North American Crop Wild Relatives

Volume 1 Conservation Strategies

Stephanie L. Greene
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Stephanie L. Greene
Karen A. Williams · Colin K. Khoury
Michael B. Kantar · Laura F. Marek
Editors

North American
Crop Wild Relatives,
Volume 1
Conservation Strategies
North American Crop Wild Relatives, Volume 1
Stephanie L. Greene
Karen A. Williams • Colin K. Khoury
Michael B. Kantar • Laura F. Marek
Editors

North American Crop Wild

Relatives, Volume 1
Conservation Strategies
Editors
Stephanie L. Greene Karen A. Williams
USDA, Agricultural Research Service USDA, Agricultural Research Service
Center for Agricultural Resources Beltsville Agricultural Research
Research, National Laboratory Center, National Germplasm
for Genetic Resources Preservation Resources Laboratory
Fort Collins, CO, USA Beltsville, MD, USA

Colin K. Khoury Michael B. Kantar

USDA, Agricultural Research Service Tropical Plant and Soil Science
Center for Agricultural Resources University of Hawaii at Manoa
Research, National Laboratory Honolulu, HI, USA
for Genetic Resources Preservation
Fort Collins, CO, USA
International Center for Tropical
Agriculture (CIAT)
Cali, Colombia

Laura F. Marek
Department of Agronomy/North Central
Regional Plant Introduction Station
Iowa State University
Ames, IA, USA

ISBN 978-3-319-95100-3    ISBN 978-3-319-95101-0 (eBook)

https://doi.org/10.1007/978-3-319-95101-0

Library of Congress Control Number: 2018953015

© This is a U.S. government work and not under copyright protection in the U.S.; foreign copyright
protection may apply 2018
Open Access Chapters 2 and 12 are licensed under the terms of the Creative Commons Attribution
4.0 International License (http://creativecommons.org/licenses/by/4.0/). For further details see license
information in the chapters.
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This book is dedicated to the custodians
of the diversity of wild plants in North
America – past, present, and future
Foreword

Reinvesting in Crop Wild Relatives in North America

Nearly any place you hike a canyon, canoe a river, climb a mountain, wade a wet
meadow, or weed a field in North America, you will come upon native plants that are
close relatives of food, fiber, animal forage and feed, industrial oil, ornamental, and
medicinal crops. On the fringes of muskegs from Hartley Bay in British Columbia
to sites in northern California, you might come upon the Pacific crab apple (Malus
fusca [Raf.] C. K. Schneid) around ancient fishing and gathering camps (Routson
et al. 2012). In the watersheds of the St. Johns and Kissimmee Rivers in Florida,
the vines of the rare Okeechobee gourd (Cucurbita okeechobeensis [Small] L. H.
Bailey subsp. okeechobeensis) trail up into the branches of wild custard apples
(Annona glabra L.) (Andres and Nabhan 1988; Nabhan 1989). Where I live and
farm in Southern Arizona just north of the Mexican border, wild chiles (Capsicum
annuum var. glabriusculum [Dunal] Heiser & Pickersgill) grow in desert canyons
not far from wild grapes (Vitis L.), walnuts (Juglans L.), passion fruits (Passiflora
L.), cassava (Manihot Mill), tomatillos (Physalis L.), and tepary beans (Phaseolus
acutifolius A. Gray var. acutifolius) (Nabhan 1990; Nabhan 1991).
To many naturalists, these plants are but botanical curiosities, worthy of con-
servation without regard to historic or current human uses. But for crop geneticists
and plant breeders, and a growing number of biodiversity conservationists, these
species are especially worthy of protection, conservation, and evaluation because
they may hold something of lasting value for the future of the way we live and eat
on this planet.
These scientific and cultural values are exactly the reason why agricultural
research and conservation management insights are not only so important but so
timely. The chapters of this book represent the first comprehensive effort to assess
wild crop genetic resources on our continent. Remarkably, this book arrives in our
hands at just the moment in North American history when many of these plants are
threatened by climate change yet also when these resources are most needed if
future generations are to adapt.

vii
viii Foreword

The wildness in these plants confers distinct advantages not found in their
domesticated cousins: tolerance to extremes of temperature and precipitation and
resistance to the pests that lay waste to agricultural fields. This is why we are
increasingly looking to the wild to strengthen our agriculture, particularly during
this period of accelerated climate change. Farmers, orchardists, ranchers, and horti-
culturalists are already suffering from shifting and often heightened frequencies of
drought, heat waves, catastrophic freezes, hurricanes, floods, and fires. And with the
changes in these abiotic stressors come other biotic impacts to our farming systems
and food security: previously unforeseen weeds, insect pests, and crop diseases that
take a long-term toll on agricultural productivity and food safety.
During eras of political, economic, and environmental stress, humanity turns
to consider a broader range of options than typically employed during “business
as usual.” This is one of those times – when agriculture is looking to draw upon
a broader and deeper gene pool of crop genetic resources as a means to re-diver-
sify and add resilience to the food plants that we depend on for survival. After
decades of focusing on a relatively small genetic base of cultivated varieties for
crop improvement, geneticists are now casting a much wider net, fortunately
enabled by a broader portfolio of diagnostic techniques, micro-propagation prac-
tices, and biotechnologies used to select and transfer genes from wild relatives
into food crops.
This is why the fact that forty-some crop wild relative species are included in the
U.S. Fish and Wildlife Service’s Endangered and Threatened Plant list is of great
concern. To make matters worse, this list is likely to grow much larger (Rogers
2015). Analyses of North American seed plants facing extinction risks exacerbated
by climate change and land use intensification (Zhang et al. 2016) would indicate
that roughly 27% of the 4600 crop wild relatives documented to occur in the U.S.
(Khoury et al. 2013) are likely to lose more than 80% of their habitat by the 2080s
and will suffer a 50% retraction of their ranges.
As I read through the names on the current U.S. list of threatened and endangered
crop wild relatives, I am struck by both their beauty and by the fragility of the plant
species they represent:
• Texas wildrice (Zizania texana Hitchc.), an aquatic perennial with high allelic
richness surviving along just a few stretches of the San Marcos River drainage of
the Edwards Plateau in Texas
• The scrub plum (Prunus geniculata R. M. Harper) of Lake Wales Ridge in
Florida, a small shrub with perfumed flowers valued by horticulturists as a showy
and fragrant ornamental, closely related to the Chickasaw plum, with a fruit of
probable hybrid origin that has been both culturally dispersed and cultivated for
well over 150 years
• The Bakersfield prickly pear cactus [Opuntia basilaris Engelm. & J. M. Bigelow
var. treleasei (J. M. Coult.) J. M. Coult. ex Toumey], with genes for drought
resistance and production of compounds which protect against adult-onset
­diabetes, both of which desperately needed by farmers and consumers on our
continent
Foreword ix

• The puzzle sunflower (Helianthus paradoxus Heiser), a halophyte of the Pecos

River in New Mexico and West Texas, which emerged from a chance hybrid of
the common sunflower (Helianthus annuus L.) and the prairie sunflower
(Helianthus petiolaris Nutt.) over 75,000 years ago, but is now far more salt
tolerant than either of its parents, and most cultivated sunflower hybrids as well
• The Okeechobee gourd, a squash relative first described by John and William
Bartram along the St. Johns River in Florida around the time of the Revolutionary
War, which has barely survived the agricultural revolution that drained the
Everglades for sugarcane production and diverted most rivers in Florida into
croplands (Nabhan 1989)
• The Oahu cowpea (Vigna o-wahuensis Vogel), a rare perennial legume that has
gone extinct on the very island in Hawaii where my daughter and grandson now
live but which tenaciously hangs on for dear life in just seven small populations
spread across four of the other Hawaiian Islands
I offer you these brief “personality profiles” to remind you that each of these
valuable and endangered crop relatives has a distinctive character. The tasks of iden-
tifying, counting, tallying, mapping, monitoring, and managing the remaining pop-
ulations of rare plant species on the verge of extinction are ever increasing. It is
worth a moment of our time now and then to remember the complex ecological and
human relationships surrounding each of these unique but declining plants.
It is important to remember that the contribution of wild relatives to crops is not
a new phenomenon. In fact, these plants have naturally exchanged genes in tradi-
tional agricultural settings for millennia. We are all beneficiaries of such serendipi-
tous crop diversification every time we sit down to eat a meal or drink a glass of
wine or cider. They have been – and continue to be – our most useful “living library,”
a set of manuals to help us maintain our food security (Gruber 2017, Khoury 2015,
and in this volume).
Several of the chapters in this book point out the importance of recognizing that
many crop relatives remain economic crops and cultural resources in their own
right. Plant breeders do not necessarily need to “improve” some of these plants to
make them acceptable to the public. For example, the fresh and dried fruits of wild
chiltepín peppers (Capsicum annuum var. glabriusculum) sell for more than USD $80
per pound in much of the U.S. Southwest and northwestern Mexico. One pound of
American wildrice from Minnesota streams and lakes that is hand-harvested and
wood-parched by Native American foragers garners prices of up to USD $17 on
Amazon. On the southern edges of the Chihuahuan Desert, consumers are willing to
pay five to ten times more for a delicious semi-cultivated potato called papita güera
(Solanum cardiophyllum Lindl) than for domesticated potatoes of exotic origin. Yet
the anciently cultivated genotypes of this species are hardly if at all represented in
most potato gene banks, including those in Mexico and the USA.
Wild apples (Malus Mill.) prized for their tartness and flavor are now included in
hard ciders in the USA and Canada. Legally collected or propagated rare food crops
like Price’s potato bean (Apios priceana B. L. Rob.), agaves (Agave L.), and cacti
command high prices in horticultural trade. Wild prickly pears (Opuntia Mill.),
x Foreword

p­ inyon nuts (Pinus L.), and ramps (Allium tricoccum Aiton) continue to attract
almost as much attention from chefs and nutritional scientists as their cultivated
counterparts do. Nevertheless, habitat fragmentation and other threats are diminish-
ing foragers’ access to these North American plants.
Beyond these direct uses, promising new applications of these plants are emerg-
ing from recent innovations in applied research. In Kansas, the Land Institute is
newly domesticating perennial wild relatives of food crops, using intermediate
wheatgrass (Thinopyrum intermedium (Host) Barkworth & D. R. Dewey) and rosin-
weed (Silphium integrifolium Michx.), a distant relative of sunflower, in their prai-
rie-adapted polycultures (Dehaan et al. 2016; Van Tassel et al. 2017). In Missouri,
botanists associated with the Missouri Botanical Garden and St. Louis University
are evaluating wild relatives of commercially important fruit tree crops for develop-
ment in their own right both as sources of food and as rootstocks, due to their hardi-
ness and resistance to emerging insect pests and plagues (Allison Miller, pers. com).
In Illinois, integrated pest management teams have experimented with the native
buffalo gourd (Cucurbita foetidissima Kunth) as a trap crop grown on the edges of
squash and pumpkin fields to reduce larval damage to these crops and increase pol-
lination efficiencies (Metcalf et al. 1980; Metcalf et al. 1982). In Arizona, our eco-
logical research in the first in situ reserve for crop wild relatives in the USA [in
Coronado National Forest] allowed us to determine how capsaicinoids and other
secondary metabolites serve as “directed” chemical defenses against Fusarium
fungi, insect pests, and seed-predating rodents in wild chiltepín peppers (Tewksbury
and Nabhan 2001; Eich 2008). It may now be possible to differentially select and
use the various capsaicinoids in the wild chile pepper arsenal for the discourage-
ment of grain storage pests, prevention of fungal contamination of seeds, treatment
of shingles, reduction of blood serum cholesterol and glucose, and management of
attention-deficit disorders (Eich 2008; Barchenger and Bosland, this volume).
Thus, crop wild relatives are extremely valuable genetic resources, yet they also
offer us their colorful and meaningful natural histories – stories of survival, if you
will, of a more diverse portfolio of plants still available to humanity.
Thankfully, as many of the chapters in this book document, the conservation and
use of wild relatives is getting more serious traction, with national and international
initiatives looking to make a significant impact in the coming years. But these
efforts are the tip of the iceberg of what is needed. As several contributions in this
volume affirm, we must continue to invest in sufficiently supporting every link in
the wild relative-food crop supply chain – from in situ conservation of natural habi-
tats in national parks and biosphere reserves to ex situ seed banks, botanical gar-
dens, and plant restoration efforts – if the entire supply and delivery system is to
function for the future. It is not enough for land grant universities to invest millions
in molecular biology laboratories if they end up closing down herbaria and cutting
budgets of campus arboreta and experimental farms in the process. As Harvard con-
servation biologist E.O. Wilson once quipped, it is the “non-sexy” and more
­descriptive sciences of systematics, ethnobotany, biogeography, and seed storage
physiology that have gotten us to where we are today.
Foreword xi

These recently emerged opportunities will not bear fruit if our funding sources
for habitat conservation and landscape management, for basic biology and seed
banking, and for horticultural innovation and biomedical research focus only on the
last few links of the wild relative-crop commodity supply chain. We not only need
to diversify the genetic base of our food supply, we also need to diversify and sus-
tain the many forms of conservation, restoration, and scholarly inquiry which
together ensure access to these crop genetic resources.
Collectively, the chapters in this remarkable book provide a valuable overview of
the best information and practices needed to safeguard and wisely use North
America’s crop wild relatives. Detailing the species native and naturalized in the
continent and related to important food, fiber, animal forage and feed, industrial oil,
ornamental, and medicinal crops, the authors outline their potential for use and
highlight the conservation needs for the species. In bringing together for the first
time this information from across the broad North American region, including
Canada, Mexico, and the USA, the book provides access to critical conservation
information for well over 600 promising plants. As this landmark volume attests,
these plants are essential elements of North America’s natural and cultural heritage.
This book becomes the model for advancing the efforts needed to better care for this
heritage for present and future generations. It provides us with operating instruc-
tions for wisely managing “our living library.”

Tucson, AZ, USA Gary Paul Nabhan

References

Andres T, Nabhan GP (1988) Taxonomic rank and rarity of Cucurbita okeechobeensis. FAO/
IBPGR Plant Genetic Resources Newsletter 75/76:71–72
DeHaan LR, Van Tassel DL, Anderson JA, Asselin SR, Barnes R, Baute G et al. (2016) A pipeline
strategy for grain crop domestication. Crop Sci 56(3):917–30
Eich E (2008) Solanaceae and Convolvulaceae: secondary metabolites biosynthesis, chemotax-
onomy, biological and economic significance. Springer-Verlag, Berlin.
Gruber K (2017) The living library. Nature 544:S8–S10
Khoury CK, Greene S, Wiersema J, Maxted N, Jarvis A, Struik PC (2013) An inventory of crop
wild relatives of the United States. Crop Sci 53(4):1496
Khoury CK (2015) The conservation and use of crop genetic resources for food security.
PhD. Dissertation, Wageningen University, Wageningen, N.L
Metcalf RL, Metcalf RA, Rhodes AM (1980) Cucurbitacins as kairomones for diabroticite beetles.
Proc Natl Acad Sci 77(7):3769–3772
Metcalf RL, Rhodes AM, Metcalf RA, Ferguson J, Metcalf ER, Lu P-Y (1982) Cucurbitacin
contents and diabrocite (Coleoptera: Chrysomelidae) feeding upon Cucurbita spp. Environ
Entomol 11(4):931–938
Nabhan GP (1989) Enduring seeds: native American agriculture and wild plant conservation.
University of Arizona Press, Tucson
Nabhan GP (1990) Conservationists and forest service join forces to save wild chiles. Diversity
6(3&4):47–48
xii Foreword

Nabhan GP (1991) Genetic resources of the U.S./Mexico border: wild relatives of crops, their
uses and conservation. In: Ganster P, Walter H (eds) Environmental hazards and bioresource
management in the U.S./Mexico Borderlands. UCLA Latin American Center Publications, Los
Angeles
Rogers K (2015) The quiet extinction: stories of North America’s rare and threatened plants.
University of Arizona Press, Tucson
Routson KJ, Volk GM, Richards CM, Smith SE, Nabhan GP, Wyllie de Echeverria V (2012)
Genetic variation and distribution of Pacific crabapple. J Am Soc Hortic Sci 137(5):325–332
Tewksbury JJ, Nabhan GP (2001) Seed dispersal: directed deterrence by capsaicin in chilies.
Nature 412:403
Van Tassel DL, Albrecht KA, Bever JD, Boe AA, Brandvain Y et al (2017) Accelerating Silphium
domestication: an opportunity to develop new crop ideotypes and breeding strategies informed
by multiple disciplines. Crop Sci 57(3):1274–1284
Zhang J, Nielsen SE, Chen Y, Georges D, Qin Y, Wang SS, Svenning J-C, Thuiller W (2016)
Extinction risk of North American seed plants elevated by climate and land-use change. J Appl
Ecol 54:303–312
Preface

Wild plants useful to food and agriculture occupy a niche frequently outside the
realm of both agricultural and natural resource professionals. The agricultural com-
munity tends to focus on a handful of domesticated species, while the natural
resource community emphasizes legislatively regulated taxa (i.e., species that are
rare, endangered, indicators of ecosystems such as wetlands, or wild species used
for timber or revegetation).
The increasing challenges to food and agricultural production due to climate
change, added to the ever-present biotic and abiotic stresses, are likely to make crop
wild relatives and other useful wild plants more of a priority to both communities.
And the combined efforts of both communities will be critical to locating, conserv-
ing, and making available these invaluable species to support agricultural produc-
tion and food and nutritional security for future generations.
The purpose of this two-volume book is to highlight the most important wild
plant genetic resources that grow in North America. We define these resources as
wild species with relevance for agriculture. These include the wild plant populations
from which domesticated varieties evolved, crop wild relatives that can be used to
improve contemporary crops, wild species that have a record of use by people, and
any other wild species with potential for future crop development. Most of the spe-
cies covered are native, but a few are introduced taxa that have become naturalized
in the region. A thorough understanding of the species that occupy North America,
including their distributions, potential value to agriculture, and conservation statuses
and needs, will give agricultural and conservation communities the basic knowledge
they need to take steps to conserve our natural heritage of useful wild plants.
The overarching goal of this book is to help ensure that these valuable but over-
looked species continue to persist, both in their natural habitats and in gene banks,
where they can be made available as resources to address compounding agricultural
and nutritional challenges. This book is authored by a broad range of experts work-
ing diligently to explore, protect, celebrate, and use crop genetic resources. They
have come together to compile the latest information on the most important North
American wild useful plants. This book focuses on Canada, the USA, and Mexico,
three countries whose combined area covers most of the continent.

xiii
xiv Preface

The first volume of the book covers topics relevant to the conservation of all wild
plant genetic resources, while the second volume focuses on specific crops and their
related wild taxa. Volume One: Conservation Strategies begins by reviewing efforts,
challenges, and opportunities to conserve food and agriculturally important wild
species from a national perspective. The first part provides not only a broad over-
view of important crop wild relatives and wild utilized species in Canada, Mexico,
and the USA but also a description of the agencies and institutes focused on con-
serving these plants, as well as the conservation and use policies followed by each
country. This section concludes with a chapter that presents Native American tribal
perspectives in the USA, providing a glimpse into the management and regulation
of plant genetic resources by Indigenous peoples through a set of case studies of
several tribal governments.
The second part of Volume One discusses various aspects of wild plant genetic
resource conservation methodologies. Managing genetic resources of wild plants
involves additional considerations beyond those required for domesticated crops.
Likewise, wild genetic resource conservation differs from managing plant species
that are rare and endangered in that a greater emphasis is placed on ex situ activities,
including storage and seed regeneration, that ensure that these resources are avail-
able for use by plant breeders and the scientific community. This section is orga-
nized in a logical sequence that begins with a chapter on current and emerging
frameworks on defining wild genetic resources, as well as a chapter on threat assess-
ments. Two subsequent chapters cover sampling strategies and collecting practicali-
ties, followed by chapters that discuss aspects of gene banking wild species,
including storage and seed increase. Volume One concludes with a chapter that
discusses public education and outreach opportunities for crop wild relatives.
There are over 20,000 plant species in North America, and all deserve a chance
to thrive. However, a small fraction of these are distinguished by their potential to
support food and agricultural production, either because they are resources that can
be used to breed more productive crops or because they have commercial or cultural
value when used directly. Many of these species are common, even weedy, and are
easily overshadowed by rare or endangered plants. Nevertheless, because of their
real or potential importance to our food and agriculture, they deserve to be recog-
nized, celebrated, conserved, and made available for use.
The Editors would like to acknowledge and thank the authors for the extensive
work they have done to compile, organize, and write their chapters. We would also
like to acknowledge the contribution and insights provided by peer reviewers of
each chapter.

Fort Collins, CO, USA Stephanie L. Greene

Beltsville, MD, USA Karen A. Williams
Fort Collins, CO, USA Colin K. Khoury
Honolulu, HI, USA Michael B. Kantar
Ames, IA, USA Laura F. Marek
Contents

Part I Conservation in North America

1 Wild Plant Genetic Resources in North America: An Overview ��������    3
Stephanie L. Greene, Colin K. Khoury, and Karen A. Williams
2 Genetic Resources of Crop Wild Relatives: A Canadian
Perspective������������������������������������������������������������������������������������������������   33
Axel Diederichsen and Michael P. Schellenberg
3 Crop Wild Relatives in Mexico: An Overview of Richness,
Importance, and Conservation Status����������������������������������������������������   63
J. Fernando De La Torre S., Rosalinda González S.,
E. Judith Cruz G., J. Manuel Pichardo G., Martín Quintana C.,
Aremi R. Contreras T., and Jorge Cadena I.
4 Conservation of Crop Wild Relatives in the USA ��������������������������������   97
Karen A. Williams and Stephanie L. Greene
5 Working with Indigenous Communities: The Original
Caretakers of Crops and Crop Wild Relatives�������������������������������������� 155
Scott M. Herron

Part II Conservation Strategies

6 The Gene Pool Concept Applied to Crop Wild Relatives:
An Evolutionary Perspective������������������������������������������������������������������ 167
Richard E. Miller and Colin K. Khoury
7 Conservation Status and Threat Assessments for North
American Crop Wild Relatives �������������������������������������������������������������� 189
Anne L. Frances, Adam B. Smith, and Colin K. Khoury

xv
xvi Contents

8 Sampling Wild Species to Conserve Genetic Diversity������������������������ 209

Sean Hoban, Gayle Volk, Kanin J. Routson, Christina Walters,
and Chris Richards
9 Practicalities of Collecting Wild Plants in North America:
Insights from the United States�������������������������������������������������������������� 229
Laura F. Marek
10 Genebank Conservation of Germplasm Collected
from Wild Species������������������������������������������������������������������������������������ 245
Christina Walters, Christopher M. Richards, and Gayle M. Volk
11 Practical Considerations for Increasing Seed Samples
of Wild Species ���������������������������������������������������������������������������������������� 281
Barbara C. Hellier
12 Public Education and Outreach Opportunities
for Crop Wild Relatives in North America�������������������������������������������� 311
Tara Moreau and Ari Novy

Appendix 1 Map Methods and Occurrence Data Sources �������������������������� 325

Common Name Index�������������������������������������������������������������������������������������� 335

Scientific Name Index�������������������������������������������������������������������������������������� 339

Reviewers

Tabare Abadie DuPont Pioneer, Johnston, IA, USA

Dani Ballesteros Kew Royal Botanical Gardens, Richmond, England, UK
Vicki Bradley USDA Agricultural Research Service, Pullman, Washington, USA
Peter Bretting USDA Agricultural Research Service, Beltsville, MD, USA
Jacques Cayouette Agriculture and Agri, food Canada, Ottawa, ON, Canada
Neil Cox The International Union for Conservation of Nature, Washington DC,
USA
Denise Culver Colorado National Heritage Program, Fort Collins, CO, USA
Luigi Guarino Crop Trust, Bonn, Germany
Edward Guerrant Portland State University, Portland, OR, USA
Eric N. Jellen Brigham Young University, Provo, UT, USA
Amanda C. Martin The Kochi Prefectural Makino Botanical Garden, Kochi City,
Japan
Allison Miller Saint Louis University, St Louis, MO, USA
Melanie Schori USDA Agricultural Research Service, Beltsville, MD, USA
Susan Stieve The Ohio State University, Columbus, OH, USA

xvii
Abbreviations

AAFC Department of Agriculture and Agri-Food Canada

ABS Access and Benefit Sharing
ABSA Access and Benefit Sharing Agreement
AFLP Amplified fragment length polymorphism
AHLV American hop latent virus
AHPA American Herbal Products Association
AMJB Mexican Association of Botanical Gardens
ANSI American National Standards Institute
ANSM Universidad Autónoma Agraria Antonio Narro
AOSCA Association of Official Seed Certifying Agencies
APGA American Public Gardens Association
ApMV Apple mosaic virus
ARS Agricultural Research Service
ASU Arizona State University
ATBI All Taxa Biodiversity Inventory
AZ Arizona
BCMV Bean common mosaic virus
BGCI Botanic Gardens Conservation International
BIA Bureau of Indian Affairs
BLM Bureau of Land Management
BONAP Biota of North America Program
BP Before present
BP&P Bayesian phylogenetics and phylogeography
Bt Bacillus thuringiensis
BTA Boyce Thompson Arboretum
CA California
CAM Crassulacean acid metabolism
CAPS Cleaved amplified polymorphic sequences
CBCN Canadian Botanical Conservation Network
CBD Convention on Biological Diversity
CBIF Canadian Biodiversity Information Facility

xix
xx Abbreviations

CCC Civilian Conservation Corps

CCD Colony collapse disorder
CCGB Canadian Clonal Genebank
CCN Conservation Center Network
CDFW California Department of Fish and Wildlife
CDI National Commission for the Development of Indigenous Peoples
CEVAMEX Valle de México Research Station
CFIA Canadian Food Inspection Agency
CGC Crop Germplasm Committee
CGN Centre for Genetic Resources
CIAT International Center for Tropical Agriculture
CICTAMEX Scientific and Technological Research Center of Avocado in the
State of Mexico
CICY Yucatan Center for Scientific Research
CIMMYT International Maize and Wheat Improvement Center
CITES Convention on International Trade in Endangered Species of
Wild Fauna and Flora
CLCuD Cotton leaf curl disease
CMS Cytoplasmic male sterility
CMV Cucumber mosaic virus
CNHP Colorado Natural Heritage Program
CNRG National Genetic Resources Center
Co County
CONABIO National Commission for the Knowledge and Use of
Biodiversity
CONANP National Commission of Natural Protected Areas
COSEWIC Committee on the Status of Endangered Wildlife in Canada
CP Coefficient of Parentage
CP Postgraduate College
CPB Cartagena Protocol on Biosafety
CPC Center for Plant Conservation
CPGR Canadian Potato Genetic Resources
CPNWH Consortium of Pacific Northwest Herbaria
CS College Station Texas
CSIRO Commonwealth Scientific and Industrial Research Organization
CSN Counter Season Nursery
CWR Crop wild relatives
DELEP Desert Legume Program
DMI Dobzhansky-Muller incompatibilities
DNA Deoxyribonucleic acid
DNR Department of Natural Resources
DOD Department of Defense
DOE Department of Energy
DOF Official Gazette of the Federation
DOT Department of Transportation
Abbreviations xxi

EC Electrical conductivity
ECOS Environmental Conservation Online System
ELISA Enzyme-linked immunosorbent assay
EMCV Mexican Plant Conservation Strategy
ENBioMex National Biodiversity Strategy of Mexico
ENGOs Environmental non-governmental organizations
ENM Ecological niche model
ESA Endangered Species Act
EUFGIS European Information System on Forest Genetic Resources
FAO Food and Agriculture Organization of the United Nations
FES-I Faculty of Higher Education Iztacala
FGP Frost grape polysaccharide
FNA Flora of North America
FRRL Forage and Range Research Laboratory
G Global
GBIF Global Biodiversity Information Facility
GBS Genotyping by sequencing
GG Germplasm Resource Information Network-Global database
GH Greenhouse
GIS Geographical Information Systems
GLIFWC Great Lakes Indian Fish and Wildlife Commission
GLS Gray leaf spot
GMP Good Manufacturing Practice
GP Genepool
GPS Global Positioning System
GRIN Germplasm Resource Information Network database
GS Genome selection
GWAS Genome-wide association study
Ha Hectare
HPLC High-performance liquid chromatography
HpLV Hop latent virus
HpMV Hop mosaic virus
HPTLC High-performance thin-layer chromatography
HRT Hormone replacement therapy
ICAMEX Institute for Training, Research, and Development in Agriculture
of the State of Mexico
ICAR Indian Council on Agricultural Research
ICRISAT International Crop Research Institute for the Semi-Arid Tropics
iDIGBio Integrated Digitized Biocollections
IGRR In Situ Genetic Resource Reserve
IICA Inter-American Institute for Cooperation on Agriculture
ILDB The International Lactuca Database
INDR Indiana Department of Natural Resources
INIFAP National Institute of Forestry, Agriculture and Livestock Research
INRA French National Institute for Agricultural Research
xxii Abbreviations

INTA Instituto Nacional de Tecnología Agropecuaria

IPGRI International Plant Genetic Resources Institute
IPK Leibniz-Institut für Pflanzengenetik und Kulturpflanzenforschung
IPR Intellectual property rights
ISO International Standards Organization
ISSR Inter-simple sequence repeat
ITIS Integrated Taxonomic Information System
ITPGRFA International Treaty on Plant Genetic Resources for Food and
Agriculture
IUCN International Union for Conservation of Nature
KBIC Keweenaw Bay Indian Community
LBOGM Law on Biosafety of Genetically Modified Organisms
LFVV Federal Law on Plant Varieties
LGCC Law on Climate Change
LGCS USDA Lactuca Germplasm Collection in Salinas
LGRCD Lactuca Genetic Resources Collection at the University of
California in Davis
MCMV Maize chlorotic mosaic virus
Mha Million hectares
MLN Maize lethal necrosis
MLS Multilateral system
MRC Major resistance clusters
MRCA Most recent common ancestor
MS Flow injection mass spectrometry
MSB Millennium Seed Bank
MSBP Millennium Seed Bank Project
MT Metric tons
MTA Material Transfer Agreement
N National
n/d Not determined
NA North America
NALPGRU National Arid Land Plant Genetic Resources Unit
NARO National Agriculture and Food Research Organization
NASS National Agricultural Statistics Service
NBSAP National Biodiversity Strategy and Action Plan
Nc Census population size
NCGC National Cotton Germplasm Collection
NCGR National Clonal Germplasm Repository
NCGR-Davis National Clonal Germplasm Repository in Davis, California
NCRPIS North Central Regional Plant Introduction Station
Ne Effective population size
NGRL National Germplasm Resources Laboratory
NGS Next-generation sequencing
NILs Near isogenic lines
NKLP Nagoya–Kuala Lumpur Protocol on Liability and Redress
Abbreviations xxiii

NLGRP National Laboratory for Genetic Resources Preservation

NLR Nucleotide binding-leucine rich repeat receptor
NM New Mexico
NMFS National Marine Fisheries Service
NMR Nuclear magnetic resonance
NOM-059 NOM-059-SEMARNAT-2010-Mexican standard that lists all
threatened native wild species
NORGEN North American Regional Network for Agricultural Research-
Genetic Resources Task Force
NP Nagoya Protocol
NPA Natural Protected Areas
NPABS Nagoya Protocol on Access and Benefit Sharing
NPGS USDA, ARS National Plant Germplasm System
NPS National Park Service
NRCS USDA, Natural Resources Conservation Service
NS SEME Novi Sad Institute of Field and Vegetable Crops
NS/S Native Seeds/Search
NSL United States Forest Service National Seed Laboratory
NUS Neglected underutilized species
OPGC Ornamental Plant Germplasm Center
OR Oregon
OSU Ohio State University
PacBio Pacific Biosystems
PANREFI National Plan of Action for the Conservation of Plant Genetic
Resources for Food and Agriculture
PBLS Pecan bacterial leaf scorch
PCA Plant Conservation Alliance
PCB Cartagena Protocol on Biosafety
PCN Plant Conservation Network
PCR Polymerase chain reaction
PEO USDA, ARS Plant Exchange Office
PGRC Plant Gene Resources of Canada
PGRFA Plant Genetic Resources for Food and Agriculture
PI Plant Introduction
PMC USDA, Natural Resources Conservation Service Plant Material
Center
PROCINORTE North American Regional Network for Agricultural Research
PRV Papaya ringspot virus
PVG Pre-variety germplasm
PVP US Plant Variety Protection
QTL Quantitative trait loci
RAPD Random amplified polymorphic DNA
RBG Kew Royal Botanical Gardens Kew
RGC16 Family of nucleotide binding-leucine rich repeat receptor
(NLR) proteins
xxiv Abbreviations

RGC2 Resistance Gene Candidate 2

RMA Research Marketing Act
RNA Research Natural Area
RNA Ribonucleic acid
S Subnational
SAGARPA Secretariat of Agriculture, Livestock, Rural Development,
Fisheries and Food
SBBG Santa Barbara Botanic Garden
SCP Species Conservation Plan
SDBG San Diego Botanic Garden
SDG Sustainable Development Goals
SDM Species distribution model
SEINet Southwestern Environmental Information Network
SEMARNAT Ministry of the Environment and Natural Resources
SID Seed Information Database
SINAREFI National Program of Plant Genetic Resources for Food and
Agriculture
SMTA Standard material transfer agreement
SNICS National Seed Inspection and Certification Service
SNP Single nucleotide polymorphisms
SNWA Southern Nevada Water Authority
SOMEFI Mexican Society of Plant Genetics
SOP Standard operating procedure
SOS Seeds of Success
SS Sequence-tagged site
SSR Simple sequence repeat
SW Southwestern
SWD Spotted wing drosophila
TES Threatened, Endangered, and Sensitive Species
TEX University of Texas
Tg Glass transition temperature
TNC The Nature Conservancy
TNPD Texas Native Plants Database
TRAP Target region amplification polymorphism
TRIPS Trade-Related Aspects of Intellectual Property Rights
TVA Tennessee Valley Authority
TX Texas
UAAAN Antonio Narro Agrarian Autonomous University
UACh Chapingo Autonomous University
UBC-BG University of British Columbia Botanical Garden
UC Davis V&E The University of California, Davis Viticulture and Enology
Department
UCBG University of California Botanical Garden
UDG University of Guadalajara
UGto University of Guanajuato
Abbreviations xxv

UMA Wildlife Conservation Management Units

UNAM National Autonomous University of Mexico
UNCCD United Nations Convention to Combat Desertification
UNESCO United Nations Educational, Scientific and Cultural
Organization
UNFAO United Nations Food and Agriculture Organization
UNFCCC United Nations Framework Convention on Climate Change
UPOV International Union for the Protection of New Varieties of
Plants
UPP Useful Plants Project
URL Uniform Resource Locator
US United States
USA United States of America
USD United States Dollars
USDA United States Department of Agriculture
USDA-ARS United States Department of Agriculture – Agricultural
Research Service
USDOI United States Department of the Interior
USFS United States Forest Service
USFWS US Fish and Wildlife Service
USFWS United States Fish and Wildlife Service
UT Utah
UV University of Veracruz
VASCAN Database of Vascular Plants of Canada
VIR Vavilov All-Russian Research Institute of Plant Industry
VNIIMK All Russian Research Institute of Oil Crops
WGD Whole genome duplication
WMV Watermelon mosaic virus
WRPIS USDA, ARS Western Regional Plant Introduction Station
WTO World Trade Organization
WUS Wild utilized species
WY Wyoming
YUNGA Youth and United Nations Global Alliance
ZWMV Zucchini yellow mosaic virus
 Part I
Conservation in North America
Chapter 1
Wild Plant Genetic Resources in North
America: An Overview

Stephanie L. Greene, Colin K. Khoury, and Karen A. Williams

Abstract North America, including Canada, Mexico, and the United States, is rich
in plant species used by humans in both ancient and modern times. A select number
of these have become globally important domesticated crops, including maize,
beans, cotton, and sunflower. Many other native and also naturalized species have
potential for use, either directly or as genetic resources for breeding agricultural
crops. However, despite increasing recognition of their potential value, deficiencies
in information, conservation, and access to the diversity in these plants hinder their
further use. This chapter provides an overview of the agriculturally relevant wild
plant resources of North America, with focus on wild relatives of globally important
major crops, as well as the wild cousins of regionally and locally important domes-
ticates. The chapter concludes by providing an overview of strategies for conserving
wild plant genetic resources, including the international regulatory frameworks
affecting policies to various degrees in Canada, Mexico, and the United States.

Keywords Germplasm · Genetic resources · Wild species · Conservation

· North America · Crop resources

S. L. Greene (*)
USDA, Agricultural Research Service, Center for Agricultural Resources Research, National
Laboratory for Genetic Resources Preservation, Fort Collins, CO, USA
e-mail: stephanie.greene@ars.usda.gov
C. K. Khoury
USDA, Agricultural Research Service, Center for Agricultural Resources Research, National
Laboratory for Genetic Resources Preservation, Fort Collins, CO, USA
International Center for Tropical Agriculture (CIAT), Cali, Colombia
e-mail: colin.khoury@ars.usda.gov; c.khoury@cgiar.org
K. A. Williams
USDA, Agricultural Research Service, Beltsville Agricultural Research Center, National
Germplasm Resources Laboratory, Beltsville, MD, USA
e-mail: karen.williams@ars.usda.gov

© This is a U.S. government work and not under copyright protection 3

in the U.S.; foreign copyright protection may apply 2018
S. L. Greene et al. (eds.), North American Crop Wild Relatives, Volume 1,
https://doi.org/10.1007/978-3-319-95101-0_1
4 S. L. Greene et al.

1.1 Introduction

Increasingly variable weather, shifting disease and pest pressures, soil degradation,
loss of arable lands, and water scarcity are not only on the horizon; they are already
our reality. Canada and the United States are experiencing higher temperatures and
more severe weather events, and wildfires (Field et al. 2007). Mexico is undergoing
an even greater range of climatic changes, including increased temperatures, espe-
cially in the north, decreased rainfall in the central regions, and more storms and
prolonged drought during the dry season (National Intelligence Council 2009).
Farmers in North America face a turbulent ride as they navigate the Anthropocene
to continue to produce a considerable portion of the food, fiber, and other plant-­
based resources utilized around the world.
An important strategy for preparing for these challenges is breeding plants that
can handle the emerging abiotic and biotic challenges. Wild plant species that are
closely related to crops are increasingly recognized as some of the most promising
genetic resources that plant breeders can turn to in their efforts to develop cultivars
adapted to more extreme conditions (Dempewolf and Guarino 2015; Dempewolf
et al. 2017; Castañeda-Álvarez et al. 2016; Zhang et al. 2017). They have already
proven their worth in breeding (Hajjar and Hodgkin 2007; Maxted et al. 2012).
Ironically, however, the very wild species being promoted as essential tools in
resolving agricultural problems are themselves vulnerable to the accelerating envi-
ronmental changes (Jarvis et al. 2008; Lira et al. 2009; Thomas et al. 2016), as well
as to the persistent threats presented by habitat modification, pollution, invasive
species, and other anthropogenic impacts (Brummitt et al. 2015).
Confounding progress on conservation of these important species is the fact
that agriculturally relevant wild plants occupy a niche that is generally neglected
by agricultural researchers, who tend to focus their attention on a handful of
crops. At the same time, agriculturally relevant wild species are often also over-
looked by habitat and endangered species conservation practitioners, who focus
on securing rare and threatened taxa and their ecosystems, rather than on safe-
guarding the intraspecific variation in frequently common, and often weedy, crop
wild relatives.
The information in this introductory chapter sets the stage for the rest of the
book. We begin by defining essential terms and concepts. We then discuss the
process of domestication, focusing on the crops domesticated in North America.
We briefly discuss the importance of wild utilized species and then focus on an
overview of the occurrence and conservation status of North American crop wild
relatives of important crops. We conclude by discussing in broad strokes the
general strategies for conserving wild plant genetic resources, including the
international regulatory frameworks affecting policies to various degrees in the
region.
1 Wild Plant Genetic Resources in North America: An Overview 5

1.2  n Agricultural Perspective on North America’s Wild

A
Flora

The number of native vascular plant species in the three countries increases from
north to south. Canada has almost 5860 (Brouillet et al. 2010), and the United States
has nearly 16,200 native plant species (Stein 2002). With 23,314 species (Villaseñor
2016), Mexico is a megadiverse country and fourth in the world for number of
native vascular plants. Given such broad diversity in species, it is perhaps surprising
that modern-day humans use a relatively small number. The United Nations Food
and Agriculture Organization (FAO) global aggregate statistics tell us that just 18 or
so plants contribute to 90% of the world’s calories and that of these, only 2, maize
and sunflower, are indigenous to North America (Khoury et al. 2014). However,
calories are not the only important component of diet. Plant species also provide
protein, fat, starch, fiber, vitamins, minerals, phytonutrients, and flavor. For exam-
ple, chili peppers, the main cultivated species that originated in North America, are
the world’s most important spice by essentially all measures. Moreover, many plant
species are used for purposes other than food. Examples of important North
American species valued for non-food uses include cotton (as a textile), echinacea
and American ginseng (as medicines), guayule and jojoba (for industrial uses), hops
and vanilla (as flavorings), and rudbeckia and phlox (as ornamentals).
Useful plants fall along a continuum that can be categorized according to the
extent that humans have influenced their form. At one end of the continuum are
domesticated species, which we call “crops.” Crops display a very considerable
suite of changes driven by selection pressures placed upon them by humans, typi-
cally including the loss of natural dispersal mechanisms; larger sizes of seeds, fruits,
or other plant parts; and the loss of dormancy. At the other end are wild species that
show no morphological evidence of human use. Both crops and wild species can be
managed by humans (e.g., wild fruit trees managed in situ by burning or annual
crops cultivated ex situ by planting and harvesting in fields), and management of
wild species can lead to domestication (i.e., Casas et al. 2007). The focus of this
book is on wild plant species with relevance for agriculture and other human uses,
which we term “wild plant genetic resources.” These species include the wild plant
populations from which domesticated varieties evolved (crop progenitors), wild
species that can be used to improve contemporary crops (crop wild relatives), wild
species that have a record of use by people (wild utilized species), and any other
wild species with potential for future crop development (new crops).

1.2.1 Wild Utilized Species

Historically, wild plant species have underpinned the diets of gatherer-hunter and
forager cultures and continue today to contribute significantly to diets, particularly
in rural regions of the developing world (Bharucha and Pretty 2010). In Mexico, it
6 S. L. Greene et al.

is estimated that 5000–7000 wild plant species were used for food and other pur-
poses (Casas et al. 1994; Caballero et al. 1998). North of Mexico, approximately
1800 species have been documented as having been used by the indigenous peoples
of North America (Moerman 2003), and Uprety et al. (2012) reported that 546
medicinal plants were used by indigenous peoples in the boreal forests of Canada.
Many of these wild food and medicinal species were adopted by early colonists in
North America (Turner and von Aderkas 2012), and foraging for wild plant species
to use as food or medicine continues to be important in North America. In recent
decades, there has been growing interest in using wild plants, especially native spe-
cies, to revegetate or restore wild lands. For example, in the United States, an alli-
ance of federal and private partners has developed the National Seed Strategy for
Rehabilitation and Restoration, driven by federal mandates to use native plant mate-
rials (Plant Conservation Alliance 2015).

1.2.2 Domestication in North America

For a select group of plant species, human use has led to domestication. Larson
et al. (2014) provide a general definition of domestication as “a selection process for
adaptation to human agro-ecological niches and at some point in the process, human
preference.” Archeological remains provide ample evidence for the domestication
of North American wild progenitors of crop plants, beginning 12,000–8200 BP in
Mesoamerica and 8200–4200 BP in temperate regions (Larson et al. 2014). A num-
ber of these domesticates have over time become globally important (Table 1.1)
(Khoury et al. 2016). A few, such as marsh elder (Iva annua L.), little barley
(Hordeum pusillum Nutt.), and devil’s claw (Proboscidea parviflora [Wool.] Wool
and Standi), have largely been abandoned (Smith 2006; Bretting 1986).
The process of domestication is driven by the interaction of environmental fac-
tors, biology, and human needs, which results in crops that range from plants that
differ only slightly from their wild ancestors to species that cannot persist without
human interaction (Larson et al. 2014; Meyer et al. 2012). Mesoamerica provides a
fine example of this (Lira 2009). Over 20 plant species have been domesticated and
have reached globally important food- crop status, including maize (Zea mays L.),
beans (Phaseolus L. spp.), chili pepper (Capsicum annuum L.), pumpkins and
squashes (Cucurbita pepo L.), cotton (Gossypium hirsutum L.), avocado (Persea
americana Mill), cacao (Theobroma cacao L.), and vanilla (Vanilla planifolia
Jacks). Within this same region, a study limited to the Tehuacán-Cuicatlán Valley,
Mexico, found that there were over 200 species currently in incipient stages of
domestication, the result of management in traditional agricultural systems (Casas
et al. 2007; Avendaño et al. 2009; Blancas et al. 2010). Table 1.1 provides a select
list of native North American domesticated species, including the approximate time
of start of domestication, and the degree to which the crop has changed from its wild
counterpart. The list demonstrates that a wide range of crops have been domesti-
cated in North America, that crops have been domesticated from the pre-Columbian
1 Wild Plant Genetic Resources in North America: An Overview 7

Table 1.1 Selected native North American domesticates

Common Domestication Comments/key
Taxon name Location Datea levelb references
Agave tequilana Agave Yucatan, 9000 Low Meyer et al.
Weber and other Mexico (2012); Colunga-­
Agave species GarcíaMarín and
Zizumbo-­
Villarreal (2007)
Amaranthus Amaranth Mexico 6000 Low Pickersgill
caudatus L., A. (2007); Janick
cruentus L., and A. (2013)
hypochondriacus
L.
Andropogon Big bluestem United States 50 Low Price et al. (2012)
gerardii Vitman
Annona cherimola Cherimoya Southern 4000 Low Casas et al.
Mill. Mexico (2007)
Apios Americana Indian bean Midwestern to 500 Low Reynolds et al.
Medik. Northeastern (1990)
United States
Asimina triloba Pawpaw Southeastern, 100 Low Meyer et al.
(L.) Dunal United States (2012)
Bouteloua Buffalo United States 50 Low Riordan and
dactyloides (Nutt.) grass Browning (2003)
Columbus
Canavalia Horse bean, Mexico 1050 Low Sauer and Kaplan
ensiformis (L.) jack bean (1969)
DC.
Capsicum annuum Chili pepper Highlands of 6000 High Meyer et al.
L. var. annuum Mexico (2012)
Carica papaya L. Papaya Southern 2000 Medium Meyer et al.
Mexico (2012)
Carya illinoinensis Pecan Southeastern 400 Medium Grauke et al.
(Wangenh.) United States (2016)
K. Koch
Casimiroa edulis White sapote Central Mexico 5000 Medium Meyer et al.
Llave & Lex. (2012)
Chenopodium Chenopod Eastern United 3700 Low No longer
berlandieri subsp. States cultivated/Smith
jonesianum (2006)
Chenopodium Huauzontle Southern 700 Low Smith (2006)
berlandieri subsp. Mexico
nuttalliae
Chenopodium Epazote Mexico 100 Low Blanckaert et al.
ambrosioides L. (2012)
(continued)
Another random document with
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The Project Gutenberg eBook of Fifteen years of
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 This ebook is for the use of anyone anywhere in the United
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Title: Fifteen years of a dancer's life

With some account of her distinguished friends

Author: Loie Fuller

Author of introduction, etc.: Anatole France

Release date: November 29, 2023 [eBook #72257]

Language: English

Original publication: United Kingdom: Herbert Jenkins Limited,

1913

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*** START OF THE PROJECT GUTENBERG EBOOK FIFTEEN

YEARS OF A DANCER'S LIFE ***
 FIFTEEN YEARS OF
A DANCER’S LIFE
WITH SOME ACCOUNT OF HER
DISTINGUISHED FRIENDS
BY
LOIE
FULLER
WITH AN INTRODUCTION
BY

ANATOLE FRANCE

“SHE OUGHT TO WRITE OUT HER MEMORIES AND HER IMPRESSIONS.”—

Alexandre Dumas

HERBERT JENKINS LIMITED

ARUNDEL PLACE, HAYMARKET
LONDON, S.W. MCMXIII
Photo Langfier
LOIE FULLER
 ORIGINALLY PUBLISHED IN FRENCH.

LONDON AND NORWICH PRESS, LIMITED, LONDON AND

NORWICH
 INTRODUCTION

I HAD seen her only as she had been seen by multitudes from
every corner of the globe, on the stage, waving her draperies in
the first light, or transformed into a great resplendent lily, revealing
to us a new and dignified type of beauty. I had the honour of being
presented to her at a luncheon of the tour du monde at Boulogne. I
saw an American lady with small features, with blue eyes, like water
in which a pale sky is reflected, rather plump, quiet, smiling, refined. I
heard her talk. The difficulty with which she speaks French adds to
her power of expression without injuring her vivacity. It obliges her to
rely on the rare and the exquisite, at each moment to create the
requisite expression, the quickest and best turn of speech. Her
words gush forth, the unaccustomed linguistic form shapes itself. As
assistance she employs neither gestures nor motions, but only the
expression of her eyes, which changes like the landscapes that are
disclosed along a beautiful highway. And the basis of her
conversation, now smiling and now serious, is one of charm and
delightfulness.
This brilliant artist is revealed as a woman of just and delicate
sensibility, endowed with a marvellous perception of spiritual values.
She is one who is able to grasp the profound significance of things
that seem insignificant, and to see the splendour hidden in simple
lives. Gleefully she depicts, with keen and brilliant stroke, the humble
folk in whom she finds some ennobling and magnifying beauty. Not
that she is especially devoted to the lowly, the poor in spirit. On the
contrary she enters easily into the lives of artists and scholars. I have
heard her say the most delicate, the subtlest things about Curie,
Mme. Curie, Auguste Rodin and other geniuses. She has
formulated, without desiring to do so, and perhaps without knowing
it, a considerable theory of human knowledge and philosophy of art.
But the subject of conversation which comes closest to her is
religious research. Should we recognize in this fact a characteristic
of the Anglo-Saxon race, of the effect of a Protestant education, or
simply a peculiarity of temperament of which there is no
explanation? I do not know. At all events she is profoundly religious,
with a very acute spirit of inquiry and a perpetual anxiety about
human destiny. Under various guises, in various ways, she has
asked me about the cause and the final outcome of things. I need
not say that none of my replies were couched in a manner to satisfy
her. Nevertheless she has received my doubts serenely, smiling at
everything. For she is distinctly an amiable being.
As regards understanding? Comprehension? She is marvellously
intelligent. She is even more marvellously instinctive. Rich in so
many natural gifts she might have become a scholar. I have heard
her employ a very comprehensive vocabulary in discussing the
various subjects of astronomy, chemistry and physiology. But it is the
unconscious in her that counts. She is an artist.
I have been unable to resist the pleasure of recalling my first
meeting with this extraordinary and delightful woman. What a rare
chance! You admire afar off, as in a vision, an airy figure comparable
in grace to those dancers whom one sees on Pompeiian wall
paintings, moving in their light draperies. Some day you discover
once again this apparition in real life, softened in colour and hidden
under those thicker robes with which mortals cover themselves, and
you perceive that she is a person of good mind and good heart, a
soul somewhat inclined to mysticism, to philosophy, to religion, a
very deep, a very cheerful and a very noble soul.
There you have to the life this Loie Fuller, in whom our Roger
Marx has hailed the chastest and most expressive of dancers,
beautifully inspired, who reanimates within herself and restores to us
the lost wonders of Greek mimicry, the art of those motions, at once
voluptuous and mystical, which interpret the phenomena of nature
and the life history of living beings.
ANATOLE FRANCE
 CONTENTS

PAGE
I. MY STAGE ENTRANCE 15
II. MY A PPEARANCE ON A REAL STAGE AT TWO YEARS AND A 20
HALF
III. HOW I CREATED THE SERPENTINE DANCE 25
IV. HOW I CAME TO PARIS 43
V. MY APPEARANCE AT THE FOLIES-BERGÈRE 51
VI. LIGHT AND THE DANCE 62
VII. A JOURNEY TO RUSSIA—A BROKEN CONTRACT 73
VIII. SARAH BERNHARDT—THE DREAM AND THE REALITY 84
IX. ALEXANDRE DUMAS 101
X. M. AND MME. CAMILLE FLAMMARION 111
XI. A V ISIT AT RODIN’S 122
XII. M. GROULT’S COLLECTION 128
XIII. MY DANCES AND THE CHILDREN 137
XIV. PRINCESS MARIE 151
XV. SEVERAL SOVEREIGNS 165
XVI. OTHER MONARCHS 184
XVII. SOME PHILOSOPHERS 192
XVIII. HOW I DISCOVERED HANAKO 207
XIX. SARDOU AND KAWAKAMI 217
XX. AN EXPERIENCE 223
XXI. AMERICAN A FFAIRS 232
XXII. GAB 250
XXIII. THE VALUE OF A NAME 267
XXIV. HOW M. CLARETIE INDUCED ME TO WRITE THIS BOOK 273
 ILLUSTRATIONS

LOIE FULLER Frontispiece

PAGE
LOIE FULLER IN HER ORIGINAL SERPENTINE DRESS 29
THE DANCE OF FLAME 59
LOIE FULLER AND HER MOTHER 75
THE DANCE OF THE LILY 93
LOIE FULLER AND ALEXANDRE DUMAS 107
M. AND MME. CAMILLE FLAMMARION, AT JUVISY 115
THE DANCE OF FLOWERS 139
THE DANCE OF THE BUTTERFLY 143
DANCE TO GOUNOD’S “AVE MARIA” 159
THE DANCE OF THE BUTTERFLY 181
LOIE FULLER IN HER GARDEN AT PASSY 193
LOIE FULLER’S ROOM AT THE FOLIES-BERGÈRE 211
GAB 253
THE DANCE OF FIRE 261
THE DANCE OF FEAR FROM “SALOME” 283
 FIFTEEN YEARS OF A
DANCER’S LIFE

I
MY STAGE ENTRANCE

“W HOSE baby is this?”

“I don’t know.”
“Well, anyway, don’t leave it here. Take it away.”
Thereupon one of the two speakers seized the little thing and
brought it into the dancing-hall.
It was an odd little baggage, with long, black, curly hair, and it
weighed barely six pounds.
The two gentlemen went round the room and asked each lady if
the child were hers. None claimed it.
Meanwhile two women entered the room that served as dressing-
room and turned directly toward the bed where, as a last resort, the
baby had been put. One of them asked, just as a few minutes before
the man in the dancing-hall had asked:
“Whose child is this?”
The other woman replied:
“For Heaven’s sake what is it doing there? This is Lillie’s baby. It
is only six weeks old and she brought it here with her. This really is
no place for a baby of that age. Look out; you will break its neck if
you hold it that way. The child is only six weeks old, I tell you.”
At this moment a woman ran from the other end of the hall. She
uttered a cry and grasped the child. Blushing deeply she prepared to
take it away, when one of the dancers said to her:
 “She has made her entrance into society. Now she will have to
stay here.”
From that moment until the end of the ball the baby was the chief
attraction of the evening. She cooed, laughed, waved her little hands
and was passed round the hall until the last of the dancers was
gone.
I was that baby. Let me explain how such an adventure came
about.
It occurred in January, during a very severe winter. The
thermometer registered forty degrees below zero. At that time my
father, my mother, and my brothers lived on a farm about sixteen
miles from Chicago. When the occasion of my appearance in the
world was approaching, the temperature went so low that it was
impossible to heat our house properly. My mother’s health naturally
made my father anxious. He went accordingly to the village of
Fullersburg, the population of which was composed almost
exclusively of cousins and kinsmen, and made an arrangement with
the proprietor of the only public-house of the place. In the general
room there was a huge cast-iron stove. This was, in the whole
countryside, the only stove which seemed to give out an appreciable
heat. They transformed the bar into a sleeping-room and there it was
that I first saw light. On that day the frost was thick on the window
panes and the water froze in dishes two yards from the famous
stove.
I am positive of all these details, for I caught a cold at the very
moment of my birth, which I have never got rid of. On my father’s
side I had a sturdy ancestry. I therefore came into life with a certain
power of resistance, and if I have not been able to recover altogether
from the effects of this initial cold, I have had the strength at all
events to withstand them.
A month later we had returned to the farm, and the saloon
resumed its customary appearance. I have mentioned that it was the
only tavern in town, and, as we occupied the main room, we had
inflicted considerable hardship upon the villagers, who were deprived
of their entertainment for more than four weeks.
 When I was about six weeks old a lot of people stopped one
evening in front of our house. They were going to give a surprise
party at a house about twenty miles from ours.
They were picking up everybody en route, and they stopped at
our house to include my parents. They gave them five minutes in
which to get ready. My father was an intimate friend of the people
whom they were going to surprise; and, furthermore, as he was one
of the best musicians of the neighbourhood he could not get out of
going, as without him the company would have no chance of
dancing. He accordingly consented to join the party. Then they
insisted that my mother go, too.
“What will she do with the baby? Who will feed her?”
There was only one thing to do in these circumstances—take
baby too.
My mother declined at first, alleging that she had no time to make
the necessary preparations, but the jubilant crowd would accept no
refusal. They bundled me up in a coverlet and I was packed into a
sleigh, which bore me to the ball.
When we arrived they supposed that, like a well-brought-up baby,
I should sleep all night, and they put me on the bed in a room
temporarily transformed into a dressing-room. They covered me
carefully and left me to myself.
There it was that the two gentlemen quoted at the beginning of
this chapter discovered the baby agitating feet and hands in every
direction. Her only clothing was a yellow flannel garment and a calico
petticoat, which made her look like a poor little waif. You may
imagine my mother’s feelings when she saw her daughter make an
appearance in such a costume.
That at all events is how I made my debut, at the age of six
weeks. I made it because I could not do otherwise. In all my life
everything that I have done has had that one starting-point; I have
never been able to do anything else.
I have likewise continued not to bother much about my personal
appearance.
 II
MY APPEARANCE ON A REAL STAGE AT TWO
YEARS AND A HALF

W HEN I was a very small girl the president of the Chicago

Progressive Lyceum, where my parents and I went every
Sunday, called on my mother one afternoon, and
congratulated her on the appearance I had made the preceding
Sunday at the Lyceum. As my mother did not understand what he
meant, I raised myself from the carpet, on which I was playing with
some toys, and I explained:
“I forgot to tell you, mamma, that I recited my piece at the Lyceum
last Sunday.”
“Recited your piece?” repeated my mother. “What does she
mean?”
“What!” said the president, “haven’t you heard that Loie recited
some poetry last Sunday?”
My mother was quite overcome with surprise. I threw myself upon
her and fairly smothered her with kisses, saying,
“I forgot to tell you. I recited my piece.”
“Oh, yes,” said the president, “and she was a great success, too.”
My mother asked for explanations.
The president then told her: “During an interval between the
exercises, Loie climbed up on the platform, made a pretty bow as
she had seen orators do, and then, kneeling down, she recited a little
prayer. What this prayer was I don’t remember.”
But my mother interrupted him.
“Oh, I know. It is the prayer she says every evening when I put
her to bed.”
 And I had recited that in a Sunday School thronged by free-
thinkers!
“After that Loie arose, and saluted the audience once more. Then
immense difficulties arose. She did not dare to descend the steps in
the usual way. So she sat down and let herself slide from one step to
another until she reached the floor of the house. During this exercise
the whole hall laughed loudly at the sight of her little yellow flannel
petticoat, and her copper-toed boots beating the air. But Loie got on
her feet again, and, hearing the laughter, raised her right hand and
said in a shrill voice: ‘Hush! Keep quiet. I am going to recite my
poem.’ She would not stir until silence was restored. Loie then
recited her poem as she had promised, and returned to her seat with
the air of having done the most natural thing in the world.”
The following Sunday I went as usual to the Lyceum with my
brothers. My mother came, too, in the course of the afternoon, and
took her seat at the end of a settee among the invited guests who
took no part in our exercises. She was thinking how much she had
missed in not being there the preceding Sunday to witness my
“success,” when she saw a woman rise and approach the platform.
The woman began to read a little paper which she held in her hand.
After she had finished reading my mother heard her say:
“And now we are going to have the pleasure of hearing our little
friend Loie Fuller recite a poem entitled: ‘Mary had a little Lamb.’”
My mother, absolutely amazed, was unable to stir or to say a
word. She merely gasped:
“How can this little girl be so foolish! She will never be able to
recite that. She has only heard it once.”
In a sort of daze she saw me rise from my seat, slowly walk to
the steps and climb upon the platform, helping myself up with feet
and hands. Once there I turned around and took in my audience. I
made a pretty courtesy, and began in a voice which resounded
throughout the hall. I repeated the little poem in so serious a manner
that, despite the mistakes I must have made, the spirit of it was
intelligible and impressed the audience. I did not stop once. Then I
courtesied again and everybody applauded me wildly. I went back to
the stairs and let myself slide down to the bottom, as I had done the
preceding Sunday. Only this time no one made fun of me.
When my mother rejoined me, some time after, she was still pale
and trembling. She asked me why I had not informed her of what I
was going to do. I replied that I could not let her know about a thing
that I did not know myself.
“Where have you learned this?”
“I don’t know, mamma.”
She said then that I must have heard it read by my brother; and I
remembered that it was so. From this time on I was always reciting
poems, wherever I happened to be. I used to make little speeches,
but in prose, for I employed the words that were natural for me,
contenting myself with translating the spirit of the things that I recited
without bothering much over word-by-word renderings. With my firm
and very tenacious memory, I needed then only to hear a poem once
to recite it, from beginning to end, without making a single mistake. I
have always had a wonderful memory. I have proved it repeatedly by
unexpectedly taking parts of which I did not know a word the day
before the first performance.
It was thus, for instance, when I played the part of Marguerite
Gauthier in La Dame aux Camélias with only four hours to learn the
lines.
On the Sunday of which I have been speaking, my mother
experienced the first of the nervous shocks that might have warned
her, had she been able to understand, that she was destined to
become the prey of a dreadful disease, which would never leave her.
From the spring which followed my first appearance at the Folies-
Bergère until the time of her death she accompanied me in all my
travels. As I was writing this, some days before her end, I could hear
her stir or speak, for she was in the next room with two nurses
watching over her night and day. While I was working I would go to
her from time to time, rearrange her pillows a little, lift her, give her