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i

Behavioral Ecology of the Eastern

Red-​backed Salamander
ii
iii

Behavioral Ecology of the

Eastern Red-​backed
Salamander
50 Years of Research

BY ROBERT G. JAEGER,

BIRGIT GOLLMANN,

C A R L D . A N T H O N Y,

CAITLIN R. GABOR

AN D

NANCY R. KOHN

1
iv

1
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 certain other countries.

Published in the United States of America by Oxford University Press

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© Oxford University Press 2016

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a retrieval system, or transmitted, in any form or by any means, without the
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above should be sent to the Rights Department, Oxford University Press, at the
address above.

You must not circulate this work in any other form

and you must impose this same condition on any acquirer.

Library of Congress Cataloging-​in-​Publication Data

Names: Jaeger, Robert, author.
Title: Behavioral Ecology of the Eastern Red-backed Salamander / by Robert G. Jaeger,
Birgit Gollmann, Carl D. Anthony, Caitlin R. Gabor, and Nancy R. Kohn.
Description: New York, NY : Oxford University Press, 2016. |
Includes bibliographical references and index.
Identifiers: LCCN 2015050051 | ISBN 9780190605506
Subjects: LCSH: Plethodon cinereus—Behavior. | Plethodon cinereus—Ecology.
Classification: LCC QL668.C274 J34 2016 | DDC 597.8/5—dc23
LC record available at https://lccn.loc.gov/2015050051

9 8 7 6 5 4 3 2 1
Printed by Sheridan Books, Inc., United States of America
v

For Professor Jane Brockmann, who initially suggested and encouraged this
book, and Professor Murray Itzkowitz, whose juxtaposition of field and
laboratory experimentation was an inspiration for our research
vi
vii

CONTENTS

List of Figures xi
Acknowledgments xiii
About the Authors xv

1. Prelude 1
1.1. Bob Jaeger meets the eastern red-​backed salamander,
Plethodon cinereus 1
1.2. An introduction to red-​backed salamanders 4
1.3. The plot of our research program 6
1.4. Comments concerning methodology and statistical paradigms 10

2. Interspecific competition between P. cinereus and P. shenandoah 13

2.1. Ecological studies 14
2.2. Behavioral experiments 18
2.3. Selected recent research by others: Interspecific competition 21

3. Intraspecific territoriality by P. cinereus 23

3.1. Definition and theory 23
3.2. Distribution and prey availability 24
3.3. Site tenacity by P. cinereus 27
3.4. Determining sex and defining behavioral patterns 28
3.5. The use of odors and dear enemy recognition 30
3.6. The expulsion of intruders 34
3.7. Testing territoriality in the forest 35
3.8. Variables that affect territorial contests 42
3.8.1. Length of residency 42
3.8.2. Body size 43
3.8.3. Body size without residency 45
3.8.4. Sex and reproductive condition 46
3.8.5. Intruder number 47
3.8.6. Tail condition 49
3.8.7. Food quantity 50
viii

viii Contents

3.8.8. Food quality 51

3.8.9. Signal honesty 53
3.9. Life history traits and territorial contests 54
3.9.1. Variation in reproductive success among females 55
3.9.2. Payoffs to and RHP of territorial owners 56
3.9.3. Resource acquisition and energy allocation 57
3.10. S easonal and geographic variation in territorial
agonistic behavior 59
3.10.1. Seasonal variation 59
3.10.2. Geographical variation 61
3.11. Selected recent research by others: Intraspecific territoriality 63

4. Foraging tactics by P. cinereus within territories 65

4.1. Foraging on live versus dead prey 66
4.2. Diet breadth 67
4.3. Optimal prey choice 67
4.4. Territorial and foraging behavioral conflicts 69
4.5. Assessing prey densities 70
4.6. Judging prey profitabilities 72
4.7. Conflicts between foraging behavior and territorial defense 74
4.8. Diet diversity and clutch size 76
4.9. Selected recent research by others: Foraging tactics 78

5. Pheromonal glands and pheromonal communication by P. cinereus 80

5.1. Early studies suggest pheromones do occur 80
5.2. Do males of P. cinereus produce territorial pheromones? 81
5.3. Do females of P. cinereus produce territorial pheromones? 82
5.4. Where are those pheromones produced in males and females? 83
5.5. Focusing on the postcloacal gland 84
5.6. What information does the postcloacal gland communicate? 86
5.7. What signals do pheromones communicate? 87
5.8. Scent matching and tail autotomy 89
5.9. Do territorial pheromones aid in homing behavior by P. cinereus? 93
5.10. Are pheromones volatile? 95
5.11. Selected recent research by others: Pheromonal communication 98

6. Interspecific territoriality and other interspecific behavioral

interactions 101
6.1. Interspecific territoriality between P. cinereus and P. shenandoah 102
6.2. Rules of engagement with juveniles of P. glutinosus 105
6.3. Plethodon cinereus in an assemblage of salamanders 108
ix

Contents ix

6.4. Ecological tests of behavioral predictions: Enclosed plots

on the forest floor 111
6.5. More ecological tests of behavioral predictions: Unenclosed plots
on the forest floor 113
6.6. Character displacement: P. cinereus versus P. hoffmani 114
6.7. Competition between P. cinereus and P. hubrichti 117
6.7.1. Habitat niche partitioning? 118
6.7.2. Microhabitat niche partitioning? 118
6.7.3. Intra-​and interspecific competition? 120
6.7.4. Intra-​and interspecific territoriality? 122
6.7.5. Summary 126
6.8. Diversity of behaviors by P. cinereus toward other species 128
6.9. Selected recent research by others: Interspecific territoriality 129

7. Intraspecific social behavior within P. cinereus 131

7.1. Interactions of adults and juveniles in the forest
and in the laboratory 131
7.2. Distributions of adult males and females 134
7.3. Microdistributions of adults and juveniles 136
7.4. Female–​female interactions 136
7.5. Male–​female behavioral interactions in the forest 138
7.6. The ESS dating game 139
7.7. Males, females, and feces 141
7.8. Females prefer larger males 143
7.9. Males and females prefer familiar opposite-​sex individuals 144
7.10. Social monogamy 146
7.11. Mutual mate guarding 149
7.12. Sexual coercion 150
7.13. Imperfect information during sexual discrimination? 152
7.14. Relationship value and conflict resolution 153
7.15. Natural versus forced partnerships 155
7.16. Females are often genetically polyandrous 156
7.17. Switching from social monogamy to social polygamy 157
7.18. Brooding behavior and neonates: Kin recognition? 160
7.19. What 3,487 uniquely marked salamanders reveal
about social relationships 162
7.19.1. Basic population ecology 163
7.19.2. Size distribution and growth rates 165
7.19.3. Sexual maturity and female fecundity 165
7.19.4. Population size 166
7.19.5. Sex ratio 166
x

x Contents

7.19.6. Intersexual associational behavior 167

7.19.7. Consequences of tail autotomy 168
7.20. A preliminary model of social organization within P. cinereus 168
7.21. Selected recent research by others: Social behavior 170

8. Predator–​prey interactions between P. cinereus and a snake 172

8.1. Can P. cinereus detect the snake visually or chemically? 173
8.2. Can the snake detect chemical cues from P. cinereus? 174
8.3. Naïve snakes recognize odors of P. cinereus 175
8.4. Tail autotomy deceives the snake 175
8.5. The snake follows the trail of P. cinereus 176
8.6. The predator–​prey evolutionary arms race 177
8.7. Selected recent research by others: Predator–​prey arms races 178

9. Cognitive ecology of P. cinereus 180

9.1. Numerical discrimination by P. cinereus 180
9.2. Both learning and heritability affect foraging ability 183
9.3. Displacement of territorial aggression 186
9.4. The impact of familiarity on salamander behavior 187
9.5. Individual recognition memory 189
9.5.1. The formation of individual recognition memory 190
9.5.2. Sensory modalities used during recognition 193
9.5.3. Do distractors inhibit individual recognition memory? 195
9.5.4. Overview 198
9.6. Selected recent research by others: Cognitive ecology 198

10. Coda: Synthesis and social behaviors by P. cinereus 200

10.1.  Behavioral variation within a population 200
10.1.1. Age 201
10.1.2. Tail autotomy 201
10.1.3. Polymorphism 201
10.1.4. Health 202
10.1.5. Bold and shy 202
10.1.6. Experimental flaws 202
10.2.  Behavioral options during contests 203
10.3.  How salamanders choose among options 204
10.4.  Definitions of social, mating, and genetic monogamy 206
10.5.  Mea maxima culpa 207

References 209
Subject Index 223
xi

LIST OF FIGURES

1.1 A young Bob Jaeger, c. 1973, in Shenandoah National

Park, Virginia 3
1.2 An adult, female red-​backed salamander, Plethodon cinereus, uncovered
in a deciduous forest at Mountain Lake Biological
Station, Virginia 5
1.3 Stacks of single salamanders in their chambers (Petri dishes)
in the laboratory at the University of Louisiana at Lafayette 10
2.1 A portion of the talus–​soil interface on Hawksbill Mountain 15
3.1 Behavioral postures used by P. cinereus during intra-​
and interspecific interactions and chemoinvestigation 29
3.2 One of our research sites at Mountain Lake Biological
Station, Virginia 35
3.3 A plot of the forest at Mountain Lake Biological Station
near where Mathis (1990a) established her censusing plots 37
3.4 A female red-​backed salamander in a Nunc bioassay dish 48
3.5 Experimental tubs similar to those used by Wise (1995)
at Mountain Lake Biological Station 58
6.1 (A) Adult Plethodon glutinosus. (B) Adult Eurycea cirrigera.
(C) Adult Desmognathus fuscus 106
6.2 Caitlin Gabor in front of the Research Building at MLBS 109
9.1 The chamber used in the numerical discrimination experiments,
redrawn from Uller et al. (2003) 181
9.2 The visual chamber used in Kohn and Jaeger (2009)
that allowed the focal salamander to see the intruder
but not chemically detect him 194
xii
xiii

ACKNOWLEDGMENTS

We thank Günter Gollmann (Universität Wien) for useful discussions while

we climbed Mount Schneeberg in the high Austrian Alps with the help of the
“Salamander” train. We also thank Terry Kohn for her assistance in typing large
portions of this book and Jenny Thibodeaux for computer assistance. Lauren
Mathews provided advice for section 10.4 and Andrea Aspbury, Douglas Fraser,
Günter Gollmann, and Sharon Wise provided critical advice as we composed
the manuscript. Nancy Kohn, Caitlin Gabor, Carl Anthony, and Bob Jaeger sup-
plied the photographs. The research reviewed here was supported by the follow-
ing universities and research stations: University of Maryland at College Park
and Shenandoah National Park (SNP; 1965–​1971); University of Wisconsin at
Madison and SNP (1971–​1974); State University of New York at Albany and SNP
(1974–​1980); University of Louisiana at Lafayette (1981–2008); and Mountain
Lake Biological Station of the University of Virginia (1981–2015). Our research
was shaped by three philosophers, K. Popper, J. Platt, and I. Lakatos, whose phi-
losophies of science led the path to the progressive research program described
herein with the ever-​fascinating eastern red-​backed salamander, Plethodon
cinereus.
xiv
xv

ABOUT THE AUTHORS

Robert G. Jaeger, Department of Biology, Utica College, Utica, New York, USA;
100 Simon Latour Road, Carencro, Louisiana, USA
Birgit Gollmann, Fakultät für Lebenswissenschaften, Universität Wien, Wien,
Austria
Carl D. Anthony, Department of Biology, John Carroll University, University
Heights, Ohio, USA
Caitlin R. Gabor (corresponding author), Department of Biology, Texas State
University, San Marcos, Texas, USA, gabor@txstate.edu
Nancy R. Kohn, Department of Biology, The College of New Jersey, Ewing,
New Jersey, USA; Department of Biology, University of Missouri-​St. Louis, St.
Louis, Missouri, USA
xvi
xvii

Behavioral Ecology of the Eastern

Red-​backed Salamander
xviii
1

Prelude

1 .1  B OB J A EG E R M E E TS THE E ASTERN RED-​B A CKED

S A L A MA NDE R , PLE TH O DO N CI NEREUS

I take the liberty here to write in the first-​person singular to convey how the
following research program began. During the late 1950s, I was an undergrad-
uate student at the University of Maryland at College Park, majoring in zool-
ogy. My initial intention was to train in ichthyology, until I realized that I was
deathly fearful of entering waters deeper than my ankles! Not many species of
fishes (especially sharks) swim in waters only 9 cm deep, so my thoughts quickly
changed to amphibians, many of which live in shallow water. Frogs and toads
had fascinated me during childhood while wandering in the forests of rural
Maryland, so I sought a faculty member who studied anurans (frogs). Luckily,
during my junior and senior years, Dr. Richard Highton hired me as a research
assistant. Dick Highton is an authority of salamandrine taxonomy, systematics,
and geographic variation with a particular interest in the family Plethodontidae,
which includes Plethodon cinereus, the eastern red-​backed salamander (hereaf-
ter the red-​backed salamander). Salamanders are not frogs, but they were close
enough for me back then. Dick Highton frequently led his flock of graduate
2

2 B ehavioral E cology of the E astern R ed - B acked S alamander

and undergraduate students on collecting trips throughout Maryland, Delaware,

Pennsylvania, West Virginia, and Virginia. He seemed to have a special passion
for red-​backed salamanders, which can be seen by his approximately 100,000
preserved specimens that now reside in the US National Museum.
I was then not interested in taxonomy, even less so after soaking my hands,
for two years, in formaldehyde while preserving Dick Highton’s salamanders.
However, I was curious about the thousands of red-​backed salamanders that we
found (and then collected) in the forests. Three observations struck me as odd.
First, many forests had far greater densities of this species than of other
co-​occurring species of caudate (tailed) amphibians. (Later, Burton & Likens,
1975, found that in a forest in New Hampshire, the red-​backed salamander
comprised 93.5% of the total biomass of all species of salamanders, with 885 to
2,367 red-​backed salamander per hectare.) I was puzzled by how one species of
salamander could maintain such large densities.
Second, while on a “good collecting day” I would find a red-​backed salaman-
der under nearly every rock in a forest, even large rocks usually housed only one
salamander, except during the spring and autumn courtship seasons when pairs
were frequently found together under rocks. This too seemed odd, because large
rocks clearly covered enough soil to house more than one or two individuals.
Third, in some forests, the geographic distribution of the red-​backed sala-
mander abruptly ended and the distribution of another same-​size congener
began. Why would two species so confront each other with very little area of
species’ overlap? This was how I first met the red-​backed salamander: puzzled
by these three observations.
As an undergraduate student, I was woefully ignorant of ecological theories
and empirical studies, because the course program then largely focused on tax-
onomy, anatomy, genetics, embryology, and physiology. So I progressed on to
the University of California at Berkeley in the early 1960s for a master’s degree.
At Berkeley, when not dodging or participating in demonstrations, I was able
to enjoy several ecology courses that expounded on the then-​current theories
of (1) population regulation, (2) intraspecific competition, and (3) interspe-
cific competition. Also, I was fortunate enough to take Peter Marler’s course in
(4) animal behavior. These four topics seemed to be pertinent, somehow, to my
previous observations of red-​backed salamanders and their densities, intraspe-
cific distributions, and interspecific interactions.
Therefore, with master’s degree in hand, I returned to College Park, Maryland,
for a doctoral degree where I worked with Dick Highton as an advisor, because
he could tell me where the salamanders were and because the department had
recently hired an ecologist and behaviorists with whom I filled out my doc-
toral committee. The resulting PhD ended with four published papers con-
cerning interspecific competition between the red-​backed salamander and the
Shenandoah salamander, which are summarized in section 2.1. However, this
3

Prelude 3

research seemed to me to be incomplete, because the interspecific ecological

interactions that I had documented seemed to have a behavioral underpinning,
which I was still too naïve to understand.
This lack of behavioral knowledge sent me to the University of Wisconsin at
Madison in 1971, blissfully supported in postdoctoral research by a three-​year
grant from the National Science Foundation to Dr. Jack P. Hailman. Jack was a
superb ethologist who taught the basics of experimental designs within a be-
havioral context to me. The postdoctoral research concentrated on phototactic
behavior of anurans, so at last I had found a niche in the biology of frogs and
toads! Yet I was also able to continue studies of the red-​backed salamander
during summers, back in Virginia at my doctoral research site (Fig. 1.1).
Finally, in 1974, armed with a total of 12 publications, it was time to earn
an honest living with a viable salary, so I obtained a position at the State
University of New York at Albany, later moving to the University of Louisiana
at Lafayette in 1981. These two universities required little teaching (one under-
graduate and one graduate seminar course per year), so much of my time could
be devoted to research with the red-​backed salamander. Most of the studies
summarized in section 2.2 through c­ hapter 10 were conducted at these two

Figure 1.1 A young Bob Jaeger, c. 1973, in Shenandoah National Park, Virginia.
4

4 B ehavioral E cology of the E astern R ed - B acked S alamander

institutions (many behavioral experiments) and at research sites in northern

New York and in Virginia (many ecological experiments). A large number of
my doctoral, master’s, and undergraduate students chose to test (often seem-
ingly wild) hypotheses concerning the behavioral ecology of the red-​backed
salamander, which led to many of the 114 publications plus three unpublished
theses and six partially unpublished dissertations discussed in this book.
Once I reached retirement in 2006, I assumed that my remaining years could
be devoted to my other pleasures: trips to operas and ballets in the United States
and Europe. While this has come to pass, research with colleagues and former
students continues to generate new manuscripts about red-​backed salamanders.
Then, in 2012, Jane Brockmann encouraged me to compose this book, integrat-
ing 50 years of research with the red-​backed salamander into a long story of
discoveries by my research group. Jane has been a dear friend since our gradu-
ate student days in the late 1960s at the University of Maryland, so I was quite
flattered by her suggestion, but not excited about writing a book alone. The ven-
ture was saved when Nancy Kohn, Birgit Gollmann, Caitlin Gabor, and Carl
Anthony offered to write some other sections of the book. The five of us hope
that this book will encourage other behavioral ecologists to discover more sur-
prises still concealed by the mysterious red-​backed salamander and, perhaps, by
other species of tailed amphibians.

1 .2   A N I N T R O DUCTI O N TO R E D -​B ACKED

S A L A MA N DE R S

The natural history, ecology, and reproductive behavior of red-​backed sala-

manders have been well summarized by Petranka (1998), Wells (2007), and
Anthony and Pfingsten (2013). Therefore, here we note only a few characteris-
tics of the species that were pertinent to our research program.
The red-​backed salamander (Fig. 1.2) is a small salamander about the thick-
ness of a pencil. Its maximum length varies geographically, but at our research
sites, fully grown adults reach approximately 45 mm snout-​to-​vent length and
90 mm total length (including its long tail). Red-​backed salamanders are lung-
less, as are all species in the family Plethodontidae, so respiration occurs ex-
clusively through skin surfaces. Consequently, plethodontid salamanders have
permeable skin requiring a moist environment, as do nearly all amphibians,
to prevent desiccation. It is this requirement that largely shapes the ecology of
the genus Plethodon, because these species are strictly terrestrial. All life stages
of the red-​backed salamander are found in the leaf litter of the forest floor
(egg, neonate, juvenile, and adult), where individuals forage for invertebrates,
but they frequently are found under rocks and logs when the leaf litter dries.
5

Prelude 5

Figure 1.2 An adult, female red-​backed salamander, Plethodon cinereus, uncovered in

a deciduous forest at Mountain Lake Biological Station, Virginia. She is ~45 mm snout-​
to-​vent length and ~90 mm total length, lying near a damp log, and in the resting and
alert (head raised) posture (front of trunk raised). Her large eyes provide enhanced
vision under dim light in the leaf litter. Note that her tail is thick (indicating stored fat
from successful recent foraging) and intact (not autotomized). Her red stripe extends
down only 40% of the tail, which probably means that the terminal 60% had been
autotomized (1+ years previously) and then regrown. Tail autotomy can occur during a
predatory attack, in which the tail, still thrashing, distracts the predator’s attention, or
during a biting attack from a territorial rival, during which the attacker usually eats the
autotomized tail. This is a striped morph of P. cinereus, but in this population individuals
vary from full-​length stripe to spotty blotches of red, orange, yellow, or tan to no stripe at
all (the leadback morph).

During extremely dry and freezing winter conditions, they move underground
following crevices in the soil; this species is not adapted to dig its own burrows.
The geographic distribution of red-​backed salamanders is enormous com-
pared to all other species of Plethodon, except for the much larger P. glutinosus
(northern slimy salamander; see Petranka, 1998, for range maps and taxonomic
controversies concerning P. glutinosus). Red-​backed salamanders are found in
most, but not all, cool, closed canopy forests throughout northeastern North
America, from southeastern Canada southward to northern North Carolina
and from the Atlantic Coast northwestward to the northern Mississippi River.
Due to the relatively low efficiency of gas exchange by cutaneous respiration,
these salamanders require cool conditions, and thus southern populations are
constrained to mountainous areas.
6

6 B ehavioral E cology of the E astern R ed - B acked S alamander

Several other species of Plethodon occur within the geographic range of red-​
backed salamanders. Some of these species have only tiny distributions (e.g.,
P. aureolus, P. hubrichti, P. nettingi, P. punctatus, and P. shenandoah), while other
species are more widely distributed (e.g., P. dorsalis, P. glutinosus, P. hoffmani,
P. richmondi, P. wehrlei, and P. welleri), according to range maps in Petranka
(1998). We were particularly interested in areas where red-​backed salamanders
overlap only slightly with other species of a similar size, and in c­ hapter 6 we
summarize our research concerning competition between P. cinereus and P. hu-
brichti, P. hoffmani, and P. shenandoah.
However, red-​backed salamanders often coexist in forests with other much
larger species of Plethodon, so we also summarize our studies, in c­ hapter 6, of
interactions between red-​backed salamanders and P. glutinosus. Many other
species of Plethodon are located outside of the geographic distribution of red-​
backed salamanders, but our research focused just on red-​backed salamanders,
their interspecific interactions, and their intraspecific social organization.

1 .3  T H E P L O T O F O U R R E SE AR C H PROGRA M

The red-​backed salamander expresses a suite of behavioral patterns that is ex-

traordinarily complex for an amphibian species (Jaeger, Gillette, et al., 2002).
This complexity of behavior has been studied since 1965 by R. G. Jaeger and
his research group leading to the publication of 114 articles thus far. We hope,
by establishing approaches and methodologies that successively reveal the
complexity of behavioral patterns in this species (e.g., the diversity of “deci-
sion rules”; Krebs, 1978), other ethologists will explore whether other species of
the caudate amphibians have evolved similarly. We make no attempt to review
the entire literature concerning P. cinereus but instead summarize our group’s
research from 1965 to 2015 with only occasional reference to publications by
others. However, ­chapters 2 through 9 end with a summary of recent research
by others to broaden the scope of our research.
While conducting our research, we attempted to follow the philosophical
guidelines (Aucoin et al., 2005) established by Popper (1959; “refutation” of
hypotheses), Platt (1964; “strong inference”), and Lakatos (1970; the “scientific
research programmes”). Popper advocated that, due to inductive reasoning, sci-
entists cannot prove anything to be true (Hume, 1748 [reprinted in 1995]); one
can only attempt to refute or falsify hypotheses and eliminate those false hypoth-
eses. Popper proposed that science progresses by forming a priori hypotheses
that are testable and refutable. If a given hypothesis is refuted, then it should be
abandoned. A limitation to falsificationism is that a single falsification may not
be enough to abandon a particular theory or research program. Given this limita-
tion, Lakatos suggested using the scientific research programme, which consists
7

Prelude 7

of replacing an old theory or hypothesis with a new model that is theoretically

progressive because it is an advance over the predecessor. According to Lakatos,
the new theory is progressive if it enables one to predict more than the predeces-
sor and is occasionally empirically progressive, in that an observation confirms
this new prediction. In so doing, a scientist’s research program progresses by re-
futing or falsifying the accepted theory, or failing to do so, as proposed by Popper.
Refutation of a null hypothesis does not mean that the one alternative hy-
pothesis is true. Scientists must consider all reasonable alternative hypotheses.
Platt (1964) proposed that a progressive research program formulates multiple
hypotheses that compete with each other. He suggested devising a critical ex-
periment where the outcome will reject at least one hypothesis and researchers
can repeat the process until the conclusions are no longer open to alternative
interpretations, thus yielding a strong inference. By making, and often rejecting,
new subhypotheses or new hypotheses (and predictions), the process is recycled.
Our investigations were composed of a progression of studies, leading from one
topic to another related topic: observations led to alternative hypotheses that led
to experiments, which then led to the next set of observations, alternative hy-
potheses, experiments, ad infinitum. We attempt to summarize our research in
this book in more or less chronological sequence to show the progression of our
hypotheses and experiments. However, we also group these experiments into
related topics (­chapters 2–​10) to tell a coherent story about P. cinereus.
The first area of research concerned interspecific competition between the
red-​backed salamander and the Shenandoah salamander (P. shenandoah, later
listed as a federally endangered species) in Shenandoah National Park, Virginia.
Plethodon cinereus is widespread in deciduous and coniferous forests of north-
eastern North America, while P. shenandoah is restricted to only three north-​
or northwest-​ facing slopes in Shenandoah National Park (the Pinnacles,
Stony Man Mountain, and Hawksbill Mountain in the Blue Ridge Mountains;
Petranka, 1998). Because they are lungless amphibians, plethodontid salaman-
ders must respire through the skin and so are susceptible to desiccation during
dry weather. To avoid desiccation between rainfalls, members of both P. cinereus
and P. shenandoah either descend into underground cavities (where prey may be
limited; Fraser, 1976) or preferentially move under rocks and logs where patches
of moisture remain for days after the last rainfall and where prey tend to aggre-
gate (Jaeger, 1970, 1979).
The parapatric distribution of these two species suggested the hypoth-
esis of interspecific competition, with P. cinereus competitively dominant to
P. shenandoah. While the latter species inhabits only three, dry, talus slopes,
P. cinereus surrounds each slope in deep, moist soil more suitable to survival
of species of Plethodon. The research on this suspected interspecific competi-
tion lasted through the late 1960s and into the early 1980s, and it is reviewed
in detail in c­ hapter 2. However, this research suggested the hypothesis that
8

8 B ehavioral E cology of the E astern R ed - B acked S alamander

P. cinereus holds interspecific territories against P. shenandoah, which led to a

detailed study of intraspecific territoriality within P. cinereus; these behavioral
experiments are reviewed in c­ hapter 3. To understand territoriality more fully,
experiments also investigated foraging tactics within territories by P. cinereus
(­chapter 4), because food seemed to be a limited resource in territorial de-
fense, and the glands and pheromonal cues used during territorial advertise-
ment and social communication (­chapter 5).
Our studies of intraspecific territoriality provided a wealth of information
about (1) olfactory and (2) visual communication among red-​backed salaman-
ders, (3) threat and submissive postures by territorial residents and intruders,
(4) combat tactics, (5) the costs and benefits of aggression, and (6) foraging
tactics within territories. Therefore, we were finally ready to return to studies of
interspecific territoriality between P. cinereus and P. shenandoah and between
P. cinereus and other species of salamanders (­chapter 6).
Our studies of territoriality led to observations of seemingly complex social in-
teractions within P. cinereus, both intra-​and intersexually. For example, observers
were surprised to see females often squash the fecal pellets of males and sometimes
carry a pellet balanced on their nasolabial grooves (Walls et al., 1989). (In plethod-
ontid salamanders, a pair of grooves lead from the upper lip to the nares, and in
the grooves, cilia move nonvolatile chemicals from the substrate into the olfactory
chamber; see “nose tapping” in Jaeger, 1984, 1986.) This observation suggested
that females sample the dietary quality of males via their squashed feces (Jaeger &
Wise, 1991). Therefore, we began an extensive series of investigations concerning
the intra-​and intersexual social behavior of red-​backed salamanders (­chapter 7).
Chapter 8 relates the predator–​prey contests between P. cinereus and a snake.
The wealth of salamandrine social behavior that we observed then led to our
last major area of research: the cognitive ecology (Dukas, 1998) of red-​backed
salamanders. This area, reviewed in c­ hapter 9, includes studies of (1) numeri-
cal discrimination of prey, (2) individual recognition memory of conspecifics,
(3) how exposure durations and separation intervals affect recognition memory
(e.g., of territorial neighbors and intruders), (4) the use of olfactory and visual
cues for recognition memory, and (5) whether individuals can recognize an
initial individual after interacting with other conspecific distractors.
We end our review with a coda (­chapter 10) that attempts to weave to-
gether the various behavioral patterns spanning from interspecific competition
(­chapter 2) to cognitive ecology (­chapter 9). In particular, we stress the large
amount of ambient information that red-​backed salamanders are able to input
and process in their neural systems. These processes lead to the complex be-
havioral responses to conspecifics, congeners, prey, and other environmental
stimuli that we report here. These cognitive-​behavioral areas of research would
be fruitful studies for other behavioral ecologists to pursue. Table 1.1 provides
short definitions of the abbreviations that we use throughout this book.
9

Table 1.1 Definitions of abbreviations used in the text

Behavior Definition
ATR All trunk raised. A threat posture
BITE Biting. Can be either a brief nip or an extended hold
Cloacal tapping. Cloaca is touched to the substrate; pheromonal
CLT
marking behavior
Chin tapping. A male touches his mental hedonic gland to the
CT
substrate; pheromonal marking used during courtship
Edge. Salamander walks along the perimeter of a chamber;
EDGE
withdrawal or submissive behavior
Flat. Salamander lies completely flat on the substrate; a submissive
FLAT
behavior
Front of trunk raised. The anterior one-​third of the body elevated
FTR
above the substrate; a resting, alert, and foraging posture
Look away. Looks directly away from the opponent; a submissive
LA
display
LT Look toward. Looks directly toward the opponent; a threat display
Move away. Moves directly away from the opponent; a submissive
MA
display
Mountain Lake Biological Station. In the Appalachian Mountains of
MLBS
southwestern Virginia
Move toward. Moves directly toward the opponent; a threat or
MT
preattack display
Nasolabial grooves. Paired grooves from the upper lip’s cirri to the
NLG
nares; chemodetection structures
Nose tapping. NLG touched to the substrate; chemodetection
NT
behavior
Postcloacal press. The postcloacal gland on the tail touched to the
PCP
substrate; pheromonal marking behavior
Resource holding potential. “The ability to persist and win a contest”
RHP
(Courtene-​Jones & Briffa, 2014)
Shenandoah National Park. Site of Hawksbill and Blackrock
SNP
Mountains in northwestern Virginia’s Blue Ridge Mountains
Snout-​to-​vent length. Measured from tip of the snout to posterior end
SVL
of the cloaca
TL Total length. Measured from tip of the snout to the tip of the tail
University of Louisiana at Lafayette. Site of many behavioral
ULL
experiments

note: Agonistic displays of Plethodon cinereus are thoroughly defined in section 3.4.
10

10 B ehavioral E cology of the E astern R ed - B acked S alamander

1 .4  C O MME N T S CO NCE R N I NG M E TH ODOLOGY

A N D S TATI STI C AL PAR ADI G M S

Plethodon cinereus is an excellent species for behavioral studies in the laboratory,

though not in its forested habitats where individuals are largely invisible except
during rainy nights. Individuals are small and thus thrive in small, transparent,
plastic chambers that are easily cleaned. Their substrates need be only damp
coffee filters or filter paper (in Petri dishes; Fig. 1.3) or paper towels (in larger
Nunc bioassay dishes), and Drosophila is a complete diet for this species as adults
but not for neonates. Room temperatures of 12o to 18oC are best for both housing

Figure 1.3 Stacks of single salamanders in their chambers (Petri dishes) in the
laboratory at the University of Louisiana at Lafayette.
Another random document with
no related content on Scribd:
THE bird picture book proved indeed a great joy to May, as Billy had
thought it would, and many were the hours she spent poring over it
during the long winter days when bad weather kept her indoors. She
was quick to learn the names of the pictured birds she did not
already know by sight, and eager to find out all about them. The
Vicar could give her more information than anyone else. All she
learnt from him she repeated to Billy.

After Christmas Billy attended the village school. He was in better

health now, and far less nervous in every way. The distressing
dreams which had haunted his sleep for so many weeks disturbed
him no longer. "'Thou shalt not be afraid of the terror by night,'" he
reminded himself if he awoke in the darkness; whilst morning and
evening he prayed Tom Turpin's prayer, "'Be not Thou far from me, O
Lord,'" and felt it was answered.

It was a dragging winter, and a cold spring. With the lengthening

days came a spell of severe frost which lasted for weeks, when no
work could be done on the land. Mrs. Brown fussed because her
husband was obliged to be idle, but he was not in the least
disturbed.

"It'll be all right, Maria," he assured her cheerfully. "By-and-by spring

will come with a rush."

Such was indeed the case. Within one week there was a great
transformation scene. The wind changed from north-east to south-
west, a gentle rain fell on the frost-dried earth; then the rain ceased,
sunshine came, and the soft stir of awakening life.

There was plenty of work to be done out-of-doors now. William

Brown had the assistance of a couple of old men from the village,
and with them he laboured from dawn till dark. On Saturdays Billy
worked in the garden too. He helped his grandfather prepare the
beds for the small seeds, making the earth as fine as possible, and
learnt the different depths the various seeds had to be sown, and not
to sow them too thickly.
"You'll make a rare good gardener one of these days," his
grandfather told him approvingly; "your heart's in the work, I see."

At last all the seeds were in the ground. The garden at Rowley
Cottage, with its trim, smooth beds, was a picture of neatness. The
gardens in the allotment field were in a like condition. And now there
was a lull in the gardening.

Billy thought the seeds were slow in growing, but his grandfather
assured him that they were doing all right.

"But how can you tell, Grandfer?" the little boy inquired. "You can't
see them."

"No, Billy," William Brown answered; "I can't see them, but I've faith
as to what's going on underground. I've seen too many springtimes
to doubt that."

"And what's going on underground, Grandfer?" asked Billy.

"Why," his grandfather said gravely, "a miracle—the miracle of the

resurrection. It's too wonderful for us to understand. We don't see
the stir of life in the spring, but we see the effect of it in everything—
in the budding trees, the flowers, the fresh green grass. It's God's
life-giving spirit wakening the world from its winter sleep. We must
have patience. We sowed good seeds; bought them from a reliable
firm, and they're all right."

And all right they were. One morning William Brown announced, at
breakfast, that the turnips and parsnips were showing above the
ground. Before going to school Billy had a look at them. What tiny
plants they were! After that, other vegetables, such as carrots, beets,
and onions, were not long in appearing, and, once above ground,
they grew apace.

One afternoon Billy arrived home to tea, carefully carrying a small

pot containing a marrow plant, a gift from the postmaster. After tea,
under the directions of his grandfather, he tilled the marrow plant out.
"Uncle John has given Harold one, too, Grandfer," he said; "so we're
going to see which of us can grow the biggest marrow. The plants
are as alike as two plants can possibly be."

"John Dingle took the prize for marrows at the vegetable show last
year," William Brown remarked.

"Yes, he told me," Billy answered, adding; "He says he should like
one of us boys to win the prize this year. We both mean to try."

Accordingly he gave his plant every care and attention, shielding it

from the too fierce rays of the sun before it had properly rooted, and
watering it regularly. He was delighted with the quickness of its
growth. When it came into flower his grandfather advised him which
flowers to take off and which to let remain. He often called at the
post office to tell John Dingle how well the plant was doing. On one
of these occasions, hearing that May and Harold had gone to the
allotment field, he followed them there.

Harold was weeding in his father's garden when Billy arrived at the
allotment field Billy examined his marrow plant, which seemed in just
the same flourishing condition as his own, and then went in search of
May, who had wandered off to the river. He found her on the river's
bank. She turned a bright, happy face to him as he approached, and,
with her finger on her lip, whispered—

"Hush! There's a dip-chick's nest! I'm watching the baby dip-chicks!"

She pointed to a clump of grasses growing beneath the overhanging

bank opposite. Close by three baby dip-chicks were disporting
themselves in the water, whilst their mother swam around them,
keeping guard.

"Aren't they sweet little things?" whispered May. "Oh, look, there's
their father! What a hurry he's in!"

Sure enough there he was, running along the bank towards his
family. Suddenly he dived off into the river. There was a great flutter
for a moment or two, then father, mother, and baby dip-chicks had all
disappeared under the overhanging bank.

"They saw us," said May regretfully. "Now they'll hide till we're gone."

"Better come away from the water," advised Billy; "you might fall in."

"Oh, I'm ever so careful!" the little girl assured him. "I do love the
river; don't you? And it's so happy to-day! Listen how it laughs and
sings! That's because it's fine weather and the sun shines. Back in
the winter it was different—so dark and deep. Billy, it must be very
wise, mustn't it? for it's come such a long, long way, from up in the
Dartmoor hills. It's going right on to the sea. Mr. Singleton told me.
He comes here sometimes, and we listen to the river together. Oh,
look, look! What's that? There, up in the sky! And what a funny
noise, Billy!"

Billy's eyes followed the direction of the little girl's pointing finger, and
saw an aeroplane coming towards them, high in the sky. It looked
like a great bird in the distance, but he recognised it as an aeroplane
at a glance, for he had frequently seen them pass over London.
Though he knew there was little likelihood of this one being an
enemy, his heart throbbed fast at the sight of it. He explained to May
what it was; then Harold rushed up to them in a great state of
excitement.

"An aeroplane! An aeroplane!" he cried. "The first I've seen! Oh, I

hope father and mother and every one in the village will see it, too!
The men say it's most likely going to Plymouth."

He referred to several allotment holders who had been busy in their

gardens a few minutes before; they were now standing with heads
thrown back watching the strange sight. Nearer and nearer it came,
then passed right over the gardens.

"It's a biplane," said Billy, rather proud of his knowledge. "I've seen
several like it before. How fast it's going! And can't we hear its
engines working plainly! That's the funny noise you spoke of, May.
Buzz-z-z-z!"

May drew a deep sigh. She was watching the aeroplane with
breathless interest, her colour coming and going fitfully. It was
momentarily growing more and more like a bird as it sailed away.
When at last it was lost to view the children left the allotment field
and hurried back to the village. Every one had been out to watch.
The aeroplane was the talk of the place, and indeed of the
neighbourhood, for days.

CHAPTER XI.
GRANNY SURPRISES BILLY.

"I'M come to tea, mother," announced Mrs. Dingle, one hot August
afternoon, as she entered the kitchen at Rowley Cottage, where Mrs.
Brown stood ironing at a table near the open window. "I shan't be in
your way, shall I? Here, you rest a bit, and let me take on your irons."

"No, thank you," Mrs. Brown answered. "You were never a good
hand at ironing starchery, Elizabeth, and I can't bear to see it done
badly. I'm doing Billy's collars—such a lot there are! This hot weather
a collar rarely lasts him more than a day."

"I suppose he makes a lot of extra work for you," remarked Mrs.
Dingle, seating herself and taking off her hat.
"You suppose?" said her mother tartly. "As if you didn't know, when
you've a boy of your own! By the way, I believe May and Harold are
about here somewhere."

"They're in the garden—I've been there. They're helping net the

plum-trees. Billy's a born gardener, his grandfather says."

Mrs. Brown nodded.

"To give him his due, he's been a great help during the fruit-picking,"
she allowed. "All his spare time out of school hours during the
summer he's spent in the garden, and now it's holidays he's there
from morning till night. Did he show you his marrow—the one he is
going to cut for the show?"

"Yes. It's a beauty. Harold has one quite as large, though."

There was a brief silence, then Mrs. Dingle said—

"Mother, have you noticed any alteration in May lately? No? Oh, I
have, and her father too! She's far less dreamy and more interested
in things in general than she used to be. The other day we were
surprised to find she's really beginning to learn to read."

"No!" exclaimed Mrs. Brown incredulously.

"Yes, mother; yes, indeed! It seems it was Billy who taught her her
letters. Don't you remember how much time she spent with him after
his illness?"

"Of course! But I never paid much attention to them—they used to sit
on the settle by the fire, looking at the pictures in some story books
he has and talking in whispers. And he taught her then? Yes? Well, I
never!"

"She can read short words now," Mrs. Dingle said, with a tremble of
joyfulness in her voice. "Fancy! Oh, I think it's marvellous—
marvellous! It's been such a trouble to John and me that our little
maid should be so different from others."

"I always said she was only backward!" cried Mrs. Brown
triumphantly.

Mrs. Dingle nodded.

"Mr. Singleton says if you can interest her in anything she can learn
all about it quicker than most children," she said eagerly; "but if you
can't it's impossible to teach her or to chain her attention. She got
interested in Billy's books because there are such wonderful stories
in them, and that made her want to learn to read. Then Billy began to
teach her—he says it wasn't so very difficult; but I don't think he quite
realises what a great matter it is—that it shows our darling's intellect
is less clouded than we thought. Oh, mother, I don't feel now that I
shall fear for the child any more! Her school-teacher's most hopeful
about her, and I—oh, I'm grateful to God from the bottom of my
heart!"

Mrs. Brown changed her iron for a hotter one, and went on ironing
silently.

"I feel we owe a great deal to Billy," Mrs. Dingle proceeded,

"because he's been so good to May. From the first she took to him.
He's always been patient with her and never laughed at her quaint
ways. Harold's always saying, 'Don't talk so silly, May!'—not that he
means to be unkind, but because he gets impatient with her. Oh, I
see Billy's companionship has been a great thing for May! John sees
it too. And you do, don't you, mother?"

Mrs. Brown nodded.

"I'm not sorry now we had to have him here," she admitted, then
added: "As boys go, he's better than most; and I'll say this for him,
he never answers me back."
"He's a dear boy!" declared Mrs. Dingle. "I often think how his poor
mother must have loved him. Ah, here is May!"

The little girl was looking in at the window.

"The boys want to know if it's nearly teatime," she said.

"Yes," replied her grandmother; "by the time they've been in and
washed their hands tea will be ready."

May disappeared, returning ten minutes later with her grandfather

and the boys. Mrs. Brown had cleared away her ironing things and
was putting the last touches to the tea-table whilst she kept an eye
on her daughter, who was measuring the tea into the tea-pot.

"That will do, Elizabeth," she said.

"All right, mother," Mrs. Dingle answered; "I haven't put one more
spoonful than you told me to."

"Elizabeth's a capital tea-maker," William Brown remarked, smiling;

"she could always put more tea in a spoon without over-filling it than
anyone I ever knew."

At that everyone laughed, even Mrs. Brown. So the meal began

merrily and went on in the same happy way.

The conversation was mostly about the flower, fruit, and vegetable
show which was to be held in the Vicarage grounds that day week.
There was to be a prize for the prettiest bunch of wild flowers, for
which May intended to compete. Billy said he would help her gather
the flowers, but she shook her head.

"No one must help me," she said, "or it wouldn't be fair."

"Our marrows will grow a lot more in a week," remarked Billy. "I do
hope one of us will get the prize, Harold."

"You mean you hope you will get it," laughed Harold.
"I meant what I said," Billy replied. "Grandfer says your marrow is a
better shape than mine," he continued, "and that will be taken into
consideration. I went around and looked at all the marrows in the
allotment gardens yesterday, and there wasn't one to beat yours,
though I did see one that came near doing it."

"Yes, I know," Harold answered; "it belongs to a man called Gibbs."

"Is that the Gibbs who was had up for poaching last winter?" inquired
Mrs. Brown. "Yes. I should have thought he was too idle to have a
garden."

"He doesn't keep it in good condition," Harold explained; "it's

generally full of weeds; but, somehow, he's managed to grow good
marrows this year, and he's mighty proud of them."

Shortly after tea Mrs. Dingle went home, accompanied by her

children and Billy.

William Brown was expecting a business letter from Exeter, and, as

there was no second postal delivery at Ashleigh during the day, and
feeling sure the letter would be at the post office, he had asked Billy
to fetch it. The Exeter letter was there, and one, bearing a London
postmark, also addressed to "Mr. William Brown." Billy took the two
letters straight back to Rowley Cottage, and gave them to his
grandfather in the garden.

William Brown read the letter he had expected first, then opened the
other, and glanced through it quickly. It seemed to be of a startling
nature, for he turned very red and uttered an exclamation of
amazement. Then he read the letter a second time, very slowly and
carefully, his face exceedingly grave. After that he thought a while.

"Why, Billy," he said at length, "you never told me your mother had
an uncle in Scotland."

"I didn't know she had," Billy answered; "that is, I remember her
telling me she had an uncle, but she'd lost sight of him and didn't
know if he was living or dead. I think she said he was a sea captain."

"Exactly. This letter is from the master of the Institution where you
stopped in London. He'd had a letter from a Captain Foster, who
says he's your mother's uncle. Captain Foster, who has left the sea
and is now living in Glasgow, has only lately learnt of your mother's
sad death, and he and his wife, who are a childless couple, are
willing to give you a home and do the best they can for you. It's a
good offer, Billy, but—well, I don't want to part with you, my boy."

"And I don't want to leave you, Grandfer!" cried Billy. "I—I—oh, it's
not to be thought of, is it?"

"I don't know," William Brown said doubtfully; "maybe this Captain
Foster can do more for you than I can. Dear me, this is most
upsetting! I think I'd better go in and tell Maria, and hear what she
has to say."

Left alone, the little boy perched himself on the edge of a wheel-
barrow to consider the situation. Would he, indeed, be called upon to
leave the home he had learnt to love? The thought that he might be
wrung his heart.

"If it rests with Granny I shall have to go," he told himself sadly; "she
will be only too glad to get rid of me—she never wanted me here."

His eyes filled with tears as he looked around the garden. How he
loved it! What happy days he had spent here with Grandfer and May!
He had been looking forward to many more such happy days, but
now, perhaps, he would be sent far away. Suddenly he jumped off
the wheel-barrow, and hurried towards the house. He would not be
kept in suspense; he would find out what was to become of him at
once.

Arrived at the back door Billy stopped, his heart sinking despairingly.
From the kitchen came the sound of Mrs. Brown's voice, loud and
indignant.
"Oh, dear, she's in a temper," thought the little boy. "Perhaps I'd
better keep out of her sight."

But he was so anxious that he could not help lingering to listen. Then
he had a most wonderful surprise.

"Nonsense!" he heard Mrs. Brown exclaim, "Perfect nonsense! What

can he do for Billy more than we can? There's nothing for you to be
so upset about that I can see! Write to the man yourself and tell him
'No!' You've the first claim on the boy, for you're his grandfather.
There's no reason why we should give him up to any relation of his
mother's now."

Billy's heart gave a bound of joy. Then Granny was against sending
him away! It was amazing, but true. He rushed into the kitchen, his
eyes a-sparkle, his face aglow with delight. Mrs. Brown appealed to
him immediately.

"You don't want to leave us, child, do you?" she asked.

"No!" Billy cried; "No, no, no! I—I, oh, the thought of it was dreadful!"

"It was," William Brown agreed. "I was only putting it to you, Maria,
that this Captain Foster might be able to do more for Billy than I can,
and—"

"And I tell you I don't believe it!" interrupted his wife. "Billy's cut out
for a gardener, and he's in his right place. I daresay this Captain
Foster would want to send him to sea. You write and tell him we can't
give the boy up."

"Oh, Granny!" cried Billy. He made a rush at Mrs. Brown, and,

clasping her around the neck, kissed her. "I thought you'd be pleased
to be rid of me," he said, "but you really want me to stay. Oh, I am
glad—I am glad!"

Mrs. Brown gave a rather embarrassed laugh. She made no answer,

but her face softened, and her eyes were a little dim as she returned
the little boy's kiss.

"Now I know what to do," William Brown remarked, adding: "We shall
all sleep the better for having decided this matter to-night."

CHAPTER XII.
CONCLUSION.

WILLIAM BROWN wrote to Captain Foster that same night, and the
first thing after breakfast the following morning Billy hastened to the
village and posted the letter. Then he went into the post office, where
he found Mr. and Mrs. Dingle in earnest conversation, both looking
unusually grave.

"Oh, Billy," the latter began, "poor Harold's in trouble. He went to look
at his marrow before breakfast and found it gone."

"Do you mean it has been stolen?" gasped Billy, aghast.

"Yes," she assented, "stolen! Oh, it's really too bad! We've told the
policeman what's happened, but it's most improbable he'll be able to
find the thief. The marrow must have been taken during the night."

"Yes," agreed John Dingle, "for it was all right last evening—I saw it
myself after dusk."

"Oh, poor Harold!" cried Billy. "Where is he?" he asked.

"Out with May somewhere," replied Mrs. Dingle. "He's dreadfully
upset about this. But what brings you to the village so early, my
dear?"

Billy explained. When he had finished his tale Mrs. Dingle looked at
her husband meaningly, and said—

"There! Now what did I say, John? Didn't I tell you mother was
growing fond of Billy?"

"Yes, you certainly did," he answered, "and this proves you were
right. You're glad to remain at Rowley Cottage, Billy?"

"Oh, Uncle John, I don't know what I should have felt if Granny had
said I must go! Of course I knew Grandfer wouldn't want me to go,
but I was so afraid Granny would. You can't imagine how glad I was
to hear she didn't like the idea of my going! She was quite upset
about it. I am so glad, so glad!"

After a little further conversation Billy left to return home. He had not
gone far from the village when he heard, someone shouting, and,
looking back, saw Harold running after him. He stopped immediately.

"Father said I should overtake you it I ran," Harold said, as he came

up. "May and I came home just after you'd left—we'd been to the
allotment field again, looking everywhere for my marrow, but of
course we couldn't find a trace of it. You've heard what's happened?"

Billy nodded.

"I'm so sorry," he said simply, his voice full of sympathy; "it's a great
shame!"

"I'd give a great deal to know for certain who's had it!" Harold cried
fiercely. "I suspect that fellow Gibbs, and I believe father does too,
though he doesn't like to say so. Gibbs believes his marrow will be
the best exhibited at the show now, but he'll be mistaken! He doesn't
know about yours, and we must take care he doesn't!"
"If mine gets the prize we'll divide the money," said Billy. "Five
shillings it'll be, won't it?"

"Yes, but—oh, do you really mean it?"

"Of course I do! Half-a-crown each will be worth having, won't it?"

"Worth having? I should think so! But—but it wouldn't be fair to you if

I took half your prize money."

"Oh, yes, it would be, because I should wish it."

The boys were walking on side by side now. There was a brief
silence, then Harold suddenly exclaimed—

"You're a real brick, Billy. I've always been nasty to you about your
gardening tools, and begrudged your having them—you must have
seen it, yet you lent me your hand-fork when I broke mine, and—oh,
it's been too mean of me! I'm sure if someone had stolen your
marrow and I thought I was going to get the prize I shouldn't offer to
divide it with you—at least, I don't think I should—"

"Oh, I expect you would!" Billy broke in. "Anyway, if I win the prize we
go shares, mind! That's agreed."

The fields on the slope of the hill behind Rowley Cottage were now
golden with corn as tall as the boys themselves, and ripening fast
under the kisses of the hot August sun. The boys raced down the
narrow foot-track behind each other, through the orchard, and into
the garden. There they found William Brown whistling light-heartedly
as he weeded his asparagus bed. He heard of the loss Harold had
had with much concern.

"I can't tell you how sorry I am, Harold," he said; "it's a great
disappointment for you. We must watch that Billy's marrow doesn't
go in the same way."

"Yes," agreed both boys eagerly, "that we must!"

But Billy's marrow was undisturbed. It continued to grow and ripen till
the morning of the show, when Grandfer cut it and the boys
conveyed it between them to the large tent which had been erected
on the Vicarage lawn, under which most of the exhibits of flowers,
fruit, and vegetables were already on the stalls.

Billy's marrow was put on a stall with a lot of others. The boys had
the satisfaction of seeing, at a glance, that it was the best marrow
there, for shape, colour, and size. An ill-kept, sullen-looking man who
was standing near saw this, too. He shot a scowling look from one
boy to the other, and moved away.

"That's Gibbs!" Harold whispered to Billy excitedly. "He's brought his

marrow, but it's smaller than yours. I thought it was, but I couldn't be
sure till I saw them together. Doesn't he look sold—and guilty?"

"Hush!" admonished Billy, "Someone may hear you. If we're right in

what we think we can't prove it, you know."

Gibbs had slunk out of the tent and disappeared. He did not return
when the exhibits were being judged, nor did he come near the show
again. Apparently his whole interest in it had gone.

The show opened at two o'clock. It was well attended, nearly every
one in the parish being present. Mrs. Brown, who seldom left home,
was there under the escort of her husband and Billy. She was in high
good humour, for Billy's marrow had won the prize; and when she
came to the stall on which the wild flowers were being exhibited,
there, in the centre, was a beautiful bouquet bearing a card on which
was written: "First Prize—May Dingle." She felt, as she said, quite
proud to be connected with two prize-winners.

The prizes were distributed by the Vicar. Next day all the vegetable
exhibits, by agreement of the exhibitors, were packed carefully and
sent off as a gift to the Fleet, whilst the flowers were returned to their
owners. May gave her bouquet to her grandmother, and for several
days it graced the round table in the middle of the parlour at Rowley
Cottage.
Corn harvest was now commencing. Billy took great interest in
Farmer Turpin's "reaper and binder," which he thought the most
marvellous piece of machinery there could possibly be. One day it
arrived to cut the corn in the fields near Rowley Cottage, and he
spent hours in watching it as it worked, gathering the corn into a
sheaf and cutting the stalks and tying them, then throwing the sheaf
out on the ground, and going through the same programme
continuously as it went on. Billy followed it till he was tired, then sat
down on a big stone near the gateway leading into the road, and
watched it from there.

So closely did it chain his attention that he failed to notice a khaki-

clad figure coming towards him, and started up in joyful surprise
when a well-remembered voice cried—

"Well met!"

The next instant he was shaking hands with Tom Turpin, back on
leave from France again.

"Is there room for two on that stone?" asked the young soldier, and,
Billy assenting eagerly, they sat down together. "I arrived home the
day before yesterday," he continued, his blue eyes looking lovingly
across the valley to Mount Farm on the opposite hill. "I can tell you
it's good to be home, my boy! How beautiful everything is! 'The
pastures are clothed with flocks; the valleys also are covered with
corn; they shout for joy, they also sing.' This is a blessed land, Billy.
Where I have come from there will be no harvest—all is desolation
and ruin. Here there is plenty, and oh, the peacefulness of it all!"

There was a note of sadness in Tom's voice, whilst his eyes had a
wistful expression in them. For a minute his face was clouded, then it
cleared, and he went on—

"Father's behind, talking to your grandfather, but he'll join me

presently. Meanwhile, tell me about yourself. I've heard of the great
marrow you grew. I wish I'd been home in time to see it. I've just
come from your grandfather's garden. It's a picture worth looking at.
Your grandfather always grew good vegetables, but this year they're
just splendid."

"I helped plant some of them," said Billy proudly.

"So I've heard," smiled Tom Turpin; "your grandfather says you've
done a lot of real hard work."

"Thanks to your tools!" exclaimed Billy. "I couldn't have done half so
much without them. I've taken great care of them, Mr. Turpin; they
are as bright as bright!"

Tom Turpin looked pleased.

"You seem remarkably 'fit,' Billy," he said; "you're rather different now
from the timid little chap I remember. You've grown a couple of
inches, I should say, and your face is almost as brown as mine. You
look happy, too."

"I am happy," Billy said earnestly. "When mother was killed I never
thought I should be happy again; but, oh, I am! And things which
used to frighten me don't frighten me any longer now. I pray your
prayer, 'Be not Thou far from me, O Lord,' and that helps me to be
brave—you understand."

"I do understand, my boy."

"I love being here," Billy continued; "I love the hills, and the river, and
the woods, and, most of all, I love Grandfer's garden. It was so
wonderful in the spring to see everything coming to life. You see I
was never in the country, in the spring, before this year. I suppose
that's why I thought it so wonderful. Everything—the river, the
woods, the grasses in the fields, seemed to be whispering—always
saying the same!"

"What was that?" Tom Turpin asked. "Is it a secret?"

"Oh, no!" Billy answered. "I'll tell you. I told little May, and she said
she could hear it, too. It was 'The gift of God is eternal life through
Jesus Christ, our Lord.' Just that, over and over again."

THE END.

LONDON: PRINTED IN GREAT BRITAIN

BY WILLIAM CLOWES AND SONS, LIMITED.

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