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A Natural History of
Bat Foraging
This page intentionally left blank
 A Natural History of
Bat Foraging
Evolution, Physiology, Ecology,
Behavior, and Conservation

Edited by
Danilo Russo
Animal Ecology and Evolution Laboratory (AnEcoEvo),
Dipartimento di Agraria, Università degli Studi di Napoli Federico II,
Portici, Italy

Brock Fenton
Department of Biology, University of Western Ontario,
London, ON, Canada
Academic Press is an imprint of Elsevier
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by the Publisher (other than as may be noted herein).

Notices
Knowledge and best practice in this field are constantly changing. As new research and
experience broaden our understanding, changes in research methods, professional
practices, or medical treatment may become necessary.
Practitioners and researchers must always rely on their own experience and knowledge in
evaluating and using any information, methods, compounds, or experiments described
herein. In using such information or methods they should be mindful of their own safety
and the safety of others, including parties for whom they have a professional responsibility.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors,
assume any liability for any injury and/or damage to persons or property as a matter of
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ISBN: 978-0-323-91820-6

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Cover: Miniopterus natalensis emerging from Gatkop Cave in Limpopo Province of South
Africa. Photo by Sherri and Brock Fenton, All Rights Reserved.
Contents

Contributors.......................................................................................... xiii
Author biographies ............................................................................... xvii

CHAPTER 1 Introduction ............................................................ 1

Danilo Russo and Brock Fenton
References.............................................................................. 5
CHAPTER 2 Foraging in the fossil record: Diet and behavior
of the earliest bats.................................................. 7
Nancy B. Simmons and Matthew F. Jones
Introduction............................................................................ 7
Dental morphology and diet ...................................................... 9
Ancient bat dentitions and hypothesized diets..............................14
Fossilized stomach contents......................................................17
Postcranial clues.....................................................................18
Evolution of echolocation ........................................................20
Vision and nocturnality............................................................22
Foraging strategies in ancient bats .............................................25
Ecological and faunal diversity in the Eocene..............................28
Acknowledgments...................................................................30
References.............................................................................30
CHAPTER 3 How the moth got its ears and other just-so
stories in the history of batemoth interactions........41
Jesse R. Barber and John M. Ratcliffe
How the moth got its ears ........................................................42
How the moth got its voice ......................................................44
How the bat got its whisper......................................................47
How the moth got its invisibility cloak .......................................49
How the moth got its tail .........................................................50
Conclusion ............................................................................51
Acknowledgments...................................................................52
References.............................................................................52
CHAPTER 4 Sensory systems used by echolocating bats
foraging in natural settings ....................................57
Clarice Anna Diebold and Cynthia F. Moss
Introduction...........................................................................57
Echolocation..........................................................................58

v
vi Contents

Sonar call structure and duty cycle ........................................58

Nonlaryngeal echolocation ...................................................60
Sonar scene analysis............................................................61
Auditory signals and ecological niche ........................................62
Aerial hawking...................................................................63
Foraging over water ............................................................64
Gleaning for prey on substrates.............................................64
Passive listening to the sounds of prey ...................................65
Insectivores and prey counter strategies ..................................65
Other bat sensory systems and specializations..............................68
Vision...............................................................................68
Olfaction...........................................................................71
Thermoreception ................................................................72
Airflow sensing for flight control...........................................73
Conclusion ............................................................................74
References.............................................................................74
CHAPTER 5 Foraging strategies of echolocating bats.................83
Hans-Ulrich Schnitzler and Annette Denzinger
Introduction...........................................................................83
Commuting behavior, search behavior, and social foraging ........83
Foraging habitats ................................................................84
Habitat types......................................................................84
Foraging modes..................................................................85
Constraints that shape the foraging strategies of bats ....................85
Echolocation......................................................................85
Flight morphology ..............................................................86
Foraging strategies of bats........................................................86
Aerial hawking foraging strategy...........................................87
Trawling foraging strategy....................................................91
The passive gleaning foraging strategy of animalivorous bats .....91
The active gleaning foraging strategy in animalivorous bats.......92
Passive/active gleaning foraging strategy of phytophagous
bats ..................................................................................93
Flutter detecting foraging strategy .........................................93
Species using more than one strategy .....................................94
Search and commuting behavior of foraging bats .........................95
Search behavior and social foraging in bats feeding on
predictable prey..................................................................95
Search behavior and social foraging in bats feeding on
ephemeral prey...................................................................99
 Contents vii

Commuting behavior in bats............................................... 100

Outlook............................................................................... 101
References........................................................................... 101
CHAPTER 6 Foraging, movements, and diet habits of
arid-zone dwelling bats........................................ 109
Irene Conenna and Carmi Korine
Introduction......................................................................... 109
Nightly movements........................................................... 110
Seasonal movements ......................................................... 111
Temporal activity.............................................................. 112
Diet habits....................................................................... 114
Conclusions ......................................................................... 115
References........................................................................... 115
Further reading..................................................................... 121
CHAPTER 7 Social foraging and information transfer ............... 123
Jenna E. Kohles and Dina K.N. Dechmann
History of the study of social information use in bats.................. 123
The cons of social foraging .................................................... 131
Other types of social foraging in bats....................................... 132
Conclusions ......................................................................... 133
References........................................................................... 133
CHAPTER 8 Insect migrations and the ecology, behavior,
and population dynamics of bats .......................... 139
Jennifer J. Krauel, Don R. Reynolds,
John K. Westbrook and Gary F. McCracken
Introduction......................................................................... 139
Introduction to insect migration............................................... 141
A. Insect migration increases the resource base for
insectivorous bats ............................................................. 142
B. Insect migrants may be disproportionately important
to bats’ diets .................................................................... 144
C. Biomass may be more important than diversity in
bat diets .......................................................................... 145
D. Insect migrations and the ecosystem services of bats
are intersecting areas of study............................................. 146
E. Insect migrations are predictable and linked to major
life history features of bats ................................................. 147
F. Behavior of bats changes in response to insect migrations.... 148
viii Contents

G. Response to insect migrations may influence impacts

of climate change on bat populations ................................... 149
Conclusions......................................................................... 149
References........................................................................... 150
CHAPTER 9 Bats as prey ........................................................ 157
Peter Mikula, Radek K. Lucan, Juan J. Pellón,
Jose W. Valdez and Brock Fenton
Introduction......................................................................... 157
Bat responses to predation...................................................... 160
Bat predation and nocturnality ................................................ 161
Predators of bats................................................................... 162
Birds .............................................................................. 162
Mammals ........................................................................ 163
Reptiles........................................................................... 164
Other vertebrates .............................................................. 165
Invertebrates .................................................................... 165
Conclusions and suggestions for further research ....................... 165
Dedication and/or acknowledgments .................................... 166
References........................................................................... 166
CHAPTER 10 Energetics of foraging bats................................. 173
Liam P. McGuire and Justin G. Boyles
Energy budgets and metabolism .............................................. 173
Energy balance ¼ energy in  energy out............................. 173
Metabolic rate: Basal, resting, torpid, and flight ..................... 175
Energy required to get food.................................................... 177
Flight and the power curve................................................. 177
Optimal flight speed theory and implications for foraging
strategies......................................................................... 180
Variation in wing loading................................................... 182
Energetic cost of terrestrial locomotion ................................ 183
Cost of echolocation ......................................................... 183
Energy in food ..................................................................... 184
Variation in diet composition .............................................. 184
Refractory content and implications for the energetics of
different diets................................................................... 184
Rapidly accessing dietary energy......................................... 187
Spatial and temporal variation in the foodscape ......................... 188
Conclusion .......................................................................... 191
Acknowledgments................................................................. 192
References........................................................................... 192
 Contents ix

CHAPTER 11 Bat migration and foraging: Energy-demanding

journeys on tight budgets ................................... 199
Christian C. Voigt, Shannon E. Currie and
Liam P. McGuire
Introduction......................................................................... 199
What species of bats migrate?................................................. 200
Resource fluctuations as a driver of migration............................ 202
The energetics of migratory flights .......................................... 203
Stopover behavior and torpor-assisted migration ........................ 204
Food for migration................................................................ 206
Flight corridors .................................................................... 208
Open research questions......................................................... 208
Acknowledgments................................................................. 209
References........................................................................... 209
Further reading..................................................................... 215
CHAPTER 12 Microbiomes of bats .......................................... 217
Melissa R. Ingala
Gastrointestinal structure........................................................ 217
Interactions between the gut microbiota and immune
development ........................................................................ 220
Host diet influences taxonomic composition of the microbiome.... 222
Interactions between microbiome functions and diet ................... 224
Beyond bacteria: the “other” bat microbiome ............................ 226
Conclusions ......................................................................... 227
References........................................................................... 227
CHAPTER 13 The diets of bats: Think outside the guild ........... 233
Elizabeth L. Clare and Phillip J. Oelbaum
Collapse of the traditional guild .............................................. 234
Are insectivores a guild?........................................................ 234
Case study 1: Antrozous pallidus, the scorpion eating bat that
pollinates cacti ..................................................................... 235
Are nectarivores a guild? ....................................................... 237
Case study 2: Glossophaga - the nectar bat that sneaks an
insect snack ......................................................................... 240
Are frugivores a guild? .......................................................... 241
Caste study 3: Widely distributed pteropodids ........................... 242
Are carnivores a guild?.......................................................... 244
Case study 4: Trachops the “frog eating” bat............................. 245
Assemblage-wide assessment of diet in bats.............................. 247
x Contents

A world with more omnivores? ............................................... 251

References........................................................................... 252
CHAPTER 14 Bioaccumulation and foraging behavior .............. 261
Daniel J. Becker, Natalia I. Sandoval-Herrera,
Molly C. Simonis and Cecilia A. Sánchez
Introduction......................................................................... 261
Dietary exposure to contaminants ............................................ 262
Dietary exposure to parasites .................................................. 264
Interactions between foraging, contaminants, and parasitism ........ 269
Direct health effects .......................................................... 269
Indirect health effects ........................................................ 272
Future research directions ...................................................... 273
Acknowledgments................................................................. 277
Glossary.............................................................................. 277
References........................................................................... 277
CHAPTER 15 Foraging-dependent ecosystem services ............. 287
Rieka Yu and Nathan Muchhala
Introduction......................................................................... 287
Arthropod suppression........................................................... 288
Pollination and seed dispersal ................................................. 292
Ecosystem disservices ........................................................... 295
Conservation recommendations............................................... 295
References........................................................................... 296
CHAPTER 16 Conserving bats and their foraging habitats ........ 305
Winifred F. Frick, Luz A. de Wit, Ana Ibarra,
Kristen Lear and M. Teague O’Mara
Introduction......................................................................... 305
Threats to bat foraging .......................................................... 308
Conservation evidence........................................................... 310
The benefits to human well-being of protecting where bats eat ..... 311
Conservation initiatives targeting bat foraging ........................... 314
Restoring and conserving healthy habitat for nectarivorous
bats, agaves, and people..................................................... 314
Bat Conservation International’s Agave Restoration
Initiative ......................................................................... 315
Gardening for bats ................................................................ 317
Conclusion .......................................................................... 317
References........................................................................... 318
 Contents xi

CHAPTER 17 Bat foraging: The next steps ............................... 327

Brock Fenton and Danilo Russo

Index...................................................................................................329
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Contributors
Jesse R. Barber
Department of Biological Sciences, Boise State University, Boise, ID, United
States
Daniel J. Becker
Department of Biology, University of Oklahoma, Norman, OK, United States
Justin G. Boyles
School of Biological Sciences, Southern Illinois University, Carbondale, IL, United
States
Elizabeth L. Clare
Department of Biology, York University, Toronto, ON, Canada
Irene Conenna
Global Change and Conservation, Faculty of Biological and Environmental
Sciences, University of Helsinki, Helsinki, Finland
Shannon E. Currie
Institute for Cell and Systems Biology, University of Hamburg, Hamburg,
Germany
Luz A. de Wit
Bat Conservation International, Austin, TX, United States
Dina K.N. Dechmann
Department of Migration, Max Planck Institute of Animal Behavior, Radolfzell,
Germany; Centre for the Advanced Study of Collective Behaviour, University of
Konstanz, Konstanz, Germany; Department of Biology, University of Konstanz,
Konstanz, Germany; Smithsonian Tropical Research Institute, Balboa, Ancón,
Panama
Annette Denzinger
Animal Physiology, Institute for Neurobiology, University of Tübingen, Tübingen,
Germany
Clarice Anna Diebold
Department of Psychological and Brain Sciences, Johns Hopkins University,
Baltimore, MD, United States
Brock Fenton
Department of Biology, University of Western Ontario, London, ON, Canada
Winifred F. Frick
Bat Conservation International, Austin, TX, United States; Ecology and
Evolutionary Biology, University of Santa Cruz, Santa Cruz, CA, United States
Ana Ibarra
Bat Conservation International, Austin, TX, United States
xiii
xiv Contributors

Melissa R. Ingala
Fairleigh Dickinson University, Madison, NJ, United States
Matthew F. Jones
Biodiversity Knowledge Integration Center, School of Life Sciences, Arizona State
University, Tempe, AZ, United States
Jenna E. Kohles
Department of Migration, Max Planck Institute of Animal Behavior, Radolfzell,
Germany; Centre for the Advanced Study of Collective Behaviour, University of
Konstanz, Konstanz, Germany; Department of Biology, University of Konstanz,
Konstanz, Germany; Smithsonian Tropical Research Institute, Balboa, Ancón,
Panama
Carmi Korine
Mitrani Department of Desert Ecology, Jacob Blaustein Institutes for Desert
Research, Ben-Gurion University of the Negev, Midreshet Ben-Gurion, Israel
Jennifer J. Krauel
Department of Biology and Microbiology, South Dakota State University,
Brookings, SD, United States; Department of Ecology and Evolutionary Biology,
University of Tennessee, Knoxville, TX, United States
Kristen Lear
Bat Conservation International, Austin, TX, United States
Radek K. Lu
can
Department of Zoology, Faculty of Science, Charles University, Prague, Czech
Republic
Gary F. McCracken
Department of Ecology and Evolutionary Biology, University of Tennessee,
Knoxville, TX, United States
Liam P. McGuire
Department of Biology, University of Waterloo, Waterloo, ON, Canada
Peter Mikula
Institute of Vertebrate Biology, Czech Academy of Sciences, Brno, Czech
Republic
Cynthia F. Moss
Department of Psychological and Brain Sciences, Johns Hopkins University,
Baltimore, MD, United States; The Solomon H. Snyder Department of
Neuroscience, School of Medicine, Johns Hopkins University, Baltimore, MD,
United States; Department of Mechanical Engineering, Whiting School of
Engineering, Johns Hopkins University, Baltimore, MD, United States; Kavli
Neuroscience Discovery Institute, Johns Hopkins University, Baltimore, MD,
United States
 Contributors xv

Nathan Muchhala
Department of Biology, University of Missouri e St. Louis, St. Louis, MO,
United States
Phillip J. Oelbaum
Department of Cell and Systems Biology, University of Toronto, Toronto, ON,
Canada
M. Teague O’Mara
Bat Conservation International, Austin, TX, United States; Southeastern
Louisiana University, Hammond, LA, United States; Max Planck Institute of
Animal Behavior, Radolfzell, Germany; Smithsonian Tropical Research Institute,
Panama City, Panama
Juan J. Pellón
Laboratorio de Ecologı́a y Conservación de Vertebrados Terrestres, Instituto de
Ecologı́a, Universidad Nacional Autónoma de México, Mexico City, Mexico;
Departamento de Mastozoologı́a, Museo de Historia Natural, Universidad
Nacional Mayor de San Marcos, Lima, Peru
John M. Ratcliffe
Department of Biology, University of Toronto Mississauga, Mississauga, ON,
Canada
Don R. Reynolds
Natural Resources Institute, University of Greenwich, Chatham, United Kingdom;
Rothamsted Research, Harpenden, United Kingdom
Danilo Russo
Animal Ecology and Evolution Laboratory (AnEcoEvo), Dipartimento di Agraria,
Università degli Studi di Napoli Federico II, Portici, Italy
Cecilia A. Sánchez
EcoHealth Alliance, New York, NY, United States
Natalia I. Sandoval-Herrera
Department of Ecology and Evolutionary Biology, University of Toronto, Toronto,
ON, Canada
Hans-Ulrich Schnitzler
Animal Physiology, Institute for Neurobiology, University of Tübingen, Tübingen,
Germany
Nancy B. Simmons
Department of Mammalogy, Division of Vertebrate Zoology, American Museum of
Natural History, New York, NY, United States
Molly C. Simonis
Department of Biology, University of Oklahoma, Norman, OK, United States
xvi Contributors

Jose W. Valdez
German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig,
Leipzig, Germany
Christian C. Voigt
Leibniz Institute for Zoo and Wildlife Research, Berlin, Germany
John K. Westbrook
Retired
Rieka Yu
Department of Biology, University of Missouri e St. Louis, St. Louis, MO,
United States
Author biographies

M.B. (Brock) Fenton received his PhD in 1969 for work in the ecology and
behavior of bats. Since then, he has held academic positions at Carleton University,
York University, and the University of Western Ontario. He has published over 250
papers in refereed journals (most about bats), as well as numerous nontechnical con-
tributions. He has written four books about bats intended for a general audience. He
continues his research on the ecology and behavior of bats, with special emphasis on
echolocation and evolution. He was inducted as a Fellow of the Royal Society of
Canada (FRSC) in November 2014. In November 2018, he began a 3-year term
as a Deputy Executive Editor-in-Chief, Canadian Science Publishing. In February
2020, he began a 3-year term as a member of the Board of Directors of Bat Conser-
vation International. Dr. Fenton is a professor emeritus from the University of West-
ern Ontario, Department of Biology, Ontario, Canada and currently serves as an
academic editor for PLoS Biology and Nature Scientific Reports.

Prof. Danilo Russo is a full professor of Ecology at Naples University Federico II

and an honorary professor at the University of Bristol, UK. He obtained his PhD in
Zoology from the University of Bristol in 2002. Currently, he serves as the head of
the Animal Ecology and Evolution Laboratory in the Department of Agriculture at
his university. His research interests encompass a wide range of subjects, including
habitat selection, resource partitioning, sensory ecology, social behaviour, evolu-
tionary biology, biogeography, and invasion ecology. While much of his research fo-
cuses on bats, he also investigates various other model organisms to address specific
questions of interest.

From 2019 to 2023, Prof. Russo chaired the Scientific Committee of the UNEP/
EUROBATS Agreement, which is responsible for the conservation of European
bat populations. Additionally, he takes great pride in his role as the editor-in-chief
of the highly esteemed zoological journal, Mammal Review. Furthermore, he is
the main proposer and chair of the Management Committee for the EU COST Action
“CLIMBATS” (CA18107), a significant initiative exploring climate change’s effects
on bat populations.

Prof. Russo’s contributions to the scientific community are exemplified by his pub-
lication record, which includes approximately 170 scientific articles in internation-
ally respected journals such as Nature Communications, Current Biology, Ecology
Letters, and Biological Reviews.

Having conducted fieldwork in diverse regions and environments across the globe,
ranging from African rainforests to the Israeli desert and European beech wood-
lands, Prof. Russo possesses invaluable firsthand experience in various ecosystems.

xvii
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 CHAPTER

Introduction

1
1
Danilo Russo1, Brock Fenton2
Animal Ecology and Evolution Laboratory (AnEcoEvo), Dipartimento di Agraria, Università degli
Studi di Napoli Federico II, Portici, Italy; 2Department of Biology, University of Western Ontario,
London, ON, Canada

Foraging is the behavior exhibited by animals searching for and acquiring food in
their environment. It encompasses activities and strategies used by animals to locate,
obtain, and consume food, involving a range of behaviors such as searching,
capturing, handling, and eating a food item. Foraging is a vital activity for animals
as it directly influences their survival, reproduction, and overall fitness (Stephens
et al., 2007). The specific foraging behaviors employed by animals can vary greatly
depending on their species, ecological niche, available resources, and evolutionary
adaptations. The study of animal foraging behavior helps us understand how animals
adapt to their environment (Dehnhard et al., 2020), make optimal foraging decisions
(Trapanese et al., 2019), and allocate their energy and time to optimize food intake
and ultimately fitness (Latty and Trueblood, 2020). Analyzing foraging provides
invaluable insights into how animals have adapted to their environment: it offers a
precious way to link individual behavior to higher organizational levels such as pop-
ulation dynamics, interspecific interactions, energy fluxes, and ultimately,
ecosystem structure and processes (Abrams, 1984; Petchey et al., 2008; Brose,
2010; Gil et al., 2018). This book is about foraging in bats, exploring this fascinating
issue from different perspectives and scales and evaluating the influence of this
crucial aspect of bat behavior not only on the ecology of these outstanding mammals
but also on the status and future of our planet and our own lives.
Bats (order Chiroptera) are a diverse group of mammals, numbering over 1460
species (Simmons and Cirranello, 2023) and, in mammals, are second only to rodents
in terms of global species richness (Fig. 1.1). This overwhelming diversity is the
result of a long evolutionary history that has led to an astonishing range of foraging
strategies used by bats in almost every habitat worldwide to acquire food. Nocturn-
ality (Rydell and Speakman, 1995) and active flight (Hedenström and Johansson,
2015) add further spatial and temporal perspectives to the natural history of bat
foraging, making bats an especially interesting case within mammals from the
ecological but also evolutionary viewpoint. To fully understand this present-day di-
versity in foraging strategies, it is therefore paramount to delve into the remote
past, studying ancient bats to unveil the origins of diverse diets and foraging strategies
in present-day species, as Simmons and Jones discuss in Chapter 2.

A Natural History of Bat Foraging. https://doi.org/10.1016/B978-0-323-91820-6.00017-6 1

Copyright © 2024 Elsevier Inc. All rights reserved.
2 CHAPTER 1 Introduction

FIGURE 1.1
A compilation of close-up images showcasing the remarkable diversity of bat species.
Arranged from top to bottom and left to right:
Row 1 Nycteris thebaica; Rhinolophus simulator; Hipposideros speoris; Desmodus
rotundus; Row 2 Molossus nigricans; Mormoops megalophylla; Diclidurus albus;
Leptonycters yerbabuenae; Row 3 Lophostoma evotis (with ears up); Lophostoma evotis
(ears down); Thyroptera tricolor; Noctilio leporinus; Row 4 Pteropus poliocephalus;
Epomophorus walbergi; Uroderma bilobatum; Centurio senex.

Bat predation also exerts powerful selective pressure that has elicited astonishing
antipredatory adaptations in certain prey species. In response to this evolutionary
challenge, bats have developed an array of behavioral and physiological counter-
measures. In Chapter 3, Barber and Ratcliffe illuminate the enthralling case of tym-
panate moths, which stands as one of the most captivating examples of the ongoing
evolutionary arms race.
 Introduction 3

In Chapter 4, Diebold and Moss explore how foraging bats use a diverse range of
sensory cues to locate food sources in various environments. Bats have evolved so-
phisticated systems, which are crucial to collect the sensory information needed to
find food, and these solve very different tasks depending on the bat’s dietary special-
ization, from insects, fruit, nectar and pollen to small vertebrates, and even blood.
The bats’ long evolutionary history has led to today’s surprising range of
foraging strategies bats use to feed on a broad range of food items: arthropods
and plants but also vertebrates such as fishes, frogs, rodents, birds, rodents, and
even other bats, not to mention the three bat species that feed on blood in the Neo-
tropics. For many of these species, echolocation plays a pivotal role in the context of
their foraging strategies. The use of echolocation and the various foraging strategies
used by bats are comprehensively covered in Chapter 5 by Schnitzler and Denzinger.
One captivating aspect of bat foraging ecology is their remarkable ability to ac-
quire food in seemingly inhospitable habitats, such as deserts. These arid regions are
home to a variety of fascinating bat species that exhibit astonishing strategies, such
as scorpion hunting, which demands the bats to be resilient to the arthropod’s
venom. In Chapter 6, Conenna and Korine delve into the spatial and temporal move-
ments of bats in the desert while also exploring their dietary preferences and adap-
tations to survive in these challenging environments.
A further characteristic that cannot be neglected is the highly gregarious nature
of bats, often forming numerous colonies made of one or more species, and the di-
versity of bat social systems, whose dynamics are certainly complex and still poorly
understood. The use of social information to increase foraging success and how it
interacts with the distribution of trophic resources is, therefore, a fascinating field
of study which we have just begun to explore and is presented in Chapter 7 by
Kohles and Dechmann.
The spatial challenges encountered by bats become particularly intriguing when
they pursue migrant insect prey. Insect migration is a captivating and still inade-
quately comprehended phenomenon that holds significant relevance to the investiga-
tion of bat foraging behavior. The predation of bats on these insects carries
substantial ecological implications and bears crucial consequences for ecosystem
services, especially considering the role of certain migratory insect species as agri-
cultural pests, as Krauel et al. highlight in Chapter 8.
In Chapter 9, Mikula et al. cover the role of bats as prey in ecosystems, illus-
trating the diverse range of bat predators, touching upon the behavioral responses
of bats to predation and the potential evolutionary consequences of predation on
bat nocturnality. The chapter remarks on the need for further research on interactions
between bats and their predators, identifying important topics for future studies.
Chapter 10, by McGuire and Boyles, focuses on the energetic aspects of bat
foraging, highlighting the significant impact flight has on their daily energy expen-
diture. The review delves into the energy costs involved in obtaining food and
accessing energy from their diet while also considering the consequences of changes
in food availability across diverse spatial and temporal conditions. By examining
4 CHAPTER 1 Introduction

both energy costs and gains, McGuire and Boyles provide valuable insights into the
foraging ecology of bats.
In Chapter 11, Voigt et al. explore the captivating realm of migratory bats, which
traverse vast distances seasonally, yet encounter significant metabolic challenges
along their journey. The chapter endeavors to unravel the pivotal role of foraging
in aiding these bats in their arduous task while highlighting the plethora of intriguing
questions that continue to captivate the scientific community in relation to this awe-
inspiring subject matter.
The study of microbiomes in animal physiology and ecology is of ever-growing
importance, as it sheds light on the intricate interactions between animals and their
associated microbial communities, influencing various aspects of their health,
behavior, and ecological roles. In Chapter 12, Ingala illustrates gut microbiomes
in bats and how these play crucial roles in bat physiology, ranging from nutrient
acquisition to immunity. Given the diverse foraging strategies and diets exhibited
by bats, from insectivory to herbivory, the composition and function of these gut
communities show remarkable variation.
Chapter 13 challenges the classical view of bat trophic guilds, revealing that ex-
ceptions previously overlooked are increasingly evident with advanced analytical
methods. In this chapter, Clare and Oelbaum stress the importance of thorough di-
etary analysis and caution against under-reporting or overgeneralizing bat diets to
safeguard their ecological significance.
In Chapter 14, Becker et al. uncover the complex interactions between bat
foraging, contaminants, and infection, shedding light on the impacts on bat health,
zoonotic risk, and conservation practices and showing how diverse foraging strate-
gies shape exposure to harmful substances and parasites.
Yu and Muchhala, in Chapter 15, unveil the crucial ecological services rendered
by bats as they consume insects, fruits, and nectar, which, in turn, contribute to pest
regulation, pollination, and seed dispersion. Delving into the varied eating habits and
foraging patterns of bats, the chapter explores their far-reaching ecological and eco-
nomic implications but also highlights the existing knowledge gaps on the economic
benefits derived from bat-dependent ecosystem services.
Bat foraging exhibits a complex web of ecological interactions, encompassing
predatoreprey dynamics and mutualistic relationships like plant pollination and
seed dispersal. Human activities pose threats to bats by modifying or disturbing
these interactions. Therefore, safeguarding and restoring bat habitats are essential
for preserving bats and promoting a One Health Strategy that addresses wildlife
health, ecosystem health, and human health interconnections. In Chapter 16, Frick
et al. emphasize these aspects, along with the importance of establishing and main-
taining robust foraging habitats that could potentially benefit local communities.
The intricate world of bat foraging offers a captivating and multifaceted lens
through which we can explore the diverse strategies and adaptations these remark-
able mammals employ to survive in a variety of environments. From the evolu-
tionary origins of their diets to the astounding antipredatory adaptations they have
developed, from the sophisticated sensory systems guiding their nocturnal hunts
 References 5

to the critical ecological roles they play, bat foraging behavior unveils a rich tapestry
of interconnected relationships and dependencies. As we investigate more compre-
hensively the complex interplay between bats and their environments, we uncover
invaluable insights into population dynamics, interspecific interactions, and the intri-
cate balance of ecosystem structure and processes. We hope that this book success-
fully sheds light on the fascinating world of bat foraging, sparking curiosity and
inspiring further research into the extraordinary ecology, behavior and evolution
of these fascinating mammals and their vital role in the broader fabric of our planet’s
biodiversity and sustainability.

References
Abrams, P.A., 1984. Foraging time optimization and interactions in food webs. Am. Nat. 124
(1), 80e96.
Brose, U., 2010. Body-mass constraints on foraging behaviour determine population and
food-web dynamics. Funct. Ecol. 24 (1), 28e34.
Dehnhard, N., Achurch, H., Clarke, J., Michel, L.N., Southwell, C., Sumner, M.D., Eens, M.,
Emmerson, L., 2020. High inter-and intraspecific niche overlap among three sympatri-
cally breeding, closely related seabird species: generalist foraging as an adaptation to a
highly variable environment? J. Anim. Ecol. 89 (1), 104e119.
Gil, M.A., Hein, A.M., Spiegel, O., Baskett, M.L., Sih, A., 2018. Social information links in-
dividual behavior to population and community dynamics. Trends Ecol. Evol. 33 (7),
535e548.
Hedenström, A., Johansson, L.C., 2015. Bat flight: aerodynamics, kinematics and flight
morphology. J. Exp. Biol. 218 (5), 653e663.
Latty, T., Trueblood, J.S., 2020. How do insects choose flowers? A review of multi-attribute
flower choice and decoy effects in flower-visiting insects. J. Anim. Ecol. 89 (12),
2750e2762.
Petchey, O.L., Beckerman, A.P., Riede, J.O., Warren, P.H., 2008. Size, foraging, and food web
structure. Proc. Natl. Acad. Sci. U. S. A. 105 (11), 4191e4196.
Rydell, J., Speakman, J.R., 1995. Evolution of nocturnality in bats: potential competitors and
predators during their early history. Biol. J. Linn. Soc. 54 (2), 183e191.
Simmons, N.B., Cirranello, A.L., 2023. Bat Species of the World: A Taxonomic and
Geographic Database, Version 1.4. (Accessed 22 August 2023).
Stephens, D.W., Brown, J.S., Ydenberg, R.C. (Eds.), 2007. Foraging: Behavior and Ecology.
University of Chicago Press.
Trapanese, C., Meunier, H., Masi, S., 2019. What, where and when: spatial foraging decisions
in primates. Biol. Rev. 94 (2), 483e502.
This page intentionally left blank
 CHAPTER

Foraging in the fossil

record: Diet and behavior
of the earliest bats 2
Nancy B. Simmons1, Matthew F. Jones2
1
Department of Mammalogy, Division of Vertebrate Zoology, American Museum of Natural
History, New York, NY, United States; 2Biodiversity Knowledge Integration Center, School of Life
Sciences, Arizona State University, Tempe AZ, United States

The Eocene brought mammals mean

And bats began to sing;

Their food they found by ultrasound

And chased it on the wing.
Pye (1968: 797)

Introduction
Living bats exhibit a greater range of diets and foraging strategies than any other
mammalian order. Extant bat lineages include species that range from strict insecti-
vores to carnivores, piscivores, frugivores, nectarivores, palynivores, granivores,
sanguinivores, and many species have mixed diets that overlap two or more of these
categories (Gardner, 1977; Freeman, 1998, 2000; Rex et al., 2010; Dumont et al.,
2012; Santana et al., 2011a,b; Clare et al., 2014; Nogueira et al., 2005; Dumont,
2003). Methods of detecting and acquiring food similarly vary across chiropteran
lineages. Most extant species are thought to obtain food by aerial hawking for flying
insect prey, but many other species are gleaners, instead plucking food items from
surfaces and branches, while yet others trawl for prey from the surface of water
(Audet et al., 1991; Barclay and Brigham, 1991; Kalko et al., 1998; Kalka and
Kalko, 2006; Santana et al., 2011a,b). Detection of food objects may depend on
echolocation, listening for prey-generated sounds, olfaction, vision, or some combi-
nation of these senses, and getting to food once detected may involve behaviors
ranging from continuous flight to perch-hunting, hovering, or even quadrupedal
walking (Audet et al., 1991; Barclay and Brigham, 1991; Kalka and Kalko, 2006;
Norberg and Rayner, 1987; Hessel and Schmidt, 1994; Thies et al., 1998; Riskin
et al., 2006; Page and Bernal, 2020). Understanding the evolutionary history of
this complex array of diets and foraging habits requires a comparative phylogenetic

A Natural History of Bat Foraging. https://doi.org/10.1016/B978-0-323-91820-6.00008-5 7

Copyright © 2024 Elsevier Inc. All rights reserved.
8 CHAPTER 2 Foraging in the fossil record

perspective, one that includes consideration of the diets and adaptations seen in the
earliest batsdthose known only from fossils.
The fossil record of bats begins in the early Eocene, approximately 55 Ma (Gun-
nell and Simmons, 2005; Brown et al., 2019). Early Eocene (Ypresian; 56e47.6 Ma)
bats are known from multiple continents including North America, South America,
Europe, Africa, Australia, Asia, and the Indian subcontinent (Gunnell and Simmons,
2005; Brown et al., 2019; Hand et al., 2016; Jones et al., 2021; Simmons et al., 2016;
Smith et al., 2012; Hand and Sigé, 2018; Tejedor et al., 2005). Most of these fossils
consist of fragments of skulls, jaws, and teeth (Gunnell and Simmons, 2005; Brown
et al., 2019; Smith et al., 2012; Tejedor et al., 2005). However, some early bat fossils
are spectacularly well preserveddparticularly those from lagerstätten in the Green
River Formation (USA: Wyoming; w52.5 Ma) and the Messel Pit (Grube Messel;
Germany: Hesse; w47 Ma) (Jepsen, 1966; Richter and Storch, 1980; Habersetzer
et al., 1994; Simmons and Geisler, 1998; Simmons et al., 2008; Habersetzer et al.,
1992). Specimens from these localities include nearly complete articulated skele-
tons, some of which even contain fossilized stomach contents (Richter and Storch,
1980; Habersetzer et al., 1992, 1994). However, even fragmentary fossils may pre-
serve important clues to foraging habits in ancient bats due to observed links be-
tween morphological and ecological traits in bats (Simmons et al., 2008, 2010,
2016; Jepsen, 1966; Simmons and Geisler, 1998; Novacek, 1985; Czaplewski and
Baker, 2022).
Living bats (crown clade Chiroptera) are currently classified in 21 families each
containing from one to over 500 species (Simmons and Cirranello, 2022). Several of
these groups have fossil records that extend back into the Eocene and/or Oligocene
including Emballonuridae, Nycteridae, Hipposideridae, Rhinolophidae, Vespertilio-
nidae, and Mormoopidae (Smith et al., 2012; Simmons and Geisler, 1998; Simmons,
2005; Ravel et al., 2014, 2016; Morgan et al., 2019). Phylogenetic relationships of
extant bats are typically assessed with molecular tools that depend on tissue samples
and DNA sequencing (e.g., Eick et al., 2005; Stadelmann et al., 2007; Agnarsson
et al., 2011), but placing fossils into evolutionary context and dating divergence
points requires analyses of phenotypic data (e.g., Simmons and Geisler, 1998;
O’Leary et al., 2013; Ravel et al., 2015, Rietbergen et al., 2023). Based on morpho-
logical comparisons, 11 extinct families of bats are currently recognized (Table 2.1).
While relationships of many of these taxa remain enigmatic, phylogenetic ana-
lyses of the better-known families (e.g., those known from more than just teeth)
have indicated that several groups clearly fall outside the crown clade and thus
may provide insights into early bat evolution that cannot be garnered from examina-
tion of only extant lineages (Gunnell and Simmons, 2005; Simmons and Geisler,
1998; O’Leary et al., 2013; Ravel et al., 2015; Rietbergen et al., 2023). Two families,
Onychonycteridae and Icaronycteridae, are thought to represent the most-basal
branch(es) in Chiroptera, with Archaeonycteridae, Palaeochiropterygidae, Hassia-
nycteridae, and perhaps Tanzanycteridae, also representing stem lineages (Gunnell
and Simmons, 2005; Simmons and Geisler, 1998; Simmons et al., 2008; O’Leary
et al., 2013; Rietbergen et al., 2023). Among the remaining, Mixopterygidae,
 Dental morphology and diet 9

Table 2.1 Extinct families of bats.

Family Age range Distribution Sources

Aegyptonycteridae Late Eocene Africa Simmons et al. (2016)

Archaeonycteridae Early Eocene Europe; India Revilliod (1917), Simmons
eMiddle and Geisler (1998), Smith
Eocene et al. (2012), Hand and Sigé
(2018)
Hassianycteridae Early Eocene Europe; India Habersetzer and Storch
eMiddle (1987), Simmons and Geisler
Eocene (1998), Smith et al. (2012)
Icaronycteridae Early Eocene North Jepsen (1966), Simmons and
America; Geisler (1998), Smith et al.
Europe; India (2012), Rietbergen et al.
(2023)
Mixopterygidae Middle Europe Maitre et al. (2008), Smith
EoceneeLate et al. (2012)
Oligocene
Necromantidae Early Eocene Africa; Sigé (2011), Hand et al.
eLate Europe (2012), Ravel et al. (2016)
Eocene
Onychonycteridae Early Eocene Europe; Simmons et al. (2008), Smith
North et al. (2012), Hand et al.
America (2015), Czaplewski et al.
(2022)
Palaeochiropterygidae Early Eocene Asia; Europe; Revilliod (1917), Simmons
eLate India; North and Geisler (1998), Smith
Oligocene America et al. (2012), Maitre (2014),
Czaplewski et al. (2022)
Philisidae Early Eocene Africa Sigé (1985), Smith et al.
eEarly (2012), Ravel et al. (2012,
Oligocene 2015)
Speonycteridae Oligocene North Czaplewski and Morgan
America (2012)
Tanzanycteridae Middle Africa Gunnell et al. (2003)
Eocene

Philisidae, and Speonycteridae are hypothesized to belong within the crown group
(e.g., Ravel et al., 2015; Maitre et al., 2008; Czaplewski and Morgan, 2012), while
Necromantidae and Aegyptonycteridae occupy more uncertain positions within Chi-
roptera (Simmons et al., 2016; Hand et al., 2012).

Dental morphology and diet

The primary clues for reconstructing diets of extinct bats come from teeth, which are
the most commonly fossilized part of the bat skeleton (Brown et al., 2019). Dental
10 CHAPTER 2 Foraging in the fossil record

morphology is highly correlated with diet in bats as in other mammals (Freeman,

1984, 1988, 1998, 2000; Santana et al., 2011b; López-Aguirre et al., 2022;
Villalobos-Chaves and Santana, 2022) (Fig. 2.1), which allows analyses of diet in

fossil taxa (e.g., Simmons et al., 2016; Horácek and Spoutil, 2012; Yohe et al.,
2015). Most extant bat species are thought to be either entirely or primarily insec-
tivorous and, like other insectivorous mammals, are characterized by a tribosphenic
dentition (Santana et al., 2011b; Freeman, 1984; Horácek and Spoutil,  2012;
Slaughter, 1970; Fenton and Simmons, 2015). This sometimes makes it difficult
to distinguish isolated fossil dental remains of bats from those of other ancient mam-
mals including Paleogene marsupials, leptictids, eulipotyphlans, adapisoricids, nyc-
titheres, and other dentally primitive therians (Hand et al., 1994, 2012, 2015, 2016;

Smith et al., 2012; Hand and Sigé, 2018; Horácek and Spoutil, 2012; Hooker, 1996;
Sigé et al., 2012).
Although the majority of extant bats are primarily insectivorous, other prey is
regularly taken by a few bat species including both fish and terrestrial vertebrates
(Norberg and Fenton, 1988; Gual-Suárez and Medellı́n, 2021). However, most
carnivorous and piscivorous bats also eat insects and other arthropods (Santana
et al., 2011a,b; Norberg and Fenton, 1988; Gual-Suárez and Medellı́n, 2021). For
this reason, researchers have long distinguished “animal-eating” or “animalivorous”
taxa from those that feed partly or entirely on plant products (Freeman, 1984, 1988,
1998, 2000; Rex et al., 2010; Santana et al., 2011a,b; Simmons et al., 2016; López-
Aguirre et al., 2022; Norberg and Fenton, 1988; Gual-Suárez and Medellı́n, 2021).
In this context, animalivory is an umbrella term that covers insectivorous, carnivo-
rous, and piscivorous species, as well as those that have mixed diets including ar-
thropods and other prey (Freeman, 1984, 1998, 2000; Santana et al., 2011a,b;
Simmons et al., 2016; Norberg and Fenton, 1988; Gual-Suárez and Medellı́n,
2021). Dental morphology in plant-eating bats including frugivores, nectarivores,
and granivores is often highly modified to the point that molar cusps and crests
cannot be easily homologized with those of animalivorous taxa (Simmons et al.,
2016; Freeman, 1988, 1995; López-Aguirre et al., 2022; Slaughter, 1970)
(Fig. 2.1). Vampire bats represent a special case with an extensively modified denti-
tion preserving barely any trace of ancestral tribosphenic traits (López-Aguirre et al.,
2022; Slaughter, 1970).
All extant animalivorous bats have a dilambdodont tribosphenic dentition
including a W-shaped ectoloph on the anterior upper molars (M1 and M2) and a
well-developed talonid on the lower molars (Freeman, 1984, 1998, 2000; Santana
et al., 2011b; Simmons et al., 2016; Slaughter, 1970; Norberg and Fenton, 1988)
(Fig. 2.1). The primitive dental formula for bats includes 3 molars, and all extant ani-
malivorous bats retain M3 although this tooth may be reduced somewhat in size and
have an incomplete N- or V-shaped ectoloph; concomitantly, the lower m3 may have
a reduced talonid (Simmons et al., 2016; Freeman, 1988, 1995; Slaughter, 1970;
Koopman and MacIntyre, 1980). Multivariate dental topographic analysis and
phylogenetic comparative methods indicate that in at least some lineages (e.g.,
Another random document with
no related content on Scribd:
suomalaisia, 3 poikaa ja 2 tytärtä norjalaisia ja 3 lappalais-poikaa,
jotka kaikki tuliwat admitteeratuiksi. Muotkan lapsia kastettiin 13;
ruumiita haudattiin 6; rippiwäkeä oli 19. Puumangin lapsia kastettiin
31; ruumiita haudattiin 40; ja rippiwäkeä oli 146, johon kuuluu myös
ulohtaammaltakin tulleita sekä Suomalaisia että Lappalaisia.
Lauantai-iltana 5 päiwä Elokuuta pidettiin rippilasten päästö, ja
Sunnuntaina jumalanpalwelus Herran Ehtoollisen kanssa ja illalla
wiimeinen Raamatun selitys. Useampana arkipäiwänäkin pidettiin
Herran Ehtoollista, kun tuli etempänä asuwaisia sitä warten luokseni,
erittäinkin Lappalaisia.

Maanantaina 7 päiwänä iltapuolella lähdin kotiapäin Wesisaaren,

Näätämöwuonon kautta, Inarin järwen ylitse Kyrön kylään, jossa, kun
oli
Sunnuntaipäiwä 13 päiwä, pidettiin Jumalanpalwelus; wielä
eteenpäin
Sompiojärwen kautta Luiron ja Kitis-jokia myöten Kemijärwelle.
Kotiin
saawuin Elokuun 26 päiwä.

Mitä ylipäätään papillisiin toimiin täällä tulee, ei aina woi

parhaimmalla tahdollakaan seurata Suomen lakeja, sillä suuri osa
asukkaista on täällä syntynyt ja elänyt, eikä siis woi mitenkään
omistaa kirjallisia todistuksia maineestaan, esteettömyydestään
awioliittoon j.n.e., kun eiwät ole missään kirjoilla. Wenäjän
wirkamiehet ottawat kirjoihinsa asewelwolliset, eiwät muita. Ainoa
wirkamies, joka käy heitä, niin sanoakseni, hoitamassa, on
Suomesta lähetety pappi; jonkatähden sitä suurella ikäwällä aina
odotetaan. Tässä katsonkin welwollisuudekseni konstateerata sitä
erinomaista ystäwällisyyttä ja sydämmellisyyttä, jolla pappia näillä
seuduin kohdellaan, ja lopuksi ilmaista sen kaikkein asukasten
hartaan toiwotuksen, että Suomen Waltio warsinkin kerran wuodessa
tahtoisi kustantaa papin matkustuksen näiden oman onnensa nojaan
jätettyin kansalaisten luo. Sywemmillä ja lewottomammilla tunteilla
kuin kutkaan muut Suomalaiset odottawat he tuon kysymyksessä
olewan Jäämeren rannikkopalasen luowutuksen ratkaisemista;
pyytäwät kumminkin, käwi tuo asia, miten käwi, sillä hartaudella,
mikä woi syntyä ainoastaan järjestetyn yhteiskunnan ulkopuolelle
muuttaneen sydämessä, saada pysyä Suomen Esiwallan muistissa.

Leppäwirroilla, Joulukuun 28 päiwä 1882

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Statistisia tietoja Suomalaisista ja Lappalaisista Muurmannilaisella

rannikolla Jäämeren luona, jokta kuuluwat ewankelis-luterilaiseen
uskontunnustukseen, tehdyt matkalla sinne kesällä 1882. (Senaatiin
annettu)

Lapsia kastettu. miespuolta / waimonpuolta

Kuolassa. . . . . . . . . — / 2
Uurassa ja Saaniwuonossa. 17 / 15
Laatsiwuonossa. . . . . . 2 / 3
Tsipna Wolokassa. . . . . — / 2
Sopuskassa. . . . . . . . — / 2
Waitokuwassa. . . . . . . 5 / 3
Kerwanassa. . . . . . . . 3 / 2
Muotkassa . . . . . . . . 4 / 9
Puumangissa . . . . . . . 19 / 12
Kakkarissa. . . . . . . . 3 / 3
Summma. . . . . . . . . 53 / 53 = 106
 Pariskuntia wihitty.

Kuolassa . . . . 1 suomalaisia kumpikin

Uurassa. . . . . 3 d:o d:o
d:o. . . . . . . 1 lappalaiset kumpikin
d:o. . . . . . . 1 suomalainen ja wenäläinen
Tsipna Wolokassa.1 norjalainen ja lappalainen
Sopuskassa. . . .1 norjalainen ja suomalainen
Waitokuwassa. . .1 suomalaisia kumpikin
Summa. . . . . .8

Haudattu.
Alle 15 wuotta miesp / waimp. / summa alla

Kuolassa . . . . 1 / 2
Uurassa . . . . . 4 / 14
Laatsiwuonossa. . 5 / 3
Kakkarissa. . . . 1 / 1
Tsipna Wolokassa. — / —
Waitokuwassa. . . 2 / 1
Muotkassa . . . . 2 / 3
Puumangissa . . . 16 / 12
[Yhteensä]. . . . 31 / 36

Yli 15 wuotta: miesp/ waimp./ summa [alle/ yli 15 v.]

Kuolassa . . . . 2 / 1 = 6
Uurassa . . . . . 8 / 6 = 32
Laatsiwuonossa. . — / — = 8
Kakkarissa. . . . — / — = 2
Tsipna Wolokassa. 3 / — = 3
Waitokuwassa. . . 9 / 1 = 13
Muotkassa . . . . 1 / — = 6
Puumangissa . . . 7/ 15 = 40
[Yhteensä]. . . . 30 / 13 = 110

Kommunikantteja.

Kuolassa . . . . 7.
Uurassa . . . . . 85.
Laatsiwuonossa. . 17.
Kakkarissa. . . . 5.
Tsipna Wolokassa. 6.
Sopuskassa. . . . 2.
Waitokuwassa. . . 14.
Kerwanassa. . . . 30.
Muotkassa . . . . 19.
Puumangissa . . . 146.
[Yhteensä]. . . . 331.

Rippikoulun käyneitä.

Suomalaisia miesp. / waimp.

Uuraassa. . . . 3 / 2
Puumangissa . . 8 / 7
Admitteerattu ilman koulua
Sopuskassa. . . — / 1
Waitokuwassa. . 1 / —
Summa . . . . .12 / 10
 Lappalaisia miesp./ waimp.
Uurassa. . . . — / 3
Puumangissa . . 3 / —
Admitteerattu ilman koulua
Sopuskassa. . . —/—
Waitokuwassa. . —/—
Summa . . . . . 3 / 3

Norjalaisia miesp./ waimp.

Uuraassa. . . . 1 / 1
Puumangissa . . 3 / 2
Admitteerattu ilman koulua
Sopuskassa. . . —/ —
Waitokuwassa. . —/ —
Summa . . . . .4 / 3

Summa miesp./ waimp.

Uuraassa. . . 4 / 6
Puumangissa. 14 / 9
Admitteerattu ilman koulua
Sopuskassa. . . — / 1
Waitokuwassa. . 1 / —
Summa . . . . 19 / 16 = 35

Muist. yksi lappalainen tyttö ei admitteerattu

Wäkiluku

Talollisia: Taloja / miesp./ waimp.

 Kuolassa. . . . . — /— /—
Saaniwuonossa . . 4 / 12 / 9
Uurassa . . . . .21 / 56 /50
Laatsiwuonossa. .7 / 14 / 17
Kakkarissa. . . .4 / 9 / 6
Tsipna Wolokassa.— / — / 1
Sopuskassa. . . .— / — / 1
Waitokuwassa. . .12 / 22 / 20
Kerwanassa. . . .8 / 20 / 15
Muotkassa . . . .10 / 23 / 24
Puumangissa . . .30 / 77 / 73
Summa . . . . . .96 taloa /
233 miesp / 216 waimp. = 449
 hyyryläsiä: miesp./ waimp.

Kuolassa. . . . .9 / 11
Saaniwuonossa . .12 / 6
Uurassa . . . . .24 / 20
Laatsiwuonossa. .3 / 3
Kakkarissa. . . .— / —
Tsipna Wolokassa.2 / 3
Sopuskassa. . . .— / 1
Waitokuwassa. . .2 / 5
Kerwanassa. . . .1 / 2
Muotkassa . . . .1 / 1
Puumangissa . . .40 / 30
Summa . . . . . .94 / 82 = 176
S:ma. . . . 449 + 176 = 625

Lappalaisia:
Taloja / miesp./ waimp.

Areskuwassa, Kassiwuonossa
ja Uurassa. . . . 4 / 12 / 15
Kakkarissa. . . . 1 / 2 / 2
Tsipna Wolokassa. — / — / 1
Mattimuotkasassa ja
Muotkassa . . . . 5 / 13 / 11
Summa . . . . . 10 taloa/
27 miesp / 29 waimp. = 56

hyyryläsiä: miesp./ waimp.

 Areskuwassa, Kassiwuonossa
ja Uurassa. . . . 3 / 4
Kakkarissa. . . . — / —
Tsipna Wolokassa. — / —
Mattimuotkasassa ja
Muotkassa . . . . — / —
Summa . . . . . . 3 / 4 = 7
S:ma. . . . .56 + 7 = 63

Syntymäpaikat

Kuolassa: perheitä/ miesp./ waimp.

Kuusamosta tulleita . . 2 / 4 / 6
Pudasjärveltä tulleita. 1 / 2 / 2
Kiannalta tulleita. . . 1 / 3 / 3
Summa . . . . . . . . . 4 perheitä/
9 miesp. / 11 waimp. = 20

Saaniwuonossa: perheitä/ miesp./ waimp.

Kuusamosta tulleita . . 3 / 10 / 6
Paawolasta tulleita . . 1 / 3 / 2
Haukiputaalta tulleita. 1 / 1 / 1
Norjan kautta tulleita. 3 / 7 / 5
Wenäjän Karjalan kautta
tulleita. . . . . . . . 1 / 3 / 1
Summa . . . . . . . . .9 perheitä /
24 miesp. / 15 waimp. = 39.

Uuraassa: perheitä/ miesp./ waimp.

Kuusamosta tulleita . . 10 / 23 / 18
Iistä tulleita. . . . . 1 / 2 / 3
Lohtajasta tulleita . . 3 / 9 / 7
Torniosta tulleita. . . 2 / 6 / 3
Sodankylästä tulleita . 8 / 21 / 22
Kemistä tulleita. . . . 2 / 6 / 5
Oulusta tulleita. . . . 2 / 4 / 3
Norjan kautta tulleita. 5 / 8 / 10
Summa . . . . . . . . 33 perheitä /
80 miesp. / 70 waimp. = 150.

Laatsiwuonossa: perheitä/ miesp./ waimp.

Muoniosta tulleita.. 2 / 4 / 3
Kiannalta tulleita.. 3 / 5 / 6
Kuusamosta tulleita. 2 / 4 / 5
Kemistä tulleita. . . 3 / 4 / 6
Summa . . . . . . . 10 perheitä /
17 miesp. / 20 waimp. = 37.

Kakkarissa: perheitä/ miesp./ waimp.

Norjan kautta tulleita 4 / 9 / 6 = 15.

Tsipna Wolokassa: perheitä/ miesp./ waimp.

Norjan kautta tulleita — / 2 / 4 = 6

Sopuskassa: perheitä/ miesp./ waimp.

Kuusamosta tulleita — /— / 2 = 2

Waitokuwassa: perheitä/ miesp./ waimp.

Kuusamosta tulleita. . .5 / 8 / 6
Simosta tulleita . . . .2 / 7 / 5
Kuolajärweltä tulleita .1 / — / 2
Kemistä tulleita . . . .4 / 3 / 8
Utajärweltä tulleita . .1 / 2 / 1
Rowaniemestä tulleita. .2 / 2 / 3
Torniosta tulleita . . .1 / 1 /—
Kontiolahdesta tulleita.1 / 1 /—
Summa. . . . . . . . .17 perheitä /
24 miesp. / 25 waimp. = 49.

Kerwanassa: perheitä/ miesp./ waimp.

Kuusamosta tulleita. . .4 / 8 / 7
Iistä tulleita . . . . .2 / 4 / 2
Simosta tulleita . . . .1 / 4 / 3
Pulkkilasta tulleita . .1 / 2 /—
Norjan kautta tulleita .3 / 3 / 5
Summa. . . . . . . . . 11 perheitä /
21 miesp. / 17 waimp. = 38.

Muotkassa: perheitä/ miesp./ waimp.

Kuusamosta tulleita. .3 / 4 / 7
Rowaniemestä tulleita.1 / 2 / 6
Iistä tulleita . . . .1 / 2 / 1
Kiannalta tulleita . .1 / 4 / 2
Kuhmosta tulleita. . .2 / 5 / 4
Kemistä tulleita . . .1 / 3 / 3
Lumijoelta tulleita. .1 / 4 / 2
Summa. . . . . . . .10 perheitä /
24 miesp. / 25 waimp. = 49.

Puumangissa: perheitä/ miesp./ waimp.

Kuusamosta tulleita . 39 / 77 / 69
Pulkkilasta tulleita. .1 / 4 /3
Kemistä tulleita. . . .1 / 4 / 2
Muoniosta tulleita. . .2 / 3 / 4
Kuhmosta tulleita . . .7 / 17 / 10
Kiannalta tulleita. . .6 / 8 / 14
Kuiwaniemestä tulleita.2 / 4 / 1
Summa . . . . . . . . 58 perheitä /
117 miesp. / 163 waimp. = 220.

Summa summarum. . . . .327 miesp. / 298 waimp. = 625

Eri seurakunnista tulleitten summat.

. . . . . . . miesp./ waimp/ summat.

Kuusamosta tulleet 138 / 126 = 264

Kiannalta. . . . . 20 / 25 = 45
Kemistä. . . . . . 20 / 24 = 44
Sodankylästä . . . 21 / 22 = 43
Kuhmosta . . . . . 22 / 14 = 36
Simosta. . . . . . 11 / 8 = 19
Lohtajasta . . . . 9 / 7 = 16
Iistä. . . . . . . 9 / 5 = 14
Rowaniemestä . . . 4 / 9 = 13
Torniosta. . . . . 7 / 3 = 10
Pudasjärweltä. . . 2 / 2 = 4
Oulusta. . . . . . 4 / 3 = 7
Paawolasta . . . . 3 / 2 = 5
Haukiputaalta. . . 1 / 1 = 2
Muoniosta. . . . . 7 / 7 = 14
Kuolajärweltä. . . — / 2 = 2
Utajärweltä. . . . 2 / 1 = 3
 Pulkkilasta. . . . 6 / 3 = 9
Lumijoelta . . . . 4 / 2 = 6
Kuiwaniemestä. . . 4 / 1 = 5
Kontiolahdesta . . 1 / — = 1
Norjan kautta. . . 29 / 30 = 59
Wenäjän Karjal.
kautta tull. . . . 3 / 1 = 4

[Summa] . . . .327 / 298 = 625

Lappalaiset owat
Norjan kautta tulleet. 30 / 33 = 63

Muistutus. Norjan kautta tulleiksi kutsutaan niitä, jotka Norjaan

muuttaneista Suomen alamaisista owat syntyneet Norjassa, mutta
eiwät norjalaistuneet.

Kaiken kaikkiaan on siis Muurmannilaisella rannikolla 688 henkeä,

jotka ewankelis-lutherilasen papin apua tarwitsewat. Tästä
summasta saattaa olla poissa joku kymmenkunta lappalaista, jotka
asuwat sisämaissa ja Petsamowuonon seutuwilla.

Leppäwirroilla Joulukuun 28 päiwä 1882. Joh. Schwatzberg.

*****

Lukijan pyydetään hywäntahtoisesti itse lukiessansa oikasemaan

ne monet korehtuuriwirheet, joita on jäänyt tähän kirjaseen. Syynä
wirheiden jäämiseen niin lukuisesti on se, että kirja on ylipainoksena
otettu sanomalehti-latomuksesta ilman tarkemman korehtuurin
lukemista.
 *** END OF THE PROJECT GUTENBERG EBOOK MATKA
JÄÄMEREN RANNALLE KESÄLLÄ 1882 ***

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