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Genetic Management of Fragmented Animal and Plant Populations
Genetic Management
of Fragmented Animal
and Plant Populations
Richard Frankham
Macquarie University and Australian Museum, Sydney, NSW, Australia

Jonathan D. Ballou & Katherine Ralls

Smithsonian Institution, Washington, DC, USA

Mark D. B. Eldridge
Australian Museum, Sydney, NSW, Australia

Michele R. Dudash & Charles B. Fenster

South Dakota State University, Brookings, SD, USA

Robert C. Lacy
Chicago Zoological Society, Brookfield, IL, USA

Paul Sunnucks
Monash University, Clayton, Vic, Australia

Line drawings by Karina McInnes, Melbourne, Vic, Australia

1
3
Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
Oxford University Press is a department of the University of Oxford.
It furthers the University’s objective of excellence in research, scholarship,
and education by publishing worldwide. Oxford is a registered trade mark of
Oxford University Press in the UK and in certain other countries
© Richard Frankham, Jonathan D. Ballou, Katherine Ralls, Mark D. B. Eldridge, Michele
R. Dudash, Charles B. Fenster, Robert C. Lacy & Paul Sunnucks 2017
The moral rights of the authors have been asserted
First Edition published in 2017
Impression: 1
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and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University Press
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Library of Congress Control Number: 2017933536
ISBN 978–0–19–878339–8 (hbk.)
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Links to third party websites are provided by Oxford in good faith and
for information only. Oxford disclaims any responsibility for the materials
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Contents

Preface xi
Acknowledgments xv
List of Symbols xix

1 Introduction 1
Why should we conserve genetic diversity? 2
What genetic problems occur in fragmented populations? 5
Can we reverse the adverse effects of population fragmentation? 6
How important is genetic management of fragmented populations? 8
How should we genetically manage fragmented populations? 11
Summary 13
Further Reading 14

Section I Genetic problems in small isolated populations 15

2 Evolutionary genetics of small populations 17
Background 18
What factors control the evolution of small populations? 18
What genetic markers are used in conservation genetics? 26
How do we measure and describe genetic diversity? 27
How much standing genetic diversity do species and populations contain? 28
How do we measure population size? 31
Why is inbreeding important in conservation? 34
Why do we use computer simulations in conservation genetics? 37
Summary 39
Further Reading 39
Software 40

3 Inbreeding reduces reproductive fitness 41

Why is inbreeding an important issue in conservation? 42
Why is inbreeding harmful? 43
What factors affect the magnitude of inbreeding depression? 44
How large are the impacts of inbreeding on total fitness? 53
Does inbreeding increase extinction risks? 54
How do we detect inbreeding depression? 58
How do we quantify inbreeding depression? 58

v
Contents

What is mutational meltdown? 62

What are the relationships between harmful mutation, drift,
inbreeding, fixation, and fitness decline? 62
Can we reverse inbreeding depression and mutational accumulation? 63
Summary 64
Further Reading 64
Software 64

4 Loss of genetic diversity reduces ability to adapt 65

Why should we be concerned about conserving the ability
of species to adapt? 66
How common is evolutionary adaptation? 67
How rapidly does adaptation occur? 67
How large are adaptive evolutionary changes? 69
What determines the ability to undergo adaptive evolution? 71
How can we measure evolutionary potential? 84
Can we restore the ability to evolve? 85
Summary 85
Further Reading 86
Software 86

5 Population fragmentation causes inadequate gene flow and increases

extinction risk 87
Why is population fragmentation important in conservation? 88
How frequently are there problems due to inadequate gene flow? 91
What are the genetic consequences of completely isolated fragments? 91
What are the consequences of gene flow among fragments? 97
What are the consequences of different fragmented population
structures? 100
Complex relationships among variables in real fragmented populations 104
How do we measure genetic differentiation among populations? 106
Summary 111
Further Reading 112
Software 112

Section II Rescue and risk 113

6 Genetic rescue by augmenting gene flow 115
What are the problems of genetic erosion in small isolated
population fragments? 116
Can we reverse inbreeding and loss of genetic diversity? 117
Does crossing have beneficial or harmful effects on fitness? 118
Are immigrant alleles at a selective advantage? 131

vi
 Contents

How large and consistent are evolutionary rescue effects? 132

Summary 134
Further Reading 134

7 Outbreeding depression is uncommon and predictable 135

Why are we concerned about harmful crosses? 136
What mechanisms generate outbreeding depression? 140
How many generations does it take to develop outbreeding depression? 149
Can we predict the risk of outbreeding depression? 150
At what generation after crossing is outbreeding depression evident? 153
Can populations recover from outbreeding depression? 154
Summary 154
Further Reading 155

8 Modified rescue and risk expectations for species with diverse mating
systems and modes of inheritance 156
Why do we need to consider diverse mating systems and modes
of inheritance? 157
How are genetic rescue and risk expectations modified for different
mating systems? 158
How can we identify mating systems? 166
How are genetic rescue and risk expectations modified for different
modes of inheritance? 169
How do we identify modes of inheritance? 175
Summary 176
Further Reading 177
Software 178

Section III Developing management decisions 179

9 Is the taxonomy appropriate? Delineating species for conservation
purposes 181
What is the role of species delineation in conservation? 182
What is our objective? 182
How do we characterize population genetic differentiation in taxonomy? 182
What problems occur with species delineations? 184
Which species definitions are appropriate for conservation purposes? 196
How do we decide whether taxonomic revision is required? 200
How should a taxonomic re-evaluation be conducted if required? 201
Conclusions 201
Summary 201
Further Reading 202
Software 202

vii
Contents

10 Determining the number and location of genetically differentiated

population fragments 203
What are our objectives? 204
How do we identify the number of genetically differentiated
populations? 205
How do we locate genetically differentiated populations in the
landscape? 209
Are current units within species suitable for genetic management
purposes? 211
What is the current recommendation when genetically differentiated
populations are identified? 214
How should we proceed when there are genetically diverged
populations? 216
How can we measure gene flow among population fragments? 217
Summary 221
Further Reading 222
Software 222

11 Are there populations suffering genetic erosion that would benefit

from augmented gene flow? 223
What are our objectives? 224
Are there guidelines for genetic rescues? 226
What questions do we need to answer? 229
Are any populations suffering genetic erosion? 229
Is there another population to which it can be crossed? 233
Will crossing result in outbreeding depression? 233
Will crossing result in worthwhile genetic rescue? 239
How do we cope with simultaneous harmful and beneficial effects
of crossing? 243
Summary 243
Further Reading 243
Software 244

12 Managing gene flow among isolated population fragments.

I. Limited information 245
Why do we need to manage gene flow? 246
How can we augment gene flow? 248
What is the objective of genetic management for isolated
population fragments? 249
Genetic management with different amounts of information 250
What can be done with little or no genetic information? 251
Genetic swamping 263
Summary 264
Further Reading 265
Software 265

viii
 Contents

13 Managing gene flow among isolated population fragments.

II. Management based on kinship 266
How does detailed genetic information aid management of gene flow? 267
What population genetic parameter should we manage? 270
How do we manage using mean kinship? 272
Population mean kinship 277
Kinships at the individual level 281
Managing gene flow at the individual level 283
When to cease? 287
What genetic monitoring is needed? 287
Integrating genetic rescue with other management considerations 288
Summary 288
Further Reading 289
Software 289

14 Global climate change increases the need for genetic management 291
What is the problem? 292
Why does global climate change increase the need for genetic
management? 294
How have species responded to global climate change? 296
What genetic management options do we have for populations that
cannot move or adapt sufficiently? 298
How do we implement genetic management to assist adaptation? 300
How do we decide what populations and species need translocation
to avoid extinction? 303
How should we genetically manage translocations to cope with
climate change? 304
Summary 310
Further Reading 311
Software 311

15 Take home messages 312

Final messages for managers of wild animal and plant populations 313

Glossary 315
References 327
Index 391

APPENDICES (available online at www.oup.co.uk/companion/frankham) A1

1 Genetic markers for conservation purposes A1
2 VORTEX simulation software for population viability analysis A7
3 How should a taxonomic re-evaluation be conducted if required? A12

ix
Preface

Importance of the topic

One of the greatest unmet issues in conservation biology is the genetic management of
fragmented animal and plant populations. Many species across the planet have frag-
mented distributions with some small isolated populations that are potentially suffering
from inbreeding, loss of genetic diversity, and elevated extinction risk (genetic erosion).
Fortunately, these effects can usually be remedied by augmenting gene flow (crossing
populations within species), but this is rarely conducted. Disturbingly, evidence of any
genetic differentiation among populations typically leads managers to conclude that the
populations should be kept isolated, thereby dooming many to eventual extinction. We
are particularly concerned with populations fragmented by human actions in the last
500 years, encompassing most of the impacts arising from a 13-fold increase in the
human population.
A paradigm shift is urgently required where the existence of genetic differentiation
among populations acts as a trigger to ask if any populations are suffering from genetic
problems. If so, are there populations to which they can be crossed to reduce inbreeding
and augment genetic diversity, and will this gene flow be beneficial or harmful?
Whether to maintain genetic isolation is a major issue for managers of wild animal
and plant populations. It is critical that managers, and those who advise them, become
aware of the issues relating to restoration of genetic diversity and contribute to the
reversal of genetic deterioration associated with small population size.

Genesis of this book

The events that triggered this book began in February 2007 during a book writing
session on the 2nd edition of Introduction to Conservation Genetics involving Jon Ballou
and Dick Frankham. When revising material on population fragmentation, we were
deeply concerned about over-emphasis in the literature on the risks of outbreeding
depression (where crossing between populations is harmful), while the benefits of
crossing between populations within species to rescue small inbred populations were
largely neglected. We subsequently outlined means to predict the risk of outbreeding
depression (thus allowing the benefits of genetic rescue to be utilized widely), recruited
Mark Eldridge, Kathy Ralls, Bob Lacy, Michele Dudash, and Charlie Fenster as collab-
orators, and subsequently published the Frankham et al. (2011) paper on the topic. That
work influenced a paper on the use of genetic translocations to cope with global climate
change (Weeks et al. 2011) co-authored by Mark Eldridge, Dick Frankham, and Paul

xi
Preface

Sunnucks among others. Our paper on implications of different species concepts for
biodiversity conservation (Frankham et al. 2012) was provoked by practical concerns
that Ingrid Porton of St Louis Zoo had about genetic management of ruffed lemurs in
light of taxonomic revisions and competing taxonomies, and also drew upon our work
on predicting outbreeding depression.
This book arose as an extension of the above studies and our previous work on
inbreeding depression, evolutionary potential, outbreeding depression, genetic rescue,
and genetic management of fragmented populations, plus our extensive involvement
in threatened species recovery. While drafting Chapter 6 on genetic rescue, Dick
Frankham identified the lack of a quantitative overview and this provoked him to
complete two meta-analyses that inform the book chapter (Frankham 2015, 2016), as
did the content of a symposium on the topic at the 2014 meeting of the Society for
Conservation Biology in Missoula, Montana.
The material on managing gene flow among isolated population fragments (optimiz-
ing gene flow) was initiated largely as an extension of the seminal Ballou & Lacy (1995)
work on genetic management by minimizing mean kinship—the recommended man-
agement strategy for threatened species in captivity. The content of this book was also
informed by a symposium on genetic management of fragmented populations at the
2011 meeting of the Society for Conservation Biology in New Zealand, organized by
authors of this book.

Purpose
This book provides the conceptual background for understanding genetic issues in the
management of fragmented wild animal and plant populations, and offers means for
making practical decisions on genetic management of population fragments. We focus
particularly on the harmful consequences of mating among relatives on offspring fitness
(inbreeding depression) that is universal among sexually reproducing non-haploid
organisms. A second major focus is on the maintenance of genetic diversity, so that
species and populations can continue to adapt to changing environments. We also
define the criteria that must be satisfied by definitions of species for conservation
purposes, and make recommendations regarding definitions that are appropriate and
inappropriate. Our goal of preserving endangered species through genetic management
can provide additional benefits beyond single species preservation, because species
interact within communities that are subject to abiotic and biotic environments, con-
tributing to ecosystem function and overall global biodiversity.

Intended audience
This book is designed for senior undergraduate and graduate students and professional
scientists in conservation and wildlife biology, and evolutionary ecology, as well as for
managers of wild animal and plant populations. We will subsequently write a shorter,
simpler Practical Guide for managers who are less familiar with conservation genetics.

xii
 Preface

To make the book accessible to a wide audience, we have assumed knowledge only of
introductory genetics and basic statistics. For the same reason, we have restricted use of
mathematics to simple algebra, even though genetic management requires a quantita-
tive approach. Readers wishing for a simple concise introduction to conservation
genetics are referred to A Primer of Conservation Genetics by Frankham et al. (2004)
or the Italian, Portuguese, or Korean translations of it.

Précis of contents
The first eight chapters of the book consist of background material and concepts, the
subsequent six chapters are concerned with genetic management (including objectives,
questions, and decision trees), and the final chapter summarizes the messages from the
prior chapters. Genetic issues in single populations are addressed prior to the more
complex issues of multiple populations.
Chapter 1 provides a general outline of the issues involved in genetic management of
fragmented animal and plant populations and its importance.
Section I provides detailed background on genetic problems in fragmented popula-
tions. This begins with a brief introduction to the evolutionary genetics of small
populations (Chapter 2) where the roles of mutation, migration, selection, and chance
(genetic drift) are described, inbreeding and its measurement considered, and popula-
tion size in the evolutionary genetics context (effective population size) defined.
Threats to small isolated populations from inbreeding depression and mutational
accumulation are covered in Chapter 3, and those from reduced ability to adapt
genetically in Chapter 4. We next address genetic problems caused by inadequate
gene flow in fragmented populations (Chapter 5).
Section II addresses risk/benefit analyses of remedies for genetic problems in frag-
mented populations. Genetic rescue by augmenting gene flow is addressed in Chapter 6.
Chapter 7 delineates the causes of outbreeding depression and considers the potential of
populations to recover from outbreeding depression through natural selection.
Chapter 8 evaluates how expectations for remedies and risk from crossing populations
need to be modified for species with diverse mating systems and modes of inheritance
(self-incompatible, self-fertilizing, asexual, polypoid, haploid, haplodiploid, etc.).
Section III of the book deals with genetic management of fragmented populations.
Chapter 9 defines the criteria that must be satisfied by a species definition used for
conservation purposes, provides recommendations on definitions that are and are not
appropriate, identifies the many pitfalls in species delineation, and recommends rem-
edies. Chapter 10 covers methods for delineating the number and locations of popula-
tion fragments. Chapter 11 presents practical guidelines (including decision trees) for
determining whether populations are suffering genetic erosion and whether they would
benefit from augmented gene flow. Chapters 12 and 13 address management of gene
flow among isolated population fragments, the former chapter based on limited infor-
mation, and the latter on extensive genetic information (using mean kinship).
Chapter 14 explores genetic management under the moving target created by global

xiii
Preface

climate change. Finally, Chapter 15 draws the prior threads together to present the Take
home messages from the book.

Format
To make the books easy to read and follow, we use a textbook format similar to that of
the Frankham, Ballou, and Briscoe textbooks Introduction to Conservation Genetics
(2nd edn, 2010) and A Primer of Conservation Genetics (2004). In particular, we use:

• main messages boxes at the start of each chapter

• list of terms for each chapter
• frequent headings and sub-headings
• small main point boxes and dot point lists throughout chapters
• ample use of figures, maps, and tables
• boxes with case studies
• example boxes
• decision trees
• line drawings by Karina McInnes
• links to software for genetic analysis and management
• List of symbols
• Glossary
• Appendices
• References and Index.

We use questions for most of the major headings within chapters, as science seeks to
answer questions. This approach was successfully trialed in undergraduate teaching,
presentations to professional scientists, and for communicating genetic concepts to
conservation practitioners.

References
Given length constraints and issues of readability, we restrict referencing to examples
and a few recent or key citations (especially meta-analyses and reviews) to allow access
to the relevant literature, except for contentious issues where we present more extensive
referencing.

xiv
Acknowledgments

We thank our partners Annette Lindsay (RF), Vanessa Ballou (JDB), Robert L. Brownell
Jr. (KR), Anne Baker (RCL), and Andrea Taylor (PS) for their forbearance and support
during the preparation of this book.
We are grateful for the support of our home institutions Macquarie University (RF),
the Australian Museum (ME and RF), the Smithsonian Institution (JDB and KR),
University of Maryland and South Dakota State University (MRD and CBF), Chicago
Zoological Society (RCL), and Monash University (PS). The Smithsonian’s National
Zoological Park and the University of Maryland kindly hosted book writing sessions
and RF thanks Jon and Vanessa Ballou and Michele Dudash and Charlie Fenster for
their hospitality during several of these sessions.
The authors thank the following for financial research support for projects relevant to
the development of ideas presented in the book during the writing period 2011–2016, as
follows:
M. Dudash: her work on inbreeding depression and its genetic basis and how mating
system evolution may be impacted was funded through the US National Science
Foundation and USDA (USA).
M. Eldridge: his group’s work has been supported by the Australian Research Council,
the Hermon Slade Foundation, the Foundation for National Parks and Wildlife, and the
Holsworth Wildlife Research Endowment.
C. Fenster: his work on the genetic and ecological consequences of population frag-
mentation has been supported by the US National Science Foundation and DOE (USA),
while that on mating system evolution and its genetic consequences has been backed by
grants from NSERC (Canada) and NFR (Norway).
R. Lacy: his work on population management has been supported by various grants
from the US National Science Foundation, the Institute of Museum and Library
Services, and the Association of Zoos and Aquariums. Much of his work to apply
genetic management to species conservation arose from collaborations within the
IUCN (International Union for Conservation of Nature) Species Survival Commission,
and its Conservation Breeding Specialist Group.
P. Sunnucks: his group’s work has been supported by diverse sources including the
Australian Research Council and the Holsworth Wildlife Research Endowment.
We thank Karina McInnes for preparing the excellent line drawings. Harry Crosby
skilfully revised figures that were based on published material and Jon Ballou prepared
the new figures in the book and revised many others.

xv
Acknowledgments

We thank M. Culver for providing us with the raw data on puma genotypes used for
the calculations in Box 13.2, J. Deitz, T. King, J. Mickelberg, and I. Porton for supplying
information, I. Porton for checking Box 9.3, and T. King for checking Box 11.2, and to
N. Hancock and P. Mahoney for commenting on Chapter 14. We thank J. Howard and
B. Pukazhenthi for supplying material for illustrations.
The staff of Oxford University Press and their outsourced production company
turned our material into this book. At Oxford University Press, we thank Ian Sherman
for championing our book proposal and for overall supervision of the project, Lucy
Nash for preparing the book contract, and Bethany Kershaw for her efficient and
cheerful work as commissioning editor. The CEO Nigel Portwood and Helen Tyas
facilitated improvements in production, liaising with the team at SPi Global led by
Cheryl Brant. We thank Cheryl and her team for the effort they put into the production
of the book and for accommodating our numerous suggestions and revisions. We are
grateful to Julie Musk for the outstanding job she did proofreading the book.

Copyright acknowledgments
We thank Cambridge University Press for permission to reproduce the following
figures: 2.3, 2.4, 2.6, 2.7, 3.1, 3.2, 3.3, 3.4, 3.8, 3.9, 3.10, 3.11, 4.1, 4.2, 4.3, 4.4, 4.5, 4.6,
5.2, 5.3, 5.4, 5.5, 5.6, 5.8, 5.10, 5.11, 5.12, 7.2, and A1, and figures within Box 2.3,
Example 2.2, Box 3.1, Chapter 4 frontispiece, Box 4.1, Box 4.2, Box 4.4, Chapter 5
frontispiece, Chapter 6 frontispiece, Box 6.3 (two), Box 8.2, Box 2.4, Chapter 9 frontis-
piece, Box 10.1, and Box 14.3 which first appeared in Frankham, Ballou, & Briscoe
(2010) Introduction to Conservation Genetics, 2nd edition, and Fig. 13.3 from the first
edition (Frankham et al. 2002).

We are grateful to the following for kind permission to reproduce copyright material:

Chapter 1 frontispiece reprinted from Oedekoven (1960), Page 184, Copyright (1960),
with permission from IOS Press; figures in Box 1.1 from figs 1 and 3 in Zhang et al. 2007
by permission of Oxford University Press; Chapter 3 frontispiece from fig 3 in Szulkin
et al. 2007, © 2014 The Authors, Journal of Evolutionary Biology © 2014 European
Society for Evolutionary Biology; Chapter 3 frontispiece (c) republished with permis-
sion of John Wiley and Sons Inc., from fig 2 in Nason and Ellstrand 1995; permission
conveyed through Copyright Clearance Center, Inc.; Fig 3.5 from fig 1a in Losdat et al.
2014, © 2014 The Authors, Journal of Evolutionary Biology © 2014 European Society
for Evolutionary Biology; Fig 3.6 from fig 4 in Szulkin et al. 2007, © 2014 The Authors,
Journal of Evolutionary Biology © 2014 European Society for Evolutionary Biology; Fig
4.2 republished with permission of John Wiley and Sons Inc., from fig 11.3 in Weber
2004, permission conveyed through Copyright Clearance Center, Inc.; Fig 4.4 reprinted
from fig 3 in Montgomery et al. 2010, Copyright (2010), with permission from Elsevier;
Chapter 5 frontispiece from fig 2 in James 1995; Fig 5.1 from fig 1 in Jump and Peñuelas
2006, Copyright (2006) National Academy of Sciences, U.S.A.; Fig 5.7 from fig 5B in
Jousimo et al. 2014, reprinted with permission from AAAS, permission conveyed
through Copyright Clearance Center, Inc.; Fig 5.9 from fig 5 in Aguilar et al. 2008,

xvi
 Acknowledgments

© 2008 Blackwell Publishing Ltd.; figures in Box 5.2 from fig 1 and 3 in Phillipsen et al.
2015, © 2014 John Wiley & Sons Ltd.; Fig 5.12 from fig 4 in Forbes and Hogg 1999,
Copyright © 1999–2016 John Wiley & Sons, Inc. All Rights Reserved; Box 5.3 from figs
1 and 3 in Major et al. 2014, © 2014 The Zoological Society of London; Fig 6.1a
republished with permission of John Wiley & Sons Inc., from fig 2 in Newman and
Tallmon 2001, permission conveyed through Copyright Clearance Center, Inc.; Fig 6.2
from fig 2 in Frankham 2015, Copyright © 1999–2016 John Wiley & Sons, Inc.; Chapter
7 frontispiece from fig 3 in Yang et al. 2004, Copyright © 2004, Kluwer Academic
Publishers; Fig 7.1 republished with permission of Oxford University Press—Journals,
from figs 3a and b in Murray and Young 2001, permission conveyed through Copyright
Clearance Center, Inc.; Fig 7.3 reprinted from fig 1 in Trends in Ecology & Evolution, 27
(4), Cutter 2012, with permission from Elsevier; Box 8.2 from fig 1 in Young and Brown
1999, © Cambridge University Press; Fig 8.2 republished with permission of John Wiley
& Sons Inc., from fig 1 in Jarne and Auld 2006, permission conveyed through
Copyright Clearance Center, Inc.; Fig 8.3 from fig 1 in Bousalem et al. 2010, © 2008
Canadian Science Publishing or its licensors, reproduced with permission; Fig 9.3
republished with permission of John Wiley & Sons Inc., from figs 1 and 3 in Bouzat
et al. 1998, permission conveyed through Copyright Clearance Center, Inc.; Box 9.1
reproduced from fig 1 from Eldridge and Close 1992, with permission from CSIRO
Publishing; Box 9.4 from fig 1 in Degnan 1993, Copyright © 1999–2016 John Wiley &
Sons, Inc.; Chapter 10 frontispiece from figs 1 and 3 in Bergl and Vigilant 2007,
Copyright © 1999–2016 John Wiley & Sons, Inc.; Box 10.2 republished with permission
of John Wiley & Sons Inc., from figs 1, 4, and 5 from Coleman et al. 2013, © 2013
Blackwell Publishing Ltd, permission conveyed through Copyright Clearance Center,
Inc.; Fig 10.3 republished with permission of Genetics Society of America, from fig 1a in
Gao et al. 2007, permission conveyed through Copyright Clearance Center, Inc.; Fig
10.4 from fig 1 in Coulon et al. 2008, © 2008 Blackwell Publishing Ltd.; Fig 10.5 from figs
1a and 1c from Bittner and King 2003, Copyright © 1999–2016 John Wiley & Sons, Inc.;
Fig 10.6 from fig 1 in Eldridge et al. 2001, Copyright © 1999–2016 John Wiley & Sons,
Inc.; Box 11.4 republished with permission of Oxford University Press, from fig 3B in
Stephen et al. 2005, permission conveyed through Copyright Clearance Center, Inc.;
Chapter 11 frontispiece from fig 1 in Dolan et al. 1999, Copyright © 2016 by American
Journal of Botany, Inc.; Chapter 11 frontispiece histogram from fig 2 in Oakley and
Winn 2012, © 2012 The Authors, New Phytologist © 2012 New Phytologist Trust; Box
11.1 image of kinked tail reprinted from Roelke et al. 1993, Copyright 1993, with
permission from Elsevier; Example 13.3 from fig 1 in Finger et al. 2011, © 2008
Blackwell Publishing Ltd.; Box 14.1 figure republished with permission of John Wiley
& Sons Inc., from figs 1 and 2 in Miller-Rushing and Primack 2008, permission
conveyed through Copyright Clearance Center, Inc.

xvii
List of Symbols

The following table contains the list of symbols used in the book, along with their
meaning and the chapters where they are mainly considered. Where the same symbol is
used for more than one purpose in the published literature, we have added an additional
letter to avoid ambiguity. Different symbols for the same variable are sometimes used in
diverse publications.

Symbol Definition Chapter

a Arbitrary genotypic value assigned to the effect on the 3, 4

phenotype of genotype A1A1, where genotype A2A2 has a
value of –a, and the two homozygotes differ in mean by 2a
A Allelic diversity, mean number of alleles per locus 2
B Haploid lethal equivalents 3, 11
C A parameter used in analyses of the impact of inbreeding 3
on fitness, where e–C is the survival of an outbred
population (often referred to as A, but we use that for
another purpose)
d Dominance deviation for the effect of the heterozygote on 3, 4
a quantitative trait: deviation of heterozygote effect from
the mean of the two homozygotes
DJ Jost’s measure of population differentiation 5
DM Absolute genetic differentiation between population 13
fragments, also referred to as the average minimum
genetic distance
DN Nei’s genetic distance 5, 13
DST Average genetic diversity between population fragments, 13
the numerator for GST
E Proportion of adaptation that is due to non-shared 7
features of the environments of two populations
ER Evolutionary rescue, ratio of evolutionary potential in a 6
crossed population compared to that in its inbred parents

xix
List of Symbols

Symbol Definition Chapter

f Relative frequency 2, 11, 12

F Inbreeding coefficient of individuals, or the mean for a 2, 3, 5, 6, 8,
population 11, 12
FIS Inbreeding within population fragments (sub- 5, 11
populations)
FIT Total inbreeding over a number of population fragments 5
FST Inbreeding due to population differentiation 5, 10, 11, 13
GA Genetic adaptation over a generation, where ΣGAt is 4, 7
cumulative genetic adaptation over t generations
GST Analogue of FST for multiallelic loci 5
0
G ST GST corrected so that it scales 0–1 (GST divided by GSTmax) 5, 11, 13
GSTmax Maximum value of GST for a particular locus (or set of 5
loci) with particular allele frequencies
H Heterozygosity (see also Ĥ, He, Ho, HS, and HT) 2, 4, 5, 6, 11
Ĥ Equilibrium heterozygosity, for example due to mutation- 8
drift equilibrium
h2 Heritability of a quantitative character 4, 6, 11, 14
ht Haplotype diversity 2
He Hardy–Weinberg equilibrium heterozygosity 2, 5, 8, 11
Ho Observed heterozygosity 2, 8, 11
hs Dominance effects of an allele on fitness, where the two 2, 8
homozygotes have fitnesses of 1 and 1 – s, respectively,
and the heterozygote 1 – hs. Thus h = 0 is dominant, 1 is
recessive, and 0.5 additive, etc.
HS Expected heterozygosities averaged across population 5
fragments
HT Expected heterozygosity across the totality of a group of 5
population fragments
I Normalized identity of genes between two populations 13
ID Inbreeding depression, difference in mean between 3
outbred and inbred offspring for a quantitative trait (e.g.
fitness)
JXY Gene identity between population fragments X and Y 5, 13
(Nei 1987), a measure of population similarity, related
to mkAB
k Number of items (e.g. population fragments, 5, 10, 13
combinations, etc.)

xx
 List of Symbols

kij Kinship (or coancestry) between two individuals: the 5, 13

probability that two randomly chosen alleles at a locus,
one from each individual, are identical by descent
KST Analogue of FST for DNA sequence data, a measure of 5
population differentiation
ln Natural logarithm, loge 3
m Migration rate, proportion of migrant alleles introduced 5, 10, 12
over a single generation, where Nm is the number of
migrants

mk Average mean kinship of individuals in a population 13
mkAB Mean kinships between populations A and B, average of 5, 13
pairwise kinships of all individuals in population A with
all in population B
mki Mean kinship of individual i, average of its kinships with 5, 13
all individuals in the population
MU Contribution to genetic adaptation from selection acting 7
on new mutations
n Sample size 2, 6
N Census population size (usually potentially reproducing 2, 8, 9, 10, 12,
adults) 14
nA Effective number of alleles 2, 5
Ne Effective population size. This determines the genetic 2, 3, 4, 5, 8, 9,
impacts of small population size 11, 14
p Frequency of allele A1 (or A) 2, 3, 4, 5, 8,
12, 13
PD Contribution to genetic adaptation from selection 7
operating on initial (pre-existing) genetic diversity
P(E) Probability of extinction of a population over a defined 10
number of generations
q Frequency of allele A2 (or a) 2, 3, 4, 5, 8, 12
q̂ Equilibrium frequency of allele A2 (a), as for example 2, 8
from mutation-selection equilibrium
QST Proportion of variation for a quantitative character due to 5
differences among population fragments (quantitative
trait analogue of FST)
r Correlation coefficient 8, 10
re Relatedness 13

xxi
List of Symbols

Symbol Definition Chapter

RST Analogue of FST that measures the proportion of 5

differences in length of microsatellite alleles among
population fragments (to incorporate evolutionary
relationships among alleles)
s Selection coefficient against a genotype at a locus 2, 8, 12
S Probability of survival 3
Sd Selection differential for a quantitative trait 4, 6
Sf Selfing rate 8
SP Number of polymorphic sites at a locus in a segment of 2
DNA
t Number of generations 3, 4, 5, 9, 11
T Outcrossing rate (1 – selfing rate) 8
u Mutation rate per generation for a locus, or nucleotide 2, 8
VA Additive genetic variation for a quantitative character 4, 11
Vm Increase per generation in additive genetic variation due 4, 11
to mutation
VP Phenotypic (total) variance for a quantitative character 4
W Mean fitness of a population or group 3
δ Measure of inbreeding depression: proportionate decrease 3
in trait mean due to inbreeding, compared to outbreeding
(usually from the impact of selfing)
Δ Change in value, usually from one generation to the next, 3, 4, 8
e.g. ΔF, Δq
ΔF Differences in inbreeding coefficient between individuals 3, 6, 8, 11
in different generations
ΔFm Differences in inbreeding coefficient between mothers of 6, 11
individuals in different generations
ΔFz Differences in inbreeding coefficient between zygotes in 6, 11
different generations
ΔGR Genetic rescue effect, measured as the proportionate 6
benefit in fitness of a crossed population compared to the
inbred parent
Δq Change in allele frequency over one generation 8, 12
ΔW Change in fitness, for example due to inbreeding 11
π Nucleotide diversity 2
Π Product of the quantities specified 12

xxii
 List of Symbols

σp2 and Variance in allele frequencies among population 2, 5

σq2 fragments for the A1 and A2 alleles, respectively. With
only two alleles at a locus, the quantities are equal
σP Phenotypic standard deviation for a quantitative character 4
(square root of VP)
Σ Sum of the following terms 2, 3, 4, 13
ΦST Analogue of FST that incorporates genealogies 5

xxiii
 Introduction CHAPTER 1

TERMS
Genetic management of fragmented populations is one of the major, largely
unaddressed issues in biodiversity conservation. Many species across the planet Critically endangered, endangered,
have fragmented distributions with small isolated populations that are potentially genetic diversity, genetic erosion,
suffering from inbreeding and loss of genetic diversity (genetic erosion), leading to genetic rescue, inbreeding, inbreeding
elevated extinction risk. Fortunately, genetic deterioration can usually be remedied depression, outbreeding depression,
by augmenting gene flow (crossing between populations within species), yet this is population, population fragmentation,
rarely done, in part because crossing is sometimes harmful (but it is possible to threatened
predict when this will occur). Benefits and risks of genetic problems are sometimes
altered in species with diverse mating systems and modes of inheritance. Adequate
genetic management depends on appropriate delineation of species. We address
management of gene flow between previously isolated populations and genetic
management under global climate change.

Brazil

1500 1845
82% 80%

1907 1952
58% 18%

Loss and fragmentation of the Atlantic

coastal forest of Brazil from 1500 to 2000
(from Oedekoven 1980) indicating that the
distributions of many species have
1973 2000 becoming increasingly fragmented. The
8% 3% percentages are extent of forest cover.

Genetic Management of Fragmented Animal and Plant Populations. R. Frankham, J. D. Ballou, K. Ralls, M. D. B. Eldridge, M. R. Dudash,
C. B. Fenster, R. C. Lacy & P. Sunnucks. Oxford University Press (2017) © R. Frankham, J. D. Ballou, K. Ralls, M. D. B. Eldridge,
M. R. Dudash, C. B. Fenster, R. C. Lacy & P. Sunnucks 2017. DOI 10.1093/oso/9780198783398.001.0001
Genetic Management of Fragmented Animal and Plant Populations

Why should we conserve genetic diversity?

.........................................................................................
Conservation of genetic diversity is one of three levels of biodiversity recommended by
the IUCN for conservation, but there has been limited activity to implement this
recommendation for wild populations
.........................................................................................
IUCN (International Union for Conservation of Nature), the premier international
conservation agency, recommends conservation of biodiversity at three levels
(McNeely et al. 1990):

• genetic diversity
• species diversity
• ecosystem diversity.

Genetics is involved in all three levels. First, genetic diversity is worthy of conservation
in its own right, because it controls the variety of life—the shape, size, color, biochem-
istry, behavior, and all other characters that define species and underpin their place in
the world. Second, adequate genetic diversity favors species persistence by defending
against harmful effects of inbreeding, maintaining fitness, and providing the ability to
evolve. Through these influences on species diversity, genetic diversity enhances eco-
system viability (Bangert et al. 2005; Lankau & Strauss 2007). More directly, genetic
diversity within species can have profound effects on the species composition of
ecological communities and higher-order ecological processes, such as nutrient cycling
(Reusch et al. 2005; Whitham et al. 2006; Hughes et al. 2008). Conservation at the
species level has received most attention, while ecosystem conservation is now receiving
increasing attention.
Genetics has received little attention in the management of threatened species in
natural habitats, despite genetic diversity being protected under legislation, such as the
United Nations Convention on Biological Diversity (UNEP 1992) and national laws in
many countries (Moyle et al. 2003; Laikre et al. 2009). For example, genetic threats to
persistence and recovery are mentioned in 63% of endangered species recovery plans in
the USA, 55% in Australian plans, and 33% in European plans (Pierson et al. 2016).
Inbreeding was commented upon only in ~7% of plans and even fewer discussed
inbreeding depression. Few plans mentioned active genetic management. Examples of
the consequences of ignoring genetic issues and poor handling of them are common
(Frankham et al. 2010). In contrast, genetics has played a large role in the management
of captive populations over the past 30 years (Ballou et al. 2010).
Efforts have recently been made to incorporate genetic considerations into the
management of wild populations, such as actions in Sweden (Laikre et al. 2008), a
Summit on Conservation of Australia’s Genetic Diversity in 2012, and creation of the
Conservation Genetics Specialist Group within the IUCN Species Survival Commission.
This book is designed to aid in this undertaking by addressing arguably the most
important issue in the conservation of genetic biodiversity, the genetic management
of fragmented animal and plant populations (Frankham 2010a, 2010b).

2
 1 Introduction

Ubiquity of fragmented populations

.........................................................................................
Most species on the planet have fragmented distributions, with some population frag-
ments genetically isolated and suffering genetic problems
.........................................................................................
Many species on the planet have fragmented distributions, much of it due to human
conversion of natural habitats over the last few hundred years. This includes many
threatened animal and plant species listed in the IUCN Red List (IUCN 2016). Many
isolated populations have reduced genetic diversity relative to historic levels (Tallmon
et al. 2004; Aguilar et al. 2008). For example, 26% of invertebrate, 29% of vertebrate, and
55% of plant species show clear evidence of inadequate gene flow among populations
(Frankham et al. 2014a).
Harmful effects of inbreeding (inbreeding depression) and loss of genetic diversity
will ultimately contribute to the extirpation of many small populations of sexually
reproducing organisms (Fenster & Dudash 1994; Moritz 1999; Frankham et al. 2010)
(see later, and Chapters 3 and 4). Box 1.1 illustrates the fragmented distribution of the
giant panda (Ailuropoda melanoleuca) and some of its genetic consequences.

Box 1.1 Fragmented distribution of the giant panda in China and its genetic
consequences
(after Lu et al. 2001; Zhang et al. 2007; Hu et al. 2010; Zhu et al. 2011)

Qinling
Giant panda habitat Xian
Mountains
N

W E

S
RUSSIA

Qionglai Minshan
Mountains Mountains
CHINA
Chengdu

INDIA
Lesser Yangtze River
Xiangling
Mountains
Liangshan
Mountains Giant panda (China)
0 100 200 km

Giant pandas in China have a geographically fragmented distribution (left map).

Genetic analyses have shown that the most geographically isolated populations in the
Qinling and Liangshan Mountains have lower genetic diversity than other larger, less
physically isolated populations, such as those in the Minshan and Qionglai Moun-
tains, based on data for 10 microsatellite loci (see Box Table below). Overall, the giant
panda has levels of genetic diversity that are similar to other bears (Zhang et al. 2007).

3
Genetic Management of Fragmented Animal and Plant Populations

Population Heterozygosity Allelic richness

Qinling (QIN) 0.486 2.58

Minshan (MIN) 0.559 3.03
Qionglai (QIO) 0.610 3.18
Liangshan (LIS) 0.366 2.30
Lesser Xiangling (XL) 0.635 3.29

The populations fall into four different genetic clusters, QIN, MIN, QIO, and LIS
plus XL as shown in the STRUCTURE plot below (Zhang et al. 2007). Bars
represent the genotypes of individual pandas. Individuals with genotypes typical
of populations other than their own are immigrants, while those with genotypes
intermediate between populations indicate gene flow. The pandas seem able to
migrate considerable distances, even across some areas that are not their primary
habitat. Consequently, there is some gene flow between the populations, but very
little into Qinling.

QIN MIN QIO LIS XL

As the term “population” is used widely in this book, we pause to define it in a

conservation genetics context.

What is a population?
.........................................................................................
A population is a group of individuals living in sufficiently close proximity that any
member of the group can potentially reproduce with any member of the other sex
.........................................................................................
There are at least 16 definitions of population (Waples & Gaggiotti 2006), but we follow
the evolutionary definition, as given above, as it is most appropriate for the issues in
this book.
There is a continuum between completely isolated populations and completely
connected ones (random mating), depending on levels of gene flow and divergence
time in generations (Fig. 1.1). We elaborate on this in Chapter 5 and subsequent
chapters.

4
Another random document with
no related content on Scribd:
 A Chinese Ritz.

Page 40

From Yünnanfu we took the ordinary route via Malong, Kütsingfu,

and the Yünnan Pass into Kweichow Province. It has been admirably
and fully described in Sir A. Hosie’s latest book On the Trail of the
Opium Poppy, so it is unnecessary for me to do it, and I shall merely
describe the things which struck us as of special interest.
The journey from Yünnanfu to Anshun took us seventeen days—a
distance altogether of about three hundred miles.[6] From Anshunfu
we struck north, through much wilder and less-frequented country, in
order to visit the haunts of aboriginal tribes, and made a wide detour,
returning to the main road near Kwei Yang, the capital. We greatly
wished to visit other tribes in the eastern part of the province, but
that was absolutely vetoed by the governor, and we were obliged to
follow the high road through Ping-yüe and Huang Ping Chow to
Chen Yüen. Here we took to the waterway, from which we did not
once swerve till we reached Shanghai. The first part of the journey,
up the Yüen Kiang, is a distance of four hundred and forty-six miles,
and one may go down it in five days, if it is in flood, with a fair
chance of getting drowned! We took ten days, but a good deal of
time was spent at places on the way. Coming up stream the journey
is long and tedious: it may extend to months instead of days. The
natural superstition of the Chinese is displayed on such a journey by
the lavish use of crackers[7] and incense to ward off evil spirits.
These superstitions will die hard: nothing less vital than genuine
Christianity can displace them.
We entered Hunan on May 14, having spent forty-two days in
Kweichow Province. The frontier is indicated by two elaborate
archways, as we saw on entering the province from Yünnan.
Although Hunan is within fifteen days’ reach of Shanghai, it has so
far no facilities for travel better than by water. It is true there is a
short railway line on its western border, but we were not encouraged
to try it, and in summer there is always a good steamship line from
the capital Changsha to Hankow—a distance of two hundred and
twenty miles. The railway is part of the projected line from Hankow to
Canton, and will be of great trade value when it is completed, as
there is no good route to connect this part of the country with the
south. We intended going from Changsha into Kwangsi Province on
account of the beautiful scenery, but unfortunately that was
impossible owing to the fighting going on between the troops of the
north and the south exactly in the region where our road lay. It might
be supposed that we could have taken an alternative route through
so vast a country, but such is not the case. If you leave the great
high road (and what a misnomer that is!) there is no way except by
devious paths through endless mountain ranges, where no
accommodation and little food would be obtainable. In a province of
83,398 square miles there appear to be only two main roads running
from north to south, and three from east to west; yet it has a
population of over twenty-one millions. The two main roads running
from north to south are near the eastern and western borders, and
all the central part of the province has none. We crossed the
province entirely by water, first in a house-boat as far as Changteh,
thence in a miserable little native steamer across the Tong Ting Lake
to Changsha; from Changsha up the Siang-kiang,[8] across the Tong
Ting on to the Yangtze, which bounds the province on the north.
We had no choice, therefore, of leaving Hunan except by going to
Hankow, and we found good accommodation on a British steamer,
the Sinkiang. There are six good lines between Changsha and
Hankow—two British, a Japanese, and several Chinese steamship
companies, whose ships run in the summer; owing to the
extraordinary subsidence of the lake in winter (see Chapter VI) it has
to be discontinued then for several months. The journey from
Changsha to Hankow takes about thirty-two hours: at Hankow we
transhipped for Shanghai on a most comfortable steamer (with nice
beds), the Nganking, belonging to Messrs. Butterfield & Swire. It is
quite easy for travellers knowing no Chinese to penetrate by this
route into the very centre of China. I am so often asked about the
possibilities of doing this that I can only recommend this as a wholly
charming and easy way of getting about and much to be preferred to
railway travelling.
Hankow itself is a big bustling cosmopolitan town, with a rapidly
increasing volume of commerce. It is a link between old and new,
and has no less than thirty-six associations, called “hangs,” for
different kinds of goods. It has its foreign concessions like the
seaports, and the important trading companies have their own
floating wharves, where the big ocean liners moor, six hundred miles
away from the coast. There are said to be 25,000 junks engaged
here in river traffic, and they connect Hankow with all the central and
western provinces, often travelling as much as a hundred miles a
day.
Hankow is a great centre of educational and missionary activity,
and many European nationalities are engaged in it. The great
viceroy, Chang Chih Tung, ardently promoted education here when
he was in office. He said in his book, China’s Only Hope (p. 61), “In
order to render China powerful, and at the same time preserve our
own institutions, it is absolutely necessary that we should utilize
Western knowledge. But unless Chinese learning is made the basis
of education and a Chinese direction is given to thought, the strong
will become anarchists, and the weak slaves.” It is most deplorable
that this is ignored by so many Chinese of the present day.[9] He
urged that old Buddhist and Taoist temples, falling into decay, be
transformed into schools, and he estimated that seven out of ten,
with their property, might well be devoted to this purpose. This is
quite in accordance with what is now being done, especially in
Northern China. He argued that the temples are national property,
and should be used for the common good.
Hankow is wonderfully situated as the internal trade centre of
China, being the meeting-point of its main railways as well as
waterways (when the former shall have been completed), linking up
north, south, east and west. There are three cities in the angles
formed by the junction of the Yangtze and the Han rivers—Hanyang,
Hankow and Wu-chang; the last was far the most important in the
past, and is the capital of Hupeh, but now Hankow rivals Shanghai in
commercial importance, and is rapidly growing. The three cities have
a joint native population of 1,150,000, of which Hankow has 800,000,
and as its native quarter was completely destroyed by fire during the
civil war in 1911, the fine old monuments of the past were destroyed
with it. Its whole interest is modern.
Between Hankow and Shanghai are several important cities which
must be most interesting to visit, several of them being treaty ports
with foreign concessions, such as Chin-kiang and Wu-hu; the
steamers stop at least fourteen times on the way. It is delightful to
spend the days sitting in comfortable chairs on deck, watching all the
varied life on the river, hearing the “honk-honk” of the wild geese
soaring overhead, or watching the wedges of ducks crossing the
river on strong wing to the big marshes, or lakes, into which it pours
its overflow.
From time to time the steamer draws to the riverside and loads or
unloads its cargo. One of the most interesting stopping-places is
Nanking; indeed, it is said to be one of the most interesting in China,
and is only about two hundred miles from Shanghai. It has an hotel
kept by an Englishman. We greatly enjoyed the river scenery of the
Yangtze: there are so many picturesque monuments in this lower
part on the numerous islands and along its banks; although it has not
the wild charm of the gorges, it is well worth making the trip. Now
there is a steam service all the way up to Chunking, so that travellers
can easily do one of the most beautiful journeys in the world at
reasonable cost, in reasonable comfort and in reasonable time,
going about a thousand miles up the finest of the great rivers of the
world.[10]
On reaching Shanghai I had to part with my travelling companion
and start the journey home alone, but as events proved my steamer
was delayed, and I had several weeks in which to visit the coastal
provinces of Chekiang, Fukien and Kwantung, and to study the
student movement, as described in the concluding chapters of this
book.
 Chapter II
The Model Governor—Yen Hsi-Shan

“Who is the true and who is the false statesman?

“The true statesman is he who brings order out of disorder;
who first organizes and then administers the government of
his own country; and having made a nation, seeks to
reconcile the national interest with those of Europe and of
mankind. He is not a mere theorist, nor yet a dealer in
expedients; the whole and the parts grow together in his
mind; while the head is conceiving, the hand is executing.
Although obliged to descend to the world, he is not of the
world. His thoughts are fixed not on power, or riches, or
extension of territory, but on an ideal state, in which all the
citizens have an equal chance of health and life, and the
highest education is within the reach of all, and the moral and
intellectual qualities of every individual are freely developed,
and ‘the idea of good’ is the animating principle of the whole.
Not the attainment of freedom alone, or of order alone, but
how to unite freedom with order is the problem which he has
to solve.
“The statesman who places before himself these lofty aims
has undertaken a task which will call forth all his powers.”—
Benjamin Jowett.

Chapter II[11]
The Model Governor—Yen Hsi-Shan
 The province of
Shansi boasts
having the best
governor in the
Chinese Empire,
and he has
accomplished in the
last ten years a
remarkable change
in the entire
province—a
province which is
considerably larger
than Great Britain.
The city of Taiyuanfu
is perhaps the most
striking evidence of
this change. The
whole place is
unrecognizable
since the days when
I first knew it in
1893. The streets
are wide and well
kept; at night they are lighted by electricity, and an efficient police
force keeps order and regulates the traffic, whereas in old days the
crowd used to fight their quarrels out in their own sweet way. The
horrible pariah dogs which infested the streets without let or
hindrance have entirely disappeared; for a dog, licence has now to
be obtained, and any unlicensed dogs are promptly destroyed. The
Governor Yen Hsi-Shan is the promoter of education in all its
manifold aspects; though not a Christian, he realizes that there must
be a radical change in morals, as well as in education, if China is to
become a strong nation, capable of taking her place among the
Great Powers.
To this end he has formed an organization called the “Wash the
Heart Society,” which strongly reminds one of the Mission of John
the Baptist, although he does not recognize the fact that repentance
is only the first step on the upward path. A large hall has been built in
a nice open part of the city, close to the city wall, but, alas! not in
Chinese style. The Governor is unfortunately under the influence of a
Teuton, who is the worst possible adviser in matters of architecture,
as well as other things. The hall is a deplorable mixture of every
conceivable style of Western art; it holds 3,000 people and services
are held there every Sunday morning, each lasting one hour, and
each for the benefit of a separate class of people—merchants,
military, students. So far there seems to be no provision for women,
but perhaps that will come later. The population is bidden to come
and reflect on its evil ways and to seek amendment of life. A special
feature of the service is a time of silence for self-examination. This
Society was started in the province of Shansi, but I found its halls in
other parts of the Empire as well, and it is a hopeful sign of the
times. The approach to the hall is by a good macadamized road, and
near by is a tea-house beside the tiny lake—the Haizabien—and a
bandstand where the élite of the city gather on summer evenings to
listen to sweet music and sip countless cups of tea.
 Yen Hsi Shan, Statesman.

Page 49

Big houses are being built by wealthy Chinese in this

neighbourhood, and there are large Government schools for girls as
well as boys. Facing the entrance to a girls’ school, which is housed
in a disused temple, we saw a list pasted up on a wall, giving the
names of successful girl students in a recent Government
examination. What an amazing contrast to the old days, when no
Government schools for girls were in existence; they only came into
being since the downfall of the old regime in 1907, but the Chinese
Ministry of Education, which based its present system on that of
Japan, is recognizing the importance of women’s education and is
encouraging it by this official recognition of success in examinations.
It is not sufficient only to give the women schools, but it is
imperative to supply them also with scope for wider culture and
congenial activities when they leave school. To this end a kind of
club, or institute, is to be started at once on ground opposite the
Governor’s hall, and it is in response to the ladies’ own request: they
have long been saying, “The men have their Y.M.C.A., why cannot
we have such a place?” and although the mission ladies have done
their best to meet the need, obviously no private house can be
adequate, not to mention the fact that Chinese ladies have too much
self-respect to be willing to be always guests of ladies with limited
incomes, to whom they can make but scant return. It is hoped that
the new hall will do much to forward the woman’s movement in
Taiyuanfu; there will be social gatherings, lectures on hygiene (for
illustration of which there is to be a complete installation of sanitary
fittings), a child-welfare department, invalid cooking, lessons on
nursing, and many other classes connected with women’s welfare.
There is room for a garden and tennis courts in order that recreation
and physical culture may be encouraged and the place made
attractive to girls as well as women. The Governor is promoting this
last matter indirectly, by putting a fine on foot-binding, which is
unfortunately still extremely prevalent. The movement that took place
some years ago in favour of a natural foot seems to have died down,
and everywhere there is foot-binding in full swing. The queue has
practically disappeared from China during the last few years, and
men wear their hair mostly rather short, while some go in for a clean
shave. I find this quite attractive when the skull is well-shaped, and if
the man is in immaculate summer garb, the effect of cleanliness is
wonderful. If the women of China were less conservative, and would
make an equally clean sweep of foot-binding, it would make an
immense difference to their health.
The Governor has encouraged physical culture not only indirectly
but directly as well, for he even went so far as to ride in a bicycle
race once in Shansi, and in 1918 public sports took place at
Taiyuanfu, which Sir John Jordan honoured by his presence, having
taken the tiresome nineteen hours’ journey from Peking for the
purpose.[12] A disused temple acted as the grand-stand and an
angle of the city wall was converted into the arena, the tiers of seats
being hewn in the base of the wall. Quite a fine sports ground was
prepared under the superintendence of one of the missionaries,
whose advice in practical matters is continually sought by the
Governor. The only matter for regret on this occasion was the
deplorable weather, for even sunny Shansi has moments when a
dusty fit of temper obscures its lustre.
One of the Governor’s most valuable new institutions is a farm for
cattle-breeding. It is just outside the city and has been successfully
started by an American and his wife. The main object is to improve
the breed of horses, cows, sheep, etc., and for this purpose stock is
being imported from the United States, whose Government has
recently supplied the necessary transport for horses, when this
difficulty of shipment arose with regard to animals already
purchased; a large number of sheep have already been imported
from Australia. Shansi is a suitable province for this experiment and
missionaries have already proved there the excellent results of
cross-breeding cows, obtaining supplies of milk of improved quality,
as well as largely-increased quantity. In a recent book on China,
highly recommended to me, an American writer states that there is
only one milk-giving cow in the Empire, and that tinned milk supplies
the rest, but evidently the traveller had not travelled far!
Another of the Governor’s institutions is a College of Agriculture
and Forestry in connexion with which there are many mulberry trees
being planted for the promotion of sericulture. This has never been
pursued with success in Shansi; hitherto only the commoner kinds of
silk have been produced, but it is considered a patriotic deed to
promote it, and the most exquisite and costly silk is now being made
in a disused temple, by Yen’s order. Perhaps the almost religious
way in which it was regarded in bygone times, when the Empress
herself took a ceremonial part in the rearing of silkworms 3,000
years ago, has caused this revival of schools of sericulture. I visited
one in the South, and after seeing all the processes was invited to
take a handful of worms away as a memento! Governor Yen has
sent 100 students to France to study textiles. Afforestation is
nowhere more needed than in Shansi, and it is to be hoped that the
Government will push this side of the work of the college. We found
such a college had been started in remote Kweichow also, cut off
from most of the new movements in China. Plantations had been
made in various parts, but they will need to be carefully guarded, as
the poverty of the inhabitants lead them to destroy ruthlessly every
twig they can for firewood; where there used to be large forests
nothing now remains of them. The genius of the Chinese race for
agriculture is so remarkable that one may well expect great results
from these colleges: the vast population has been able in the past to
produce food more or less according to its needs, but when there is
a dearth of rain, or other cause producing bad harvests, there is at
once terrible scarcity, and the application of Western knowledge and
agricultural implements ought to be of considerable value.
There are in some parts such as Chekiang as many as four
harvests per annum, and no sooner is one reaped than the land is
prepared for the next. The introduction of new trees, vegetables,
etc., would add greatly to the wealth of the country, and with its
unrivalled climate and soil there is every reason to promote the
multiplication of agricultural colleges.
One of the most noticeable changes in Taiyuanfu is the complete
absence of the beggar of hideous mien, who dogs the steps of
strangers in every other city in China, and who seems to be the most
immovable feature of life in the East. He was an integral part, one
had been taught to believe, of the social fabric, and as hallowed as
the very temple itself; yet Taiyuanfu has the glory of having solved
this difficult problem. All the male beggars have been collected into
the splendid old temple of Heaven and Hell to be taught a trade, so
as to be able to earn a living, and they are not dismissed until they
are capable of doing so. They seemed quite a jolly crew, and were
hard at work in various buildings, though others of these were closed
for the New Year. The most interesting part of the institution was the
town band which has been formed out of the younger part of the
beggar population. They were summoned to play for our
amusement, and they ended by playing for their own. The
performance was most creditable, especially considering that the
band was only seven months old; if there was some defect in tune,
there was an excellent sense of rhythm, which I have found lacking
in many bands of long standing at home; and it was really fascinating
to see the gusto with which they all played. The band has already
taken part in various town functions, and is making itself useful. The
music is, of course, Western, as are the instruments. Chinese
musical instruments do not give enough sound, as a rule, for large
gatherings.
The rules of the workhouse seem good, and the inmates can earn
money (five dollars per month) so as to have something in hand
when they leave. The women’s department is in another part of the
city and we had not time to visit it. It is very noticeable how the
temples are being everywhere used for such useful purposes, for the
housing question is here, as at home, a serious problem. No doubt it
would be good from a practical point of view that these buildings
should be replaced by new ones, built for the purpose, but the loss of
beauty would be incalculable. The temple of Heaven and Hell has
glorious turquoise-blue roofs and handsome tiles and large
medallions of green pottery on the walls. It is the most beautiful of all
the temples, in my opinion, though the Imperial Temple, where the
Dowager Empress stayed on her historic flight to Sianfu, is also very
fine. This is now used as a school for the teaching of the new script,
which is a simplified form of the Chinese character. It was devised in
1918 by the Ministry of Education in order to make literacy easier for
the population. The ordinary Chinese boy takes three or four years
longer than the Western boy to learn to read. When the old system
of education was abolished in 1907 a new one had to be devised,
but it is an extremely difficult thing to carry out such a reform
throughout so vast an empire. Governor Yen Hsi-Shan is an ardent
promoter of the scheme, and he has established the school for the
express purpose of promoting it. Every household in Taiyuanfu is
required to send at least one member to study the script; in order to
make it easier, there are a few characters, with their equivalent in the
old script, put up outside many of the shops, so that people may
learn it as they go about their business. Not only so, but all over the
city may be seen notice boards with the two scripts in parallel
columns, and these boards have generally some one studying them,
not infrequently with notebook in hand. Many of the schools are
teaching it, but it is difficult to add this to an already well-filled time-
table. It may be of interest to know that it is “a phonetic system
containing thirty-nine symbols (divided into three denominations, viz.
twenty-four initials, three medials and twelve finals).”—North China
Daily News.
This is one part of an important movement for the unification of the
language, the importance of which can only be fully realized by those
who have travelled widely in China. Not only every province varies
from every other province, but also every district from every other
district. There are sixty-four dialects in Fukien alone. The unification
of the Empire would be greatly promoted by the unification of the
language, and this has been frankly recognized by the Ministry of
Education, which has issued a notice to that effect:
“We recognize that because of the difference between our
classical and spoken language, education in the schools makes slow
progress, and the keen edge of the spirit of union both between
individuals and in society at large has thereby been blunted.
Moreover, if we do not take prompt steps to make the written and the
spoken language the same quickly, any plans for developing our
civilization will surely fail.
“This Ministry of Education has for several years made positive
advances in promoting such a National Language. All educationists,
moreover, throughout the country are in favour of a change by which
the teaching of the national spoken language shall take the place of
the classical language. Inasmuch, therefore, as all desire to promote
education in the National Language, we deem it wise not to delay
longer in the matter.
 “We therefore now order that from the autumn of this current year,
beginning in the primary schools for the first and second years, all
shall be taught the National Spoken Language, rather than the
National Classical Language. Thus the spoken and written
languages will become one. This Ministry requests all officials to take
notice and act accordingly.”
It is not sufficient, as we all know, merely to issue such an order.
Governor Yen has taken various practical ways of enforcing it.
Posters with large script characters have been widely set up,
exhorting the people to study the script, and a daily paper is issued
in it. He has had 2,500,000 copies printed of a simple script primer,
and has published at a nominal price, and in vast numbers, various
educational books, such as What the People Ought to Know,[13]
New Criminal Laws of the Republic, and Handbook for Village
Leaders. The last-named is of special importance in view of the fact
that by his order reading- and lecture-rooms have been established
in all the cities and large villages of the province, where lectures and
talks are given from time to time on various subjects of interest to the
people. A regular educational campaign may be said to have been
inaugurated by Yen. On every post and wall in the remotest villages
may be seen maxims inculcating honesty, diligence, industry,
patriotism and military preparedness.
 Temple of Heaven and Hell, Workhouse.

Page 53

An important new book which Governor Yen has recently

published, is called What Every Family Ought to Know, and is a
description of what he conceives to be a good home and the
happiness which results from it. “If we desire to have a good home,
virtue is of first importance,” he says, but alas! he gives no clue as to
how it is to be achieved.
The chief rules for family life are, (i) Friendliness, (ii) Magnanimity,
(iii) Dignity, (iv) Rectitude, (v) Diligence, (vi) Economy, (vii)
Cleanliness, (viii) Quietness. He makes the Head of the House
responsible, as setting the example, and exhorts him to repentance
(if he falls short) before God and his ancestors. The whole book is
eminently practical, and he recommends what would be a startling
change of immemorial custom, that the son should not marry until he
is grown up and able to support a wife in a home of his own—
namely, not under his father’s roof. This is an innovation which is
beginning to be seen elsewhere, as the result of foreign intercourse.
As a writer, Governor Yen is concise and practical: he has
completely broken away from the old Chinese classical style. His last
work is written, like all his books, in simple mandarin instead of in
beautiful classical mandarin, so that every one may be able to
understand it. This is the more noteworthy, because the additional
cost entailed was $5,400 per leaf; he states this fact in the preface of
What the People Ought to Know.
His one object appears to be the uplift of the people in every way,
and he believes in God and in righteousness. As an index of his view
of life it may be interesting to quote a few of the forty Family Maxims
which form the concluding chapter of his above-named book.

“Unjust wealth brings calamity.”

“Vitiated air kills more people than prison.”
“To be cruel to one’s own is to be worse than a beast.”
“Of people who lack a sense of responsibility—the fewer
the better.”
“If your conscience tells you a thing is wrong, it is wrong:
don’t do it.”
 “The experience of the uneducated is much to be preferred
to the inexperience of the educated.”
“The wise are self-reliant, the stupid apply to others.”
“There is no greater calamity than to give reins to one’s
desires, and no greater evil than self-deception.”

Governor Yen, it will be seen, from his words as well as from his
deeds, is a clear-sighted, independent thinker, and he believes in
religious liberty. His reforms deal with a wide range of things—
opium-smoking, narcotics, polygamy, infanticide, early marriages,
early burial, gambling, training and morals of the troops, compulsory
free education for boys, the introduction of uniform weights and
measures, alteration in legal affairs. All these and other matters have
within the last five years occupied his thoughts and been practically
dealt with—no small achievement, especially when the insecurity of
his position and lack of trained men to carry out his projects is taken
into consideration.
As will be readily understood, all these enterprises cost money,
and taxation is never looked on kindly by the taxed, so there is some
discontent among the people of Shansi, and the Central
Government, instead of showing satisfaction at the prosperity and
good government of the province, which is in striking contrast to that
of so many others, has taken the opportunity of threatening to
impose a Civil Governor in Shansi—that means a heavy squeeze,
and in consequence, the stoppage of many of the Governor’s
schemes. He is continually threatened by those who would like to
see him out of the way, and is consequently rarely seen, and then
strongly guarded.
The system of having military governors is extremely bad, but in
the case of an exceptional man like Yen it has worked well, and the
Government saved its “face” by uniting the civil and military
governorship in his one person. At the present time the Government
has ordered the military governor of Shensi to retire in favour of
another Tuchun. He refuses to do so, and his various military friends
are all hurrying to the rescue. It is estimated that there are one and a
half million soldiers in China, largely unpaid, so that they are glad of
any excuse to loot and pillage. Feng Yu Hsiang has been sent up to
Shensi by the Government to compel the Tuchun to leave, and has
carried out the work with brilliant success. He has in vain been
demanding money to pay his troops, while turbulent, unscrupulous
generals have been receiving large sums to prevent them from
committing excesses.
The Tuchuns have been encouraging opium-growing in order to
get funds, and now there is hardly a province where it is not done
more or less openly. Governor Yen has set his face against it, but
smuggling goes on all the time, mainly from Japan, and morphia is
also becoming increasingly popular. No wonder Young China is
clamouring for the suppression of the Tuchuns and disarmament:
there can be no peace in China till this is done.
One of the most interesting places in the city is the model gaol,
which was planned and carried out by Mr. Hsü, who studied in Japan
and has progressive views. It covers a considerable space of ground
and is entirely one-storied; it is in the shape of a wheel, with many
spokes radiating from the centre. The entrance is charming, as
unlike as it is possible to imagine to any English prison. Within the
gates is a lovely garden, for Chinese are first-rate gardeners, and the
prisoners raise all the vegetables necessary for the inmates, and a
grand show of flowers to boot. An avenue of trees leads to the
offices, and when we were there in February we saw beautiful little
trees of prunus in full bloom on the office table! All the prisoners
have to work at useful trades, and if it were not for their fetters it
would be difficult to imagine one was in a prison at all. The
workshops were bright and airy; every one looked well cared for and
not unhappy. A feature of the workrooms was the boards on which
all tools were hung up when not in use, each tool being numbered
and outlined on the board, so that it should be hung on its own peg.
Every kind of trade was in full swing, and the work is so well
executed that there is never any lack of orders. Certainly one would
be only too glad to have things made under such good conditions.
The sleeping accommodation was excellent: the cells and beds of
remarkable cleanliness and comfort; no one could object to them.
The bath-house was of some interest. All the inmates have to
undergo a weekly bath on Sundays, in batches of ten at a time, and
their clothes are also kept thoroughly clean. The kitchen looked most
attractive, and the rice and soup, which form the staple of their food,
compared favourably with what one sees in the inns. The prisoners,
too, are allowed as much as they like at their two daily meals.
Throughout the Army there are no more than two meals a day. The
place of punishment looked uncommonly like a theatre stage, and
one cannot but hope that soon all executions will take place within
the prison precincts instead of in public; but as Europe has not yet
learnt to do this, one cannot be surprised that China has not.
After inspecting the Delco Engine, which provides light for the
whole place, we went to visit the women’s prison, which is within the
same enclosure as the men’s, though separated by a wall. It was
very much smaller in extent but equally well kept, and even, I must
add, attractive. The matron was a pleasant-faced, comely woman,
and her own room quite a picture. The white-curtained bed, pretty
coverlet, vase of flowers, and various little treasures suggested a
home, and as she took us round, it was easy to see that she was
happy in her work. We passed through the dining-room, where the
tables were spread with clean cloths, and bowls and chopsticks were
ready for the forthcoming meal. The prisoners were only about thirty
in number, and were busy making mattresses and clothing, knitting
and crocheting. It was suggested that they should sing a hymn,
which they did with evident pleasure, and some of them talked with
the missionary, who comes to see them once a week. The matron is
not a Christian, but finds the singing and reading does them so much
good that she has taken to learning and to teaching them herself.
The missionaries were originally invited by the master to come and
speak to the prisoners, and it is now a regular custom. One woman
who is in for murder has become quite a changed character, and her
term has been shortened in consequence of her good behaviour.
Some were in for opium-smoking, which is here a punishable
offence, while in other parts of the empire it is frankly encouraged.
The prisoners are allowed to have a visitor once a month, but no
complaints are allowed to be made. Visits are stopped if this