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 ENCYCLOPEDIA OF
EVOLUTIONARY BIOLOGY
 ENCYCLOPEDIA OF
EVOLUTIONARY BIOLOGY

EDITOR IN CHIEF

RICHARD M. KLIMAN
Cedar Crest College, Allentown, PA, USA

VOLUME 1
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16 17 18 19 20 10 9 8 7 6 5 4 3 2 1
EDITOR IN CHIEF

Richard M. Kliman, PhD, is Professor of Biological Sciences at Cedar Crest College in

Allentown, Pennsylvania. He received his BA from Colby College in biology and music. His
graduate work at Wesleyan University focused on quantitative genetics of circadian rhythms
and photoperiodism in the Djungarian hamster, Phodopus sungorus. As a postdoctoral fellow
at Rutgers University and Harvard University, he studied molecular evolution and popu-
lation genetics. Prior to Cedar Crest College, he taught at Radford University in Virginia and
Kean University in New Jersey. He has also served as a program director in the Division of
Environmental Biology at the US National Science Foundation (NSF).
Kliman’s research interests center on questions in molecular evolution, including the
evolution of codon usage bias in a variety of organisms; speciation and natural history; and
ecology and conservation. Much of this work has relied on population genetics/genomics
and bioinformatics approaches. He has also collaborated with Cedar Crest colleague John
Cigliano on an Earthwatch-supported “before-after-control-impact” study on the effects of a
new marine reserve in Belize on queen conch populations. His research in evolutionary and
ecological genetics has been supported by the US National Institutes of Health and by
Conservation International.
Kliman has served on the editorial boards of Genetica and The Journal of Molecular Evolution. He has been deeply involved in
evolution education, helping to coordinate “Undergraduate Diversity at SSE/SSB,” an NSF-supported program to bring a diverse
group of undergraduates to the annual Evolution research conference. He was a lead editor of population/quantitative genetics
and evolutionary genetics for Nature Education/Scitable at its inception. He is a member of the Education and Outreach Committee
of the Society for the Study of Evolution, and editor of the society’s peer-reviewed educational resource, the EvoEd Digital Library.

v
SECTION EDITORS

Hiroshi Akashi is a Professor of Evolutionary Genetics at the National Institute of Genetics,

Japan. He worked with Marty Kreitman for his PhD in Ecology & Evolutionary Biology from
the University of Chicago (1996) and with John Gillespie as a postdoctoral fellow at UC
Davis. He has been a faculty member at the University of Kansas (1998–2000), Penn State
University (2000–2008), and NIG (2009–present). Akashi’s research focuses on inferring
causes of genome evolution, especially weak selection, from within and between species
sequence variation. His studies of codon usage employed population genetic methods to
detect natural selection acting at its limit of efficacy and identified a phenotypic basis of
natural selection (translational accuracy) from sequence comparisons in Drosophila. Exten-
sions of this work revealed constraints related to biosynthesis that act globally on com-
positional properties of microbial proteins. The interplay of weak evolutionary forces
appears to shift frequently among closely-related species and current interests include tests
of adaptive changes in protein/DNA composition.

Tim Coulson’s primary interest is in creating better links between the fields of ecology and
evolution. He does this by developing theory, parameterising models for field and laboratory
systems, making predictions from these models, and, where possible, testing these predic-
tions with experiments. He works on a range of systems, from bulb mites within the la-
boratory, to guppies living in streams in Trinidad, to wolves in Yellowstone. His motivation
to do this comes from observations that ecological and evolutionary change can be observed
occurring on similar time scales, yet ecological theory typically ignores evolutionary pro-
cesses and vice versa.
Tim was awarded his PhD in plant ecology from Imperial College, London, in 1994. He
moved on to research genotype-by-environment interactions as Natural Environment Re-
search Council (NERC)-funded post-doc at the Institute of Zoology in London. He remained
at the Institute on a fellowship where he developed models to investigate the economic and
life history consequences of a range of population management strategies. In 2000 he moved
to the University of Cambridge, where he briefly lecturered in the Zoology department. In
2004 he moved back to Imperial College London as a senior lecturer where he started
developing models that allow the simultaneous investigation of the dynamics of life history, populations, and quantitative
characters. In 2007 he became Professor of Population Biology at Imperial College London. He left Imperial in 2013 to take up his
current position as Professor of Zoology at the University of Oxford. He is also a Professorial fellow of Jesus College, Oxford.

Andrew Forbes

vii
viii Section Editors

Rosemary Gillespie is a Professor at the University of California, Berkeley, where she also
holds the Schlinger Chair in Systematics. She is Past President of the International Bio-
geography Society and Trustee and Fellow of the California Academy of Sciences, and serves as
Associate Editor for Molecular Ecology. Gillespie was born and educated in Scotland, receiving
her BSc in Zoology from Edinburgh University in 1980. She came to the US to conduct
graduate work on the behavioral ecology of spiders at the University of Tennessee. After her
PhD she spent several months at the University of South in Tennessee, and then started work
at the University of Hawaii in 1987, initially as a postdoc, and then in 1992 as Assistant
Professor in Zoology and Researcher in the Hawaiian Evolutionary Biology Program. It was
during her first year in Hawaii that she discovered an adaptive radiation of Tetragnatha
spiders. She left Hawaii in 1999 to join the faculty at the University of California in Berkeley,
where she continues her research focus on the islands of the Pacific, Hawaii in particular,
using islands of known age and isolation to assess the combined temporal and spatial
dimension of biogeography and determine patterns of diversification, adaptive radiation,
and associated community assembly.

David Guttman received his PhD from Stony Brook University in 1994 working with Daniel
Dykhuizen on questions related to the role and importance of recombination in structuring
genetic diversity in bacterial populations. He followed this with a postdoc in molecular
evolution with Brian and Deborah Charlesworth at the University of Chicago, and a second
postdoc at the University of Chicago with Jean Greenberg to gain experience in the fields of
molecular plant pathology and plant-microbe interactions. He started his faculty position at
the University of Toronto in 2000, and is currently a Professor in the Department of Cell &
Systems Biology (CSB). He is also the Associate Chair for Research in CSB, founder and
Director of the University of Toronto Centre for the Analysis of Genome Evolution &
Function, and Canada Research Chair in Comparative Genomics. He has served as the Chair
of the American Society for Microbiology, Division R (Evolutionary and Genomic Micro-
biology), and was the PLoS Pathogens Section Editor for Bacterial Evolution & Genomics.
Dr. Guttman runs a highly diverse research program generally focused on bacterial
evolutionary genomics, with three major foci: (1) the evolution of host specificity and
virulence in plant pathogenic bacteria; (2) microbial comparative genomics; and (3) studies
of the human and plant-associated microbiome. He is best known for elucidating and
linking evolutionary and mechanistic processes that determine the course and fate of bac-
terial infections, and characterizing the impact of genetic variation on the balance between
disease and immunity.

Norman A. Johnson, the section editor for Applied Evolution, is an evolutionary geneticist
and author. He received his PhD from the University of Rochester in 1992 and did post-
doctoral research at the University of Chicago. His research interests have generally focused
on aspects of speciation, specifically those related to the genetics and evolution of hybrid
incompatibility: sterility, inviability, or other reduction of fitness in hybrids between species.
Dr. Johnson, an adjunct professor in the Biology Department at the University of Massa-
chusetts at Amherst, has taught classes there, as well as at Hampshire College, the University
of Texas at Arlington, and the University of Chicago.
Dr. Johnson also has a long-standing commitment toward improving the communi-
cation of science in general and evolutionary biology in particular to other scientists, edu-
cators, and the public at large. He is the author of Darwinian Detectives: Revealing the Natural
History of Genes and Genomes (Oxford University Press: 2007), a book geared to general
audiences that shows how biologists use DNA sequence data to make inferences about
evolutionary processes. He also was the lead organizer for a working group on communi-
cating human evolution at the National Evolutionary Synthesis Center (NESCent).
 Section Editors ix

Laura Kubatko received a PhD in Biostatistics from The Ohio State University (OSU) in
1999. After seven years on the faculty at the University of New Mexico, she returned to OSU
in the Fall of 2006, and is now Professor of Statistics and of Evolution, Ecology, and
Organismal Biology at OSU. Laura served as an Associate Director of the Mathematical
Biosciences Institute at OSU from 2013–2015. At OSU, she is a Faculty Affiliate of the
Initiative in Population Research, and a Faculty Affiliate in Translational Data Analytics
(TDA@OSU). She holds appointments as an Affiliate Faculty Member at the Battelle Center
for Mathematical Medicine at Nationwide Children’s Hospital in Columbus and as an Ad-
junct Research Scientist at Lovelace Respiratory Research Institute in Albuquerque, NM.
Laura’s research interests are in statistical genetics, with a focus on the development of
statistical methods for inferring phylogenies from molecular data. Her recent work in this
area concentrates on bridging the gap between traditional phylogenetic techniques and
methodology used in population genetics analyses, primarily through the application of coalescent theory to species-level
phylogenetic inference. She develops and distributes several software packages for phylogenetic inference, and has been an active
member of the Society of Systematic Biologists. She has served as an Associate Editor for the journal Systematic Biology since 2007.

Amy Litt has been studying plant evolution and diversity since her PhD on floral structure
and evolution in the neotropical plant family Vochysiaceae, known for its beautiful but
unusual flowers many of which have only one petal and one stamen. While completing her
PhD in plant systematics and morphology in the joint City University of New York/New
York Botanical Garden Plant Sciences program under Scott Mori and Dennis Stevenson, she
became interested in the molecular basis of plant diversity. She did her post-doc in the
developmental genetics lab of Vivian Irish at Yale University on the evolution of a family of
transcription factors involved in flower development, and she continues to study the func-
tional evolution of this gene family currently. After one year on the faculty of University of
Alabama, she moved back to The New York Botanical Garden as Director of Plant Genomics,
where she developed her research program studying the evolution of plant form along two
paths: studying evolutionary changes in genes to see how those changes affected flower and
fruit form; and identifying the genes that underlie differences in form among closely related
species. Dr. Litt also served as a program director in Plant, Fungal, and Microbial Devel-
opment and Evolutionary Development at the National Science Foundation. She recently moved to the University of California at
Riverside, where she continues to study the genetic basis of plant diversity.

Maria E. Orive is a professor of evolutionary genetics in the Department of Ecology and

Evolutionary Biology at the University of Kansas. Her research in theoretical population
genetics aims to develop mathematical models that provide a conceptual framework for
exploring important questions in evolutionary biology and analytical tools for demographic
and genetic data. Her work has considered levels of selection and mutation in organisms that
reproduce both sexually and asexually, the relationship of population structure and life-
history attributes to gene flow and genetic diversity, and models of within- and between-host
pathogen and symbiont population dynamics. Orive received her BS from Stanford Uni-
versity and her PhD from the University of California at Berkeley. After spending two years as
a postdoctoral researcher in genetics at the University of Georgia, she was an NSF-NATO
Postdoctoral Fellow at the University of Edinburgh. Her research has been funded by
multiple grants from NSF and NIH. In 2007–2008, she was the Carl and Lily Pforzheimer
Foundation Fellow at the Radcliffe Institute for Advanced Study (Harvard University), and
has served as the University Faculty Ombudsman for the University of Kansas since 2007.
x Section Editors

Daniel Ortiz-Barrientos is an Associate Professor in evolutionary genetics in the School of

Biological Sciences at The University of Queensland, Brisbane, Australia. During his scientific
career he has investigated the ecological and genetic basis of speciation both in plants and
animals. His current research program explores the early stages of speciation, the molecular
basis of parallel speciation, and the interplay between recombination and natural selection
during the origin of new species. His research funds come from The Australian Research
Council. He is married to Antonia Posada, and is the father of three energetic and
beautiful kids.

Claudia Russo was born in Leeds, England, but has lived in Rio de Janeiro, Brazil since she
was two years old.
Claudia has an academic major in Ecology from Universidade Federal do Rio de Janeiro
completed in 1989, and finished her Master’s thesis in 1991 on population genetics of two
actiniid species of sea anemones with different reproductive strategies, under the supervision
of Associate Professor Antonio Mateo Sole-Cava. Her PhD dissertation was on the di-
versification of drosophilids and on the use of a known phylogenetic tree to estimate the
reliability of tree building methods. The dissertation was completed in 1995 under the
supervision of the Evan Pugh Professor Masatoshi Nei who recently received the prestigious
Thomas Hunt Morgan Medal. Her graduate degrees were obtained as a student at the
Genetics Program from the Universidade Federal do Rio de Janeiro and as a visiting scholar
at the Pennsylvania State University (1992–1995).
Claudia is currently the Head of the Genetics Department at the Federal University of Rio
de Janeiro, having been a member since 1997. Claudia has supervised 13 Master’s disser-
tations, eight PhD theses and seven post-docs, of which eight are now Assistant Professors at
universities in Brazil and abroad. She has published 42 academic papers that have been cited over 1,200 times. Her h-index is 14.
Since 2012, Claudia has been a member of the editorial board, and an associate editor of the Molecular Biology and Evolution
journal. Since 2012 she has been a council member for the Pan American Association of Computational Interdisciplinary Sciences
and since 2009 for the Brazilian Association for the Advancement of Science.
Claudia’s general academic interests are on key aspects of animal phylogenetics, including their diversification patterns in time
and space. She has worked with various metazoans groups but more prominently on marine sponges, sea anemones, arthropods,
passerine birds, and mammals. Claudia has also published on the use of known phylogenetic trees to estimate the efficiency of
phylogenetic methods in recovering and rooting those trees. More recently, she has developed some interesting hands-on edu-
cational tools for evolutionary biology practices in the classroom.

Karen E. Sears is an evolutionary developmental biologist whose primary research goal is to

determine how developmental variation within a species produces congenital mal-
formations in humans, and among species generates new evolutionary forms in mammals.
Dr. Sears earned her PhD from the University of Chicago, did postdoctoral research at in the
Howard Hughes Medical Institute (HHMI) lab of Dr. Lee Niswander, and joined the faculty
of the University of Illinois at Urbana-Champaign. At Illinois she holds positions as an
Associate Professor in the Department of Animal Biology, a Faculty Member in the Institute
of Genomic Biology, and an Affiliate of the Program in Ecology, Evolution and Conservation
Biology and the Department of Cell and Developmental Biology. She is also the President of
the Pan American Society for Evolutionary Developmental Biology. She has authored or co-
authored over 35 publications including first-authored publications in Nature, Proceedings of
the National Academy of Sciences, and Evolution. She has served as a principal investigator
on multiple, nationally-funded research projects, and presented invited seminars at more
than 30 institutions and symposia. She is routinely ranked among the top 10% of Illinois professors for her teaching, and was a
featured scientist in the PBS/HHMI documentary “Your Inner Fish.”
 Section Editors xi

Vassiliki “Betty” Smocovitis is Professor of the History of Science in the Department of

Biology and in the Department of History at the University of Florida. Her areas of expertise
include the history of evolutionary biology, genetics and systematics and the history of
botany. She is best known for her contributions to understanding the historical event known
as the “evolutionary synthesis” and in gaining greater understanding of the origins of the
discipline of evolutionary biology. She has published extensively on both the intellectual
and social aspects of the history of evolutionary biology including a history of the Society for
the Study of Evolution, a history of the Darwin Centennial of 1959, and the integration of
botany, genetics, and anthropology into the evolutionary synthesis. She was the contributor
to the Oxford Bibliographies entry on Charles Darwin at over 25,000 words and the entry on
the modern synthesis. She is the author of Unifying Biology: The Evolutionary Synthesis and
Evolutionary Biology (Princeton: Princeton University Press, 1996).

Nina Wedell is a professor of evolutionary biology with research interests focused on the
evolutionary ecology of sex. She has worked extensively on various aspects of sexual selec-
tion and sexual conflict, in particular on the role of selfish genetic elements in reproductive
biology. Nina is the Academic lead for the Behaviour research group at the University of
Exeter.

Jason Wolf is Professor of Evolutionary Genetics in the Department of Biology & Bio-
chemistry and The Milner Centre for Evolution at the University of Bath. His research is
unified with a special focus given to understanding the influence that frequently ignored or
under-appreciated sources of genetic variation have on the genotype-phenotype relationship
and how this, in turn, influences evolutionary processes. He integrates theoretical, compu-
tational and empirical quantitative and population genetic techniques to achieve this goal.
He is particularly interested in understanding the evolutionary consequences of various types
of interactions, including gene interactions (epistasis), parent-offspring interactions and
social interactions. He received a PhD from the University of Kentucky, after which he
was a postdoctoral researcher at Indiana University and a US National Science Foundation
Postdoctoral Fellow at Washington University School of Medicine. Prior to moving to the
University of Bath he held positions at the University of Tennessee and the University of
Manchester. He won the Dobzhansky Prize from the Society for the Study of Evolution, a
Young Investigator’s Prize from the American Society of Naturalists and the Scientific Medal
from the Zoological Society of London.
LIST OF CONTRIBUTORS

LT Ackert Jr. DA Baum

Drexel University, Philadelphia, PA, USA University of Wisconsin, Madison, WI, USA
JD Aguirre MS Bergen
Massey University, Auckland, New Zealand Rutgers University, New Brunswick, NJ, USA
RC Albertson M Berger
University of Massachusetts, Amherst, MA, USA Institute of Hygiene, University of Münster, Münster,
AC Algar Germany
University of Nottingham, Nottingham, UK V Berger
RG Allaby Universite ́ Claude Bernard Lyon 1, Villeurbanne, France
University of Warwick, Coventry, UK AJ Betancourt
SH Alonzo Vetmeduni Vienna, Vienna, Austria
University of California, Santa Cruz, CA, USA
LW Beukeboom
L Altenberg University of Groningen, Groningen, The Netherlands
The Konrad Lorenz Institute for Evolution and Cognition
JP Bielawski
Research, Klosterneuburg, Austria
Dalhousie University, Halifax, NS, Canada
DM Althoff
Syracuse University, Syracuse, NY, USA BK Blackman
University of California, Berkeley, CA, USA; and
JM Á lvarez-Castro University of Virginia, Charlottesville, VA, USA
University of Santiago de Compostela, Lugo, Galiza,
Spain H Blackmon
University of Minnesota, Saint Paul, MN, USA
C Amemiya
Benaroya Research Institute, Seattle, WA, USA JL Blois
University of California, Merced, CA, USA
M Ansdell
University of Hawai’i at Mānoa, Honolulu, HI, USA JD Bloom
Fred Hutchinson Cancer Research Center, Seattle, WA,
WS Armbruster USA
University of Portsmouth, Portsmouth, UK
A Blumberg
APA Assis
Ben-Gurion University of the Negev, Beer-Sheva, Israel
Universidade de São Paulo, CEP São Paulo, Brazil
J Blumenstiel
A Aswad
University of Kansas, Lawrence, KS, USA
University of Oxford, Oxford, UK
JR Auld DT Blumstein
West Chester University, West Chester, PA, USA University of California, Los Angeles, CA, USA

CF Baer E Bolund
University of Florida Genetics Institute, Gainesville, FL, Uppsala University, Uppsala, Sweden
USA C Bonenfant
SCH Barrett Universite ́ Claude Bernard Lyon 1, Villeurbanne, France
University of Toronto, Toronto, ON, Canada
S Borish
G Barshad California State University East Bay, Hayward, CA,
Ben-Gurion University of the Negev, Beer-Sheva, Israel USA
J Bast JW Boughman
University of Lausanne, Lausanne, Switzerland Michigan State University, East Lansing, MI, USA

xiii
xiv List of Contributors

BW Bowen BSW Chang

Hawaiʻi Institute of Marine Biology, University of University of Toronto, Toronto, ON, Canada
Hawaiʻi at Mānoa, Kāneʻohe, HI, USA
MA Charleston
LM Boykin University of Tasmania, Hobart, TAS, Australia
The University of Western Australia, Perth, WA,
Australia Q Chen
Rutgers University, New Brunswick, NJ, USA
PLR Brennan
Mount Holyoke College, South Hadley, MA, USA J Chifman
Wake Forest School of Medicine, Winston-Salem, NC,
JC Briggs USA
Oregon State University, Corvallis, OR, USA
P-A Christin
ED Brodie III University of Sheffield, Sheffield, UK
University of Virginia, Charlottesville, VA, USA
JA Clack
TJ Brodribb University Museum of Zoology, Cambridge, UK
University of Tasmania, Hobart, TAS, Australia
FM Cohan
RM Brown Wesleyan University, Middletown, CT, USA
University of Kansas, Lawrence, KS, USA
AA Comeault
LT Buck University of North Carolina, Raleigh, NC, USA
Natural History Museum, London, UK; and University
of Roehampton, Whitelands College, London, UK JK Conner
Michigan State University, Hickory Corners, MI, USA
JG Burleigh
University of Florida, Gainesville, FL, USA BMA Costa
Universidade de São Paulo, CEP São Paulo, Brazil
T Burmester
University of Hamburg, Hamburg, Germany T Coulson
University of Oxford, Oxford, UK
LF Bussier̀ e
University of Stirling, Stirling, UK BA Counterman
Mississippi State University, Mississippi State, MS, USA
R Butlin
University of Sheffield, Sheffield, UK; and University of R Covas
Gothenburg, Strömstad, Sweden CIBIO-InBio, University of Porto, Campus Agraŕ io de
Vairão, Vairão, Portugal
A Caballero
Universidad de Vigo, Vigo, Spain TP Craig
University of Minnesota Duluth, Duluth, MN, USA
RM Calhoun
Western University, London, ON, Canada D Crowson
University of Toronto, Toronto, ON, Canada
U Candolin
University of Helsinki, Helsinki, Finland AD Cutter
University of Toronto, Toronto, ON, Canada
AC Carnaval
The City College of New York, New York, NY, USA; V Daubin
and The Graduate Center of the City University of Centre National de la Recherche Scientifique (CNRS),
New York, New York, NY, USA Universite ́ de Lyon, Villeurbanne, France
A Caro-Quintero PD Polly
University of Texas, Austin, TX, USA Indiana University, Bloomington, IN, USA
RL Carroll FR Davis
Redpath Museum, McGill University, Montreá l, QC, Florida State University, Tallahassee, FL, USA
Canada
GK Davis
F Casey Bryn Mawr College, Bryn Mawr, PA, USA; and The
Park Ave, Merced, CA, USA College of New Jersey, Ewing, NJ, USA
 List of Contributors xv

MN Dawson S Edwards
University of California, Merced, CA, USA Harvard University, Cambridge, MA, USA
AM Dean SP Egan
Laboratory of Microbial Evolution, College of Ecology Rice University, Houston, TX, USA
and Evolution, Guangzhou, PR China; and
BioTechnology Institute, St. Paul, MN, USA Y Eglit
Dalhousie University, Halifax, NS, Canada
JH Degnan
University of New Mexico, Albuquerque, NM, USA IM Ehrenreich
University of Southern California, Los Angeles, CA,
S Devillard USA
Universite ́ Claude Bernard Lyon 1, Villeurbanne, France
E Eizirik
A Dhar ̂ cias, PUCRS, Porto Alegre, Brazil
Faculdade de Biocien
University of Washington, Seattle, WA, USA
S Engen
S Dhole Centre for Biodiversity Dynamics, Trondheim, Norway
University of North Carolina at Chapel Hill, Chapel
Hill, NC, USA JR Etterson
University of Minnesota Duluth, Duluth, MN, USA
GP Dietl
Paleontological Research Institution, Ithaca, NY, USA; WJ Ewens
and Cornell University, Ithaca, NY, USA The University of Pennsylvania, Philadelphia, PA, USA
MR Dietrich CM Fagan
Dartmouth College, Hanover, NH, USA Keene State College, Keene, NH, USA
U Dobrindt DJ Fairbairn
Institute of Hygiene, University of Münster, Münster, University of California, Riverside, CA, USA
Germany; and Interdisciplinary Center for Clinical
Research (IZKF), University of Münster, Münster, R Faria
Germany Institute of Evolutionary Biology (Universitat Pompeu
Fabra-CSIC), Barcelona, Catalonia, Spain; and CIBIO,
E Domingo Centro de Investigac- ão em Biodiversidade e Recursos
Centro de Biología Molecular "Severo Ochoa" (CSIC- Genet́ icos, InBIO, Laboratoŕ io Associado, Universidade
UAM), Consejo Superior de Investigaciones Científicas do Porto, Vairão, Portugal
(CSIC), Madrid, Spain; and Centro de Investgación
Biomédica en Red de Enfermedades Hepáticas y TE Farkas
Digestivas (CIBERehd), Barcelona, Spain University of Connecticut, Storrs, CT, USA
K Donohue JL Feder
Duke University, Durham, NC, USA University of Notre Dame, Notre Dame, IN, USA
M Douhard A Feduccia
Universite ́ Claude Bernard Lyon 1, Villeurbanne, France; University of North Carolina, Chapel Hill, NC, USA
and Universite ́ de Sherbrooke, Sherbrooke, QC, Canada
EJ Feil
MP Dunn University of Bath, Bath, UK
St. John’s University, Jamaica, NY, USA
P Fields
I Dworkin University of Basel, Basel, Switzerland
McMaster University, Hamilton, ON, Canada
JL Fitzpatrick
D Dykhuizen University of Manchester, Manchester, UK
Stony Brook University, Stony Brook, NY, USA
CI Fraser
E Edeline Australian National University, Canberra, ACT,
Sorbonne Universités/UPMC Univ Paris 06/CNRS/ Australia
INRA/IRD/Paris Diderot Univ Paris 07/UPEC/Inst.
d’Ecologie et des Sciences de l’Environnement – Paris JV Freudenstein
(iEES-Paris), Paris, France The Ohio State University, Columbus, OH, USA
Subject Classification List
Animal Diversification
Amniotes, Diversification of
Amniotes, the Origin of
Animal: What is an Animal?
Bird Flight Origins
Birds, Diversification of
Cambrian Explosion: A Molecular Paleobiological
Overview
Complexity, the Role of Oxygen in Evolution of
Homo, Diverification of
Insects and Ecdysozoa, Diversification of
Land Animals, Origins of
Land Vertebrates, the Origin and Evolution of
Lophotrochozoa, Diversification of
Mammalian Diversification
Mammals Everywhere
Mammals, Origin of
Metazoans, Origins of
Vertebrates, the Origin of
Applied Evolution
Basic Science and Evolutionary Biology
Conservation Biology, Evolution and
Evolution and Agriculture I. The Evolution of
Domestication
Evolution and Agriculture II. Evolutionary
Applications to Breeding
Evolutionary Computation
Evolutionary Medicine: I. An Overview and
Applications to Cancer
Evolutionary Medicine II. Use of the Comparative
Method and the Animal Model
Evolutionary Medicine III. Mismatch
Evolutionary Medicine IV. Evolution and Emergence
of Novel Pathogens
Human Life Histories, Evolution and
Invasive Species, Evolution and
Pest Management, Evolution and
Philosophy, Evolutionary Biology and
Responses to Climate Change, Evolution and
Security, Evolution and
Coevolution
Antagonistic Interspecific Coevolution
Coevolutionary Fitness Landscapes
Coevolution, Introduction to
Commensalism, Amensalism, and Synnecrosis
Cospeciation
Ecological Fitting and Novel Species Interactions
in Nature
Endogenous Retroviruses and Coevolution
Endosymbiotic Theory
Geographic Mosaic of Coevolution
Intraspecific Coevolutionary Arms Races
Microbiome
Mitochondrial and Nuclear Genome
Coevolution
Mutualism, the Evolutionary Ecology of
Predation and Parasitism
Sequential Speciation
Symbiosis, Introduction to
Evo-Devo
Adaptive Radiations: Insights from Evo-Devo
Cellular Behaviors Underlying Pattern Formation
and Evolution
Developmental Biases on Morphological
Evolvability
Developmental-Genetic Toolkit for Evolutionary
Developmental Biology
Developmental Mechanisms Controlling Cell Fate,
Evolution of
Developmental Paleontology and Paleo-Evo-Devo
Developmental Plasticity and Phenotypic
Evolution
Ecological Evolutionary Developmental Biology
Gene Networks Driving Development, Conservation
and Evolution of
Genome Evolution’s Role in Developmental
Evolution
Genotype to Phenotype: Insights from Evo-Devo
Model Systems: The Key Roles of Traditional and
New Models in Evolutionary Developmental
Biology
Modularity and Integration in Evo-Devo
Novel Structures in Animals, Developmental
Evolution of
Novel Structures in Plants, Developmental
Evolution of
Phylogenetic Approach to Studying Developmental
Evolution: A Model Clade Approach
Regulatory and Coding Changes in Developmental
Evolution, Roles of
xxvii
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lain for years within the thick volume. They might have
been there when Professor Lavers bought the encyclopædia.
Michael knew that he bought many of his books second-
hand. Besides, if he had left the notes there, they would
have been missed, and questions asked and search made.
Evidently no one had missed the money.

Why, then, Michael asked himself, need he proclaim what he

had found? If a man found a sum of money, he had a right
to keep it till some one claimed it, and could prove that it
was his. If Mrs. Lavers came and told him that she had lost
this money, he would restore it to her at once. But till then
he had a perfect right to retain it. At any rate, he would do
nothing hastily. He would wait and see what happened. So
Michael locked the notes within his desk and tried to go on
with his work as usual. But this was not easy. He could not
forget that the notes were there, though he tried hard to do
so. Nor, in spite of the many excellent arguments by which
he strove to persuade himself that he was acting rightly,
could he get the better of an uneasy sense that he was
swerving from the path of rectitude.

A week passed by, and the roll of notes remained within

Michael's desk. He was still trying to persuade himself that
he was justified in retaining them, and he still found that
the voice of conscience would not endorse his arguments.
There were moments when that voice would say to him that
his action in keeping money that was not his was little
better than stealing.

One evening Michael Betts, goaded by these irritating

suggestions, started to walk up Gower Street, with the half-
formed intention of calling at Mrs. Lavers' house and asking
to see her, that he might tell her what he had found in the
old encyclopædia. The struggle within him was still so
strong, the love of gain contending so fiercely with the love
of integrity, that it is probable he would in any case have
turned back when he reached Mrs. Lavers' house. But when
he came to the door he found the steps littered with straw
and paper; there was no light in the windows, and when he
lifted the knocker, its fall resounded hollowly through the
now empty house. Mrs. Lavers and her children had gone
away, and the house was no longer a home.

Michael's feeling was one of relief.

"Now there's an end to the matter," he said to himself.

"She's gone away, I don't know where, and it's not in my
power to tell her. It's clear she knows nothing about the
money, and she does not want it. I have therefore a perfect
right to keep it."

And hugging this thought to his heart, and congratulating

himself on the lucky thing he had done when he purchased
the professor's library, Michael turned homewards.

As he approached his shop, he saw a girl standing beneath

the lamp-post at the corner. She was a girl about fifteen
years of age, with bright eyes showing beneath the thick
black fringe which covered her forehead. Her cheeks were
flushed by the cold wind, which, however, she did not seem
to mind as she stood there. A large white apron covered her
dark gown; she wore a violet woollen shawl crossed over
her chest, and a hat with many feathers was on her head.
She turned and looked curiously at Michael as he passed
her, eyeing him so intently that he was conscious of her
scrutiny, and resented it. As he was opening the door of his
shop, she came to the head of the steps and called out in
loud, clear tones:

"I say, are you Michael Betts?"

"That is my name, certainly," replied Michael, with dignity,
"but I cannot see what business that is of yours."

"Maybe not. And yet, p'raps it is my business. Maybe I know

more of you than you think for, Michael Betts."

"Then you know I'm a respectable man, and have nothing

to say to a girl like you," returned Michael angrily.

"Respectable, indeed!" cried the girl hotly. "I don't know

about your being so mighty respectable, but I'd have you
know, other folks can be respectable besides yourself. You
might practise a little civility along with your respectability."

Michael closed his door sharply, cutting short this tirade.

The girl made an angry, defiant gesture in his direction, and
then ran off.

"The hussy!" said Michael to himself. "How did she get my

name so pat, I wonder? I hope none of the neighbours
heard her calling it out. Anyhow, they all know me for a
respectable man."

Suddenly with the thought his face flushed, and a pang of

shame smote him. Was he indeed a respectable man?
Would they respect him if they knew how he had kept the
notes? Could he say that in that instance he had acted
perfectly on the square? Alas! His conscience convicted him.
He was no longer satisfied of his own probity. He had given
his self-respect in exchange for those fifty pounds.

CHAPTER VI
 AN UNWELCOME ENCOUNTER

MICHAEL paid the fifty pounds into his bank, and had the
satisfaction of seeing them entered in his pass-book. He
was so much richer than he had expected to be, yet
somehow he did not feel richer, but poorer. He had rather
the feeling of one who had suffered loss. There was a stain
upon his conscience, and a weight upon his mind; yet, with
the strange perversity of human nature, he would not own
this to himself. He still professed to believe himself justified
in keeping the money he had found. He clung to it, and
liked to think how it had swelled his balance at the bank,
even whilst he knew that he should be filled with shame, if
any one should ever learn how he had come by that money.

In Michael's lonely life there was no one save Mrs. Wiggins,

the charwoman, to observe how he lived, and mark the
variations in his moods. She began at this time to observe a
change in the bookseller. He had never been what she
would call a "pleasant-speaking" gentleman. She had
always found him short of speech, irritable, and disposed to
snub her whenever she attempted to inform him as to the
gossip of the neighbourhood; but now he was positively
surly in his manner towards her, and so quick of temper
that it was almost more than she could "put up" with.

One morning, she thought she was giving him intelligence

in which he could not fail to be interested, when she said:

"I've 'eard where that lady, the professor's widow, as you

bought so many books of, 'as gone to live."

Her words startled Michael, and he turned his eyes on her

without speaking. She took his silence as an encouragement
to proceed.
"She's gone to live in Clarendon Gardens. That's not a very
nice place, is it? And even there, she 'as only part of a
'ouse. It seems she's quite poor now she's lost 'er 'usband.
She's sent away 'er servants, and keeps only a bit of a girl
now. She washes and dresses the children 'erself, and does
most of the cooking—and she such a lady, too! It's 'ard,
ain't it? I 'eard it all from my cousin, who works for 'er
landlady."

"I wish to goodness your cousin would mind her own

business, or that you would keep her gossip to yourself!"
exclaimed Michael angrily. "How can I help it, if Mrs. Lavers
is poor and has to do without servants? I've done what I
can for her in buying her books. What more can I do?"

"Oh, lor me! Nobody would expect you to do anything for

her," exclaimed Mrs. Wiggins, with unconscious satire. "I
just thought you might like to know about the poor lady.
You need not turn so fiercely on me, Mr. Betts, for giving
you a piece of news."

"Keep your news till it's wanted, and mind your own
business," responded Michael crossly.

Mrs. Wiggins took up her dust-pan and brush, and retreated

with much clatter into the back premises, muttering words
which were not complimentary to her employer.

"Do, indeed! I'd like to see 'im ever do anything for anybody
besides 'isself, the close-fisted old curmudgeon. I'd be sorry
for a mouse that had to live on 'is leavings. The idea of 'is
turning on me like that, as if a body couldn't speak about
nothin'."

It was impossible for Mrs. Wiggins to understand how her

words had for Michael the force of an accusation. She did
not know of the secret consciousness which they awakened.
"So, then, it was a poor woman whom he had defrauded,"
said the voice of Michael's conscience.

"Defrauded! What an absurd idea!" the voice of his other

self responded. "There was no fraud in the matter; the
money had never been Mrs. Lavers'. A man had a right to
keep what he found, unless he knew it to belong to some
one else."

But reason as he might with himself, the information

imparted by Mrs. Wiggins had disturbed Michael's mind.
An incident which occurred a few days later further
destroyed its peace.

Business had taken Michael to the south side of the river,

and he was returning late in the evening across one of the
bridges on his way back to Bloomsbury, when his attention
was arrested by the appearance of a man who stood leaning
against one of the parapets and coughing violently. Michael
started, and a thrill ran through him. He had so long
thought of his brother as dead, that the sight of this man,
who bore so striking a resemblance to him, affected him
almost with terror, as though he conceived him to be a
ghost.

To be sure, he had grey hair, and Frank's had been brown

when Michael last saw him, and his form was pitifully bent
and wasted; but still the resemblance was there, and so
strong that Michael involuntarily stood still as he saw him,
whilst his heart began to beat more painfully than was
pleasant. The same instant the man ceased coughing; he
lifted his head and saw by the light of the gas-lamp on the
parapet above him the man who stood at his side. A low cry
of wonder—and was it of pleasure?—escaped him. He
moved a step nearer, exclaiming eagerly:

"Michael! Michael!"

"You don't mean to say it is you?" exclaimed the other, in

tones which expressed no pleasure at the meeting. "You,
Frank, after all these years! I thought you were dead."

"And perhaps hoped that I were," returned the other,

retreating a step or two, whilst an air of hopelessness came
over him again. "Well, it's no wonder. I'm not a brother you
can be proud of."

Michael looked at him for a moment ere he made reply. The

man appeared thin, and cold, and ill; but his was not one of
the most abject-looking of the forms to be seen abroad in
London. His clothes, though worn and threadbare, were
decently tidy.

"You've only yourself to thank for being what you are,"

Michael said. "Drink and gambling and bad company bring a
man to this."

"I've given up the drink, and gambling too, thank God!" said
his brother. "I've been a teetotaler for more than a year,
Michael."

"I'm glad to hear it," replied Michael, his tone implying that
he doubted the statement. "But if that is so, how do you
come to be in such low water? How do you live?"

"I can scarcely tell you how I live," returned the other. "I
shouldn't live at all, if it were not for my little girl."

"Your little girl!" exclaimed Michael. "You don't mean to say

that you've been so foolish as to marry?"

"I married many years ago, and I had one of the best of
wives, though, God forgive me! I was often a brute to her.
It was foolish of her to take me, no doubt, but I could never
regret it. Whilst she lived, things were better with me; but
when she died I went all wrong again. And now, when I fain
would live a different life, I can't find any one willing to give
me a chance."

"You could surely get work, if you exerted yourself."

"Could I? You don't know how hard it is to get work in

London. And who would employ a man like me, when there
are plenty of big, strong fellows to be hired? I haven't the
strength to lift or carry. I'm good for very little now. I was in
the Infirmary for three months. They patched me up a bit
there, but I'm not much the better. If it were not for my
little girl, I shouldn't be living now. She works at the match-
making, and she keeps me more often than I keep her,
bless her!"
A feeling of cold disgust was creeping over Michael Betts.
There was no room for pity in his heart. He was conscious
only of intense annoyance that such a being as this should
be his brother.

"I'm sorry for you," he said loftily, "but it's your own fault.
You made your bed, and you must lie upon it. I did what I
could for you years ago, and ill you repaid me for what I
did. Now I must wipe my hands of you. But here is a shilling
for you."

He held out the coin as he spoke, but the chilled fingers of

the other made no attempt to take it. The shilling fell from
them to the pavement, and the ill-clad man did not pick it
up.

"I don't want your money, Michael," he said hoarsely. "I

could not help speaking to you when I saw you by my side;
but don't think that I should ever have sought you out or
asked you to do anything for me again. I know too well that
I deserve nothing from you."

"That is well," said Michael coldly. "I must confess that had
you come, you would not have found a welcome from me,
since the last time you were at my place, you left me with
good cause to regret your visit."

He turned on his heel without giving another glance at his

unhappy brother.

"Michael, Michael," the weak voice called after him.

But Michael strode on and paid no heed. He did not slacken

speed till he was the length of several streets from the river.
Then he paused and drew a deep breath as he wiped the
heat from his forehead.
"To think that he should turn up again!" he said to himself.
"And I thought he was dead! But no, here he is again, and
in the same deplorable plight. A pretty sort of a brother!"

And Michael knew that in his heart, he wished that his

brother were dead.

"Of course, it's the same old story," he continued. "He's a

sorry scamp, and always will be. It's all gammon about his
becoming a teetotaler and trying to lead a better life. I don't
believe a word of it. No, that won't go down with me. I only
hope he'll keep away from me, as he says!"

Thinking thus, Michael arrived at his shop. It could hardly

be called his home. He drew out his key, unlocked the door,
and went in. He struck a match and kindled a lamp which
he had placed just inside the door. Seen by its dim light the
shop looked gloomy indeed. He passed through to the little
room at the back. This, too, was littered with books. The
fire had fallen out, and the place looked unhomelike and
cheerless. Michael shivered. He became conscious of
weariness and depression. He placed the lamp on the table
and looked about for some means of rekindling the fire. The
flame flickered feebly. Mrs. Wiggins had forgotten to
replenish the oil.

As he looked about him in the dim light, Michael's eyes fell

on the old leathern armchair which stood beside the
fireplace. He had known that ancient piece of furniture as
long as he had known anything. His mother, as her health
grew feeble, had sat in it constantly, and now, as Michael
glanced at it in the uncertain light, it suddenly appeared to
him that he saw her form seated again in the old armchair.
Most vividly, he seemed to see her sitting there with her
white muslin cap resting on the dark hair which had been
untouched by silver when she died, her little grey shawl
upon her shoulders, and her hands busied with the socks
she was perpetually knitting for her sons. Only an instant, it
was ere the lamp flame leaped up, and he knew that he had
been deceived by his fancy, but that instant left its
impression. He sank on to a chair, shaken in body and mind.

How his mother would have grieved if she could have

foreseen what Frank's future would be! Her darling son, a
poverty-stricken tramp wandering about in search of a job!
And he, Michael, had promised her that he would always be
good to Frank. Had he kept that promise? Was he keeping it
when he turned away from his brother on the bridge, telling
him that he had made his bed and he must lie on it?

Michael sat for some minutes absorbed in painful thought.

Then, resolutely turning his back on the old armchair which
had awakened such unwelcome reflections, he began to
pace to and fro the floor, for his limbs had grown benumbed
by the chill of that fireless room.

"No," he said, half aloud, "I have not failed in my duty

towards my brother. I have done all that could be expected
of me. No one would do more. I've helped him again and
again, only to be repaid by the basest ingratitude. Now, I
will do no more."

CHAPTER VII
IN THE GRIP OF PAIN
MICHAEL went to bed that night feeling thoroughly chilled in
body and miserable in mind. Sleep would not come to him,
nor could he get warm, though he put all the wraps he
could find upon his bed. As he turned and tossed upon the
mattress throughout the night, unable to find ease, the
form of his brother as he had seen him on the bridge was
ever before his eyes. What a wretched thing Frank had
made of his life! It was all his own fault, for he had had a
good chance when he was young. And then to think of his
marrying, when he had not enough to keep himself! What
improvidence!

Michael wondered what the little girl was like of whom his
brother had spoken. With the thought, the image of the
professor's sunny-faced, winsome little daughter rose
before his mind. But it was not likely that his niece was at
all like her. A girl who worked at match-making! Well, it was
hard on a respectable, hard-working man to have relatives
of such a description. Michael wished that he had taken
another way home than the way that had led him across
that bridge. He had been so much more comfortable under
the persuasion that his brother was dead.

When Michael woke from the brief sleep that visited him
towards dawn, it was past the hour at which he usually
rose. But when he would fain have bestirred himself in
haste, he found it impossible to do so. His back and limbs
seemed to have grown strangely stiff, and when he tried to
move, an agonising pain shot through them. He struggled
against the unwelcome sensations, and did his best to
persuade himself that he was suffering only from a passing
cramp. But the pain was terrible. He felt as if he were held
in a vice. How to get up he did not know; but he must
manage to do so somehow. It was necessary that he should
get downstairs to open the door for Mrs. Wiggins. Setting
his teeth together and often groaning aloud with the pain,
he managed at last to drag himself out of bed and to get on
his clothes. It was hard work getting downstairs. He felt
faint and sick with pain, when at length he reached the
lower regions. It was impossible to stoop to kindle a fire. He
sank into the old armchair and sat there bolt upright, afraid
to move an inch, for fear of exciting fresh pain, till he heard
Mrs. Wiggins' knock. Then he compelled himself to rise, and
painfully dragged himself forth to the shop door, where he
presented to the eyes of the charwoman such a spectacle of
pain and helplessness as moved her to the utmost
compassion of which she was capable.

"Dear me! Mr. Betts, you do look bad. It's the rheumatics,
that's what it is. I've 'ad 'em myself. Is it your back that's
so very bad? Then it's lumbago, and you'd better let me
iron it."

"I'll let you do nothing of the kind!" cried the old man
angrily. "Do for pity's sake keep away from me; I can't bear
a touch or a jar. Make haste and light me a fire, and get me
a cup of tea. That's all I want."

"You ought to be in bed, that's where you ought to be," said

Mrs. Wiggins. "Just let me help you upstairs, now do, and
then I'll bring you a cup of tea, all hot and nice."

"How can I go to bed?" he asked impatiently. "Who is to

look after the shop if I go to bed?"

"Oh dear! That's a bad look out. Have you no one to whom
you could send to come and take your place? Have you no
brother now who would come to you?"

"Of course I have not!" he cried, annoyance betraying him

into a quick movement, which was followed by a groan of
pain. "I do wish you would attend to your business, and not
ask me stupid questions."
"Stupid or not, you're not fit to stand in that shop to-day.
Why, you couldn't lift a book without wincing. Ah me! It's
bad enough to be lonesome when you're well, but it's sad
indeed when you're ill to have no one to do a thing for you."

"Do be quiet," he cried; but Mrs. Wiggins, excited by the

sight of his suffering, was not disposed to hold her tongue
till she had fully relieved her mind. She began to suggest
one patent remedy after another, and showed a remarkable
acquaintance with all the quack medicines of the day. But
Michael refused to try any of them. He had hardly ever been
ill in his life, and he did not in the least know what to do
with himself, or how to bear his pain.

It grew worse as the day wore on, and though Mrs. Wiggins
made him a good fire, and he sat over it, he could not get
warm. It was hopeless to think of attending to business. He
was obliged to give in at last, and allow the shop door to be
closed, whilst he was ignominiously helped up to bed by
Mrs. Wiggins.

"Now you'd better let me send for a doctor," she said.

"No, indeed," he replied with energy. "I want no doctor yet.

You don't suppose I can afford to send for a doctor every
time I have an ache or pain?"

"Maybe not," she said, "but it seems to me you're pretty

bad now."

"Folks don't die of rheumatics," he said.

"Oh, don't they?" she returned. "I've known a many cases

in which they 'ave. Rheumatics is no joke. They're apt to
seize on the 'eart, don't you know?"

And with this comforting reflection she left him.

As Michael lay there in pain and misery, he was reminded of
a childish voice, which had said:

"I should think you would cross soon, Mr. Betts, for you are
so very old."

Could it be that he was drawing near to the hour when he

would have to cross that river of death?

The pain grew worse. From shivering, he passed into

burning fever. Mrs. Wiggins felt very uneasy when the time
came for her to go home.

"I don't like to leave you, Mr. Betts, I don't indeed," she
said. "I can't think it's right for you to be all alone in this
house. If you was to be took worse—"

"I shall not be worse," he said hoarsely; "the pain can't be

worse than it is now, and it would not make it any better to
have some one else in the house."

"But I wish you'd let me stay with you," she suggested.

"Just let me go and tell my 'usband, and come back
directly."

"No, no, no," he said, for he was weary of her attentions.

"Take the house key with you, and lock the door on the
outside, and come as early as you can in the morning. But
first bring a big jug of cold water, and set it here beside the
bed. I'm so thirsty, I could drink the sea dry, I believe."

"Ah, you've got fever, that's what you've got," replied Mrs.
Wiggins. "Well, I suppose you must have your way."

So she did what he told her, and then went home.

But she was so impressed with the fact of his being very ill
that she bestirred herself unusually early the next morning,
and was at Mr. Betts' shop quite an hour before the time at
which she generally appeared. She unlocked the door and
let herself into the shop. Already the place seemed to have
a deserted look. The dust lay thick on the books. Mrs.
Wiggins went quickly up the steep staircase and knocked at
the door of the attic, which was Michael's bedroom. She
knocked, but there was no response to her knock. She
knocked again more loudly, but Mr. Betts did not bid her
enter, only she could hear his voice talking in strange, far-
off tones. After a little hesitation she turned the handle and
entered the room. Michael Betts lay on the bed, his face
flushed with fever, his brows contracted with pain, his eyes
wild and dilated. He was talking rapidly and incoherently.

"Well, Mr. Betts," she said, as she approached the bed, "and
'ow do you find yourself this mornin'?"

But he paid no heed to her words. They fell on unconscious

cars. He went on talking rapidly; but she could not
understand what he was saying. As she bent nearer, she
could catch a few words now and then; but there seemed
no connection in them.

"The river—it's cold and deep—there's a little girl on the

other side—Oh, the pain—the awful, burning pain!—Oh,
water—give me water—there's water in the river—I don't
care if he is my brother. Give me water—water, I say. What
are you telling me about the money?—It's mine—I have a
perfect right to it. Oh, this pain! The water—the river."

"Lor' bless me! He's right off 'is 'ead," said Mrs. Wiggins;
"'e's in a raging fever. It's no good speakin' to 'im. I must
just fetch a doctor, whether 'e likes it or not."
A little later, a doctor stood beside Michael's bed. He
pronounced it a severe case of rheumatic fever, made some
inquiries respecting the circumstances of his patient,
prescribed for him and departed, saying that he would send
a nurse to look after him, since he needed good nursing
more than medicine. The doctor showed his wisdom in so
acting, for had Michael been left to the tender mercies of
Mrs. Wiggins, well-meaning though they were, he would
probably never have risen from his bed. As it was, he had a
hard struggle ere the force of life within him overcame the
power of disease. He was very ill, and at one time, the
medical man had but faint hope of his recovery.

He was confined to his bed for weeks, and the little book-
shop remained closed the while, for Michael was far too ill
to give any directions as to what should be done about the
business. After the fever left him, he was as weak as a
baby: too weak to care about anything, so weak that every
effort was painful, and he felt as if he had not the heart to
struggle back to life again. Yet he shrank from the thought
of death, and one of the first questions he asked his nurse,
when he was able to think and speak connectedly, was if
she thought he would recover.

"Yes," she said cheerfully, "you've turned the corner now.

All you need is feeding up. Every day will see a change for
the better now, if you're good and do as I tell you."

"I'll try," said Michael, quite meekly. "You've been very good
to me, nurse."

"I should be a poor kind of nurse, if I hadn't been good to

you," she replied. "It's my business to look after people
when they're ill, and I have to take the greatest possible
care of them, or things would go seriously wrong."
"I never had any one do as much for me as you've done,"
said Betts; "not since I lost my mother, I mean."

"Then you never had a wife?"

"No. I've never had time to think about getting one."

The nurse laughed.

"You're a strange man," she said; "but now eat some of this
jelly."

"It is good," said Michael; "really I don't know as I ever

tasted anything better."

"It's real, strong calf's-foot jelly, and it was made by a lady

on purpose for you."

"That can't be," said Michael, looking at her in surprise;

"there's no lady would make jelly for me. You must be
making a mistake."

"Indeed I am not. You have more friends than you think.

She came here and gave it into my very hands, so I must
know. She said she'd heard from the doctor that you were
ill, and she felt sorry for your being all alone. She said she'd
done business with you, and when she saw the shop shut
up, she asked about you."

"Well, I never!" said Michael. "I can't imagine who it could

be."

"She said her name was Lavers."

"What?" exclaimed Michael in amazement. "What name did

you say?"

"Lavers—Mrs. Lavers."
Michael gave a groan.

"What is the matter?" asked the nurse, turning to look at

him; "have you the pain again?"

"No," he muttered, "not that sort of pain; but I wish she

had not done it."

"Why, you ungrateful man!" exclaimed the nurse.

Michael made no reply. A hot flush of shame was dyeing his

cheeks, and mounting to his forehead. The nurse observed
it with some anxiety. She took his hand to feel his pulse.
Was the fever about to return?

CHAPTER VIII
THE BURDEN MAKES ITSELF FELT

"MR. BETTS," said the nurse, three days later, as she came
into the room, "that lady is downstairs, and she wants to
know if you would like to see her."

"What lady?" asked Michael, though he thought he knew.

"The lady who made you the jelly—Mrs. Lavers, of course."

"Oh no," said Michael, shrinking down and drawing the

bedclothes closer about him, as if he would fain hide
himself; "I don't want to see her. I can't have her coming
here. Tell her so, please."