonishing work from the eighteenth, nineteenth, twentieth,
and twenty-first centuries. Most of these paintings are housed in an annex to the Kew Library, with
selections on display in the Shirley Sherwood Gallery of Botanical Art and the Marianne North Gal-
lery.
10. The Berlin collection was acquired in 1911 from Paul Kuegler, a senior staff physician in the
German navy. He probably obtained the collection in Japan, toward the end of the nineteenth cen-
tury (Lack, 1999). The Koishikawa Botanic Garden holds a set of twenty-five boards in the same style.
Harvard University Museum also has a small collection of boards brought to New England by Edward
Sylvester Morse, who was the first professor of zoology at Tokyo University between 1877 and 1879.
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There is also a small private collection in London. The Koishikawa Botanic Garden, or Koishikawa
Shokubutsuen, was founded as the Koishikawa Medicinal Herb Garden in 1684 by the Tokugawa
Shogunate, and was the birthplace of modern Japanese research in plant science following the Meiji
Restoration. It contains one of the most famous of all ginkgo trees: see Chapter 9.
11. Chikusai Kato prepared sketches of ginkgo for Koishikawa-shokubutsuen-somoku-zusetsu
(Illustrations of the trees and herbs in the Koishikawa Botanical Gardens), edited by Keisuke Ito
(1802–1901) and Hika Kaku (1786–1862); see Ito and Kaku (1881–1883). Kaku’s elder brother studied
under Siebold. Keisuke Ito gave Siebold fourteen collections of dried plants, including a specimen of
ginkgo, to bring back from Japan.
12. The way that rays are cut in preparing a piece of timber is often responsible for the distinctive
“grain” of certain high-quality woods. In ginkgo the rays are not sufficiently thick to impart a distinc-
tive grain. For more on ray cells and their development in ginkgo wood, see Barghoorn (1940, 321).
13. Extreme cork production occurs in the cork oak, the main source of corks for wine bottles. The
high suberin content of cork oak bark also makes it more pliant and water resistant than the bark of
other trees.
14. The dead, water- conducting cells, the so- called tracheids, reach up to two to three thou-
sandths of an inch wide and a tenth to three-tenths of an inch long in the trunk and nearly four-
tenths of an inch long in roots. The specialized water-conducting cells (vessel elements) in the wood
of many flowering plants may be an order of magnitude longer, up to about fourteen inches (Sperry
et al. 2006; Wilson and Knoll, 2010). The tiny valves in the water- conducting cells of ginkgo, the so-
called torus-margo structures associated with the pit membranes, improve water conductivity by
minimizing hydraulic resistance while also preventing the formation of embolisms; see Hacke et al.
(2004) and Pitterman et al. (2005).
15. Eleven annual rings are visible in the small branch that makes up the top left-hand corner of
the ginkgo board in the Kew xylotheque. The rings on the other branches are obscured by tool marks
and varnish (Mark Nesbitt, Royal Botanical Gardens, Kew, personal communication).
16. The different uses of ginkgo wood are listed by Hori and Hori (1997).
17. Ojiya is a small town known as an important center for the breeding of koi carp. It gained
notoriety as the epicenter of a fatal, magnitude 7.2 earthquake in October 2004. Li Shizhen recorded
the use of ginkgo wood by Daoist shamans in his A.D.1596 herbal Bencao Gangmu, writing, “The
wood of the tree is white with fine texture and lasts a long time. Alchemists carve chops with the wood
saying that such thing is good to summon the spirits” (Shizhen and Xiwen, 2003).
18. The legacy of carved wooden Buddhas, mokujikibutsu, that Mokujiki Shonin left across Japan,
deeply moved Yanagi Sōetsu, the philosopher and founder of the mingei (folk art) movement in
Japan. Yanagi retraced Mokujiki’s original route, cataloguing the mokujikibutsu, which he described
as “simple,” “natural,” and “ego-less” in their beauty and tradition; see Kibuchi (1997). See also the
drawing that introduces Part VII.
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7. Sex
1. Epigraph: Dawkins (1976, xxi).
2. Camerarius (1665–1721) reported his results on reproduction in plants in his 1694 publication
De sexu plantarum epistola. The botanists John Ray (1627–1705) and Nehemiah Grew (1641–1712) were
among the earliest fellows of the Royal Society, the U.K. national academy of science.
3. According to Science and Civilization in China, the earliest reference to ginkgo comes from
the Ko Wu Tshu Than (Simple discourses on the investigation of things), written by Lu Tsan-Ning, a
“learned monk.” This is a series of short statements about natural phenomena, written about A.D.
980; see Needham (1986, 491) and Chapter 24.
4. The evolutionary transition from a species composed entirely of hermaphrodite individuals
to species with separate male and female individuals (dioecy) is best documented among flowering
plants. In many cases remnant or vestigial organs from the other sex remain in the flower, suggesting
that the separation into male and female plants occurred relatively recently in evolutionary history.
Based on observations like these, Darwin (1876, 1877) developed his initial ideas on the evolutionary
advantages of separate male and female plants, which have been further elaborated by later evolu-
tionary biologists (e.g., Charnov et al., 1976; Lloyd, 1982). Dioecy, monoecy (separate male and female
flowers on the same plant), and dichogamy (separation in the timing of maturation of male and
female parts of the same flower) in flowering plants are all effective in promoting cross-pollination
between individuals, which increases the genetic variation in the next generation upon which natural
selection can act.
5. For description of the development of ginkgo pollen cones, see Liu et al. (2006). See Chris-
tianson and Jernstedt (2009) for details on the position of pollen cones and seed-bearing structures
in ginkgo.
6. Ginkgo pollen grains probably do not remain viable for long after they are shed, but under
laboratory conditions they can remain alive for up to sixteen months (Newcomer, 1939). Under ster-
ile conditions Tulecke (1954) achieved a germination rate of 35 to 45 percent for ginkgo pollen after
storage for two years.
7. Andrew Leslie’s estimate is based on observations of trees growing outside the Hinds Geo-
physical Laboratory at the University of Chicago with the following estimated parameters: a single
pollen sac contains about twenty thousand pollen grains; each pollen cone contains about seventy-
seven side branches, each with two pollen sacs, or about three million grains per cone; each short
shoot produces about seven cones, and there might be around 17,500 short shoots on a forty-foot tree.
8. Often in trees with separate sexes, males begin to produce pollen slightly before females pro-
duce ovules, which is consistent with theoretical predictions: see Lloyd and Webb (1977).
9. After pollination, the dried mucilaginous residue from the pollination drop seals the micro-
pyle, and pollen development and ultimately germination proceed within the sealed cavity (Lee,
1955). For more on how pollination drops function in conifers, see Takaso (1990), and for video
imagery of pollination and fertilization in ginkgo and cycads, see the Tokyo Cinema film The Sea in
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n o t e s t o pa g e s 5 7 – 6 2

the Seed. A pollen tube is formed once the pollen has been taken into the micropyle, and in ginkgo
and cycads the tube appears to be modified for nutrient uptake. It penetrates the nutritive tissues in
the central part of the ovule, forming extensive branching networks of fine haustoria among the cells;
see Friedman and Gifford (1997) and Chapter 9.
10. Occasionally seed-bearing stalks have only a single seed at their tip or several, but two is the
norm. The development of more than one mature embryo in a single seed occurs in about 2 percent
of ginkgo seeds and also occurs occasionally in conifers: see Cook (1902, 1903), Buchholz (1920), and
Berlyn (1962) for additional information. For a photograph of two seedlings emerging from a single
ginkgo seed see Stuppy et al. (2009, 24).
11. For more on the development of the embryo of ginkgo, see Lyon (1904). The fleshy outer layer
appears to inhibit germination if it is not removed (Rothwell and Holt, 1997). Ginkgo differs from
most conifers in having hypogeal germination: the cotyledons remain embedded in the nutritive tis-
sue of the seed, often underground. Only the lower parts of the cotyledons project from the seed shell;
see Seward and Gowan (1900, 116), Chick (1903).

8. Gender
1. Epigraph: Angelou (1990).
2. An important question is why there is apparently so little change in the evolutionary history
of ginkgo, as well as in other classic examples of so-called stasis. Standard explanations suggest that
either ginkgo has tracked the same environmental conditions for more than 200 million years or the
same form (perhaps in spite of genetic variation) has been maintained by strong stabilizing selec-
tion. However, a third possibility is that there is some kind of strong, inbuilt constraint during devel-
opment that has kept ginkgo more or less unchanged for a long period of time. Potentially, all three
factors may be at work.
3. Joseph Franz von Jacquin (1766–1839) succeeded his father, Nikolaus Joseph von Jacquin
(1727–1817), as professor of botany and director of the botanical garden at the University of Vienna.
The family lived in a house on the Rennweg, near the modern Institute of Systematic Botany of the
university, close to the Belvedere Palace. Along with his brother and sister, Joseph von Jacquin was
taught the piano by Mozart. The composer, part of his father’s aristocratic circle, regularly visited the
Jacquin home. Mozart’s “Kegelstatt Trio,” dedicated to the Jacquin family, was first performed at their
home in August of 1786 by Franziska, Joseph Jacquin’s sister.
4. Nikolaus von Jacquin had connections to the palace at Schönbrunn, for which he worked as
a plant collector from 1755 to 1759. An account of early ginkgos in Europe is provided by Loudon
(1838); see also Jacquin (1819) and Chapter 30.
5. August Pyramus De Candolle was the first to recognize ovule-bearing shoots on a ginkgo in
Europe at Bourdigny, a village about six miles outside Geneva, in 1814. See also Chapter 30. The Bour-
digny tree was cut down in 1866 by a new owner of the estate; see Wilson (1920, 56). Jacquin was the
first botanist to use grafting for a scientific purpose. He published the results from his grafting experi-
ment in the same year that Goethe first published his ginkgo poem (Jacquin, 1819).
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6. The slightly delayed development of the female branch is consistent with theoretical predic-
tions: see Lloyd and Webb (1977).
7. The experiments carried out by Hugo de Vries at the University of Amsterdam (commemo-
rated by a plaque in the garden) were some of the most important ever done in a botanical garden.
The term genes was introduced by the Danish botanist Wilhelm Johannsen in 1909. Johannsen also
was the first to use the terms phenotype and genotype.
8. Theodor Boveri (1862–1915) worked in Germany while Walter Sutton (1877–1916) worked in
the United States. Thomas Hunt Morgan (1866–1945) received the Nobel Prize in 1933 for his role in
linking chromosomes with heredity. The research of Nettie Stevens (1861–1912) at Bryn Mawr College
and of Edmund Beecher Wilson (1856–1939) at Columbia was conducted on the mealworm, the lar-
vae of a species of beetle (Tenebrio molitor), which has the advantage of having relatively large chro-
mosomes that are easily observed.
9. Human males and females both have twenty-two pairs of matching chromosomes in all their
cells, but in the twenty-third pair, the sex chromosomes, the two chromosomes differ in the male.
Females have two matching copies of chromosome twenty-three, which are designated XX. Males
have one long chromosome, the X chromosome, and one short chromosome, the Y chromosome.
10. The first chromosome count for ginkgo was reported by the Japanese scientist Ishikawa
(1910).
11. The suggestion of an XY-type sex determination system in ginkgo was based on several early
studies that reported a satellite on the arm of chromosome eleven (Tanaka et al., 1952; Newcomer,
1954). Later research found various satellites associated with various chromosomes in male and
female plants (e.g., Ho, 1963; Chen et al., 1987). For more on the chromosomes of ginkgo see Hizume
(1997).
12. A similar situation occurs in certain animals where the sex of offspring is determined after the
embryo has begun to develop and depends on environmental conditions. For example, in many liz-
ards and turtles higher temperatures during a critical phase in the development of the embryo favor
the production of females, while in alligators they favor the production of males.
13. The story of the graft on the Old Lion at Kew is recounted in Bean (1973). See Crane (2006) for
the report of seeds on the same tree. The example in the Jena botanical garden is cited by Melzheimer
and Lichius (2000).
14. “Witches’ brooms,” dense masses of shoots growing from a single point often high up in the
canopy of a tree, can be caused by damage of various kinds—for example, by pests or diseases, as
well as by human interference. The result is a kind of cancer, an abnormal proliferation of growth,
probably caused by a local loss of control of normal development in that part of the plant. In the Cave
Hill tree the result has been to change the sex of that part of the tree. The Japanese botanist Seiichiro
Ikeno, who had earlier been the first to observe swimming sperm in cycads, also noted that male
ginkgos can occasionally produce seeds; see Ikeno (1901), Miyoshi (1931).
15. Observations at the Blandy Experimental Farm are reported by Santamour et al. (1983a).
16. Lloyd and Webb (1977).
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9. Seeding
1. Epigraph: Shakespeare (1623c), act 1, scene 3. The Great Kanto Earthquake struck on Septem-
ber 1, 1923, and the most destructive air raid on Tokyo took place on March 10, 1945. Information
about the Koishikawa Ginkgo was provided by Tetsuo Ohi-Toma, Koishikawa Botanical Gardens,
University of Tokyo (personal communication). The ginkgo at the Koishikawa Botanical Garden
was visited by His Majesty the Emperor of Japan and Her Majesty the Empress of Japan in 2006 (His
Majesty the Emperor of Japan, 2007).
2. The Koishikawa Ginkgo was planted around 1680, and an attempt was made to fell it in August
1868. The scars from the axe cuts made at the time were visible until about fifty to seventy years ago.
Part of the motivation in trying to fell the tree may have been profit (Tetsuo Ohi-Toma, University
of Tokyo, and Toshiyuki Nagata, Hosei University, personal communications; see also Primack and
Ohkubo, 2008).
3. The University of Tokyo has undergone various name changes over the years, from Tokyo Uni-
versity (1877–1886), to Imperial University (1886–1896), to Tokyo Imperial University (1896–1948),
and finally to the University of Tokyo (1948–present). Ryokichi Yatabe, the first Japanese graduate
of Cornell University, became the first professor of botany and curator of botanic gardens at Tokyo
University. Jinzo Matsumura was the second professor and curator. Keisuke Ito, a student of Siebold,
oversaw the garden in the late nineteenth century and retired from the university in 1886. Appointed
at the age of seventy-five, he was given a special title and did not have to teach (Toshiyuki Nagata,
Hosei University, personal communication).
4. Stopes studied at Tokyo Imperial University during the time (1897–1935) when the Department
of Botany was based at the Koishikawa Garden. The garden is now part of the Department of Bio-
logical Sciences, which is located on the main campus (Toshiyuki Nagata, Hosei University, personal
communication). Marie Stopes’s visit to Japan was funded by a fellowship from the Royal Society. The
studies of Cretaceous petrified fossil plants from Hokkaido, initiated by Stopes and Fujii (1910) in the
course of that fellowship, continue today through Harufumi Nishida at Chuo University, Tokyo; see
Nishida (1991) for a review. Stopes and Fujii became involved in an ill-fated love affair. Fujii, who was
married, feigned leprosy to break it off. Stopes later published their letters under the pseudonym of
G. N. Mortlake as Love Letters of a Japanese; see Mortlake (1921) and Hall (1977).
5. This list of significant books was compiled in 1935 by asking a number of American academics
to name the twenty-five most influential books of the previous fifty years (Hall, 1977). More on Marie
Stopes can be found in several biographies; see Hall (1977), Briant (1962); for selections of Stopes’s
writing, see Stopes (1918), Garrett (2007).
6. According to some sources, the University of Manchester attempted to retract its offer when
administrators realized that Stopes was a woman, but she nevertheless was appointed. Marie Stopes
was as flamboyant as she was brilliant. Bill Chaloner, one of my paleobotanical mentors, recalls meet-
ing her in 1952 at the Geological Society of London. After he had explained some of his research to
her, she exclaimed loudly so all could hear, “Ah dear boy, that is wonderful! Of course, paleobotany
was my first love!” (Chaloner, 2005).
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7. Details of sexual reproduction in ginkgo, based on the material sent from Vienna, were pub-
lished by Strasburger (1892). German students of plant science still study from a revised version of
the massive textbook that Strasburger wrote. Hirase worked at the botanical laboratory of the College
of Science of the Tokyo Imperial University. Soon after his discovery of motile sperm in ginkgo, the
Department of Botany was moved to the Botanical Garden at Koishikawa, where it remained from
1897 until 1935 (Toshiyuki Nagata, Hosei University, personal communication).
8. Hirase had been appointed by Ryokichi Yatabe as the first professor of botany at the Imperial
University, Tokyo. Initial observations on fertilization in ginkgo were published by Hirase (1895a, b),
but the key paper recognizing the motile sperm appeared in 1896 (Hirase, 1896). According to Singh
(2006, 236), Hirase began his career as a technical illustrator, but taught himself botanical techniques
and began to study ginkgo fertilization and embryo formation in 1893. He made his discoveries while
preparing microscope slides of ginkgo ovules, when he observed a peculiar ellipsoid body with an
attached coiled band in the pollen tube. He correctly noted that it might be a spermatozoid in a lec-
ture on April 25, 1896, and continued cutting and examining ovules until at last finding motile sperm
a few months later on September 9. An excellent film of the swimming sperm, and many other aspects
of the biology of ginkgo, is available in The Sea in the Seed.
9. Not long after Hirase made his discovery, his most important mentors left the Imperial Uni-
versity. Hirase left soon after. Because he lacked formal botanical training, the strictly hierarchical
system may have made his position awkward. For the rest of his career Hirase worked as a school-
teacher; see Ikeno and Hirase (1897) and Nagata (1997) for more on Hirase and his discovery.
10. Marie Stopes’s observations on ginkgo sperm came exactly twelve years after Hirase’s observa-
tion: see Stopes (1910, 218). Using the dates recorded in Stopes’s journal, Professor Toshiyuki Nagata,
a former director of the Koishikawa Garden, collected ginkgo seeds from the same and other trees
on September 9, 1997. He found that fertilization within each tree was broadly synchronized, but that
there were differences in timing among trees.
11. The work by Oliver and Scott was published in 1903 and 1904; for more on its significance see
Andrews (1980). Since Oliver and Scott’s work many different kinds of fossil seed plants have been
grouped together as seed ferns, but studies of the relationships among living and fossil seed plants
show that seed plants are a very heterogeneous and unnatural group of diverse relationships (e.g.,
Crane, 1985; Doyle and Donoghue, 1986; Hilton and Bateman, 2006).
12. Aberrant ovule-producing leaves in ginkgo were first described by Shirai (1891) and then by
Fujii (1896). When fertilized, the ovules developed into seeds, although these remained smaller than
those of normal trees. The leaves bearing ovules and pollen sacs were smaller than normal leaves, and
those leaves bearing viable seeds were even smaller still. The significance of such aberrations is still
debated.
13. See Favre-Ducharte (1958) and Eames (1955). At EWHA University, Seoul, South Korea, I col-
lected nearly mature seeds that fell from several trees well before the normal time for fertilization.
The seeds were left outside for several months, and a few produced viable embryos. More detailed
work would be needed to securely establish that fertilization occurred after the seeds were shed.
14. Liz Jaeger, councilor for Whitton (personal communication). Masamichi Takahashi, Niigata
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University (personal communication; see also Kochibe, 1997). Ginkgolic acid may also be called gink-
goic acid.

10. Resilience
1. Epigraph: Gandhi (1961, 133).
2. In Utah, a single clonal colony of quaking aspen called Pando is considered to be the world’s
heaviest, and by some measures, oldest organism. With more than forty-seven thousand stems grow-
ing over 107 acres, the plant weighs an estimated six thousand tons. Though the average stem age is
130 years old, according to some estimates, genetically identical plants—parts of the same clone—
may have existed ten thousand or more years ago. See also Chapter 24.
3. Chi-chi has also been translated as “nipples” in Japanese. Fujii (1896) investigated the internal
anatomy of ginkgo chi-chi and showed that close to the point of attachment to the parent shoot each
contains an embedded short shoot, the buds of which keep growing so as to maintain their position
on the surface of the downward growth. These buds have the potential to burst into life when they
reach the ground, or even before, and produce new upward-growing shoots. For more on the devel-
opment of chi-chi see Barlow and Kurczyńska (2007).
4. For more on the development of ginkgo lignotubers see Del Tredici (1992a).
5. See the drawing that introduces Part II for an illustration of the Kitakanegasawa Ginkgo.

11. Origins
1. The founding collections of the Swedish Museum of Natural History were the collections of
the Royal Swedish Academy.
2. Alfred Nathorst (1850–1920) retired in 1919; see Seward (1921) and Andrews (1980) for brief
biographies.
3. Nathorst’s studies of the Spitsbergen fossils are revised and discussed by Kvaček et al. (1994).
See Schweitzer and Kirchner (1995) for additional information on Ginkgo cordilobata and the other
fossil plants with which it occurs. The Stockholm collections include about thirty of Nathorst’s
ginkgoalean specimens (Else Marie Friis, Swedish Museum of Natural History, personal communi-
cation).
4. In hindsight, Mackie recalled discovering a piece of plant-bearing chert around 1880, more
than thirty-five years before he first announced his discovery; see Mackie (1913, 225) and Trewin
(2004).
5. The Rhynie Chert formed in the Early Devonian (Kenrick and Crane, 1997).
6. Clubmosses are still native to Scotland, and Asteroxylon is especially similar to the fir club-
moss, a plant that is common in the Scottish Highlands.
7. See Kenrick and Crane (1997) for an overview and analysis of the early fossil record of plants
on land.
8. The closest living relatives of land plants are the freshwater “charophycean green algae,” which
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n o t e s t o pa g e s 8 9 – 9 4

include the stonecrops. This provides additional support for the idea that the land was colonized by
plants from freshwater rather than directly from the sea.

12. Ancestry
1. Epigraph: According to a letter written by Huxley to his friend Dr. Dryster about two months
after the debate, his response was: “If then, said I, the question is put to me would I rather have a
miserable ape for a grandfather or a man highly endowed by nature and possessed of great means of
influence and yet who employs these faculties and that influence for the mere purpose of introduc-
ing ridicule into a grave scientific discussion, I unhesitatingly affirm my preference for the ape.” For
a complete account of the Wilberforce-Huxley encounter and the associated controversies, see Jensen
(1988).
2. The Southern African paleoflora from the Devonian to the Cretaceous is described by Ander-
son and Anderson (1985). See Anderson and Anderson (1983, 1989, 2003, 2008) and Anderson et al.
(2007) for descriptions of and context for fossil plants from the Molteno Formation.
3. For a review of ancient ginkgolike fossils from North America see Ash (2010). For an early
occurrence of ginkgo species from the early Middle Triassic of Australia see Holmes and Anderson
(2007).
4. The two most informative of the three specimens of Trichopitys heteromorpha studied by
Florin (1949) from Lodève, France, were formerly in the collection of the École Nationale Supérieure
des Mines de Paris but are now at the University of Lyon. The other is in the Natural History Museum
in London. However, Florin did not examine Saporta’s original material, which is in the Natural His-
tory Museum in Paris. The specimen of Trichopitys illustrated by Taylor et al. (2009, 745) is from
another locality at Montpellier (Hans Kerp, University of Münster, personal communication) and is
preserved in a different way. Even though shoots with multiple ovules are occasionally produced in
living ginkgo, the ovules rarely develop to maturity. The largest number of mature seeds I have seen
on a single shoot is three.
5. See Meyen (1988, 344–346) for his interpretation of Saporta’s original material of Trichopitys.
6. For treatment of Permian fossils from Argentina similar to Trichopitys see Archangelsky and
Cúneo (1990). More likely candidates for Permian members of the ginkgo lineage are plants that pro-
duced Permian Sphenobaiera-like leaves, some of which have leaf cuticles and resin similar to ginkgo.
Many of these Sphenobaiera leaves have been described under different names such as Ginkgophyton
and Ginkgophytopsis (Zhou Zhiyan, Nanjing Institute of Geology and Palaeontology, personal com-
munication; see also Zhou, 2009).
7. See Meyen (1984).
8. Kerpia has some similarities to pinnate leaves assigned to Psygmophyllum. The fossil seed clus-
ters were assigned the genus Karkenia; see Naugolnykh (1995, 2007) and Chapter 15 for more on Kar-
kenia.
9. In ginkgo, fusion among the veins on the leaf, which creates reticulations, does occur but is
relatively rare; see Chapter 4.
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n o t e s t o pa g e s 9 4 – 1 0 1

10. Glossopterid leaves (Glossopteris) have been discovered in Antarctica, Australia, India, South
Africa, and South America. Continental drift is often referred to as the theory of plate tectonics.
11. The remarkable occurrence of probable sperm in glossopterid fossils was reported by Nishida
et al. (2004), based on pollen grains preserved inside a fossil ovule. The illustrations show what ap-
pear to be the bases of flagellae arranged in a spiral as in living ginkgo.
12. Anderson and Anderson (2003) and Anderson et al. (2007) suggest that the seed-bearing
structures Kannaskoppia may be similar to fossils previously described as Petriellaea (Taylor et al.
1994). Although fossils of Petriellaea are much more fragmentary than fossils of Kannaskoppia, some
details of their structure are better understood, and there are two to six small ovules inside each
recurved cuplike structure. Each ovule is triangular in cross section. Petriellaea is quite different
from the seed-bearing structures of ginkgo but the differences may be possible to reconcile (see also
Meyen, 1984); in this regard the recurved ovules of Karkenia may be significant.

13. Relationships
1. A synopsis of Hennig’s ideas appeared in English in 1965 (Hennig, 1965). Hennig’s book was
translated into English mainly by Rainer Zangerl, a specialist in fossil fishes and one of my predeces-
sors as chairman of the Department of Geology at the Field Museum in Chicago; see Hennig (1966).
Something of the flavor of the vigorous and often vituperative debates around the development of
cladistics, and the personalities involved, can be gleaned from Hull (1988).
2. The group of plants defined by the production of wood (secondary xylem) is termed ligno-
phytes.
3. These numbers are the number of potential rooted phylogenetic trees, based upon the most
recent common ancestor of all the associated entities. The formula to calculate the number of rooted
trees for n different organisms is: (2n−3)!/(2(n’²)×(n−2)!). For further explanation see http://www
.scientific-web.com/en/Biology/Evolutionary/PhylogeneticTree.html. For three taxa the number of
rooted trees is 3, for four taxa it is 15, for five taxa it is 105, for six taxa it is 945, for seven taxa it is
10,395, for eight taxa it is 135,135, for nine taxa it is 2,027,025.
4. Cladistic analysis is a method used to build these so-called phylogenetic diagrams, or clado-
grams (sometimes called phylogenetic trees) and to test which provides the simplest explanation of
the data.
5. A classic early paper that used molecular data to develop a new understanding of the relation-
ships among a wide range of flowering plants was coordinated by Mark Chase at the Royal Botanical
Gardens, Kew; see Chase et al. (1993). For an overview of our current understanding of relationships
among major groups of angiosperms, based mainly on DNA data, see Stevens (2008); for more on
the angiosperm fossil record see Friis et al. (2011). For a formal classification of angiosperms based
on recent phylogenetic discoveries see APG (2009).
6. The Gnetales include three superficially different but fundamentally similar kinds of plants;
the genus Ephedra, sometimes referred to in North America as Mormon tea, the genus Welwitschia,
a bizarre plant that is well known in Namibia and produces only two leaves in its entire life; and the
 303
n o t e s t o pa g e s 1 0 3 – 1 0 6

genus Gnetum, a tree or climber of tropical forests with leaves that look very like those of angiosperms
(flowering plants).
7. The first paper to apply cladistic analysis to the study of relationships among the major groups
of living plants was Parenti (1980). This was followed by a more detailed treatment of living seed
plants by Hill and Crane (1982) and later by studies that included living and fossil seed plants in the
same cladistic analysis (Crane, 1985; Doyle and Donoghue, 1986). There have been many cladistic
analyses since, but the real need is to understand a broader range of fossil plants in greater detail. For
example, there remains much potential for paleobotanical collecting in the cherty layers of the Mol-
teno Formation and for comparison of the Molteno seed plants with increasingly well-known fossils
of similar age from Antarctica (John Anderson, personal communication); see also Taylor and Taylor
(2009) for fossil seed plants from Antarctica.

14. Recognition
1. Epigraph: Wieland (1768, canto II).
2. The work carried out by Zhou Zhiyan at Reading under Harris’s supervision resulted in the
description of a strange conifer, Stalagma samara (Zhou, 1983). The details of the life and work of
Thomas M. Harris (1903–1983) in this chapter lean heavily on the biography written by his student
and one of my own paleobotanical mentors William G. Chaloner (1985).
3. Harris’s work on fossil plants from Yorkshire covered mosses, liverworts, clubmosses, and
ferns (Harris, 1961); cycads and various extinct seed plants (Harris, 1964); an important extinct group
of seed plants, Bennettitales (Harris, 1969); ginkgo and its possible relatives (Harris et al., 1974); and
conifers (Harris, 1979).
4. Harris’s B.Sc. was from the University of London. At that time the University of Nottingham
did not have its own degree-awarding authority. For more on H. S. Holden, including his work in
forensic science, see Andrews (1980). Paleobotany first took root in Manchester when William C.
Williamson was appointed to the Chair of Natural History at Owens College in 1851, which eventually
became the Victoria University of Manchester in 1880 (one of the two institutions that later merged
to form the modern University of Manchester). In addition to Williamson other influential paleo-
botanists who have been associated with the University of Manchester over more than 150 years have
included Marie Stopes, Ernest Weiss, William Lang, John Walton, Isabel Cookson, and Joan Watson.
For a full account of the history of paleobotany in Manchester, see Watson (2005).
5. Sir Albert Charles Seward (1863–1941) served as vice chancellor of Cambridge University in
1924 and 1925; see Andrews (1980, chapter 6) for an engaging account of Seward’s life and work.
Francis Darwin (1848–1925) was also a Cambridge don and a specialist in plant physiology.
6. The overused “abominable mystery” quotation is from a letter that Darwin wrote to Joseph
Dalton Hooker on July 22, 1879. Hooker was then director of the Royal Botanic Gardens, Kew (Dar-
win and Seward, 1903). See Friedman (2009) for a modern discussion of the background to this fa-
mous quotation and its meaning.
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n o t e s t o pa g e s 1 0 6 – 1 1 6

7. See Chaloner (1985) and Andrews (1980) for the background to Harris’s work on fossil plants
from East Greenland.
8. Thor G. Halle (1884–1964) was appointed as an assistant in the Swedish Museum of Natural
History in Stockholm in 1913. He succeeded Alfred Nathorst and became professor and director of
the Department of Palaeobotany in 1918. Harris’s comment on his first encounter with Lauge Koch is
from Chaloner (1985).
9. Harris described fourteen species of fossil ginkgolike leaves from East Greenland. The seeds
that he believed to be associated with Ginkgoites taeniata he named Allicospermum xystum; Harris
(1935).
10. The coastline of northeastern England around Whitby provided a suitably foreboding back-
drop for Bram Stoker’s Dracula.
11. The possible ginkgo pollen cone from Scalby Ness was collected and first described by Han
van Konijnenburg–van Cittert (1971); see also Harris et al. (1974).

15. Proliferation
1. I was an undergraduate in the Department of Botany at the University of Reading from 1972
to 1975, a Ph.D. student from 1975 to 1978, and then a temporary lecturer there from 1978 to 1981.
2. The first publication on the ginkgolike fossils from Yima is by Zhou and Zhang (1988).
3. See Zhou and Zhang (1989) for the first detailed account of these fossils. Zhou and Zhang
noted that in living ginkgo there are occasionally aberrant ovule-bearing stalks in which as many as
ten young ovules are borne on distinct side branches; see also Florin (1949).
4. For additional information on the leaves of Baiera hallei and the associated seed-bearing struc-
tures, Yimaia recurva, see Zhou and Zhang (1992).
5. Yimaia qinghaiensis is preserved in a paper coal from the Lucaoshan coal mine, Qinghai Prov-
ince, northwestern China. It is known from deeply divided leaves with very narrow segments, and
also from seeds and seed-bearing structures. The seeds are a little smaller than those of Yimaia re-
curva, and there are fewer seeds at the tips of the seed-bearing structures, but they are otherwise
very similar; for more on this fossil plant see Wu et al. (2006). Zhou has also described a third
Yimaia species, Yimaia capituliformis, from Daohugou, Inner Mongolia; see Zhou et al. (2007). With
hindsight we now recognize also that fossil plants very similar to Yimaia have been described from
Europe. Black (1929) described seeds and seed-bearing axes associated with the leaf he described as
Baiera gracilis. Harris et al. (1974) reassigned Baiera gracilis to Baiera furcata. Similar material from
Germany was described by Schenk (1867) and Kirchner (1992).
6. Archangelsky (1965), Del Fueyo and Archangelsky (2001), Zhou et al. (2002).
7. Since Archangelsky’s original description, there have been discoveries of Karkenia seed-
bearing structures in the Northern Hemisphere. For example, Karkenia asiatica, from the Upper Ju-
rassic of Bureya, Russia (Krassilov, 1970), is very similar and is associated with leaves very like those of
Ginkgoites tigrensis (named Sphenobaiera umaltensis). About six reasonably well-understood species
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n o t e s t o pa g e s 1 1 6 – 1 2 2

of Karkenia are now known across Europe and Asia. They differ mainly in the size and number of
seeds that they bear and in the details of the leaves with which they are associated, but all are funda-
mentally similar. For further treatment of Karkenia-like plants and a review, see Krassilov (1970) and
Zhou (2009).
8. A case in point is the abundant leaves of Ginkgo australis in the Koonwarra Fossil Bed in Vic-
toria, Australia, which date from about 125 million years ago (Drinnan and Chambers, 1986). They
are always deeply divided in two but are variable in the degree of dissection; some leaves have only
four leaf segments, while others are divided into as many as sixteen. Ginkgo australis is very similar to
Ginkgo rajmahalensis from the Jurassic of northeast India and Ginkgoites ticoensis from the Early Cre-
taceous of Tico, Argentina. No ovulate organs other than Karkenia have been linked to the ginkgo-
like leaves found in the southern continents, including at the Koonwarra locality; see Drinnan and
Chambers (1986). Several species of Sphenobaiera and Ginkgo leaves are associated with male catkins,
although the associated ovulate organs are quite different from those of ginkgo; see Anderson and
Anderson (1989, 2003); Holmes and Anderson (2007); Anderson et al. (2007); Zhou (2009).
9. Archaefructus, an intriguing early flowering plant from the Jehol Biota, is especially fascinat-
ing and controversial. The U.S. television documentary series Nova built a whole program around
the origin of flowers, and Archaefructus was a central part of that story; see Lewis (2007). The Yixian
Formation is in western Laoning Province, China.
10. For more on the significance of Ginkgo apodes see Zhou and Zheng (2003). For a complete
description and illustration of Ginkgo apodes and its associated seed-bearing shoots, see Zheng and
Zhou (2004).

16. Winnowing
1. Epigraph: Murdoch (1970, 170). A ginkgo leaf is included in the recent portrait of Iris Murdoch
by Tom Phillips. He recalls wanting “a bit of nature to be present,” and he soon discovered they were
both enthusiasts for “the world’s oldest tree”; see Phillips, “Portraits: Dame Iris Murdoch.”
2. Zhou and Wu (2006).
3. The twenty-two species of fossil ginkgo leaves recorded by Zhou in the Early Cretaceous of
China include species assigned to the two genera Ginkgo and Ginkgoites.
4. For more on the diversification and vegetational expansion of flowering plants in the Creta-
ceous, see Friis et al. (2011). See Kvaček et al. (2005) for a description of Nehvizdyella, a strange puta-
tive ginkgo relative from the Late Cretaceous of the Czech Republic.
5. Peter Del Tredici cites the well-developed olfactory lobes and jaws adapted to opening hard
seeds as evidence that multituberculates were more likely candidates for ginkgo dispersers than the
dinosaurs or early birds of the time: see Del Tredici (1989); but see also Van der Pijl (1982), Janzen
and Martin (1982), and Tiffney (1984). Biomechanical analysis of multituberculate jaws suggests that
they were unlikely to have been effective dispersers of ginkgo seeds (Wall and Krause, 1992).
6. Angiosperm fossils are relatively rare in the Horseshoe Canyon Formation, but seeds, wood,
 306
n o t e s t o pa g e s 1 2 3 – 1 3 1

leaf impressions, and pollen indicate the presence of early relatives of sassafras, katsura, plane (“syca-
more”), alder, sweet gum, and other trees. For more on plant fossils from the Horseshoe Canyon For-
mation, Alberta, see Aulenback (2009).
7. K-T boundary (Cretaceous-Tertiary boundary = Cretaceous-Paleogene boundary). Of 130
Cretaceous leaf species appearing at multiple levels in the Hell Creek and Fort Union Formations,
only 29 are also present in the Paleocene; an estimated 30 to 57 percent of the flora present in the final
five million years of the Cretaceous became extinct at the boundary; see Wilf and Johnson (2004). The
presence of ginkgo leaves below the K-T boundary is clearly documented based on fossils collected
from seven different localities in the Hell Creek Formation of North Dakota; see Johnson (2002). See
Zhou and Wu (2006) for more on the decline of the ginkgo in the mid-Cretaceous.
8. Following rapid diversification, the diversity of grazing horses reached its acme (about sixteen
species) between about 15 million and 18 million years ago. Today there is only one, or under some
interpretations two, extant species; see MacFadden and Hulbert (1988).

17. Persistence
1. Epigraph: Carlyle (1858, 286). I had corresponded with David Dilcher while I was at the Uni-
versity of Reading, and he had brought me to the United States to work with him on fossil plants as
a postdoc in his laboratory.
2. David Dilcher and Steven Manchester were the first to publish on Almont fossil plants (Man-
chester and Dilcher, 1982). They described fossil fruits of Cyclocarya, a special kind of Asian wingnut,
the wheel wingnut, and found only minor differences from the fruits of the single living species, which
is native to the rich deciduous forests of central China. The fossils I described from southern England
that have a counterpart at Almont were assigned to the fossil genus Palaeocarpinus; see Crane (1981).
3. For our preliminary account of the Almont flora, see Crane et al. (1990).
4. The fossil ginkgo from Almont has now been studied in detail by Zhou and his colleagues
(Zhou et al., 2012) and formally named Ginkgo cranei.
5. For more on living fossils, see Eldredge and Stanley (1984) and Schopf (1984). Lingula, the
clamlike organism, is a brachiopod, not a mollusk.
6. See Crane et al. (1990, Fig. 28a) for the possible ginkgo pollen catkin. A fragment of a ginkgo
pollen catkin, very like that of the living species, is known from the Horseshoe Canyon Formation
(Rothwell and Holt, 1997). Zhou et al. (2012) note minor differences between the cuticles of Ginkgo
cranei and living ginkgo.
7. See Royer et al. (2003). Taken together, the distribution of ginkgo fossils through time sup-
ports the idea that the genus has, since the Cretaceous, preferred growing in warm temperate climates
with moist hot summers and cold winters; see Del Tredici (2000), Tralau (1968), and Uemura (1997).
 307
n o t e s t o pa g e s 1 3 2 – 1 3 9

18. Prosperity
1. Epigraph: Attributed to Goethe, but the specific source is not confirmed. For details on the
paleobotanical career of John Starkie Gardner (1844–1930), see Andrews (1980, 372). See British Eo-
cene Flora, vol. 1, for the early collaborative work of Gardner and Ettingshausen on fossil ferns (Gard-
ner and Ettingshausen, 1879–1882); vol. 2, on fossil gymnosperms, was published by Gardner alone
(Gardner, 1883–1885).
2. See Andrews (1980) for commentary on the paleobotanical work of Constantin von Etting-
shausen (1826–1897).
3. See Andrews (1980) for commentary on the paleobotanical work of James Scott Bowerbank
(1797–1877), Eleanor Reid (1860–1953), and Marjorie Chandler (1897–1983).
4. In a series of publications during the 1960s, Marjorie Chandler expanded and revised the
original London Clay Flora from 1933 and placed it in the context of other Eocene fossil floras from
Southern England; see Chandler (1961, 1962, 1963, 1964). Additional revisions have since been made
by Professor Margaret Collinson of Royal Holloway, University of London, and Dr. Hazel Wilkin-
son of the Jodrell Laboratory at Kew. For more on the importance of work done on the London Clay
flora, see Crane and Carvell (2007). The seed described by Bowerbank (1840) and assigned to Ginkgo
(see also Gardner and Ettingshausen, 1879–1882) was reexamined by Reid and Chandler (1933), and
reassigned to the flowering plant family Icacinaceae.
5. The London Clay is known today to contain more than 500 kinds of plants and about 350
named species. For additional information on the London Clay and the fossil plants that have been
described from it, see the classic works of Reid and Chandler (1933) and later reviews and updates by
Chandler (1961) and Collinson (1983).
6. The exposed columnar basalts that create Fingal’s Cave were part of the inspiration for the
music of the Norwegian Romantic composer Edvard Grieg.
7. For the geological observations of the Duke of Argyll see Duke of Argyll and Forbes (1851).
8. For further details on the Paleocene fossil plants from the Isle of Mull, see Boulter and Kvaček
(1989).
9. In 1995, the Messel Pit Fossil Site became a UNESCO World Heritage Site. For details on the fos-
sil Panama hat palm, see Smith et al. (2008); for more on the Messel flora see Collinson et al. (2012).
Ginkgo orientalis appears in several Paleocene localities from Eastern Europe; see Samylina (1967).
10. Well-preserved fossil fish are particularly common, earning one especially rich bed the name
“split fish layer.” Millions of fish fossils have been collected from these lake beds, which have been a
major source of fossils for commercial collectors; see Grande (1984, 2013).
11. For more on the fossil flora of the Clarno Formation see Manchester (1981) and Wheeler and
Manchester (2002).
12. Wes Wehr was a gifted artist and poet as well as a dedicated paleobotanist and collector of fos-
sil plants. Fossil floras similar to those from the Pacific Northwest, which also contain ginkgo leaves,
are known on the other side of the Pacific, for example, from Fushun (Liaoning Province), Huachian
 308
n o t e s t o pa g e s 1 3 9 – 1 4 8

(Jilin Province), and Yilan (Heilongjiang Province) in northeastern China; see Endo (1942), Man-
chester et al. (2005), He and Tao (1997). Ginkgo is also known from fossils of Late Cretaceous age in
this region (Sun et al., 2007).
13. Ginkgo is reasonably common in the Middle Eocene formations of northern Washington State
and British Columbia at Driftwood Creek, Quesnel, Horsefly, Tranquilo, McAbee, Quilchena, Prince-
ton, and Republic localities; see Mustoe (2002).
14. Ginkgo was among the fossil plants encountered by Alfred Nathorst, the paleobotanist on
several of Otto Nordenskiold’s expeditions. Nathorst collected ginkgo at several locations, includ-
ing most notably from Spitsbergen at 80° North Latitude. Nathorst also traveled on Nordenskiold’s
remarkable Vega Expedition that sailed from Sweden across the northern coast of Asia and down to
Japan through the Bering Strait in 1883. For more on the Eocene fossil flora of Ellesmere Island, see
McIver and Basinger (1999).

19. Constraint
1. Epigraph: Nietzsche (1896, 98).
2. Minus 20° Fahrenheit is equivalent to minus 29° Celsius, and minus 45–50° F corresponds to
minus 43–46° C. The coldest temperature on record in Chicago is minus 27° F (minus 33° C), with a
wind chill of minus 83° F (minus 64° C) on January 20, 1985.
3. For more on freezing resistance in North American trees, see Sakai and Weiser (1973). At
minus 40° the Fahrenheit and Celsius scales converge.
4. For more on the effect of climate on plant growth and productivity see Skre (1990), Dahl
(1990), and Melillo et al. (1993).
5. The fate of the ginkgo seedlings was relayed by Wolfgang Stuppy (Royal Botanic Gardens,
Kew, personal communication).
6. The Arnold Arboretum, one of the world’s great collections of living trees, is located in Jamaica
Plain, just outside Boston. It has a long history of sending expeditions all over the world to bring trees
back for cultivation. The collections are especially rich in material from China, and strong collabora-
tions with Chinese botanists gave Peter Del Tredici the opportunity to work on the timing of sexual
reproduction in ginkgo trees growing in a near wild situation; see Del Tredici (2007).
7. While there is no natural period of dormancy, retention of the outer fleshy layer on the seed
does inhibit germination; see Rothwell and Holt (1997). Cold stratification, although not required,
improves the evenness of germination and may improve the overall germination percentage; see Holt
and Rothwell (1997), Rothwell and Holt (1997), and Del Tredici (2007).
8. This account of the process from fertilization to germination, and the factors that cause varia-
tions in this process, is based on Del Tredici (2007).
9. Other manifestations of the temperature sensitivity of ginkgo include earlier bud burst and
leaf drop in response to warming temperatures and a difference of forty days in bud burst and leaf
drop in southern versus northern Japan (Matsumoto et al., 2003).
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n o t e s t o pa g e s 1 4 8 – 1 5 5

10. The existence of warm winters at high latitudes during the Eocene is also consistent with the
presence of crocodiles among the high-latitude fauna at that time.
11. Forty-three degrees F is equivalent to 6° C.
12. Recent research found that the number of hours of chilling at temperatures below 38° F
(3.5° C) was an important factor in determining the tree’s height and shoot development; see Wilson
et al. (2004).
13. Forty-one degrees F is equivalent to 5° C.
14. It seems to me that ginkgo never looks quite as robust, luxuriant, and fresh in northwest
Europe as it does in Seoul or Chicago. Ginkgo seems to like more warmth during the growing season,
perhaps even needs great warmth to really flourish, but it also needs a cold winter, just so long as it is
not overly tough. Similarly, ginkgo in southern California and other dry places rarely seems as robust
as it is in places with a more pronounced winter.

20. Retreat
1. Epigraph: Attributed to Groucho Marx, but the specific source has not been confirmed.
For a description of the Paleogene ginkgolike leaf fossils from Tasmania, see Hill and Carpenter
(1999).
2. For a description of the Paleocene ginkgo leaves from the Isle of Mull, see Boulter and Kvaček
(1989, 34–39). Ginkgo is missing, for example, from the well-studied fossil flora from Kreuzau, and
also from the exceptionally preserved Pliocene assemblage of fossil plants recovered from the fill of an
ancient sinkhole at Willershausen near Göttingen, in Germany. The Willershausen flora is especially
rich, “more than thirty thousand specimens have been collected, representing at least 130 species of
plants in 77 genera,” but so far those plants do not include ginkgo; see Ferguson and Knobloch (1998).
The occurrence of ginkgo in the Miocene flora from Frankfurt, Germany, is unusual in the context
of fossil floras of the time. The Frankfurt specimen was described as Ginkgo adiantoides by Florin
(1936a) but renamed Ginkgo florinii by Samylina (1967).
3. For more on the Selárdalur flora in Iceland, see Denk et al. (2011); see also Akhmetiev et al.
(1978). For more on ginkgo fossils from southeastern Europe see Kovar-Eder et al. (1994, 2006) and
Denk and Velitzelos (2002). For the work of Royer et al. (2003) see Chapter 17.
4. Only one ginkgo leaf has been recovered among the many thousand leaf fossils collected from
the Florissant fossil flora (Bret Buskirk and Herb Meyer, Florissant Fossil Beds National Monument,
personal communication). The occasional occurrence of ginkgo leaves in the Oligocene Ruby River
fossil flora from southwestern Montana (see Becker, 1961) also indicates that the geographic distri-
bution of ginkgo at this time was complex. See also Chaney and Axelrod (1959), Schorn et al. (2007),
and Wolf (1987).
5. At the same time, much farther south, Miocene ginkgo is present in the Cedarville flora of
northwestern Nevada and neighboring California, about two hundred miles inland from the present-
day coastline.
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6. See Scott et al. (1962) and Wheeler and Dilhoff (2009) for additional information on the Middle
Miocene woods from Vantage, Washington. There is a hint of a pattern of increasing restriction in
eastern Asia similar to that seen in western North America. There seems to be a decline of ginkgo in
China through the Cenozoic. Ginkgo is present at several different Eocene sites and continues into
the Oligocene in far eastern sites along the Russian border. Ginkgo is missing from the fossil record
of China from the Early Miocene, about twenty million years ago, but persists in southeastern Russia
along the Sea of Okhotsk. Ginkgo is also known from several fossil occurrences in Japan, during the
Pliocene and Pleistocene, between about two million and five million years ago, for example in the
fossil floras from Hoshiwara, Hiradoguchi, and Daiwa in southern Japan. For a thorough account of
the Cenozoic distribution of ginkgo in East Asia, see Uemura (1997). Ginkgo, however, is far from
ubiquitous at this time. It is missing, for example, from the classic Mogi flora that Nathorst collected
in Honshu when Nordenskiold’s Vega expedition spent the summer there in 1879.

21. Extinction
1. Epigraph: From Jimmy Cliff ’s album Jimmy Cliff, 1969, Trojan Records. For more information
on the Early Pliocene fossil vegetation in southern Europe, see Kovar-Eder et al. (2006).
2. The Willershausen flora is especially rich; more than 130 species have been collected, repre-
senting more than one hundred different kinds of plants. See Straus (1967); Ferguson (1967); Fergu-
son and Knobloch (1998).
3. Like ginkgo, the Caucasian wingnut has been reintroduced by people into many of the places
where it once grew; there are large specimens of the Caucasian wingnut at Kew, for example, that date
from the late nineteenth century. The nearest native populations are in the Caucasus, with its closest
relative, a similar species native to China.
4. The gomphotheres may have persisted until as recently as six thousand years ago in present-
day Colombia; see Rodríguez-Flórez et al. (2009). For a complete list of now-extinct large herbivores
of Central America, see Janzen and Martin (1982, 21).
5. For a popular discussion and elaboration of Janzen and Martin’s idea, see Barlow (2002).
6. Known foragers of the seeds of living ginkgo include the catlike Paguma larvata in China and
the Japanese badger Nyctereutes procyonoides. Rothwell and Holt (1997) note the improved germina-
tion rates of seeds scarified by passing through the digestive tract of badgers.
7. By the end of the Pliocene, ginkgo had disappeared from the fossil record everywhere except
perhaps for a small area of southern Japan; see Uemura (1997).
8. Castiglioni had visited North America between 1785 and 1787, and also had consulted Flora
Virginica, published by Gronovius in 1739 and 1743, and Thunberg’s Flora Japonica published in 1784:
see Spongberg (1993). Asa Gray at Harvard was a frequent correspondent and staunch supporter of
Darwin in North America. Darwin’s letter to him on “botanical geography” was written on Octo-
ber 12, 1856. New information that Gray had at his disposal included Siebold’s Flora Japonica as well
as specimens brought back from the Rodgers-Ringgold Expedition (1853–1856), also known as the
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North Pacific Exploring and Surveying Expedition, a United States scientific and exploring project
with the broader purpose of finding shorter trade routes for merchant ships in the Pacific; see Cole
(1947). Also available were specimens brought back from Japan by Charles Wright.
9. Gray (1859, 422) listed about 580 Japanese species “which have particular relatives in other
and distant parts of the northern temperate zone,” along with the corresponding plants in the floras
of Europe; central and northern Asia; western North America; and eastern North America. To ex-
plain the widely separated but highly similar floras of eastern Asia and eastern North America, Gray
suggested that before the glacial epoch, the flora of the North Temperate Zone had been relatively
homogeneous and that regional extinction during the Ice Ages resulted in greater losses from western
North America and Europe. In some cases the impact of regional extinction was less pronounced. The
sweet gum, for example, has widely separated remnants not only in eastern Asia and eastern North
America, but also in southeastern Europe.

22. Endurance
1. Epigraph: Thoreau (1862, 517).
2. See Chapter 29 for additional information on initial Western encounters with ginkgo.
3. The Royal Horticultural Society estate where Robert Fortune worked as a gardener is now
Chiswick House. The first Opium War lasted from 1832 to 1842. Fortune’s four trips to China—in
1843–1846, 1848–1851, 1853–1856, and 1858–1859—are recounted in Fortune (1847, 1852, 1857, 1863).
Fortune’s trip to Japan from 1860 to 1862 was one of the first by a Western botanist following the
Treaty of Amity and Commerce (the Harris Treaty), which was signed on July 29, 1858, and expanded
trade to five major Japanese ports and allowed for diplomatic exchange.
4. Fortune (1847, 118). See Chapter 30 for more information on bonsai ginkgo.
5. Père Jean Marie Delavay (1834–1895); Père Paul Guillaume Farges (1844–1912); Père Jean Pierre
Armand David (1826–1900). Père David arrived in Canton in 1862. His journeys into the interior were
as a missionary priest, but he devoted much energy to studies of native plants and animals. His suc-
cess was emulated by his contemporaries Père Delavay and Père Farges. Their collective contributions
resulted in the introduction of hundreds of new plants into Europe, including the butterfly bush,
many species of rhododendron, and the spectacular dove tree. They also sent thousands of botani-
cal specimens back to the Muséum national d’Histoire naturelle in Paris. Their work often carried
a significant price in personal hardships; Delavay contracted bubonic plague, which eventually cost
him his life. David at various times suffered from typhus, smallpox, leprosy, rabies, cholera, plague,
dysentery, and malaria. For further information on the contributions of the French missionary bota-
nists see PlantExplorers.com (1999–2012). The dove tree is sometimes more descriptively called the
handkerchief tree.
6. Augustine Henry writes about the classification, fossil history, varieties, distribution, uses, and
history of ginkgo, also providing a list of important ginkgo trees in the British Isles. Of the seeds he
wrote, “The nuts are sometimes eaten boiled or roasted, but are not much thought of,” and plates
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21–23 show photographs of old ginkgos; see Elwes and Henry (1906, 55–62). For more on the career
and contributions of Augustine Henry (1857–1930), see Nelson (1983). Wilson received guidance from
Henry in his efforts to re- collect the much sought-after dove tree, which had originally been de-
scribed by and named for Père David.
7. Ernest Henry Wilson (1876–1930) was appointed at the Arnold Arboretum in 1927. Tragically,
after surviving many adventures in China, including having his leg crushed by boulders in an ava-
lanche, he was killed in an automobile accident in Worcester, Massachusetts, at the age of fifty-four.
8. Wilson began his career as an apprentice gardener at a local nursery near his home in Chip-
ping Campden, but after working briefly at the Birmingham Botanic Gardens, and also at Kew, he
accepted a position with the horticultural firm of James Veitch and Sons as their collector of Chinese
plants. Wilson traveled to China via the United States, stopping in Boston to visit Charles Sargent at
the Arnold Arboretum, and arrived in Hong Kong on June 3, 1899. On his first visit Wilson gathered
thirty-five cases of living plants, which he sent home as bulbs, corms, rhizomes, and tubers as well
as seed. There were also herbarium specimens of nearly a thousand plant species, many of which are
in the Herbarium at Kew. Wilson returned to China as a collector for the Arnold Arboretum in 1907,
1908, and 1910. Between 1911 and 1915 he was in Japan, and in 1917 and 1918 in Korea and Formosa. He
also traveled widely in the Southern Hemisphere, collecting plants in Australia, New Zealand, India,
South America, and Africa. Wilson ended his career directing the Arnold Arboretum at Harvard Uni-
versity. His remarks on ginkgo are recorded in Wilson (1913, 45). Wilson’s claims regarding the link
between ginkgo and Buddhism (Wilson, 1920) were later challenged by Li (1956) among others; see
Chapter 26.
9. For an overview and discussion on the significance of the ginkgos growing on Tianmu Moun-
tain, see Del Tredici (1990, 1992b) and Del Tredici et al. (1992).

23. Relic
1. Nikolai Vavilov is the father of modern studies of crop diversity. He accumulated more than
200,000 collections of crop plant seed from around the world and identified the areas in which the
domestication of many crops took place. These collections were protected during the twenty- eight-
month Siege of Leningrad by Vavilov’s colleagues, twelve of whom starved to death while surrounded
by the edible seeds in the collection. Vavilov was arrested in 1940 after disputing the pseudoscience
encouraged by Stalin’s regime and died of malnutrition in prison in 1943. For additional information
on centers of crop diversity and crop origins, see Vavilov (1992). For a biography of Vavilov, his career,
and his persecution see Pringle (2008).
2. Most of our modern studies of DNA depend on being able to replicate large quantities of DNA
from small samples using the polymerase chain reaction technique (PCR). Kary Mullis received the
Nobel Prize for this discovery in 1993, and the process is well described in “The Polymerase Chain
Reaction”; see Mullis et al. (1994).
3. Fan et al. (2004).
4. For the RAPD study see Fan et al. (2004).
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5. The DNA for the second kind of study was obtained from the chloroplasts in the leaves. For
more on this so-called restriction fragment length polymorphism (RFLP) study see Shen et al. (2005).
6. More precise characterization of the DNA fragments would certainly be possible using the
sophisticated techniques of modern molecular biology but is beyond the resolution of these two ap-
proaches.
7. See Gong et al. (2008a, b) and Zhao et al. (2010).
8. Based on low genetic variability among forty ginkgo trees from the population on Tianmu
Mountain, Wu et al. (1992) concluded that the trees had probably originated from those planted near
the old temple. Subsequent work has revealed greater genetic variation supporting the possibility that
both Jinfo and Tianmu Mountain may preserve relic ginkgo populations; see Gong et al. (2008 a, b)
and Zhao et al. (2010). It is hard to exclude completely the possibility that some of the genetic varia-
tion in particular populations resulted from ancient people bringing together ginkgos from multiple
sources, but at Jinfo and Tianmu Mountain this seems unlikely.
9. Ginkgo is not the only “living fossil” to come from China; in the mid-1940s, the dawn redwood
was found in a small population of large trees along small streams and slopes of northeastern Sichuan
Province, only four years after the genus had been described first from fossils. Like ginkgo, the dawn
redwood once had a much more extensive former range, covering parts of North America and Asia,
and was on the verge of extinction within its native habitat; see Merrill (1948). The dawn redwood is
now a popular street tree, especially in China; see Chapter 35.

24. Antiquity
1. Epigraph: Emerson (1883, 478). The large tree at Jinfo Mountain, Naquan County in Chong-
qing Municipality, was partially destroyed by fire in the 1960s but survives through natural resprout-
ing (Fan et al., 2004). For more on old ginkgo trees in China, see Li et al. (1999).
2. For a list of old, large ginkgos in different Chinese provinces, see Lin et al. (1995).
3. The small hamlet of Li Jiawan is a little more than sixty miles west of Guiyang, the capital of
Guizhou Province; see Xiang et al. (2009). There has been much discussion about the possible fire
resistance of ginkgo (e.g., Handa, 2000), which has sometimes been attributed to water retention in
its leaves, or even to the fire-retardant properties of its sap. To view what happens when ginkgo and
maple leaves are burned side by side, see Japan Probe, “Ginkgo Trees Protect Shrines and Temples
from Fire.” Best known of the six ginkgo trees that survived within a mile of where the atomic bomb
exploded at Hiroshima is the ginkgo on the grounds of the Hosenji Temple. Only about half a mile
from the epicenter of the destruction, it survived and leafed out again, after having been stripped by
the blast. The new temple built around it celebrates the hope of renewal that the tree embodied amid
terrible devastation. For more on the ginkgos that survived the Hiroshima bomb, see Hageneder
(2005) and Kwant, “A-bombed ginkgo trees in Hiroshima, Japan.”
4. The Li Jiawan Grand Ginkgo King has been described as “five-generations-in-one-tree,” based
on its successive episodes of resprouting; see Xiang et al. (2009).
5. The estimate for the age of the Li Jiawan Grand Ginkgo King is the maximum calculated by
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Xiang et al. (2009). Ages of ginkgo trees that are believed to be more than two thousand years old are
cited by He et al. (1997). Trunk diameters (DBH) are based on Lin et al. (1995).
6. Coast redwoods, the tallest of all trees, have trunks up to 370 feet tall and 24 feet in diame-
ter. A slice across the trunk of a giant sequoia in the Natural History Museum in London, which is
about 14 feet across, is estimated to be 1,335 years old; the beginnings of Islam, the spread of Bud-
dhism into Japan, the fall of major civilizations, the Black Death of Europe, the birth of Shakespeare,
and other historic landmarks are marked against their corresponding rings. That we felled so many
of these spectacular trees, while knowing their great antiquity, must rank among the greatest acts of
human hubris. The giant sequoia, known as the Mark Twain Tree, was felled in 1891 just to prove the
existence of such massive trees to disbelievers in the East. The American Museum of Natural History
in New York City and other museums hold slices of its trunk. The greatest age so far recorded from a
bristlecone pine is from a specimen nicknamed Prometheus that was mistakenly felled by a graduate
student researcher in eastern Nevada in 1964. Based on both radiocarbon dating and its tree rings, it
had lived for at least 4,862 years and possibly more than 5,000; see Ferguson and Graybill (1983). A
potentially older, but much less spectacular, tree is a Norway spruce in the Dalarna region of Sweden.
Though the tree is only sixteen feet tall, its rooting system has been dated at around 9,550 years old;
see Kullman (2005).
7. Humboldt alludes to the baobab in describing another impressive tree, the Dragon Tree of
Tenerife in the Canary Islands: “Among organic creations, this tree is undoubtedly, together with the
Adansonia or baobab of Senegal, one of the oldest inhabitants of our globe”; see Humboldt and Bon-
pland (1852, 62). The provocative age suggested by Adanson approached that calculated by Bishop
Ussher for the age of the Earth. David Livingston was among those enraged; see Wickens and Lowe
(2008). Rather than indicating especially great age, the massive trunks of baobabs reflect an ability
to store water as a buffer against drought in the dry environments where they grow. In the Limpopo
Province of South Africa, an enormous hollow baobab with a diameter of thirty-five feet at the ground
and a thirteen-foot-tall hollow chamber in its base has been converted into a pub. It is younger than
the 6,000 years claimed by the proprietor’s website (www.bigbaobab.co.za): radiocarbon dating of
samples from one of the internal cavities gave an age of 1,060 years plus or minus 75 (Adrian Patrut,
“Babeş-Bolyai” University, personal communication). For the use of radiocarbon dating to determine
the age of the ancient baobab from Namibia, see Patrut et al. (2007). Radiocarbon dating relies on the
fact that the carbon atom in each molecule of carbon dioxide captured by photosynthesis may be of
one of two kinds: Carbon-12 or Carbon-14, which occur at a known ratio in the atmosphere. Carbon-
12 is stable, but Carbon-14 begins to decay to Carbon-12 at a known rate as soon as it is incorporated
into the tree. Because the ratio of Carbon-12 to Carbon-14 in the atmosphere is known, along with
the exact rate of decay, precise measurement of the amount of Carbon-12 and Carbon-14 in a piece of
wood can be used to calculate its age. Eventually the amount of Carbon-14 in a sample becomes too
small to measure accurately, and this sets a limit of about 60,000 years on how far back radiocarbon
dating will provide an accurate age.
8. For beautiful photographs of Jōmon-sugi on Yakushima see Pakenham (2002). When Ernest
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Henry Wilson recorded a stump of a Japanese cedar on Yakushima that was fourteen feet in diameter
(Sargent, 1913), it caused a sensation.
9. Dendrochronology takes advantage of variation in the thickness of different annual rings in
the same trunk to match and cross-correlate among woods from different sources. It serves the needs
of archaeologists interested in dating wood fragments, as well as scientists interested in the history of
climates over the past few thousand years. For a description of dendrochronology and its application
to determining the age of the bristlecone pine, see Ferguson (1968).
10. Minamoto no Yoritomo established the Kamakura bakufu, the first government in Japan con-
trolled by the samurai, twelve years later.
11. In the spring of 2010, to the distress of the temple and many Japanese people, the great Tsuru-
gaoka Big Ginkgo blew down in a storm after heavy rains. While it was possible to count the rings
from the outer part of the trunk, attempts to arrive at a definitive age were thwarted by decay at the
center (Toshiyuki Nagata, Hosei University, personal communication 2011).
12. The Huiji Temple, also known as the Huiji Yuan Monastery and Xiangquan si, or Fragrant
Spring Temple, is located just to the north of Tangquan town in the Pukou district of Nanjing. It
was founded in the Liu Song Dynasty in the fifth century (420–479). Zhaoming, also known as Xiao
Tong, reputedly read there and took baths in the hot spring. About 1821–1850, unusually, a Confucian
academy called Yinghua, or Quintessence Flower, was established in the temple. The temple was de-
stroyed during the Taiping Rebellion between 1850 and 1864.
13. See Needham et al. (1996, 581). For most of his life, Joseph Needham worked at Cambridge
University, but he also spent much time in China. His early career included distinguished contribu-
tions as an embryologist and biochemist, for which he was elected to the Royal Society in 1941. He
spent most of the 1940s in China, and on his return he devoted his life to the study of East Asian
science and culture. So far, seven volumes of Science and Civilization in China, in twenty-four parts
(fifteen written or cowritten by Needham), have appeared. A dawn redwood stands in memory of
Joseph Needham, his wife Dorothy Needham, and his companion Gwei-Djen Lu-Needham outside
the Needham Research Institute in Cambridge, where the work that he began is carried on. For addi-
tional information on Needham’s life and career see Cullen (1995). For a popular account, see Win-
chester (2008). As Menzies notes, it may also be significant that the most important early sources of
botanical information from China, the Shih Ching, Ērh Ya, and Nan Fang Tshao Mu Chuang (Records
of the plants and trees of the southern regions), written by Chi Han around A.D. 300, make no men-
tion of ginkgo.

25. Reprieve
1. Epigraph: Pandit and Nagarjuna (1977, 66). See Chapter 24 for more on the ginkgo at the
Fu-Yen Ssu Monastery. Purported representations of ginkgo in early Chinese art from the fourth to
eighth centuries (e.g., see Kwant, “Seven Worthies of the Bamboo Grove”) are not sufficiently diag-
nostic to be fully reliable and require more detailed study. Tso Ssu (253–307) was a prominent poet of
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the Jin Dynasty (265–420), though only fourteen of his poems survive; see Mair (2001) for more on
T’ai-k’ang poetry.
2. Simple Discourses on the Investigation of Things, a series of short statements about natural phe-
nomena that was written about 980, includes the statement “Let male and female ginkgo trees grow
near one another, then fruit will form”; see Chapter 8 for more on this and on gender in ginkgo. The
poems about ginkgo were exchanged between Ouyang Hsiu (1007–1072) and the poet Mei Yao-Chēn
(also known as Sheng-Yü; 1002–1062); see Needham et al. (1996, 581).
3. See Needham et al. (1996) for a history of Chinese agriculture, which remains an important
area of research.
4. The Yuan Dynasty spanned from 1271 to 1368.
5. A comparison of height and girth of old ginkgo trees in different Chinese provinces supports
the idea that the old ginkgos of Guizhou are on average both taller and wider than those of Anhui and
Zhejiang. Sichuan Province, bordering Guizhou to the north, has even larger ginkgos by both mea-
sures; data compiled from Lin et al. (1995). The Anhui origin of ginkgo is cited by Menzies from the
Pen Ts’a Kang Mu, or the Great Pharmacopoeia (1596, p. 1801), quoting the Jih Yung Pēn Tshao. The
quote from Shihuazonggui is from He et al. (1997, 374). Beginning in the Yuan Dynasty other names
have also been used; pei yen, “white eye”; pei kuo, “white fruit”; ling yen, “spirited eye”; jen hsing, “nut
apricot”; others such as kung sun shu, “grandfather-grandchild tree,” appeared later. These records,
from the Song Dynasty (960–1279) and Yuan Dynasty are reported in He et al. (1997, 374) and Li
(1963, 92).
6. The oldest ginkgo trees in South Korea have been designated as natural monuments by the
Cultural Heritage Administration; see Invitation ForestOn “Story of forest: Old gigantic trees in
Korea.”
7. The villagers of Yeongwol believe that a giant snake resides inside the tree, keeping animals
and insects away, and that young children who fall from the tree will not be injured. For descriptions
and images of the Yeongwol Ginkgo, Duseo-myeon Ginkgo, and Yongmunsa Ginkgo, see Invitation
ForestOn, “Story of forest: Old gigantic trees in Korea.” The Anbulsa Temple in North Korea is in
Tonghung-ri, Kumya County, South Ham-Yong Province.
8. A list of old ginkgo trees in Japan and their associated legends is given by Hori and Hori (1997).
According to legend, the Yongmunsa and Zenpukuji ginkgo trees in Korea and Tokyo, respectively,
were both grown from sticks planted in the ground by priests wishing for the prosperity of their
temples. In another legend, an official accompanying Emperor Kao Tsung as he moved from Kaifeng
to Hangchow in the south of China in 1127 picked a branch of ginkgo and stuck it into the ground,
declaring that if it lived, he would settle there; the branch grew into a huge tree adorned with many
chi-chis. The Senbon Ginkgo (“One Thousand Ginkgo Trees”) in Japan allegedly all sprouted from
the trunk of a grand ginkgo that was struck by lightning; see Primack and Ohkubo (2008), Handa
(2000), Li (1963), and Kwant, “An Old Chinese Legend.” At Ubagami Shrine in Miyagi Prefecture, the
large female tree is called Uba—“wet nurse”—Ginkgo. Emperor Shōmu ruled from 701 to 756. The
Heian period began in 794 and ended in 1185.
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9. See Chapter 24 for more on the Tsurugaoka Big Ginkgo. The Nara period in Japan is usually
regarded as beginning in 710 and ending in 794.
10. For more on the cultural history of ginkgo in Japan, see Hori and Hori (1997).
11. These ancient trees in Japan are reported by Li (1963).
12. See Hori and Hori (1997).
13. The Kamakura period began in 1185 and ended in 1333; and the Muromachi period began in
1336 and ended in 1573.

26. Voyages
1. Epigraph: Fuller (1732, #1850). The Korean National Maritime Museum is the responsibility of
the Ministry of Culture and Tourism; see www.seamuse.go.kr.
2. In 1994 the Marine Antiques Preservation Center became the National Maritime Museum of
South Korea. The remains of the Shinan Ship, and much of its cargo, can be seen at the National Mari-
time Museum. Since the excavation of the Shinan Ship the museum has led the excavation of other
wrecks discovered around the Korean coast.
3. For a description and diagrams of the ship’s structure and construction, see Green (1983); for
descriptions and photographs of the ship, its cargo, and the underwater excavation see Kim (2006a,
b, and c).
4. The ginkgo nut recovered from the Shinan Ship is on display at the National Museum of Korea
in Seoul, along with ceramics and other materials recovered from the wreck.
5. Among the objects from the Shinan Ship were more than twenty inkstones used in calligraphy
and day-to-day writing; a bone die, a little smaller but otherwise identical to dice of today; and several
examples of lacquerware. Glass objects included delicate hairpins, buttons, beads, and pieces of small
glass bottles. There were also metal bottles, mirrors, incense burners, balance weights, cups, lamp oil
containers, scoops, gongs, small cymbals, plates, wine cups, boxes, spoons, chopsticks, shovels, locks,
wine dippers, candlesticks, bells, kitchenware, containers, coal basins, cauldrons, pots, instruments,
soup bowls, astronomical instruments, Buddhist statues, figurines, acupuncture needles, and rings.
The more than three hundred pieces of raw metal recovered, including tin and white copper sheet,
would have been destined for manufacturing of all kinds and for making metal alloys. Stored as bal-
last in the hold of the boat were also about eight million Chinese coins, the heaviest component of
the cargo. They had been held together by strong strings threaded through the holes in their center.
Together these coins weighed nearly twenty-seven tons and included about seventy different kinds,
all from China, that were in circulation from the first to the fourteenth century. Imported Chinese
coins were widely used as currency in medieval Japan. Japanese coins were not minted at this time.
Poor-quality coins may have been destined to be melted down and made into Buddhist statues and
other luxury items.
6. See Seyock (2008). The temples in Fukuoka and Kyoto still have several ancient ginkgos
nearby. The trade between China and Japan documented by the Shinan Ship continued through
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Japan’s period of self-imposed isolation. Through the seventeenth century previously scattered
Chinese communities were consolidated at Nagasaki, which then became the main center through
which foreign trade was conducted and controlled. Many aspects of Chinese culture were introduced
and became assimilated into Japan through this bridgehead. For example, in 1654 Yin Yuan, a high-
ranking Buddhist priest, arrived at the Kofukuji Temple in Nagasaki and introduced the teachings
of Huan Bo, a particular sect of Zen Buddhism, which later was taken up by the Shogun and many
powerful Japanese lords.
7. That ginkgo was present in Japan by the 1300s is also consistent with the discovery of ginkgo
leaves pressed inside the pages of books from the fourteenth century at the Kanagawa Provincial
Museum. For reasons to do with the history of these collections, the leaves are unlikely to have been
placed there recently. The practice of pressing ginkgo leaves between the pages of books continues
today. The leaves are thought to have insect repellent properties. See Chapter 25 for a brief review of
early written records of ginkgo in Japan. For more details see Hori and Hori (1997).
8. For a list of other artifacts and animals that take their name from ichou in Japan see Hori and
Hori (1997). The Azuchi-Momoyama period began in 1573 and ended in 1603.

27. Renewal
1. Epigraph: Proust (1919). Engelbert Kaempfer (1651–1716) has been called “the first Interpreter
of Japan” (Brown, 1992).
2. For Kaempfer’s full quote about the occurrence of ginkgo in Japan see Chapter 28.
3. See Evelyn (1664, 194); Campbell-Culver (2006) raises the possibility that the large trees in
China described by John Evelyn may be ginkgo. Given that early contact with China was mainly in
the subtropical south of the country, the possibility that they are figs seems more likely.
4. Early reports of Portuguese traders in the 1540s attracted the interest of Jesuit missionary
entrepreneurs, who saw opportunities to advance both commercial and spiritual interests. In a letter
to attract the interest of private shipowners and traders, the missionary Francis Xavier wrote: “Get
someone of them, and that you may tempt his palate with a foretaste of the gains to be gathered in
Japan—which happen now to be so serviceable to religion . . .”; see Newitt (2005, 135). A key early
convert to Christianity in Japan was the feudal lord Omura Sumitada. With the help of the Portu-
guese, he established the trading port at Nagasaki in 1571 and by the time other European traders
followed, Nagasaki was already well established. The European influence is still evident in Nagasaki’s
modern-day churches and specialty shops that sell the popular Portuguese sponge cake castella. The
Japanese word tempura may have been derived from the Portuguese tempero, but alternatively it may
be derived from the Portuguese words for temple—templo and têmpora—referring to the days when
fish and vegetables are eaten instead of meat.
5. Clavell (1975).
6. The Dutch East India Company is often referred to as the VOC (Vereenigde Oost-Indische
Compagnie). The Shimabara Rebellion began not far to the east of Nagasaki when tens of thousands
of peasants and their allies, many of them Christians, rose up against the local authorities. The rebel-
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lion was finally put down at the fall of Hara Castle in April 1638, but in the aftermath the existing ban
on Christianity was strictly enforced and the Portuguese were finally expelled. The Dutch, who had
assisted the victorious Tokugawa Shogunate, were moved from Hirado and began trading from De-
shima in 1641. As part of the self-imposed policy of national isolation (sakoku), the only other foreign
ships allowed to come to Japan were from China.
7. See Chapter 6 for more on Siebold and his time in Japan.
8. Quotations from Kaempfer (1690–1692). Kaempfer, like many early European explorers in
Japan, refers to the Shogun as emperor.
9. See Stearn (1948) for an account of Kaempfer’s travels, and also the thorough Kaempfer forum;
see Michel (2009). On his return to Europe, Kaempfer first gained formal medical qualifications from
the University in Leiden. In 1694 he returned to Lemgo to start his medical practice, and in December
1698 Friedrich Adolf, Count of Lippe-Detmold, appointed him as his personal physician. Kaempfer
married in 1700, when he was almost fifty, and two daughters and a son were born soon after. All
three died in infancy. The full title of his book is Amoenitatum exoticarum politico- physico- medi-
carum fasciculi V: quibus continentur variae relationes, observationes & descriptiones Rerum Persicarum
& Ulterioris Asiae, multa attentione, in peregrinationibus per universum Orientem, collectae / ab auctore
Engelberto Kaempfero. It was published by a printer in his hometown of Lemgo in northern Germany.
10. Thunberg’s botanical collections from Japan and elsewhere are preserved at the University
of Uppsala. At the end of the seventeenth century, the Dutch East India Company went bankrupt
and was dissolved. Trading continued with the oversight of the Dutch government. Only a few ships
visited each year, but the Dutch at Deshima amused themselves by playing badminton and billiards
and brewing beer. Their presence had a lasting impact on both Europe and Japan. In the eighteenth
century, Deshima became less isolated and found itself at the mercy of political conflict on the other
side of the world. During the Napoleonic Wars, when the Netherlands came under control of the
French, the tiny colony became vulnerable to the British, who began to prey on Dutch shipping. In
1808 the British warship Phaeton entered Nagasaki harbor under a Dutch flag, but on learning that
no Dutch ships would be arriving, it left before Japanese reinforcements arrived. A few years later,
Deshima was completely cut off from European contact when the British occupied the Dutch base at
Batavia in 1811. However, with the help of its Japanese partners and the steady leadership of the Dutch
Opperhoofd Hedrik Doeff, Deshima survived and trading resumed in 1814.
11. The porcelain ginkgo dish is illustrated as Figure 184 in the Ohashi catalogue, RD30.6, H8.0,
BD15.2; see Ōhashi (2006); also the drawing on p. 175.

28. Naming
1. Epigraph: Linnaeus (1751).
2. For more on the etymology of ginkgo see Hori and Hori (1997). The Song Dynasty began in
960 and ended in 1279. The Yuan Dynasty began about 1271 and ended in 1368.
3. Kaempfer owned two copies of the Kinmo Zui. One is a first edition from 1660, the other is a
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slightly different later edition published in 1668. Both are now part of Kaempfer’s Japanese Library
in the British Library.
4. That the second g in ginkgo comes from Germany rather than from the East is an intriguing
possibility, but the Kaempfer scholar Wolfgang Michel points out that Kaempfer transcribed with a y
other Japanese words, including other plant names, that contain the Japanese syllable “kyo” or “kyō.”
Michel thinks it more likely that Kaempfer made a mistake: see his research notes (Michel, 2009).
An alternative hypothesis suggests that Kaempfer accurately followed the pronunciation of his inter-
preter, Genomon Imamura (Van der Velde, 1995), and transcribed “ginkyo” as it would have sounded
in the regional dialect spoken in Nagasaki at that time. The word for strawberry, for example, ichigo,
is still pronounced “itzingo” in the Nagasaki dialect, which is the way that Kaempfer spelled it in his
Amoenitatum Exoticarum. (Toshiyuki Nagata, Hosei University, personal communication).
5. For more on Kaempfer’s collections of manuscripts, drawings, notes, maps, books, and other
materials, including forty-nine woodcut books, see Hüls and Hoppe (1982).
6. Sloane was born at Killyleagh, County Down, on April 16, 1660. He arrived in London at the
age of eighteen and took up the study of medicine a year later. In 1683 he traveled to Paris with the
British physician-botanist Tancred Robinson and attended the botanical lectures of Tournefort. Later,
he traveled to Montpellier and took his medical degree in Orange, in southern France, before return-
ing to London in 1684. For more on Sloane’s life, acquaintances, and collections, see Dandy (1958)
and MacGregor (1994). The so-called Glorious Revolution, by which King James II was deposed by
his daughter Mary and her husband William of Orange, was facilitated by many shared elements of
British and Dutch culture. Dutch craftsmen, scholars, artists, and merchants had long integrated into
Britain, and their influence is apparent in many Dutch-derived English terms, such as yacht, sketch,
and landscape. For more on the Glorious Revolution and links between the Dutch and British at this
time see Jardine (2008).
7. Sloane was in Jamaica from 1687 to 1689 (Sloane, 1696; 1707–1725) and recorded the uses there
of chocolate, which was mixed with water into a drink. Sloane found it more palatable when mixed
with milk, and his recipe was taken over by Cadbury Brothers as “Sir Hans Sloane’s Milk Choco-
late.” The Royal Society, the U.K. national academy of science, had been founded in 1660, during the
time of King Charles II. In its second Royal Charter, granted by King Charles II in 1663, the society is
named the Royal Society of London for Improving Natural Knowledge. Among its founders were the
architect Sir Christopher Wren, the chemist Robert Boyle, the physician and philosopher John Locke,
the experimentalist, microscopist, and astronomer Robert Hooke, and the early anatomist and plant
physiologist Nehemiah Grew, one of the first to use the microscope to observe structural details of
animals and plants. Also among the founding fellows were the botanists John Evelyn and John Ray.
Ray, in particular, was a major influence on Sloane. Sloane was president of the Royal Society from
1727 to 1741 and became one of the most influential patrons of science of the time.
8. Sloane’s botanical collections also include thirteen thousand samples of seeds, fruits, gums,
waxes, bark, and curiosities stored in small sealed boxes with a glass top and bottom to allow study
of the contents (Sloane’s collection of “Vegetables and Vegetable Substances”). Kaempfer’s material
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from Japan is in Volume 211 in the Sloane Herbarium, between a collection of plants from Italy made
by the German botanist Schröter and a collection of plants from the Carolinas and other parts of
eastern North America made by Mark Catesby, author of the Natural History of Carolina, Florida and
the Bahama Islands. Specimens of ginkgo collected by Kaempfer are on two pages. On folio 91 are
specimens from three kinds of plants, among them small sprigs of ginkgo long shoots and a single
leaf. On folio 103 are nine different plants, one of them a single ginkgo leaf, nicely laid out, perhaps
from a seedling. Beneath it, in ink, in Kaempfer’s spindly hand in Latin, is written “Folium Itsjo ar-
boris nucifera, folio adiantino.” For more on the Sloane collections in the Natural History Museum
in London, see Dandy (1958) and Trustees of the British Museum (1904).
9. For Salisbury’s study of Sloane’s collections see Salisbury (1817). A detailed catalogue of
Kaempfer’s plants in the Sloane Herbarium is provided by Hinz (2001).
10. John Gaspar Scheuchzer (1702–1729) received his doctor of medicine degree at Cambridge in
1728. His father, an acquaintance of Sloane’s, was well known for his work in natural history. His uncle
Johann Scheuchzer (1684–1738) was professor of mathematics and chairman of physics at Zürich, and
was also knowledgeable about botany.
11. See Kaempfer (1690–1692, p. 181). By the time Amoenitatum Exoticarum was published in 1712,
Kaempfer had changed his spelling to “Gín an.”
12. For more on William Adams see Chapter 27. The account of his life in Japan that he wrote in
two letters between 1611 and 1617 was reprinted from the papers of the Hakluyt Society; see Adams
(1896). Swift’s Gulliver’s Travels was published in 1726, just one year before The History of Japan.
Sloane died on January 10, 1753, and his will led directly to the establishment of the British Museum
by an act of Parliament on June 7, 1753. He is also remembered in the name of Sloane Square, which
stands at the end of the King’s Road in Chelsea, not far from the Chelsea Physic Garden, where Sloane
studied, and which was established by the Worshipful Company of Apothecaries in 1673; see Minter
(2000). As the size and diversity of the British Museum collections expanded, in the 1880s Sloane’s
natural history objects (including those of Kaempfer) were transferred to the newly created British
Museum (Natural History), now the Natural History Museum, in South Kensington. Kaempfer’s
books eventually became part of the founding collections of the newly created British Library in 1972.

29. Resurgence
1. Epigraph: Shakespeare (1623b, act 3, scene 2).
2. See also Jarvis (2007) for more on Linnaeus and the significance of his work. Early in his life
perhaps the closest Linnaeus came to encountering ginkgo was in 1736, when at the age of twenty-
nine he visited Sloane in London. It was a brief social call; Sloane was almost fifty years his senior, and
Linnaeus had no time to consult Sloane’s vast collection in detail. In retrospect, Linnaeus’s encounter
with Sloane was a passing of the torch from the great plant collector of one generation to the great
collector and classifier of the next.
3. Linnaeus named a genus of ginger Kaempferia.
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4. Ellis (1770).
5. Ellis’s correspondence with Linnaeus is cited in Loudon (1838, 1: 77), who inserts the botanical
names in parentheses.
6. The quotation from Ellis to Linnaeus appears in Smith (1821, 207). Siren lacertina, the greater
siren, is an eel-like amphibian named by Linnaeus in 1766; it occurs along the Atlantic and Gulf
coastal plains of eastern North America, south to southern Florida.
7. The ginkgo at the Mile End Nursery was described as the oldest and “most handsome” in En-
gland; see Lyman (1885) and Loudon (1838). Dr. John Hope lived from 1725 to 1786. For discussion of
Hope’s observations in London see Harvey (1981).
8. See Kaigi (1930). Linnaeus’s collections and texts are now scattered among various institu-
tions, including the Institut de France in Paris, the British Library in London, the Botanical Museum
in Uppsala, and the Swedish Museum of Natural History in Stockholm. The bulk of his collections
and the most important were sold in 1784 to James E. Smith and are the founding collections of the
Linnean Society of London; see Blunt and Stearn (2001) for further details. The specimen preserved
from the Uppsala plant is filed as Specimen 1292.2 in the Linnaean Herbarium, which is preserved in
the vaults of the Linnean Society of London beneath Burlington House on Piccadilly.
9. A possible source of Gordon’s living material of ginkgo is the tree that still survives in the
botanical garden at Utrecht, which was probably in cultivation by then. See Chapter 30 for more on
the Utrecht tree.
10. The “Chinese boy” portrait is of Wang-y-Tong, page to the Duchess of Dorset. Sir Joshua Rey-
nolds painted the portrait in about 1776. The painting is now in a private collection. Another version
of the portrait is in the Reynolds Room at Knole House in Sevenoaks, Kent (Martin Postle, Paul Mel-
lon Centre for Studies in British Art, personal communication).
11. Bradby Blake’s intentions “to procure the seeds of all trees, shrubs, roots, fruits, flowers, &c.
&c. which that great empire produces, and are used either for promoting commerce, or useful to
mankind” are described in the letters of Henry Laurens (Laurens, 1980), who succeeded John Han-
cock as the fifth president of the Continental Congress. Laurens was imprisoned in the Tower of Lon-
don during the Revolutionary War. He was the owner of Mepkin Plantation (now Mepkin Abbey),
one of the largest plantations in South Carolina. Bradby Blake died in Canton on November 16, 1773.
News of his death arrived in England in May 1774, and after his death, Blake’s name was removed
from candidacy by the Royal Society, where John Ellis had introduced it just a few years before. For
details on the porcelain sent from China by Blake, see Corbeiller (1974). The papers of the British East
India Company are housed at the British Library in the India Office Records.
12. Ellis may have had Bradby Blake in mind when in 1770 he wrote his Directions for Bringing
over Seeds and Plants from the East Indies and other Distant Countries in a State of Vegetation, which
was written for “Captains of Ships, Surgeons and other Curious Persons”; see Ellis (1770). This was
written nearly sixty years before the 1829 invention of Wardian cases revolutionized the transport of
live plant material. Ellis also urged plant collectors to carefully examine seeds before buying them to
be sure they were fresh and not “decayed.”
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30. Gardens
1. Epigraph: Pollan (1991, 64). In Mahayana Buddhism, “Buddha nature” is the intrinsic poten-
tial for reaching enlightenment that exists in every sentient being. The Lotus Sutra, one of the most
widely revered scriptures of the Mahayana Buddhism that permeates the Buddhist traditions of
China, Korea, and Japan, declares “bodhisattvas each of these, I call the large trees.” For more on
“Buddhist ecology,” see Swearer (2001).
2. For more on early Jesuit missionaries in China see Hsia (2009). Dimensions of the Geetbets
ginkgo tree are from Kwant, The Ginkgo Pages. It is possible that this old female tree was overlooked
because, with no male trees nearby, seed was not being produced and it would not have been obvi-
ously female. The first female ginkgo to be recognized in Europe failed to produce seed until cuttings
were grafted onto male trees at other gardens; see Loudon (1838, 2096).
3. After the French Revolution, the Jardin du Roi became known as the Jardin des Plantes. Ehr-
hart (1787) also notes ginkgo growing at the Royal Gardens of Herrenhausen in Hannover, Germany.
4. This quotation is from Loudon (1838, 2099), who also reviews other early ginkgos planted in
Europe.
5. For more on the introduction of Japanese plants into European gardens, see Farrer (2001) and
Crane and Saltmarsh (2002).
6. Lord Bute was close to Princess Augusta, who founded the gardens at Kew in 1759. He was also
a mentor and early prime minister (1762–1763) for the young King George III.
7. For discussion of the provenance of the Old Lions, see Fergusson (2006). See Chapter 29 for
more on James Gordon.
8. For more on the early cultivation of ginkgo in Europe, see Loudon (1838). The Hortus Botani-
cus at Leiden was founded in 1590 to instruct medical students at the University of Leiden on the
healing properties of plants.
9. Pétigny’s ginkgo at the Jardin des Plantes in Paris was first grown in a pot and overwintered
in the greenhouse. It was planted outdoors by André Thouin in 1792, but its growth was stunted.
Another ginkgo obtained by layering from one of the four other Pétigny seedlings was planted sub-
sequently. The story behind the l’arbre aux quarante écus is from Loudon (1838, 2096).
10. Pierre Marie Auguste Broussonet (1761–1807) was a French naturalist and honorary member
of the Royal Society; these plant exchanges are cited by Loudon (1838, 2096) and Wilson (1920, 56).
11. Loudon (1838, 2096).
12. Details of the first ginkgo to produce seed in Europe are given by Loudon (1838) and Wilson
(1919, 147). The female tree was apparently brought to the Bourdigny Estate in 1790 by an English
plant collector named Blakie who had stayed there while collecting plants in the Alps in 1776. After-
ward he often sent surplus plants from his stock to the proprietor of Bourdigny, M. Paul Gaussen
de Chapeau-Rouge. The ginkgo was part of a collection of trees and plants that Blakie was raising at
Monceau for the Duke of Orleans, and was probably grown from stock imported from England. How
the ginkgo obtained by Blakie came to Britain is uncertain (Loudon, 1838, 2097).
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13. Hamilton’s admiration for English woodlands is quoted by Madsen (1989, 14).
14. Hamilton’s prominence as an early North American collector of living plants is reflected in
his extensive collections of exotic plants. Among other acquisitions, he had three hundred silver firs
and five hundred Portuguese laurels shipped from Europe for Woodlands (Smith, 1905). When his
greenhouses were catalogued in 1809, they contained nearly ten thousand plants of five thousand to
six thousand species collected at great trouble and expense from all around the world (Oldschool,
1809, 507). For Hamilton’s correspondence with his secretary, see Smith (1905, 144).
15. When last measured in 1981, Hamilton’s ginkgo was nearly sixty-eight feet tall (Del Tredici,
1981, 155). It was felled in the mid-1980s, along with a nearby female of the same vintage, after the
caretaker’s dog grew sick from gorging on ginkgo seeds (Madsen, 1989, 23). For more on Collinson
see Dillingham and Darlington (1851). John Bartram (1699–1777) is sometimes considered the father
of American botany; Linnaeus called him the “greatest natural botanist in the world.” Bartram’s Gar-
den, established around 1728, is the oldest surviving botanical garden in North America. David Ho-
sack is better known as the doctor who tended Alexander Hamilton’s fatal wound following his duel
with Vice President Aaron Burr in 1804. The exact date of the ginkgo’s planting at the Vanderbilt Man-
sion Estate is unknown, but is estimated as between 1799 and 1835, by either Hosack or Dr. Samuel
Bard (Dave Hayes, Roosevelt-Vanderbilt National Historic Sites, personal communication). Hosack
was also the founder of Elgin Botanical Garden, a garden mainly of native plants, which he estab-
lished in New York City in 1770 on a site now occupied by Rockefeller Center.
16. The elder Michaux was tasked with searching out new species of trees through which France
could rebuild its forests. Important for shipbuilding, forests had been decimated by overharvesting
through nearly a century of warfare with England. Michaux discovered and described more than
three hundred different kinds of plants and shipped more than ninety cases of American stock and
seeds back to France. Michaux’s reputation was tarnished by work he did delivering messages for
Citizen Genêt, the new French minister to the United States. Genêt, to the outrage of President Wash-
ington, attempted to rally American citizens in taking arms against Spain; see Williams (2004). Coth-
ran (1995) discusses Michaux’s garden in Charleston. François Michaux published an account of his
travels, Michaux’s Travels to the West of the Alleghany Mountains, in 1805. The garden that his father
founded in Charleston no longer exists, although several fine, but younger, ginkgo trees can still be
found there, for example, outside the Charleston Historical Society and at the nearby Middleton Place
Estate.
17. Hawke and Rush (1971) describe the parade celebrating the ratification of the Constitution. For
more on the horticultural interests of the founding fathers of the United States, see Wulf (2011).
18. Henry Clay was one of the most influential nineteenth- century American politicians. He
played a significant role in leading the nation to war in 1812, but was also dubbed the Great Com-
promiser for his work during the 1820s and 1830s in negotiating agreements during the nullification
crisis and on the issue of slavery; see Heidler and Heidler (2010). Clay reputedly acquired ginkgos as
seedlings sent from Japan to Washington, D.C.; see Claxton (1940). The ginkgos on his estate (Ash-
land in Lexington, Kentucky) were planted roughly at the time of the Civil War; see Ashland (2012).
19. See Falconer (1890) for the quotation from Smith. The article was published in The Garden,
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a British publication, which explains why the cost is given in British pounds. The value of £1 in 1890
would be the equivalent of about $120 today. About fifty years earlier, Loudon (1838, 2100) gives the
cost as follows: “Plants, in the London nurseries, are from 1s 6d to 5s, according to the size; female
plants, 5s each. At Bollwyller, plants are 5 francs each; and at New York, 2 dollars.” For more on the
ginkgo trees at the Kentucky Military Institute see Falconer (1890) and Del Tredici (1981, 157).
20. Patricia Colleran, who worked in Giannini Hall on the Berkeley campus, won first prize in a
campus photo and writing contest for her poem about the tree, which is available online: see Kell
(2005).
21. The ginkgo haiku by Sem Sutter was published in the University of Chicago Magazine, April 30,
2007.
22. Santamour et al. (1983b) list eighty horticultural varieties of ginkgo, but Begović Bego (2011)
cites at least 220. For a list of ginkgo cultivars based on nut size and shape see Santamour et al. (1983b,
91). A selection of ginkgo cultivars are planted in the Bamboo Garden at the Royal Botanic Gardens,
Kew.
23. The leaves of Saratoga recall the frayed margins of several fossil species from the ginkgo lin-
eage, including Ginkgoites telemachus and several species of Sphenobaiera; see Anderson and Ander-
son (2003). The Japanese practice of creating bonsai was first brought to the attention of the West by
Engelbert Kaempfer, the first Western botanist to describe ginkgo.

31. Nuts
1. Epigraph: Shakespeare (1623a, act 3, scene 2). To a botanist, the so-called ginkgo nut is strictly
a seed and is not to be confused with the true nuts (single-seeded, indehiscent fruits) of almond,
hazel, and many other flowering plants.
2. The quotation from Ouyang Hsiu is cited in Li (1963).
3. The Chun Chu Chi Wen, from the Song Dynasty, is cited by Foster and Chongxi (1992). Del
Tredici (1991) comments on the production nuts from a single large ginkgo. The estimate of dried nut
production in China is from He et al. (1997).
4. Excavations of a Neolithic settlement in Jordan (ca. 6,760 B.C.) recovered a large basket of
carbonized pistachio nuts that would have weighed around forty pounds when fresh (Henry and Ser-
vello, 1974); see Rosengarten (1984) for more on the history of nuts and people. Huntley and Birks
(1983) describe the changing postglacial forest composition of Northern Europe. Abrams and No-
wacki (2008) summarize the likely impact of Native Americans on the pre-Columbian vegetation of
North America.
5. Botanically there is little difference between nuts and other convenient “naturally packaged”
plant foods, such as grains. Nuts were hard to come by in the early 1600s, and the word came to signify
a source of pleasure, as in the eighteenth-century phrase “to be nutts upon.” Originally this meant to
be “very fond of,” but in mid-nineteenth-century American English it came to mean “crazy.” See On-
line Etymology Dictionary, “nuts,” www.etymonline.com/index.php?term=nuts. The edible portion
of ginkgo nuts has a starch content of 68 percent, a very low fat content of 3 percent, and an energy
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value of only 940 calories per pound. By comparison, nuts of pinyon pine have a fat content of 60
percent and an energy value of 2,800 calories per pound. For every 100 grams dry weight, ginkgo nuts
contain approximately 403 calories, 10.2–10.5 percent protein, 3.1–3.5 percent fat, 83 percent carbohy-
drates, 1.3 g fiber, 3.1–3.8 g ash, 11 mg calcium, 327 mg phosphorus, 2.6 mg iron, 15 mg sodium, 1,139
mg potassium, 392 mg beta-carotene equivalent, 0.52 mg thiamine (B1), 0.26 mg riboflavin (B2), 6.1
mg niacin, and 54.5 mg ascorbic acid. For more on the nutritional value of nuts, see Duke (1989) and
Kris-Etherton et al. (1999).
6. The original classification of ginkgo nuts (Tsen, 1935) recognized three varieties: the normal
so-called variety typica, which includes the Meihe-Yinxing group or plum-stone-shaped ginkgo, with
a round seed; the huana, or “finger-citron” ginkgo, with an elliptic or oblong seed; and the apiculata,
the Maling-Yinxing or “horse’s bell–shaped” ginkgo, with a small point at the top of the seed.
7. The young embryo developing in the seed is bitter to the taste and is sometimes removed by
pushing it out with a toothpick.
8. Occasionally access to ripe ginkgo seeds becomes a source of conflict. In one Oakland neigh-
borhood a novice herbalist planning to harvest seeds from the female tree on her property called the
police when a neighbor beat her to the crop (Kemba Shakur, Urban ReLeaf, personal communica-
tion).
9. See Crosby (2008). Beware “ginkgo” seeds available on the Internet from Cameroon. These are
imposters, most often the similar “meats” of gnetum, a very different but equally interesting plant, a
climber from tropical forests. Gnetum seeds are often ground and fried into crackers called emping,
which are enjoyed in traditional Indonesian and southeast Asian cuisine.
10. The medicinal uses of raw ginkgo nuts appear in the Bencao Gangmu, or The Compendium of
Materia Medica, written by Li Shizhen (1518–1593) during the Ming Dynasty; see also Hori and Hori
(1997), Foster and Chongxi (1992, 256), and Chapter 34. Yoshimura et al. (2006) report on the content
of ginkgotoxin in raw and cooked seeds.
11. Many nuts and seeds used traditionally for oil production are finding specialty markets, but
ginkgo is not, so far, among them. Watermelon seed oil, traditionally used in West Africa, is a popular
emollient in cosmetic lotions. Oil from pumpkin seeds is a culinary delicacy in Austria, Slovenia, and
Hungary. Argan oil, produced from a tree endemic to the desert valleys of southwestern Morocco, is
an important constituent of high-end cosmetics. On a much larger scale, the African oil palm yields
an estimated 117.5 million tons of palm oil and 2.1 million tons of palm kernel oil per year, from more
than sixteen million acres of plantations in southeast Asia alone.
12. Hori and Hori (1997). Shogun Ashikaga Yoshiteru reigned from 1536 to 1565. The traditional
pairing of ginkgo nuts with sake is described by Hageneder (2005).
13. The Edo period in Japan began in 1603 and ended in 1868. The source of the quotation is Morse
(1917, 365).
14. For the putative ability of ginkgo to counteract the effects of alcohol in animals see Duke
(1997).
15. For more on the ginkgo at the Tsurugaoka Hachiman-gū shrine in Kamakura, see Chapter 24.
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32. Streets
1. Epigraph: Written by S. Kupka and E. Castillo and recorded by Tower of Power on its first
album, East Bay Grease, 1970—San Francisco Records/Atlantic Records. For more on the urban for-
ests of the United States see McPherson (2003). For recent statistics on the decline of urban tree cover
in the United States see Nowak and Greenfield (2012). For a short perspective on the importance of
nature in cities see Crane and Kinzig (2005).
2. For more on Trees for Cities, see www.treesforcities.org. The late Nobel Peace Prize laureate
Professor Wangari Maathai, who raised global awareness of the importance of trees through her work
as founder of the Greenbelt Movement in Kenya, was the patron of UNEP’s Plant for the Planet: The
Billion Tree Campaign.
3. Wolf (2003) reports on the economic benefits of street trees in business districts.
4. For an explanation of how i-Tree Streets calculates benefits, see i-Tree Tools (2010).
5. Inventories of urban street trees were conducted by the USDA Forest Service Center for Urban
Forest Research. The number of ginkgo trees in New York has increased since the 2007 inventory. The
MillionTreesNYC initiative, a campaign to plant and care for one million trees in New York City over
a decade, pledged in 2008 to plant 5,190 additional ginkgos in the ten-year period. Of the projected
$82.40 in benefits gained from every ginkgo planted, the aggregated values are ranked as follows: aes-
thetic/other: $38.98; rainfall interception: $20.13; energy savings: $19.40; air pollutants neutralized:
$3.39; total CO2 sequestered: $0.50. In the i-Tree system, aesthetic value accounts for nearly half of
the total estimated value of ginkgo as a street tree.
6. For more on the effects of trees on driving speeds see Naderi et al. (2008), and for effects on
stress response in drivers, see Parsons et al. (1998); for more on the study of hospital patients, see
Ulrich (1984); see also Kuo (2003), Kuo and Faber Taylor (2004), and Kuo and Sullivan (2001).
7. For more on the calculated benefits of urban trees, see USDA (2003). Gerhold (2007) describes
the origins and history of urban forestry practice. In five years the Urban Resources Institute is work-
ing to plant about five thousand trees in New Haven, including an estimated 150 ginkgos. With an eye
to long-term disease resistance in the urban forest as a whole the group plants no more than 5 percent
of any one kind of tree. Similarly, in Oakland, California, Urban ReLeaf hires at-risk youths to plant
trees in their own communities, putting them to work alongside scientists from the Center for Urban
Forest Research to collect data on the many benefits that trees provide. For more on urban forestry
as a tool for community development and green job creation, see Walsh (2011) and Pullen (2005).
8. There is also a long tradition of planting plane trees along the roads of France. Often attributed
to Napoleon, the practice apparently goes back to Henri IV and has recently drawn the ire of some
segments of French society who point to the role of trees in traffic fatalities (Economist, “The Killer
Trees”).
9. For more on the history of garden design, and the history of linking trees with public health
in cities, see Gerhold (2007).
10. The quotation is from Thone (1929). Unfortunately, the celebrated ginkgo allée near the De-
 328
n o t e s t o pa g e s 2 3 7 – 2 4 4

partment of Agriculture has long since been uprooted; many turned out to be female as they matured.
Only a handful of large ginkgo trees, perhaps of that vintage, remain in that part of the National Mall,
for example, near the northwest corner of the Department of Agriculture and not far away near the
edge of the Tidal Basin.
11. Chlorpropham is more commonly used to prevent sprouting in potatoes; see US EPA (2002).
12. See Brown (2006) for more on municipal efforts to protect heritage trees, including those on
private properties. Information on the New York City urban forest is from Andrew Newman (Project
Coordinator, MillionTreesNYC, personal communication).
13. Urban forest statistics for Manhattan are from Peper et al. (2007). Ginkgo is so much part of
the streets of New York that Disneyworld uses ginkgo in their re-created New York City streetscape
in the Hollywood Studios theme park.
14. For more information on ginkgo in Japanese landscapes, see Handa et al. (1997). The use of
ginkgo as a street tree in South Korea is on a similarly massive scale. In central Seoul in particular,
ginkgo is by far the most common street tree.
15. “Nasty, brutish and short,” see Quammen (1998, 71), who borrows this famous phrase from
Hobbes. Gangloff ’s quotation is from Brown (2006).
16. Studies in Britain show that ginkgo can thrive in hot summers and may benefit from warmer
conditions, provided that sufficient water is available (Fieldhouse and Hitchmough, 2004). See Bock-
heim (1974) and Shober and Toor (2009) for more on the properties of urban soils.
17. Injured ginkgo trees are sometimes said to be slow to isolate (compartmentalize) injured parts
of the tree, but the arborist who worked on the Everett ginkgo reported, “The tree had compartmen-
talized better than any tree I’d ever seen, and I’ve been in this business for 46 years”; see Mallow
(2008).

33. Pharmacy
1. Epigraph: Aristotle, 350 B.C., in Ogle (1912).
2. This folktale is from the information sign at the Myeoncheon Ginkgo. In another folktale, Lon
Po Po, the Chinese version of Little Red Riding Hood, three children outsmart a wolf that tries to eat
them by tricking it into falling to its death from a ginkgo tree; see Cai (1994).
3. According to the doctrine of signatures, a plant “provides clues” to its medicinal efficacy
through its resemblance to parts of the human body. It was long believed that the chi-chi of ginkgos
were the result of nutritional excess and that nursing mothers would benefit from an infusion made
from the shavings. Marks from such shavings are evident on some old trees, for example, at Jounichiji
Temple in Japan; see Himi City, “The Jounichiji Ginkgo.”
4. The Chinese term for the meat from a single ginkgo nut, bai guo, means literally “white fruit.”
Ma huang is the common name applied to several species of Ephedra that are native to the western
United States. For a description of the many uses of ginkgo nuts in traditional Chinese medicine see
Foster and Chongxi (1992, 257). Ginkgo nuts are crushed and applied topically to clear the complex-
 329
n o t e s t o pa g e s 2 4 4 – 2 4 6

ion. Flavonoid-rich extracts from the leaves have shown activity as free radical scavengers, which af-
fect pigmentation. This supports the earliest documented use of the leaf extract in Chinese traditional
medicine to treat freckles; today an extract is recommended as an alternative treatment for vitiligo;
see Soni et al. (2010), Zhu and Gao (2008).
5. The first chemical investigation of ginkgo (Peschier, 1818) described butyric acid from the
seeds. K. Nakanishi, who worked on early studies to characterize the structure of chemical com-
pounds in ginkgo, once quipped that the ginkgolide molecule “is a diterpene with an aesthetically
beautiful cage skeleton. . . . Because the Ginkgolides can be obtained in several large polymorphic
crystalline forms, and because of their extreme stability, we finally thought they might be nice as pen-
dants. However, they are not to be licked because they will be very bitter tasting” (Nakanishi, 2000).
Ginkgolide B was first synthesized by Elias Corey of Harvard University, who received the Nobel Prize
for chemistry in 1990 for his novel approach to making complex organic molecules by working back-
ward from the desired product.
6. According to Loudon (1838, 2098), “Thunberg says that even the fleshy part of the fruit is eaten
in Japan, though insipid or bitterish; and that, if slightly roasted, skin and all, it is not unpalatable.
Some of the fruit which ripened in the botanic garden of Montpelier were tasted by M. Delille and
M. M. Bonafous of Turin, who found their flavor very like that of newly roasted maize.” Lyman (1885)
notes, “The juice of the thick pulp outside the nut is very astringent and is used in making a some-
what waterproof, tough paper, and a preservative black wash for fences and buildings.”
7. Naxi recipes provided by Selena Ahmed, Tufts University (personal communication, 2011).
Ginkgotoxin (methoxypyridoxine) and its derivatives are made through the vitamin B6 pathway
involving genes (PDX1 and PDX2) that are necessary for seedling growth and survival in the model
plant Arabidopsis. A single piece of raw ginkgo nut contains about eighty micrograms of ginkgo-
toxin. Ginkgotoxin suppresses the formation of GABA, essential for proper function of the brain and
the central nervous system, inducing repetitive seizures, and also interferes with the body’s uptake
of vitamin B6. In Japan about 27 percent of the cases of ginkgo nut food poisoning result in deaths,
probably because of the increased susceptibility of children. The negative effects can be counteracted
by supplementing the diet with foods high in B vitamins; see Gengenbacher et al. (2006), Wada et al.
(1985), and Wada (2000).
8. For estimates on global use of ginkgo leaf extract, see Pérez (2009). Ben Cao Pin Hue Jing Yaor
was written by Liu Wen-Tai in 1505.
9. Flavonoids are part of a class of chemical compounds known as polyphenols, and their ability
to scavenge free radicals is thought to give them helpful antibacterial, antiviral, antitumor, anti-
inflammatory, and antiallergenic activity (Robberecht and Caldwell, 1983).
10. Yoshitama (1997) provides a comprehensive overview of the flavonoids of ginkgo.
11. The extract is standardized to 22–27 percent flavonoids and 65–67 percent terpenes. Ginkgolic
acids, the allergenic compounds in ginkgo nuts that are also present in the leaves, are kept below
5 parts per million. For more on the extraction process, see Juretzek (1997) and Sticher et al. (2000).
For more on the history of commercialization of ginkgo extract, see van Beek (2000, 316).
 330
n o t e s t o pa g e s 2 4 7 – 2 5 5

12. Before the reform of the German health care system in 2004 most herbal medicines were
deemed reimbursable by public health insurance. However, the reforms abolished reimbursement for
all herbal medicines except for standardized preparations of ginkgo, Saint John’s wort, and mistletoe;
see Bonakdar (2010, 96). For more on the market for ginkgo leaf extract, see Jensen et al. (2002) and
Diamond et al. (2000).
13. Global sales of ginkgo extract exceed $1 billion annually; see van Beek (2000). In 1997 annual
sales broken down by country were $280 million in Germany, $200 million in the rest of Europe, $205
million in the United States, and $200 million in Asia (Pérez, 2009). Most of these products use a
50:1 ratio by weight: fifty pounds of leaf are reduced to one pound of the resulting extract. For use as
a dietary supplement, the recommended daily dose, according to The Physician’s Desk Reference for
Herbal Medicines, is 120 mg of dry extract, in divided doses; see Diamond et al. (2000) and Chabrier
and Roubert (1988).
14. The standardized purified extract from ginkgo leaves is commonly referred to as EGb 761. Side
effects are few; in a study of 739 patients, 2.6 percent reported gastrointestinal upset, 0.9 percent ex-
perienced headaches, 0.4 percent reported sleep disturbance or dizziness, and 0.3 percent suffered
skin eruptions. While there are some reports of antiplatelet activity leading to increased bleeding in
those taking anticoagulants, a recent review concluded that there is scant evidence that ginkgo poten-
tiates the effects of such drugs; see Bone (2008). Ginkgo treatments are from Letzel et al. (1996) and
Diamond et al. (2000). For potential contraindications, see Medline Plus (2011).
15. For a careful assessment of the likely efficacy of ginkgo extract for enhancing cognitive func-
tions, see Gold et al. (2003) and Gold et al. (2002). See also the Ph.D. thesis by York (2006) for a meta-
analysis of more than a thousand clinical studies.

34. Risk
1. Epigraph: Darwin (1859, 392–393). Raup and Stanley’s book Principles of Paleontology (Raup
and Stanley, 1971) was a landmark in the development of paleontology and helped increase recog-
nition of its relevance to evolutionary theory (Sepkoski and Ruse, 2009). The book has now been
revised by two of Raup’s students (Foote and Miller, 2007). The Chicago School of Paleontology
is carried forward by my former colleagues in the Department of the Geophysical Sciences, Kevin
Boyce, Michael Foote, Dave Jablonski, Sue Kidwell, Michael LaBarbera, and Mark Webster, as well
as by many students of the Chicago program who are now scattered across the United States and the
world.
2. Jack Sepkoski’s compilation of data continued even as he was already analyzing it. Raup and
Sepkoski (1984) analyzed data from 3,500 families. Raup and Sepkoski (1986) analyzed data on nearly
11,800 genera.
3. According to some commentators, we are currently in the middle of a sixth mass extinction,
of similar magnitude to those of the past, which is driven by people and their increasing planetary
footprint. This may be an apt description of the current biodiversity crisis, but the data we have today
 331
n o t e s t o pa g e s 2 5 5 – 2 5 8

on extinctions relate mainly to land-dwelling vertebrate animals, such as birds and mammals, rather
than to marine invertebrates on which the idea of the “big five” extinctions is mainly based. Raup and
Sepkoski (1982, 1983) analyzed the long-term effects of the different extinctions, and the nature of
the subsequent recoveries. They also noted some regularity in the interval between twelve extinction
events, which pointed toward periodic extinction peaks on a roughly twenty-six-million-year cycle
(Raup and Sepkoski, 1984). Together with growing evidence in the early 1980s that the K-T extinction
had been caused by an asteroid impact, this led to the idea that all twelve extinctions may have had
an extraterrestrial basis—for example, periodic showers of comets caused by predictable astrophysi-
cal phenomena. Despite initial interest in this idea, the likelihood of the twelve extinctions having a
similar cause is now discounted by most paleontologists.
4. See Chapter 23 for more on possible relict living populations of ginkgo in China.
5. Luis Alvarez and his son Walter were the first to suggest that an asteroid impact may account
for the extinctions at the K-T boundary; see Alvarez et al. (1980).
6. Gould (1986) posed this question and later pursued a similar theme in book form (Gould,
1989).
7. The IUCN’s Red List ranks species under different categories of threat that range from criti-
cally endangered through endangered to vulnerable. For additional details on the categories and the
formal criteria on which they are based see www.iucnredlist.org.
8. According to the IUCN, Père David’s deer (Elaphurus davidianus) has been extinct in the wild
since 2008; the scimitar oryx (Oryx dammah) has been extinct in the wild since 2000. A global assess-
ment of the state of plant diversity is more difficult to accomplish than for vertebrate animals because
of the much larger number of species involved. However, an important initial unbiased evaluation
was made through a Sampled Red List Index Assessment presented at the 2010 Conference of the
Parties to the Convention on Biological Diversity held in Nagoya, Japan. The results show that the
world’s plants are just as threatened as the world’s mammals and more threatened than the world’s
birds; one in five of the species sampled is threatened with extinction. For a summary of this joint
study by the Royal Botanic Gardens, Kew and the Natural History Museum, under the auspices of
the IUCN, see Kew (2010). For more on threatened conifers, see the IUCN Conifers Status Survey and
Conservation Action Plan.
9. The World Wide Fund for Nature is known as the World Wildlife Fund in the United States;
both are generally abbreviated to WWF. For more details on the Living Planet Index see WWF (2010).

35. Insurance
1. Epigraph: From Joni Mitchell’s album Ladies of the Canyon, “Big Yellow Taxi,” words and
music by Joni Mitchell, © 1970 (Renewed) Crazy Crow Music. All Rights Administered by SONY/ATV
Music Publishing. The Yale School of Forestry and Environmental Studies, the oldest graduate for-
estry program in the United States, was founded in 1901. From its inception, the school strongly in-
fluenced the practice of American forestry, with the first five chiefs of the U.S. Forest Service coming
 332
n o t e s t o pa g e s 2 5 9 – 2 6 3

from its graduates and faculty. Aldo Leopold graduated from the Yale School of Forestry and Envi-
ronmental Studies in the Class of 1909.
2. In a similar way to ginkgo, the Monterey pine survives today in only three native stands, all of
which are threatened by pitch canker infection. As a timber tree it is planted over at least seven mil-
lion acres worldwide (Conifer Specialist Group, 1998).
3. Despite stringent protection, in 2005 some of the Wollemi pines were found to be infected with
Phytophthera cinnamomi, a potentially devastating pathogenic fungus probably introduced by hikers
who had found their way into the restricted area. For more information see Salleh (2005).
4. The Global Trees Campaign is a partnership between Fauna and Flora International, Botanic
Gardens Conservation International, UNEP World Conservation Monitoring Centre, and others: see
www.globaltrees.org.
5. For additional information on the drautabua and its conservation status see Farjon and Page
(1999).
6. Currently only about one in one hundred seedlings of the Mulanji cypress survive, includ-
ing those in cultivation. Clement Chilima of the Forestry Research Institute of Malawi described re-
cent management and conservation interventions for this and other plant species from Malawi at
the October 2009 conference Plant Conservation for the Next Decade: A Celebration of Kew’s 250th
Anniversary. The fourth species of Widdringtonia, the mountain cedar, is relatively widespread from
southern Malawi down to the Cape.
7. See Bartram (1791). The estimate of when the ancestor of the Hawaiian cotton tree arrived in
Hawai’i is based on Seelanan et al. (1997). For additional information on some of the plant species no
longer found in the wild but part of the collections of the Royal Botanic Gardens, Kew, see Kew, n.d.
8. For a review of ex situ plant conservation including effective integration with in situ conserva-
tion, see Guerrant et al. (2004).
9. The premise is that there is only so much money to go around; funds spent on one kind of
conservation will not be available to be spent on other conservation initiatives. However, my experi-
ence is that different funders have different priorities; funds available for one purpose are often not
available for another. A case in point was the funding provided for the Millennium Seed Bank project
by the British government, which resulted in the ex situ conservation of twenty-four thousand plant
species in ten years. This was possibly the largest single effort ever undertaken to conserve plant di-
versity. It drew on new funds that were unavailable for in situ conservation.
10. Many modern botanic gardens are increasingly moving along a similar trajectory to that fol-
lowed by zoos over the past fifty years from menageries to proactive conservation organizations.
11. I thank my Yale colleague Michael Donoghue for helpful discussion of extinction as a loss of
knowledge.
 333
n o t e s t o pa g e s 2 6 4 – 2 7 1

36. Gift
1. Epigraph: Hyde (1983, 101).
2. The Convention to Combat Desertification was opened for signature in October 1994 and
entered into force on December 26, 1996. Official information on the Convention on Biological Di-
versity can be found at www.cbd.int.
3. Rachel Carson’s book Silent Spring (Carson, 1962) is credited with helping to launch the envi-
ronmental movement. The plight of Chico Mendes and the Amazonian rubber tappers was brought
to international attention in 1990 with a series of films called A Decade of Destruction by the British
filmmaker Adrian Cowell, which gave greater visibility to the destruction caused by thousands of for-
est fires in the Amazon.
4. An assessment of plant diversity in the Atlantic rainforest is provided by Murray-Smith et al.
(2009).
5. Biological diversity is treated as national patrimony of particular countries under the CBD, but
it is self-evident that the distributions of species do not respect national borders, that the boundaries
of countries only rarely have a basis in biological or ecological reality, and that sensible conservation
strategies will require integration rather than a country-by-country approach.
6. CBD Article 15 recognizes the sovereign rights of states over their natural resources. On Octo-
ber 29, 2010, the Nagoya Protocol on Access and Benefit-sharing (ABS) was adopted by the Confer-
ence of the Parties to the CBD, following eleven meetings in six years by the Ad Hoc Open-ended
Working Group on ABS. The Nagoya Protocol aims to provide a transparent legal framework for the
fair and equitable sharing of benefits arising out of the utilization of genetic resources: see www.cbd
.int/abs.
7. Official information on the International Treaty on Plant Genetic Resources for Food and
Agriculture can be found at www.planttreaty.org. Conditions by which amendments to the Treaty
may be made are elaborated in “Article 23—Amendments of the Treaty”; Article 23.3 specifies that
amendments shall be made only by consensus.
8. Perhaps the kindest thing that can be said about the CBD is that it has helped keep biodiversity
on the international policy agenda, and at least for plants, one positive outcome was the adoption of
the Global Strategy for Plant Conservation (GSPC) by the Conference of the Parties at their meeting
in The Hague in the spring of 2002. This had the helpful effect of focusing the world’s botanists, espe-
cially those in the botanical garden community, on the common goal of conserving plant diversity.
The global strategy emerged from an appeal by Peter Raven, then director of the Missouri Botanical
Garden, at the International Botanical Congress held in Saint Louis in 1999, which was followed by a
meeting in the Canary Islands that resulted in the Gran Canaria Declaration. Additional information
on the global strategy can be found at www.cbd.int/gspc. The Global Strategy for Plant Conservation
is now into its second ten-year iteration, the original 2010 deadline for the completion of the first
round of GSPC goals having now passed.
9. Since 1996 the Forum on Religion and Ecology, organized by my Yale colleagues Mary Evelyn
 334
n o t e s t o pa g e s 2 7 1 – 2 7 6

Tucker and John Grimm, has highlighted the important roles that religions play in helping guide how
people interact with their environment; for additional information see www.religionandecology.org.
10. For additional information on the philosophy of Thomas Berry see Berry (2009). For more on
the theory of biophilia, see Kellert and Wilson (1993).
11. See National Academy of Sciences (2011, 1).

37. Legacy
1. Epigraph: Hyde (1983, 182).
2. Tony Kirkham updated the classic The Pruning of Trees, Shrubs, and Conifers, first published
by George Ernest Brown in 1977 (Brown and Kirkham, 2004).
3. On his 1908 expedition for the Arnold Arboretum, Wilson photographed a large ginkgo at the
village of Lengji in the valley of the Tung River. The caption to his photograph, preserved in the ar-
chives of the Arnold Arboretum, records the tree at an elevation of three thousand feet. Wilson esti-
mated it as eighty feet tall, and his photograph clearly shows the small shrine placed in the branches
near the base of the tree. Tony Kirkham and Mark Flanagan revisited the same ginkgo and photo-
graphed it in the summer of 2001; see Flanagan and Kirkham (2010).
4. The fall of the great ginkgo at the Tsurugaoka Hachiman-gū Shrine in Kamakura, Japan, was
reported in the Economist; see “Japan’s favourite tree: An Easter story from Japan” (2010). For more
detail on the restoration of the Mizufuki Ginkgo see Handa (2000, 32). The Japanese concern for old
ginkgo trees also comes through in the story of the Kubi Kake Ginkgo, the “Head-Stake Ginkgo,”
sometimes called the “Over My Dead Body Ginkgo,” in Hibaya Park. Rather than allowing the tree
to be felled for road expansion, Dr. Seiroku Honda, the designer of Hibaya Park, Tokyo, embarked
on a project of more than three weeks to move the tree more than a quarter of a mile to safety. He is
reputed to have said, “I will have it transplanted even if my head is put on a stake.”
5. See Chapter 24 for more on the Huiji Temple.
6. John Montagu (1690–1749) married the daughter of the Duke of Marlborough and served
under him at the Battle of Culloden. The elms that he planted were decimated by Dutch elm disease;
many of the lindens (“limes”), which were a smaller component of the avenues, still survive.
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 Illustration Credits

Frontispiece—Drawing by Pollyanna von Knorring, based on a photograph taken at the Royal Bo-
tanic Gardens, Kew.
Part I opener—Drawing by Pollyanna von Knorring, from the Economic Botany Collection of the
Royal Botanic Gardens, Kew.
Pp. 6, 11, 13, 19, 33, 36, 44, 49, 77, 112, 150, 163, 169, 181, 224, 231, 232, 238, 239, 243, 247, 269—Photo-
graphs by Peter R. Crane.
Part II opener—Drawing by Pollyanna von Knorring, from photographs in Hori and Hori (2005).
P. 28—Scanning electron micrograph courtesy of Zhou Zhiyan and collaborators.
P. 39—From Siebold and Zuccarini (1835–1870), vol. II, plate 136.
P. 47—Photograph courtesy of Andrew McRobb, © Royal Botanic Gardens, Kew; artwork attributed
to Chikusai Kato, 1878, from the Economic Botany Collection at the Royal Botanic Gardens, Kew.
P. 50—Photograph by Elisabeth Wheeler.
Pp. 58, 145—Photographs by Nancy Hines.
P. 69 (top)—Photograph courtesy of Masaya Satoh.
P. 69 (bottom)—Film still from The Sea in the Seed, Tokyo Cinema.
P. 71—Photograph courtesy of Marie Stopes International.
Part III opener—Drawing by Pollyanna von Knorring, redrawn from Zhou and Zhang (1989, 1992)
and from Zhou (2009).
P. 83—Photograph by Peter R. Crane, from material at the Swedish Museum of Natural History,
Stockholm.
Pp. 87, 99, 102, 121—Andrew B. Leslie.

362
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i l l u s t r at i o n c r e d i t s

P. 95—Drawing by Pollyanna von Knorring, redrawn from Anderson and Anderson (2003), 288.
P. 105—Photograph courtesy of Meinte Boersma.
P. 109—Photograph by Ghedoghedo, from material at the Paläontologische Museum, Munich.
P. 115—Photograph by Andrew N. Drinnan.
P. 117—Andrew B. Leslie, based on Drinnan and Crane (1989).
P. 126—Image courtesy of Steven Manchester, paleobotanical collection of the Florida Museum of
Natural History, University of Florida, Gainesville.
P. 127—Image courtesy of Zhou Zhiyan, from material at the Field Museum, Chicago.
P. 135—Illustration by Francisco Manuel Blanco (1778–1845), from material at the Real Jardín Botá-
nico of Madrid.
Part IV opener—Drawing by Pollyanna von Knorring, based on Hori and Hori (1997, 391).
P. 153—Andrew B. Leslie, compiled and redrawn from Tralau (1967).
P. 166—Photograph by Ernest Henry Wilson (1914), © 2006, President and Fellows of Harvard Col-
lege, Arnold Arboretum Archives, Jamaica Plain.
P. 172—Drawing by Andrew B. Leslie.
Part V opener—Drawing by Pollyanna von Knorring, based on material from Kyushu Ceramics
Museum, Japan.
P. 192—Drawing by Pollyanna von Knorring, based on a model at the National Maritime Museum,
Mokpo, Korea.
P. 197—Image from Isaac Titsingh from material at National Library of the Netherlands, The Hague.
P. 202—Photograph by Ashley DuVal from material of the Mertz Library at the New York Botanical
Garden, New York.
P. 212—Image by permission of the Linnean Society of London.
Part VI and VII openers—Drawings by Pollyanna von Knorring, based on photographs by Peter R.
Crane.
P. 260—Illustration courtesy of Jim Xerogeanes.
P. 274—Photograph courtesy of Andrew McRobb, ©Royal Botanic Gardens, Kew.
This page intentionally left blank
 Index

Note: Page numbers in italics refer to illustrations.

academia, 19–20 Amazon, 40, 149, 333n3

Adam and Eve, 10 American beech (Fagus americana), 8
Adams, John, 222 American Civil War, 21, 241
Adams, William, 198, 208 American elm (Ulmus americana), 35
Adanson, Michel, 179, 314n7 American Museum of Natural History, New
Afghanistan, 82, 83, 88, 89, 90 York, 98
Africa, 10, 89–90, 116, 117, 124, 179, 261–262, amphibians, 118, 257
265 Amsterdam, 236
African oil palm (Elaeis guineensis), 326n11 Anbulsa Temple ginkgo, 187
age, 177–183, 314n6 ancestry, 89–96
agriculture, 12, 14, 29, 268 Anderson, John and Heidi, 89–91, 94
Ainosho, 188 Andes, 169, 265
Alaska, 144, 148 Angel Oak, 14, 287n17
alcohol, 232 Angelou, Maya, 60
alder (Alnus spp.), 306n6 animals, 14, 15, 30, 62, 81, 83, 84, 109, 117, 122–
Allicospermum xystum, 304n9 124, 129–130, 137, 138, 158–159, 160, 254, 255,
Almond (Prunus dulcis), 231, 325n1 256–257, 259, 267, 268, 297n12, 310n6, 331n3,
Almont fossils, 125–126, 126, 127, 127, 128–131, 331n8
163, 306nn2–4 Antarctica, 5, 117, 261

365
 366
index

antiquity, 177–183 Banks, Sir Joseph, 16, 17, 66, 207, 210, 220,
apple (Malus pumila), 35, 162, 225, 272 288n4
apricot (Prunus armeniaca), 192, 289n16 banyan (Ficus benghalensis), 10
Archaefructus, 305n9 baobab (Adansonia digitata), 10, 179, 314n7
Archaeopteris, 84 Barbados, 10
Archangelsky, Sergio, 115–116 bark, ginkgo, 36, 48–49, 49, 50, 275
Arctic, 5, 139–140, 148 Bartram, John, 221, 222, 262, 324n15
Ardtun Head, 136–137 Bartram, William, 262
argan (Argania spinosa), 326n11 Bartram ginkgo, 221
Argentina, 115, 116 basalt, 136–137
Argyll, Duke of, 219 Basinger, Jim, 139
Aristotle, “On the Parts of Animals,” 242 bay tree (Laurus spp.), 136, 208
Arizona, 91 bearded fig (Ficus citrifolius), 10
aroids, 122, 137 beech (Fagus spp.), 49, 157, 227
art, ginkgo, 20–21, 39, 45–46, 47, 51, 175, 184, Beijing, 3, 22
194, 201, 202, 203, 251, 252, 293n9, 294n11, Beijing Botanic Garden, 21
294n18, 315n1 Belgium, 218
Art Nouveau, 20 Berlin-Dahlem Botanic Garden, 46
Arts and Crafts movement, 20 Berry, E. W., 137
ash (Fraxinus spp.), 10, 15, 157, 272 Berry, Thomas, 271
Asia, 5, 11, 143, 152, 156, 157, 161, 164, 168, 187, betel (Piper betle), 192
190–194, 195, 198, 265, 273, 276, 311n9. See Bible, 10, 81, 111, 168, 286n7
also specific countries big leaf hydrangea (Hydrangea macrophylla),
aspen (Populus spp.), 75 293n3
Asteroxylon, 85, 86, 300n6 biological diversity, 257, 262–271, 330n3, 331n8,
Atlantic cedar (Cedrus atlantica), 8, 9 333nn5–8; conservation of, 264–271
Aulenback, Kevin, 122 birch (Betula spp.), 139, 157
Australia, 9, 15, 20, 91, 94, 115, 116, 117, 143, 152, birds, 9, 83, 117, 158, 257, 267, 305n5
207, 259–260, 261, 265, 269–270 blackberry (Rubus fruticosus), 158
autumn, 32–33, 33, 34, 50, 54, 62, 72, 230 black locust (Robinia pseudoacacia), 219
Avatia, 90 black tupelo (Nyssa sylvatica), 9
azalea (Rhododendron spp.), 243 Blaise Castle ginkgo, 214
Blake, John Bradby, 213–214, 322nn11–12
“bad luck,” 259, 260, 262, 270 Blandford, Marquess of, 8, 285n1
Baiera, 120, 121 Blandy Experimental Farm, 64
Baiera furcata, 304n5 Bodhi tree, 10, 21, 286n8
Baiera gracilis, 304n5 Bok Ji-gyeom, 243
Baiera hallei, 114 Bones, Tom, 138
bald cypress (Taxodium spp.), 155 bonsai, 225, 325n23
 367
index

Borneo, 10, 277 camellia (Camellia spp.), 222

Boscobel, 10 Camerarius, Rudolf Jakob, 54, 61, 295n2
Bourdigny ginkgo, 296n5, 323n12 Canada, 22, 90, 122, 139, 144
Bournemouth fossils, 132–133, 135, 137 Candomblé, 10
Boveri, Theodor, 62, 297n8 carbohydrates, 28, 29
branches, ginkgo, 31, 34, 35–36, 36, 37, 38, 41, carbon, 14, 28, 29
46, 48, 51, 75–78, 85, 91, 118, 228, 229, 291n1; carbon dioxide, 28, 29, 30, 31, 85, 287n18, 314n7
chi-chi, 25, 75–76, 77; grafted, 61–64 Carboniferous, 87, 103, 121
Brazil, 10, 149, 265 Carlyle, Thomas, Chartism: Past and Present,
breasts, 243–244 125
bright-green cave-moss (Cyclodictyon laetevi- Carolina mahogany (Persea borbonia), 221
rens), 288n2 carotenoids, 33, 290n7
bristlecone pine (Pinus longaeva), 77, 179, 183, Carson, Rachel, 265, 333n3
276, 314n6 Castiglioni, Luigi, 161, 310n8
Broussonet, Pierre Marie Auguste, 220, 323n10 Castle Rock fossils, 138
Brown, Capability, 17 castor oil (Ricinus communis), 54, 61
Brown, Robert, 207 Cathay silver fir [yin shan] (Cathaya argyro-
Brudvig, Gary, 27 phylla), 157, 173
Buddhism, 5, 10, 12, 42, 173, 217, 323n1; ginkgos Caucasian wingnut (Pterocarya fraxinifolia),
and, 6, 12, 21, 51–52, 189, 193–194, 217, 251, 157–158, 160, 310n3
252, 288n9, 294n18 Cave Hill ginkgos, 223
buds, ginkgo, 35, 37, 38, 76, 78, 145, 145, 151, Cavendish Laboratory, 20
255, 300n3 cedar of Lebanon (Cedrus libani ), 10, 21
bulbs, 73 celery (Apium graveolens), 231
Bulgaria, 156 cell division, 48–51
Burgess Shale, 84 Cenozoic, 160, 254, 310n6
Burkina Faso, 12, 287n13 Central America, 158, 159, 265, 310n4
burls, 75 ceramics, 175, 176, 189, 196, 201–203, 213; trade,
Bute, Lord, 219, 323n6 190–194, 201–203
butyric acid, 244–245, 329n5 Chambers, Sir William, 17, 21
Chandler, Marjorie, 134, 135, 307n4
cabbage palmetto [palmetto] (Sabal palmetto), Charles II, King of England, 10, 320n7
10, 286n9 chawan mushi, 233
Café Marron (Ramosmania rodriguesii ), 262 chemicals, 244–246, 329n5
California, 7, 9, 11, 144, 179, 223, 235, 240, cherry (Prunus avium), 55
309n14, 327n6 chestnut (Castanea sativa), 227, 233
cambium, 48–50, 292n7 Chiang Kai-shek, 18
Cambrian, 87 Chicago, 8, 20, 22, 37, 72, 98, 128–129, 138, 144,
Cambridge University, 20, 106, 224, 224 224–225, 240, 253, 275, 276
 368
index

chi-chi, 25, 26, 75–76, 77, 167, 188, 217, 225, 243, 234–238, 238, 239, 239, 240–241. See also
243, 300n3, 328n3 specific cities
Chichi Ginkgo, 187 Clanwilliam cedar (Widdringtonia cedar-
China, 2, 5, 8, 10, 15, 18, 20, 104, 105, 111–124, burgensis), 262
144, 157, 162–173, 177–179, 190–194, 196, Clarkia fossils, 155, 157
212–214, 260, 273, 311n3; ceramics, 190–194, Clarno fossils, 138–139
213; climate, 139, 144–147, 149; edible nuts, clay, 134
226–231, 231, 232; ginkgos, 1, 2, 17–19, 21, Clay, Henry, 222–223, 324n18
22, 32, 51, 54–55, 76, 77, 79, 80, 111–124, 131, Cliff, Jimmy, “Many Rivers to Cross,” 156
139, 144–149, 160–169, 169, 170–173, 177–179, Clifford, George, 209–210
182–186, 188, 189, 196, 201, 204–205, 225, climate, 135–140, 143–151, 152, 154, 155, 306n7,
226–231, 242–246, 260, 270, 273, 275, 286n9, 308n9, 309nn10–14; change, 14, 15, 261, 264,
288n8, 310n6, 313nn3–5, 315n13, 316nn5–8, 277, 287n18; cold, 144–151, 255; extinction
328n2, 334n3; herbal medicine, 242–246; and, 157–161, 255, 261; growth cycle and,
Jinfo Mountain, 171, 172, 173, 177, 178, 186, 146–151
313n8; literature, 184–186; Opium Wars, climbing, tree, 9–10
164, 165; temples and shrines, 21, 166, 178, Clinton, Bill, 14, 287n18
182, 183, 217, 243, 275; Tianmu Mountain, cloning, 62, 238
166–167, 169, 169, 170–172, 172, 173, 178, 186, clothing, 194
313n8; trade, 190–196, 317nn5–6; wild popu- clubmoss, 85, 87, 99, 107, 122, 300n6, 303n3
lations and genetic diversity, 162–172, 172, coal, 14
173; Yima fossils, 79, 80, 113–115, 116, 119, coast redwood (Sequoia sempervirens), 9, 154,
120 179, 286n3, 314n6
China fir (Cunninghamia lanceolata, Cunning- coelacanth, 129–130
hamia konishii ), 122, 155, 157, 173 Collinson, Peter, 221, 222
Chinese date (Ziziphus jujuba), 231 Colorado, 138, 154
Chinese swamp cypress (Glyptostrobus pen- Columbia University, 62
silis), 157 commemorative trees, 21–22
chlorophyll, 29–30, 31, 32, 33, 290nn4–5 commerce, 12, 14, 15, 20, 22, 42, 43, 164
chloroplasts, 29–30, 31, 32 Conan Doyle, Sir Arthur, The Lost World, 3
chocolate, 207, 320n7 Confucianism, 5, 21, 217, 289n16
Christianity, 10, 196–197, 227, 271, 318n4, 318– conifers, 4, 50, 70, 91, 92 97, 98, 99, 101, 102,
319n6 107, 109, 118, 122, 132, 157, 179, 228, 257, 261
chromosomes, 54, 62, 63, 297nn9–11 conservation, 12, 17, 241, 258–263, 287n18–19,
Ciennich Tree, 196 332n2–10; of biological diversity, 264–271;
cilia, 70 ex situ, 260–263, 270; in situ, 262
cinnamon (Cinnamomum aromaticum), 192 constraint, 143–151
cities, 6–7, 15, 230, 234–241, 327nn 1–8, continental drift, 94, 302n10
328nn11–16; soils, 240–241; street trees, Convention on Biological Diversity, 264–271
 369
index

Convention on International Trade in Endan- decline and survival, 7, 123, 141–173; constraint,
gered Species, 265 143–151; endurance, 162–167; extinction, 156–
Coolidge, Calvin, “Have Faith in Massachu- 161, 253–257; relic, 168–173; retreat, 152–155
setts,” 27 Del Tredici, Peter, 76, 146, 147, 148, 167, 227,
cork, 49–50, 294n13 305n5, 308n6
cork oak (Quercus suber), 294n13 Denk, Thomas, 154
Costa Rica, 158 Denmark, 144
Council Oaks, 11, 286n10 desertification, 264, 333n2
Courtney, Jeff, 149 Deshima, 5, 43–45, 197, 197, 198–203, 219, 232,
crepe myrtle (Lagerstroemia indica), 222 319n6, 319n10
Cretaceous, 118, 121, 122, 123, 133, 152, 160, 254, Detroit, 21
255, 306n7 Devonian, 86, 87, 103
crocus (Crocus spp.), 73 de Vries, Hugo, 62, 297n7
croton (Croton tiglium), 192 Dhammapada, The, 42
crown, ginkgo, 31, 35, 37 dictionaries, 188, 205
cultivars, 225, 325n22 Dilcher, David, 125–128, 131, 137, 138, 306nn1–2
cultivation, 184–189, 259–263 Dinlinsi Temple ginkgo, 178
cultural history, 10–12, 18–23; early, 184–189; dinosaurs, 5, 83, 109, 117, 122, 159, 254, 305n5
naming, 204–208; renewal, 195–203; trade, dipterocarp, 277
190–194 disease, 6, 15, 151, 236–237, 243–249, 261, 273,
currant (Ribes spp.), 139 276, 297n14, 332n3
cuticle, 31, 85 diversity, 119–124, 225, 257, 262–263, 312n1; bio-
cuttings, 75, 211, 219, 222, 225, 237–238, 275 logical, 257, 262–263, 264–271, 330n3, 331n8,
Cycadofilicales, 71 333nn5–8; genetic, 168–173, 268
cycads, 68, 70, 71, 85, 97, 99, 101, 102, 107, 109, divi-divi (Caesalpinia coriaria), 158
257, 303n3 DNA, 20, 32, 54, 59, 63, 98, 101, 168–173, 312n2,
Cyclocarya brownii, 306n2 313nn5–6
cylinder. See cambium dove tree [handkerchief tree] (Davidia involu-
cypress, 4, 122 crata), 122
dragon tree (Dracaena draco), 314n7
Dalai Lama, 21 drautabua (Acmopyle sahniana), 261
Daoism, 5, 21, 51, 294n17 duck’s foot, 185, 188, 204, 205
Darwin, Charles, 4, 19, 53, 55, 60, 62, 86, 106, Dutch East India Company, 42, 43, 197–201,
161, 256, 285n3, 295n4, 310n8; Origin of 209, 318n6, 319n10
Species, 62, 253 Dutch elm disease, 276, 334n6
David, Père Jean Pierre Armand, 165, 257, 311n5 dwarfing, 165
Dawkins, Richard, The Selfish Gene, 53
dawn redwood (Metasequoia glyptostroboides), Early Cretaceous, 115, 117, 120
15, 122, 140, 157, 260, 313n9 Early Jurassic, 83, 89
 370
index

Early Miocene, 154 Everett ginkgo, 241, 328n16

Early Paleocene, 136 evolution, 9, 30, 40, 50, 55, 60–61, 70, 78, 159,
Early Permian, 91 253–257, 295n4, 296n2; ancestry and, 89–96;
Early Triassic, 91, 120 gender and, 60–65; natural selection,
Earth Summit (1992, Rio de Janeiro), 264 60–61, 86, 225, 295n4; origins and, 81–88;
Easter Island, 262 persistence and, 125–131; proliferation and,
ecology, 12–15, 158, 258–263 111–118; prosperity and, 132–140; “random”
Edo, 43, 45, 52, 200, 201 events, 256; recognition and, 104–110; re-
Edwards, Dianne, 130 lationships and, 97–103; winnowing and,
Eemian interglacial, 157–158 119–124. See also fossils; origin and prehis-
egg, 53, 54, 56, 57, 59, 63, 68, 70, 87, 88 tory; specific periods
Egypt, 10, 227, 236 ex situ conservation, 260–263, 270
Ehrhart, Frederick, 218 extinction, 7, 15, 91, 92, 102, 122–124, 156–161,
Elizabeth I, Queen of England, 10 253–257, 261–263, 271, 311n9, 330–331n3;
Elizabeth II, Queen of England, 273 future and, 253–257; Great Dying, 255; in-
Ellis, John, 210–214, 219, 322nn5–6, 322nn11–12 surance against, 258–263; K-T, 254, 306n7,
elm (Ulmus spp.), 139, 157, 276, 334n6 331n3; regional, 156–161; risk of, 253–257
Emancipation Oak, 11, 286n10 extracts, 7, 242, 245–249, 329nn5–11, 330nn12–15
Emberger, Louis, 72
embryo, 53–59, 63, 64, 68, 72, 88, 146–148, 229, Falkowski, Paul, 271
296nn10–11, 326n7 female ginkgo, 53–57, 58, 59, 60–65, 67, 72, 78,
Emerson, Ralph Waldo, “Works,” 177 129, 164, 185–186, 215, 216, 218, 220, 227, 229,
endurance, 162–167 237, 297n11, 323n2; ban, 237–238; “leaky,” 65;
energy, 27–34, 37, 40, 41, 59, 146 smell, 237–238
English oak (Quercus robur), 8, 10 ferns, 70–72, 84, 85, 87, 99, 107, 109, 122, 132,
Eoasteria, 90 137, 303n3
Eocene, 133, 136, 137, 138, 148, 153, 154 fertility, 243
ephedra [ma huang, Mormon tea] (Ephedra fertilization, 66–68, 69, 70–72, 78, 87, 88, 146,
spp.), 244, 302n6, 328n4 147, 295n9, 299nn10–13; self-, 55, 60
epidermis, 31 Field Museum, Chicago, 98, 128, 138
Ettingshausen, Baron Constantin von, 132–133 fig tree (Ficus spp.), 10, 196
eucalypt (Eucalyptus spp.), 9, 146 fir (Abies spp.), 140
Europe, 5, 6, 36, 43, 61, 62, 70, 81, 114, 129, 136, fish, 139, 233, 244, 257, 307n10
137, 143, 144, 150, 153, 154, 156, 157, 159, 160, fishtail palm (Caryota spp.), 225
161, 164, 165, 189, 198, 199, 201, 209, 214, Five Generations Ginkgo, 169
217–221, 236, 237, 246, 247, 255, 309n14, Flanagan, Mark, 273
311n9, 323nn2–12. See also specific countries flavonoids, 246, 329n4, 329nn9–11
Evelyn, John, 320n7; Silva: a Discourse of Forest Florin, Rudolph, 91, 92, 301n4
Trees, 196 Florissant fossils, 154, 309n4
 371
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flowering plants, 99, 101, 102, 106, 121, 122, 153, Gallows Oak, 11
285n4, 295n4, 302n5 Gandhi, Mahatma, Non-Violent Resistance
food, 189, 245; ginkgo nuts, 226–231, 231, 232, (Satyagraha), 73
232, 233 Gangjin-gun ginkgo, 186
Forrest, George, 165 gardens, 217–225
Fortune, Robert, 164–165, 311n3 Gardner, John Starkie, 132–133, 135, 137
Fort Union Formation, 123, 306n7 Geetbets ginkgo, 218, 323n2
fossils, 4, 14, 28, 70–71, 82–83, 83, 84–88, gender, 53–57, 59, 60–65, 78, 185–186, 216, 218,
89–96, 153, 153, 154–155, 160–161, 187, 253, 220, 237, 295n4, 295nn7–8, 297nn6–14; ban,
260, 263, 302nn11–12, 305nn3–6, 306nn1–7, 237–238; determination, 60–65; deviations,
307nn4–12, 308nn12–14, 309nn2–4, 310nn2– 63–65. See also female ginkgo; male ginkgo
8; Almont, 125–127, 126, 127, 127, 128–131, genetics, 62–63, 225, 297n7, 313nn5–8; diversity,
163, 306nn2–4; ancestry and, 89–96; ex- 168–173, 268
tinction and, 156–161, 253–257; Greenland, Geneva, 220–221
106–108, 110, 304n9; living, 129–131; Lon- George III, King of England, 16, 17, 66, 274
don Clay, 134–136, 138, 307nn4–5; Molteno, Germany, 20, 43, 64, 67, 68, 98, 137, 143, 153,
89–90, 94, 303n7; origins and, 82–88; per- 154, 156–157, 199, 206, 291n2, 309n2; medici-
sistence and, 125–131; proliferation and, 111– nal use of ginkgo, 246–247, 247, 330nn12–13;
118; prosperity and, 132–140; recognition Weimar ginkgo, 150
and, 104–110; relationships and, 97–103; germination, 54, 59, 78, 147–148, 160, 295n9,
Rhynie Chert, 84–88, 90, 93; winnowing 296n11, 308n8
and, 119–124; Yima, 79, 80, 113–115, 116, 119, giant sequoia (Sequoiadendron giganteum), 8,
120; Yorkshire, 108–109, 109, 110, 113. See 9, 179, 183, 314n6
also specific sites gift, 264–271
France, 40, 91, 143, 153, 165, 236, 246, 247, Gilbert and George, Ginkgo pictures by, 20–21,
324n16, 327n7; ginkgos, 219–220 289n15
Franklin, Benjamin, 222, 262 ginau, 208
Franklinia (Franklinia alatamaha), 262 ginger (Zingiber officinalis) 122, 192, 210, 321n3
Frederick, Prince of Wales, 17 ginkgo (Ginkgo biloba), 10, 15, 22, 39, 81, 102,
French Botanical Garden, 222 202, 212; ancestry, 89–96; antiquity, 177–183;
Friis, Else Marie, 82 constraint, 143–151; decline and survival,
fruit, ginkgo, 3, 58, 72 141–173; endurance, 162–167; energy, 27–34;
fuel, 12, 15 extinction, 156–161, 253–257; future, 251–
Fujii, Kenjiro, 67, 68, 71 277; gardens, 217–225; gender, 60–65; gift,
Fuller, Thomas, Gnomologia, 190 264–271; growth, 35–41; history, 175–214;
Fuquan City ginkgo, 178 insurance, 258–263; introduced to Western
future, 251–277; gift, 264–271; insurance, 258– science, 195–214, 217; legacy, 272–277; nam-
263; legacy, 272–277; risk, 253–257 ing, 22, 38, 81, 185, 188, 204–208, 211–212;
Fu-Yen Ssu Monastery, 184 nuts, 226–233; origin and prehistory, 79–
 372
index

ginkgo (Ginkgo biloba) (continued) Glover, Thomas, 42

140; origins, 81–88; persistence, 125–131; Gnetales, 99, 101, 102, 302n6
pharmacy, 242–249; proliferation, 111–118; gnetum (Gnetum spp.), 303n6, 326n9
prosperity, 132–140; recognition, 104–110; Goethe, Johann Wolfgang von, 6, 22–23, 38,
relationships, 97–103; relic, 168–173; re- 132, 143, 150
newal, 195–203; reprieve, 184–189; re- Golden Gate Park, 7
silience, 73–78; resurgence, 209–214; re- gomphotheres, 159, 310n4
treat, 152–155; risk, 253–257; seeding, 66–72; Gordon, James, 210–212, 214, 219, 220
sexual reproduction, 53–59; streets, 234– Goüan, Antoine, 220
241; use, 215–249; voyages, 190–194; wild Gould, Stephen Jay, 84, 256
populations, 162–173; winnowing, 119–124 grafts, 61–64, 225, 228, 296n5
Ginkgo adiantoides, 309n2 grape (Vitus spp.), 226
Ginkgo apodes, 118, 305n10 grape family, 136, 149, 154
Ginkgo australis, 115, 305n8 grasses, 153
Ginkgo cordilobata, 83, 83, 88, 89, 90 Gray, Asa, 161, 310n8, 311n9
Ginkgo cranei, 121, 126, 127, 306n4 Great Britain, 3–4, 5, 8, 10, 12, 16–18, 21, 46, 61,
Ginkgo florinii, 309n2 67–68, 70, 104–106, 108, 127, 132–137, 153,
Ginkgo huttoni, 109, 110 157, 164, 165, 198, 206–214, 265, 272–273,
Ginkgoites, 120, 121 276, 320nn6–7, 328n15; ginkgos, 4, 6, 16, 21,
Ginkgoites koningensis, 90 61, 64, 66, 108–110, 131–136, 137, 146, 177,
Ginkgoites matatiensis, 90 210–214, 218–219, 224, 224, 234, 238, 240,
Ginkgoites muriselmata, 90 273, 274, 322n7; London Clay fossils, 134–
Ginkgoites taeniata, 107–108, 304n9 136, 138, 307nn4–5; Yorkshire fossils, 108–
Ginkgoites telemachus, 90, 325n23 109, 109, 110, 113
Ginkgoites ticoensis, 305n8 Greece, 143, 154, 156, 236
Ginkgoites tigrensis, 115–116, 304n7 Greenland fossils, 106–108, 110, 304n9
ginkgolic acids, 72, 244, 248 Green River Fossil Basin, 138, 139
Ginkgo Museum, 247 Grew, Jeremiah, 54, 295n2
Ginkgoopsida, 92–93, 94 growth, 35–41, 59, 76, 78, 146, 180; climate and,
Ginkgo orientalis, 307n9 146–151
Ginkgo Petrified Forest State Park, 155 guanacaste (Enterlobium cyclocarpum), 158–159
Ginkgo rajmahalensis, 305n8 Guo Moruo, 18
ginkgotoxin, 245, 326n10, 329n7
Ginkgo yimaensis, 79, 80, 114, 115, 116, 118, 123 Haeinsa Buddhist Temple, 51
Giverny, 7 Hales, Stephen, 61
glacial advances, 157–158, 255 Halle, Thor, 106–107, 108, 114, 304n8
Global Trees Campaign, 261 Hamel, Hendrick, 186
glossopterids, 94, 95, 96, 97, 103, 255, 302nn10– Hamilton, Martin, 64
11 Hamilton, William, 221–222, 324nn14–15
 373
index

Hamilton Woodlands, 221–222, 324nn14–15 history, 175–214, 262; antiquity, 177–183; nam-
Hammerton, John, 272 ing, 204–208; renewal, 195–203; reprieve,
Hamshaw-Thomas, Hugh, 108 184–189; resurgence, 209–214; voyages,
handkerchief tree [dove tree] (Davidia involu- 190–194
crata), 122 Holden, H. S., 106
Hargraves, Matthew, 35 Holland, 42, 43, 197–200, 201, 209, 218–219,
Harris, Tom, 104–105, 105, 106–110, 111, 114, 236, 319n10
115, 125, 303nn2–4, 304nn8–9; The Yorkshire Holocene, 158
Jurassic Flora, 106, 108 Hong Kong, 164
Hartz, Nikolaj, 106 Hooker, Joseph Dalton, 11, 286n10, 303n6
Harvard University, 33; Arnold Arboretum, 76, Hooker, William Jackson, 16–17, 45, 73, 286n10,
146–147, 223, 308n6, 334n3; Herbaria, 46 288n4, 293n9
harvest, 229–230 Hooker Oak, 11, 286n10
Hawai’i, 137, 144, 262 Hope, John, 211
Hawaiian cotton tree (Kokia drynarioides), 262, Hori, Shihomi and Terumitsu, 182, 187, 188,
332n7 205, 206
hazel (Corylus spp.), 127, 137, 231, 325n1 hornbeam (Carpinus spp.), 15, 122, 127
hazelnut, 192 horse, 123–124, 306n8
heartwood, 48 Horseshoe Canyon Formation, 122, 305n6,
heath (Erica spp.), 221 306n6
height, 40–41 Hōryō Ginkgo, 187, 275
Hell Creek Formation, 122, 123, 306n7 Hosack, David, 222, 324n15
hemlock (Tsuga spp.), 277 Hoxnian interglacial, 157
Hemsley, William Botting, 165 Huiji Temple ginkgos, 182, 243, 275, 315n12
Hennig, Willie, 98–101, 302n1 Humboldt, Alexander von, 43, 179, 314n7
Henry, Augustine, 165–166, 311–312n6 Humboldt Redwoods State Park, 9
Henry, Christie, 4 Hyde, Lewis, The Gift, 264, 272
herbal remedies, 7, 242–249. See also medici- hydrangea (Hydrangea spp.), 293n3
nal use hydrogen, 28, 30
hermaphrodites, 55, 295n4
Heungju Temple, 6 Ice Ages, 5, 187, 255, 260
Hickey, Leo, 131 Iceland, 137, 153–154
hickory (Carya spp.), 157, 227 Idaho, 155, 157
Hida-Kokubunji, 21 identity, 16–23
Hideyoshi, Toyotomi, 197 Ikeno, Seiichiro, 68–70, 71, 297n14
Hinduism, 10 illustrations, first Western, 201, 202
Hirase, Sakugoro, 68, 69, 70–72, 94, 223, Imperial University, Tokyo, 70, 232, 298n3,
299nn7–10 299nn7–9
Hiroshima, 22, 178, 289n19, 313n3 India, 116, 117, 164, 199, 235
 374
index

Indonesia, 199 Japanese maple (Acer japonica), 225

insects, 109, 118, 139, 163, 237, 267 Jardin de Roi, Paris, 218, 323n3
in situ conservation, 262 Jefferson, Thomas, 222
insurance, 258–263 Jehol Biota, 117
International Treaty on Plant Genetic Re- Jianshangou Bed, 118
sources for Food and Agriculture, 268 Jien, Eukan-sho, 182
International Union for the Conservation of Jinfo Mountain ginkgos, 171, 172, 173, 177, 178,
Nature (IUCN), 256–257 186, 313n8
Ireland, 136 Jingdezhen ceramics, 193
iron pyrite, 134 Johnson, Kirk, 32, 123, 138
ironwood, 157 Jōmon-sugi, 180
Italy, 18, 143, 219, 236 Jonichiji Temple, 21
Ito, Keisuke, 46, 66, 298n3 Juan Fernández Islands, 262
i-Tree, 235–236, 327n4 Jurassic, 83, 108, 113, 118, 120, 121, 125, 133, 160,
255
Jacquin, Joseph von, 61–62, 63, 68, 296n3 Jyonichiji Ginkgo, 187
Jacquin, Nikolaus Joseph von, 61, 296nn3–4
Jamaica, 206, 207, 213 Kaempfer, Engelbert, 22, 43, 44, 195–208, 209,
Janzen, Dan, 158–159 210, 217, 230–231, 232, 319n9, 320nn4–8,
Japan, 4, 5, 17–18, 20, 42–46, 66–70, 164, 325n23; Amoenitatum Exoticarum, 201, 202,
180–182, 190–197, 212, 223, 273, 299nn7–9, 205–206, 208, 209, 210, 218, 214, 319n9;
318nn4–6; ceramics, 175, 176, 201–203; collection, 207–208; The History of Japan,
edible nuts, 226–232, 232, 233; ginkgos, 19, 199–201, 208, 230
21, 22, 25, 26, 31–34, 36, 51–52, 64, 66–68, kana, 205
75, 76–78, 160, 161, 164, 165, 166, 175, 176, Kannaskoppia, 94–95, 95, 97, 103, 121, 302n12
177, 180, 181, 182, 187–189, 193–194, 201, Kannaskoppianthus, 95, 95
204–208, 215, 216, 226–233, 239, 243–245, Kannaskoppifolia, 94–95, 95, 97, 103
251, 252, 275, 289n18, 291n12, 294n10, 310n6, Karkenia, 102, 115, 116, 120, 121, 302n12, 304–
315nn11–12, 316n8, 318n7, 334n4; herbal 305n7
medicine, 242–245; literature, 187–189; Karkenia asiatica, 304n7
Meiji government, 66–67; temples and Karkenia incurva, 116, 123
shrines, 21, 51–52, 166, 180, 181, 182, 187, 193– Kato, Chikusai, 46, 294n11; portrait of ginkgo,
194, 217, 232, 251, 252, 275, 313n3, 315nn11–12, 47
317n6; trade, 42–44, 190–203, 317–318n6; katsura (Cercidiphyllum japonicum), 137, 157,
West and, 195–203 306n6
Japanese cedar (Cryptomeria japonica), 51, 180, Katsushika Hachiman Shrine, 21
315n8 Kawahara, Keiga, 46
Japanese elm (Zelkova serrata), 219 Kentucky, 64, 137, 139, 222–223
Japanese fig (Ficus erecta), 188 Kerpia, 93, 301n8
 375
index

Kew Arboretum, 273 Latin America, 137, 149

Kew Economic Botany Collection, 17–18, 46 laurel (Laurus spp.), 101, 137
Kew Gardens. See Royal Botanic Gardens, Kew leaves, ginkgo, 1, 2, 5, 18–20, 22, 23, 27–34, 35,
Kew Library, 45, 293n9 46, 59, 62, 74, 120, 141, 142, 145, 175, 176,
Kidston, Robert, 84 188, 194, 201, 202, 203, 211, 217, 225, 244,
Kingdon-Ward, Frank, 165 255, 288nn8–9, 290nn1–9, 305n8; ancestry,
King of Donting Mountain, 229 89–96; autumn, 32–33, 33, 34; blade, 27,
Kinmo Zui, 205, 319n3 30, 37, 38; climate and, 145, 145, 146; drop,
Kirchner, Martin, 83, 90 33, 33, 34, 50; extract, 7, 242, 245–249,
Kirkham, Tony, 272–273 329nn5–11, 330nn12–15; flavonoids, 246; fos-
Kirkland, Larry, 19–20 sil, 83, 83, 85, 90–97, 107–109, 109, 110–115,
Kitakanegasawa Ginkgo, 25, 26, 76–78 115, 116–124, 125–126, 126, 127–131, 132–136,
Koch, Lauge, 107 138, 155, 305n3, 309n4; growth, 35–36, 36,
Koishikawa Botanical Garden, 46, 66–68, 293– 37–38, 39, 40–41; long-shoot, 35–38, 39, 59,
294n10, 298n4 292n6; medicinal uses, 242–249; motifs and
Koishikawa Ginkgo, 67, 68, 298n2 logos, 19, 19, 20; origin of, 83–87; photo-
koompassia (Koompassia spp.), 286n6 synthesis, 27–34; pressing, 318n7; seedling,
Koonwarra fossils, 115, 117, 305n8 59, 76; short-shoot, 35–36, 36, 37–38, 39,
Korea, 6, 18–19, 20, 273; edible nuts, 226, 228; 292n6; stalk, 27, 30, 37; stomata, 28, 28, 85,
ginkgos, 6, 6, 11, 11, 12, 13, 18–19, 19, 21, 32, 290n3; veins, 27, 28, 30, 290n1; water con-
41, 49, 51, 163, 164, 177, 186–187, 188, 226, tent, 28–31
238, 239, 243, 275, 288n10, 316n6, 328n13; Lebanon, 10
herbal medicine, 242, 243; shrines and legacy, 272–277
temples, 6, 19, 21, 51, 187, 275; trade, 190– Leiden ginkgo, 219, 323n8
196 L’Enfant, Pierre, 237
Kyoto, 21, 193, 194, 275 Leopold, Aldo, 258
Kyoto Protocol, 264 Leslie, Andrew, 56
Kyushu, 36, 193, 195 licorice (Glycyrrhiza glabra), 244
lignotubers, 76
lacquerware, 18, 51 Li Jiawan Grand Ginkgo King, 178–179, 182,
lakes, 139 187, 313nn3–5
Lang, William H., 84Lan Mao, Dian Nan Ben lily bulbs (Lilium spp.), 231, 232
Cao, 245 linden [lime] (Tilia spp.), 276, 334n6
Lao Tzu, 21 Lingering Garden, 288n8
Late Cretaceous, 120, 121, 154, 308n12 Lingula, 129–130
Late Devonian, 88, 254 Linnaean Society of London, 211, 213, 214,
Late Miocene, 154, 156 322n8
Late Paleocene, 126, 127 Linnaeus, Carl, 22, 38, 81–82, 161, 201, 206,
Late Triassic, 89, 95, 120 209–214, 218, 321nn2–3, 322nn5–8; ginkgo
 376
index

Linnaeus, Carl (continued) maize (Zea mays), 54, 61

and, 22, 38, 81, 209–212, 212, 213–214; Phi- Major Oak, 10
losophia Botanica, 204; Species Plantarum, male ginkgo, 53–57, 59, 60–65, 66, 78, 185–186,
210 215, 216, 218, 220, 221, 237–238, 297n11,
literature, ginkgo, 182–189, 208; early, 184–189 297n14; “leaky,” 65
live oak (Quercus virginiana), 237, 286n10 mammals, 117, 122, 137, 139, 158, 159, 163, 256–
Living Planet Index, 257 257
living tree, 25–78; energy, 27–34; gender, Manchester, Steve, 128, 131, 138, 306n2
60–65; growth, 35–41; resilience, 73–78; Man’yōshū, 187–188
seeding, 66–72; sexual recognition, 53–59; Mao Zedong, 18, 111
stature, 42–52 maple (Acer spp.), 157, 313n3
Loddiges, Conrad, 219 Mark Twain Tree, 314n6
Lombardy poplar (Populus nigra), 221 Martin, Paul 158–159
London, 3, 43, 46, 72, 82, 98, 132–136, 164, 206– Marx, Groucho, 152
214, 219, 234, 238, 276 Massachusetts, 146–148
London Clay fossils, 134–136, 138, 307nn4–5 Mathew, Brian, 73–75, 76
London plane (Platanus x acerifolia), 240 Matsumura, Jinzo, 67
Long, John Luther, 42 McIver, Elizabeth, 139
Longquan ceramics, 192 McPherson, Greg, 235, 236
long shoots, 35–36, 37–38, 39, 59 medicinal use, 7, 12, 17, 66, 189, 192, 242–249,
Loti, Pierre, Madame Chrysanthemum, 42 326n10, 328nn3–4, 329nn5–11, 330nn12–15
lotus (Nelumbo nucifera), 101, 230, 231 Mediterranean fig (Ficus carica), 10
Loudon, John, 214, 218, 219 Meiji-Jingu Park, 33–34, 291n12
Louis XIV, King of France, 236 Mei Yao-Chēn, 184–186, 226
Low Countries, 5, 218 Mendel, Gregor, 62, 63
lumber, 12, 14, 15 Mendes, Chico, 265, 333n3
lychee (Litchi chinensis), 192 Menzies, Nicholas, 182–183, 184, 185
merchandise, ginkgo, 20
Mackie, William, 84 meristems, 59
Madagascar, 265 Mertz, LuEsther, 16
Madama Butterfly (opera), 42 Mesozoic, 91, 111, 114, 116, 120, 160
Madison, James, 222 Messel fossils, 137, 307n9
magnolia (Magnolia spp.), 8, 15, 55, 101, 122, Mexico, 90, 149
136, 154, 211 Meyen, Sergei, 91–93, 94
mahogany (Swietenia spp.), 15 Michaux, André, 222, 324n16
ma huang [ephedra, Mormon tea] (Ephedra Middle Eocene, 133, 308n13
spp.), 244, 302n6, 328n4 Middle Jurassic, 80, 114, 120
maidenhair fern (Adiantum spp.), 22, 165 Middle Miocene, 154
Maidenhair tree, 165 Middle Ordovician, 85
 377
index

Middle Triassic, 91 Munmyo Confucian Shrine, 19, 21

Millennium Development Goals, 266 Murdoch, Iris, A Fairly Honourable Defeat,
mimosa (Mimosa spp.), 222 119
Minneapolis, 235
Miocene, 153, 154, 155, 309n5, 310n6 Nagasaki, 42–44, 46, 195–197, 197, 198–203,
Missouri, 22, 289n19 318n4
mistletoe (Viscum album), 330n12 naming, 22, 38, 81, 185, 188, 204–208, 211–212
Mitchell, Alan, 59 Nanjing Institute of Geology and Palaeontol-
Mitchell, Joni, “Big Yellow Taxi,” 258 ogy, 111, 112
Miyoshi, Manabu, 67 Nara Prefecture, 166
Mizufuki Ginkgo, 275 Nathorst, Alfred, 82, 106, 108, 308n14
Mokujiki Shonin, 51–52; ginkgo wood statues National Academy of Sciences, 19
by, 251, 252, 294n18 National Geography Society, 259
Molteno fossils, 89–90, 94, 303n7 Natural History Museum, London, 98, 133, 134,
monkey puzzle (Araucaria araucana), 8 137, 207, 210, 238
Montagu, second Duke of, 276, 334n6 natural populations, 162–167
Montana, 122, 309n4 natural selection, 40, 60–61, 86, 225, 295n4
Monterey pine (Pinus radiata), 332n2 Naugolnykh, Serge, 93
Montpellier, 220, 221 Needham, Joseph, Science and Civilization in
moonseed family (Menispermaceae), 136 China, 182, 184, 315n13
Morgan, Thomas Hunt, 62, 63, 297n8 Nemerov, Howard, 34
Mormon tea [ephedra, ma huang] (Ephedra Neogene, 120, 121
spp.), 244, 302n6, 328n4 Nevada, 309n5, 314n6
Morocco, 9 New Haven, 236, 327n6
morphology, 23 New Jersey, 14, 21
Morse, Edward, 232–233 New South Wales National Parks and Wildlife
Morton, Joy, 22 Service, 259, 269–270
Morton Arboretum, 22, 289n20 New York, 3, 6, 18, 98, 223, 230, 234, 235–239,
moss, 85, 87, 98, 99, 122, 275, 303n3 270, 327n4, 328nn11–12
motifs and logos, ginkgo, 19, 19, 20, 175, 194, New York Botanical Garden, 16, 223
202–203 New York Horticultural Society, 222
Moule, A. C., 205 Nietzsche, Friedrich, Nietzsche contra Wagner,
mountain cedar (Widdringtonia nodiflora), 143
332n6 Nigatake Ginkgo, 187, 244
Muir, John, 9 Nishi Honganji Temple, 275
Mulanji cypress (Widdringtonia whytei ), 261– nitrogen, 28
262, 332n6 Noble, David, 259
mulberry (Morus spp.), 54, 208, 244 Nongso, Seonsan, 18–19, 288n10
Mull, 135, 136, 153, 154, 309n2 Norse, 10
 378
index

North America, 5, 6, 36, 70, 90, 137, 138, 144, proliferation, 111–118; prosperity, 132–140;
152, 154, 156, 157, 160, 161, 237, 255, 311n9. recognition, 104–110; relationships, 97–103;
See also specific countries winnowing, 119–124. See also evolution; fos-
Northcott, Kenneth, 4 sils; specific periods
North Dakota, 122, 123, 125–131 origins, 81–88
Northern Hemisphere, 152, 153, 156 Osaka University, 19
North Pole, 5, 139, 140 Otomo-no-Yakamochi, 188
North Sea, 109 Ouyang Hsiu, 184–185, 226–227
Northwest Forest Plan, 287n18 ovules, 56–57, 59, 63, 65, 68, 78, 114, 296n10,
Norway maple (Acer platanoides), 221 299n12, 302n12, 305n8
Nung Sang Chi Yao, 185–186 oxygen, 28, 29, 30, 85, 240
nuts, ginkgo, 17, 122, 149, 184, 185–186, 193–
194, 208, 220, 225, 226–233, 242, 325nn1–5, Pacific Northwest, 14, 138, 154–155, 277, 307n12
326nn5–12, 328n4, 329nn6–11; edible, 226– pagoda tree (Styphnolobium japonicum), 219
231, 231, 232, 232, 233; harvest, 229–230; Pakenham, Thomas, 9, 286n4
toxicity, 245 Palaeocarpinus, 306n2
Nuttall, Thomas, 161 Paleocene, 123, 154, 306n7
nypa (Nypa fruticans), 135, 135, 136 Paleogene, 120, 121
Paleozoic, 92
oak (Quercus spp.), 10–11, 12, 14, 15, 49, 137, palmetto [cabage palmetto] (Sabal palmetto),
139, 154, 155, 157, 227, 237, 276 10, 286n9
Oak Grove, 14 palm family (Arecaceae), 101, 136, 137, 138
oil, 12, 230–231, 326n10 Panama hat palm (Carludovica palmata), 137
oil palm [African] (Elaeis guineensis), 14 Pandit, Saskya, “Elegant Sayings,” 184
Ojiya Temple, 51–52, 251, 252, 294n17 Paris, 43, 91, 143, 218, 220, 236, 323n9
Okawachi Kiln, 175, 176, 203 Pau Brasil (Caesalpinia echinata), 10
Oldfield, Sara, 261 pea (Pisum sativum), 62
Old Lion, Kew, 4, 6, 16, 21, 61, 64, 66, 177, 214, peach (Prunus persica), 192
219, 220, 273, 274, 285n2, 297n13 peanut (Arachis hypogaea), 227, 231
Oligocene, 153, 154, 310n6 peepal [pipal] (Ficus religiosa), 10
Oliver, Daniel, 165 Permian, 91–92, 94, 97, 120, 121, 123, 254, 255
Oliver, F. W., 70–71, 299n11 Perry, Commodore, 42, 45
OneTree Project, 12 Persia, 199, 201
Ono, Yoko, 21, 289n18 persistence, 125–131
orchards, 228–229 pests, 151, 237, 261
Ordovician, 87, 254 Pétigny ginkgo, 220, 323n9
Oregon, 138–139, 154, 155 Petriellaea, 302n12
origin and prehistory, 4–5, 7, 79–140; ancestry, Philadelphia, 221, 235, 236–237
89–96; origins, 81–88; persistence, 125–131; phloem cells, 48
 379
index

phosphorous, 28, 32 Proust, Marcel, À la recherche du temps perdu,

photosynthesis, 27–34, 40, 85, 146, 290nn2,4–5, 195, 318n1
314n7 Psygmophyllum, 301n8
pine (Pinus spp.), 4, 18, 55, 140, 228, 231 pumpkin (Cucurbita pepo), 326n11
pinyon pine (Pinus subsection Cembroides
and Nelsonianae), 326n5 Qing Yuan Lu, 193
Pisa ginkgo, 219, 220 quaking aspen (Populus tremuloides), 300n2
pistachio (Pistachio vera), 208, 214, 227, 325n4 Quin, John, 17–18, 51, 288n6
plane [sycamore] (Platanus spp.), 101, 155, 219,
306n6, 327n7 radiocarbon dating, 179, 314nn6–7
plate, ginkgo-decorated, 175, 176, 203 rainfall, 144, 151, 155, 235, 327n4,
Pleistocene, 171, 310nn6–7 rain forest, 9, 14, 265, 266, 277, 286n6, 333n4
Pliocene, 153, 154, 156, 160, 309n2, 310nn1,6–7 RAPD technique, 170, 312nn3–4
poetry, 18, 22–23, 34, 184–188, 223, 291n14, Raup, Dave, 253–256, 259, 330nn1–3
292n5, 296n5, 307n12, 315–316n1, 316n2, Ray, John, 54, 295n2, 320n7
325nn20–21 red maple (Acer rubrum), 32, 240
poisoning, 245, 329n7 Reid, Eleanor, 134, 135, 307nn3–5
poison ivy (Toxicodendron radicans), 72, 231, Rember, Bill, 155
244 Remy, Winfried, 84
poison oak (Toxicodendron diversilobum), 72, Republic fossils, 139, 308n13
244 restaurants, 20, 22, 230, 233
Pollan, Michael, Second Nature, 217, 323n1 Rhynie Chert, 84–88, 90, 93, 300n5
pollen, 53–59, 65, 68, 70, 71, 72, 78, 88, 94, rice (Oryza sativa), 52, 185, 213
95, 96, 130, 131, 146, 157–158, 295nn4–9, Richmond Gardens, 17
306n6; catkin, 65, 90, 127, 131, 305n8, 306n6; rings, 50, 51, 178, 179, 180, 183, 294n15, 314n6,
cones, 36, 55–56, 61, 64, 65, 93, 130, 146, 315n9, 315n11
220, 295n5, 295n7, 304n11; tube, 70, 296n9, Robin Hood legend, 10, 11
299n8 Robinson Crusoe cabbage tree (Dendroseris
pollination, 53–59, 61, 65, 68, 70, 71, 72, 78, litoralis), 262
146, 147, 148, 292n3, 295nn4–9; drop, 57, 68, Rock, Joseph, 165
295n9; self-, 55, 60, 65 rose (Rosa spp.), 139
pollution, 6, 15, 235, 264, 327n4 Rouen, 219–220
pomegranate (Punica granatum), 226 Royal Botanic Gardens, Kew, 3–4, 6, 8, 9, 11,
poppy, 101 12, 16–18, 21, 45, 46, 51, 61, 64, 70, 73, 146,
Portuguese laurel (Prunus lusitanica), 324n14 151, 165, 177, 214, 219, 220, 221, 242, 259,
Portuguese trade, 10, 196–198, 318n4, 319n6 262, 269, 272–273, 274, 276, 285n2, 286n10,
potato (Solanum tuberosum), 19, 169, 226, 232, 288nn3–6,8, 292n9, 293n9, 294n15, 297n13,
328n10 302n5, 303n6, 307n4, 310n3, 312n8, 323n6,
prehistory. See origin and prehistory 325n22, 331n8, 332nn6–7
 380
index

Royal Society, 207, 208, 210, 221, 295n2, 298n4, climate and, 146–148; dispersal, 158–160,
315n13, 320n7, 322n11 255, 305n5; fertilization, 66–72, 146, 147,
Royer, Dana, 131, 154, 290n3, 309n3 295n9, 299nn10–13; germination, 54, 59, 78,
Ruan Ye, 186 147–148, 160, 295n9, 296n11, 308n8; harvest,
rubber (Hevea brasiliensis), 14–15, 157, 333n3 229–230; medicinal use, 242–249; pollina-
Russia, 92, 93, 144, 154, 199, 304n7, 310n6 tion, 53–59, 146, 147, 148, 295nn4–9; stalks,
129
Sago cycad (Cycas revoluta), 68 self-propagation, 74–75, 76
Saint Helena ebony (Trochetiopsis ebenus), 262 Senbon Ginkgo, 316n8
Sakuzaemon, Takahashi, 44, 45 Seoul, 6, 11, 19, 21, 49, 163, 164, 238, 239, 299n13,
Salisburia adiantifolia, 39, 165, 208, 210 309n14, 317n4, 328n13
sandalwood (Santalum spp.), 192 Sepkoski, Jack, 253–254, 330n2, 331n3
San Francisco, 7, 9, 123, 140, 230, 235, 240, sessile oak (Quercus petraea), 40
286n3 Seward, A. C., 106–107, 296n11, 303nn5–6,
Santi, Giorgio, 219 385n3,
sap, 40, 48, 292n8, 313n3 sexual reproduction, 41, 53–59, 61, 68, 70, 75,
Saporta, Gaston de, 91, 301n4, 301n5 87–88, 147, 295nn4–9, 308n6
sassafras (Sassafras spp.), 306n6 shade, 131
Scalby Ness, 109, 109, 110, 304n11, Shakespeare, William, 314; As You Like It, 226,
Scheuchzer, John Caspar, 208, 321n10 325n1; Henry IV, Part 2, 209, 321n1; Macbeth,
Schopf, Tom, 129–130, 306n5 66, 298n1
Schweitzer, Hans Joachim, 82–83, 90, 300n3 Shanghai, 77, 111, 165, 167, 170
Scotland, 84, 133, 136, 137, 153, 154, 164, 211, Shaolin Zheng, 118
300n6 Shen Nung, 242
Scott, D. H., 70–71, 299n11 Sheppey fossils, 134–136
Scott, Robert Falcon, 94 Sherwood Forest, 10
seasonality, 143–151 Shinan Ship 190–192, 192, 193, 196, 205,
seed ferns, 71, 299n11 317nn2–6
seedlings, ginkgo, 59, 76, 78, 83, 107, 146–147, Shintoism, 5, 21, 217, 288n9,289n17
177, 222, 228, 255; wild populations, 162–163, short shoots, 35–36, 36, 37–38, 39, 46, 55, 56,
163, 164–167 94, 114, 290n1, 292n6, 295n7, 300n3
seed plants, 101, 102, 273; ancestry of, 89–96; Siebold, Philipp Franz von, 38, 39, 42–44,
origins of, 81–88; relationships among, 44, 45–46, 66, 197, 198, 200, 203, 207, 232,
97–103 292n1, 293nn2–5, 293n8, 294n11, 298n9,
seeds, ginkgo, 5, 17, 18, 19, 46, 54–59, 61, 64–65, 319n7; Flora Japonica, 38, 39, 45–46, 293n3,
66–72, 78, 84, 90, 91, 113–118, 127, 134, 135, 293n8, 310n8
163–164, 186, 191, 192, 213–214, 223, 227– silica, 84, 85, 122, 128, 138, 155, 286n6
233, 255, 290n4, 299nn10–13, 304nn3–7, silk tree (Albizia spp.), 222
326nn6–11; Almont fossils, 127, 127, 128, 129; Silurian, 85, 87, 129, 130, 254
 381
index

silver apricot, 46, 57, 185, 188, 204, 205, 214, stonecrops (Chara spp.), 301n8
289n16 stone pine (Pinus pinea), 8–9
silver fir (Abies alba), 324n14 Stopes, Marie, 67–68, 70, 71, 298nn4–6,
Sloane, Sir Hans, 206–208, 320nn6–8, 299n10, 303n4
321nn2,8–10,12; collection, 207–208, 320– Strasburger, Eduard, 68, 299n7
321n8 strawberry (Fragaria x ananassa), 320n4
smell, ginkgo, 3, 72, 159, 224, 229, 237–238, strawberry tree (Arbutus unedo), 8
244–245, 288n13, 289n15 streets, 234–241, 327nn1–8, 328nn11–16
Smiley, Jack, 155 Strobel, Scott, 51
Smith, J. E., 207–208, 214 sucker shoots, 77, 77, 78
Smith, Selena, 137, 307n9 sugar maple (Acer saccharum), 8, 292n8
snowdrop (Galanthus spp.), 73 summer, 32, 36, 50, 54, 72, 144, 146, 148, 150,
soil, 38, 74, 76, 143, 148, 151; urban, 240–241 151
Solander, Daniel, 207 Sung Kyun Kwan University, 19, 19, 21
South Africa, 89–90, 91, 94, 95, 262, 302n10, sunlight, 28, 29, 30, 31, 32, 40, 86, 141, 142,
314n7 148–149
South America, 116, 117, 152, 158, 265, 302n10, Sutton, Walter, 62, 297n8
312n8 swamp cypress (Taxodium distichum), 154, 240
South Carolina, 10, 14, 144, 213, 222, 286n9, swamp gum (Eucalyptus regnans), 9, 122
287n17 Sweden, 81, 82, 91, 106–107, 144, 190, 199,
southeast Asia, 136, 192, 195, 201, 265, 326n9, 308n14, 314n6,
326n11 Swedish Museum of Natural History, 4, 82–83,
Southern Hemisphere, 91, 94, 116, 149, 152, 107, 300n1,300n3, 304n8, 322n8,
312n8 sweet gum (Liquidambar styraciflua), 155, 240,
Spain, 227, 324n16 306n6, 311n9
sperm, 53, 54, 68–70, 87, 302n11; swimming, 68, sweetsop family, 136, 138
69, 70, 299nn8–9 Swift, Jonathan, Gulliver’s Travels, 208, 321n12
Sphenobaiera, 120, 121, 301n6, 305n8, 325n23 sycamore (Platanus spp.), 101, 155, 219, 306n6
Sphenobaiera umaltenis, 304n7 Sze, H. C., 114
spores, 85–88, 99
spring, 36–37, 40, 50, 53, 147, 151 tamarisk (Tamarix spp.), 221
spruce (Picea spp.), 51, 314n6 Tamba Kokubunji Temple, 21
Stalagma samara, 303n2 Taoism, 217, 243
Stanley, Steve, 253, 306n5, 330n1 Tasmania, 9, 152, 309n1
stature, 42–52 teak (Tectona spp.), 15
steppe, 152–153 temperature, 143–151, 308n9, 309nn10–14
Stevens, Nettie, 63, 297n8 Temple of the Reclining Buddha, 21, 289n16
Stockholm, 81, 82, 106, 144, 199 Tennessee, 137, 139
stomata, 28, 28, 85, 290n3 Texas, 10, 22
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textiles, 194 Truman, Harry, 22, 289n19

Thoreau, Henry David, “Wild Apples,” 162, trunk, ginkgo, 40–41, 48–50, 74–75, 77, 77, 78,
311n1 166, 179, 180, 229
Thouin, André, 218, 219–20, 323n9 Tsan-Ning, Ko Wu Tshu Than, 184, 295n3
Thunberg, Carl Peter, 43, 44, 197, 198, 201, Tso Ssu, 184, 315–316n1
203, 207, 232, 293n8, 319n10, 329n6; Flora Tsurugaoka Big Ginkgo, 180, 181, 182, 187, 275,
Japonica, 201, 310n8 315n11, 317n9, 334n4
Tianmu Mountain ginkgos, 166–167, 169, 169, Tsurugaoka Hachiman-gū Shrine, 180, 181, 232,
170–172, 172, 173, 178, 186, 312n9, 313n8 233, 275, 315n11, 334n4
Tico fossils, 115–116, 305n8 Turner, Rudi, 125–126
Tiffney, Bruce, 159–160, 305n5 twigs, ginkgo, 20, 31, 35–36, 36, 37, 46, 48–50,
timber, 12, 14, 15, 51, 191, 261, 272, 286n6, 114, 201, 291n2
294n12, 332n2
time, 1–7 Ubagami Shrine, 21, 316n8
Titsiugh, Isaac, 201 Uba Ginkgo, 187, 316n8
Tōfukuji Temple, 193 umbrella pine (Sciadopitys verticillata), 157
Tokugawa Ieyasu, 197–198 United Nations, 264
Tokyo, 3, 19, 21, 22, 31, 33–34, 43, 66–68, 180, United Nations Environment Programme,
194, 200, 233, 239, 291n12, 298n1, 316n8 Plant for the Planet: The Billion Tree Cam-
Tokyo University (Tokyo Imperial University, paign, 234, 327n2
University of Tokyo), 46, 47, 66–68, 70, 71, United Nations Framework Convention on
232, 293n10, 298nn3–4, 299nn7–8 Climate Change, 264, 287n18
tomato, 169 United States, 10–12, 14, 19, 85, 90, 91, 122–
topophytic effect, 228 131, 137–139, 143, 144, 265, 276, 277, 291n3,
Toromiro tree (Sophora toromiro), 262 324nn16–18; Almont fossils, 125–126, 126, 127,
toxicity, 230, 245, 329n7 127, 128–131, 163, 306nn2–4; Clarno fossils,
tracheids, 290n8, 294n14 138–139; climate, 139, 146–149, 152; ginkgos,
trade, 42, 43, 164, 186, 189, 190–194, 196–198, 19, 20, 21–22, 33, 34, 64, 123, 125–126, 126,
318nn4–6; Deshima, 43–45, 197–203, 319n6, 127, 127, 128–131, 137–139, 146–149, 154–155,
319n10; Dutch, 42–43, 197–201, 318n6, 221–223, 275, 308n13, 309nn4–6; herbal
319n10; Shinan ship, 190–192, 192, 193, 196, medicine, 247, 248; Republic fossils, 139;
317nn2–6 street trees, 234–241. See also specific states
Treaty Oak, 11 United States Botanic Garden, 223
tree of heaven (Ailanthus altissima), 221 University of California, Berkeley, 14, 223,
trees, 8–15 287n17, 291n2, 325n20
Trees for Cities, 234, 327n2 University of Chicago, 4, 22, 56, 129, 224–225,
Triassic, 91, 92, 93, 95, 97, 121, 124, 125, 254, 255, 253, 276, 295n7, 325n21,
301n3 University of Reading, 8, 104–105, 108, 111,
Trichopitys heteromorpha, 91–94, 95, 97, 103, 304n1, 306n1,
120, 121, 123, 301nn4–6 University of Tokyo. See Tokyo University
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(Tokyo Imperial University, University of Weimar ginkgo, 150

Tokyo) welwitschia (Welwitschia mirabilis), 285n3,
University of Uppsala, 81, 199, 211 302n6
University of Vienna, 61, 62, 68, 296n3 Western science, ginkgo introduced to, 195–
University of Virginia, 64 214, 217
Unseld, Siegfried, Goethe und der Ginkgo, 4, Wettstein, Richard von, 68
289n21, 291n11, 292n5 whales, 19
Upper Triassic, 120 wheel wingnut (Cyclocarya and Pterocarya
use, 215–249; gardens, 217–225; nuts, 226–233; spp.), 126, 306n2
pharmacy, 242–249; streets, 234–241 Whitfield ginkgo, 214
Utah, 138, 300n2 Wieland, Martin, Muscarion oder die Philoso-
Utrecht ginkgo, 218–219, 322n9 phie der Grazien, 104, 303n1
Wilberforce, Samuel, 89, 301n1
variants, 225 wild population, 156, 162–167; genetic diversity
variation, 60–61, 225 and, 168–173
vascular plants, relationship of, 98–99 Wilf, Peter, 123, 306n7
Vaughan, Bill, 14, 287n16 Willershausen fossils, 156–157, 309n2, 310n2
Vavilov, Nikolai, 169, 312n1 willow (Salix spp.), 18, 75, 225
Venezuela, 40 willowmore cedar (Widdringtonia schwarzii ),
vernalization, 149 262
Vienna, 61, 68, 219, 296n3, 299n7 Wilson, Edmund Beecher, 63, 297n8
Virginia, 11, 64 Wilson, Ernest Henry, 165–166, 166, 273,
voyages, 190–194 312nn6–8, 315n8, 323nn10, 323nn12, 334n3
Wing, Scott, 131
walnut (Juglans spp.), 15, 139, 157, 192, 231 winnowing, 119–124
Wang Shi Yuan, Master of the Nets Garden, 1, winter, 32, 34, 36, 37, 38, 41, 48, 50, 51, 59, 72,
2, 18 74, 144–146, 147, 148, 149, 151, 255, 275,
Washington, D.C., 19, 22, 82, 138, 223, 237, 291nn1–2, 306n7, 309n10, 309n14
288n12, 324n18, 327–328n9 Wisconsin, 34
Washington, George, 222, 237, 324n16 witches’ brooms, 64, 297n14
Washington Oak, 14 witch hazel (Hamamelis spp.), 139
Washington State, 139, 154, 155, 308n13, 310n6 Wollemi pine (Wollemia nobilis), 15, 259–261,
water, 14, 28–31, 70, 85, 86, 87, 88, 98–99, 131, 260, 262, 269, 269, 270, 273, 332n3
134, 143–145, 145, 148, 151, 186, 235, 240, 255, wood, ginkgo, 17–18, 37, 40, 41, 46, 47, 48–50,
292nn7–9, 294n14; rainfall, 151, 235; trans- 50, 51–52, 90–91, 130, 138, 155, 251, 252,
port, 38–41, 48, 50, 50, 61, 235 275, 292n10, 294nn12–18; rays, 48, 292n8,
watercress (Nasturtium officinale), 233 294n12; statues, 251, 252, 294n18
water lily, 101 World Heritage Sites, 18, 307
watermelon (Citrullus lanatus), 326n11 World War II, 66, 98, 106, 239
Wehr, Wes, 139, 307n12 World Wide Fund for Nature, 257, 331n9
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Wright, Frank Lloyd, 20, 275, 288n13 yin shan [Cathay silver fir] (Cathaya argyro-
Wyoming, 125, 138 phylla), 157, 173
Yinwoshan Mountain, 118
xylem, 48, 302n2 Yixian Formation, 117–118, 305n9
Yongmunsa Ginkgo, 11, 11, 12, 13, 41, 131, 187,
Yakushima, 180, 314–315n8, 275, 287n12, 316nn7–8
Yale School of Forestry and Environmental Yongmunsa Temple, 11, 11, 12, 13, 275, 287n11
Studies, 236, 258, 331–332n1 Yorkshire fossils, 108–109, 109, 110, 113, 303n3
Yale University, 27, 31, 33, 33, 35, 51, 131, 236,
332n11, 333n9 Zenpukuji Temple, 21, 316n8
yam (Dioscorea spp.), 231 Zhang Bole, 113, 114, 115, 304nn2–4
Yasukuni Shrine, 21, 289n17 Zhaoming, 182, 315n12
Yatabe, Ryokichi, 67, 298n3, 299n8 Zhejiang University, 19, 170, 171
Yeongwol ginkgo, 187, 316n7 zhōng rǔ, 26, 76, 243
yevaro (Eperua purpurea), 40, 292n9 Zhou Zhiyan, 80, 104–105, 108, 110, 111–112,
yew (Taxus spp.), 4, 272 112, 113–118, 119–124, 125, 160, 301n6, 303n2,
Yggdrasil, 10 304nn3–5
Yima fossils, 79, 80, 113–115, 116, 119, 120, 304n2 Zhuanchengzi Bed, 118
Yimaia, 102, 114, 120, 121, 304n5 zoos, 259, 262, 332n10
Yimaia capituliformis, 304n5 Zuccarini, Joseph Gerhard, 38, 293n8; Flora
Yimaia qinghaiensis, 304n5 Japonica, 38, 39, 45–46
Yimaia recurva, 79, 80, 114, 116, 123, 304nn4–5