The Little Owl

Population Dynamics, Behavior and Management of

Athene noctua
Second Edition

The basic biology of owls is poorly understood compared to that of other bird species. The
Little Owl, Athene noctua, is one of the best models for biological and conservation research.
Though widespread across Europe, Asia and North Africa, populations of the Little Owl are
now in decline, making studies of its behavior and ecology all the more important. This
extensively revised and updated second edition features substantial new long-term data on
population dynamics, behavioral observations and breeding biology of the Little Owl. The
authors discuss its wide-ranging ecology, genetics, subspecies and population status by
country. In addition, they outline a research strategy and monitoring program. Exceptional
illustrations of all 14 subspecies cover embryonic and chick development, feather growth and
molt, including high-quality drawings presenting concrete management suggestions. Whilst
being an invaluable resource for academic researchers, its accessible and straightforward
style will also appeal to amateur ornithologists and enthusiasts.

Dries Van Nieuwenhuyse is a life-long owl researcher and statistician active in ecological
method development and publication. He is Professor of Performance Management at
EHSAL Management School in Brussels and thought leader in fact-based decision-making.
He has authored five books on the impact of technology and statistics on the decision-
making processes of organizations, and in particular brings his skills as a statistician to his
ornithological work.

Ronald van Harxen is Chairman of the Dutch Little Owl Working Group (STONE), and has
been active in the research and conservation of the Little Owl in the Netherlands and
internationally for more than 43 years. His research focuses on breeding biology and
population dynamics within nestbox populations. He received several awards including
the Special Achievement Award in 2016 from the World Owl Hall of Fame, International
Owl Center, Houston, USA.

David H. Johnson is Executive Director at Global Owl Project, USA. He is currently working
on a “12-year demographic study of the Burrowing Owl (Athene cunicularia) 2010–2022.”
He has worked in natural resource conservation for 42 years and has written two previous
books on owls, wildlife and fisheries.

Published online by Cambridge University Press

Published online by Cambridge University Press
 The Little Owl
Population Dynamics, Behavior and
Management of Athene noctua
Second Edition

DRIES VAN NIEUWENHUYSE

EHSAL Management School, Brussels, Belgium

RONALD VAN HARXEN

Dutch Little Owl Working Group (STONE)

DAVID H. JOHNSON
Global Owl Project, USA

ILLUSTRATED BY JORIS DE RAEDT

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www.cambridge.org
Information on this title: www.cambridge.org/9781009100151
DOI: 10.1017/9781009103725
First edition © D. Van Nieuwenhuyse, J.-C. Génot and D. H. Johnson 2008
Second edition © Dries Van Nieuwenhuyse, Ronald van Harxen, and David H. Johnson 2023
This publication is in copyright. Subject to statutory exception and to the provisions
of relevant collective licensing agreements, no reproduction of any part may take place
without the written permission of Cambridge University Press and Assessment.
First published 2008
Second edition 2023
Printed in the United Kingdom by TJ Books Limited, Padstow Cornwall
A catalogue record for this publication is available from the British Library.
Library of Congress Cataloging-in-Publication Data
Names: Nieuwenhuyse, Dries van, author. | Harxen, Ronald van, author. | Johnson, David
H., author.
Title: The little owl : population dynamics, behavior and management of Athene Noctua /
Dries van Nieuwenhuyse, EHSAL Management School, Brussels, Belgium, Ronald van
Harxen, Dutch Little Owl Working Group (STONE), David H. Johnson, Global Owl Project,
USA ; illustrated by Joris De Raedt.
Description: Second edition. | Cambridge, United Kingdom ; New York, NY, USA :
Cambridge University Press, 2023. | Revised edition of: The little owl : conservation,
ecology and behavior of Athene noctua / Dries van Nieuwenhuyse, Jean-Claude Génot,
David H. Johnson. 2008. | Includes bibliographical references and index.
Identifiers: LCCN 2022056296 (print) | LCCN 2022056297 (ebook) | ISBN 9781009100151
(Hardback) | ISBN 9781009108249 (Paperback) | ISBN 9781009103725 (epub)
Subjects: LCSH: Little owl–Conservation. | Little owl–Ecology. | Little owl–Behavior.
Classification: LCC QL696.S83 N54 2023 (print) | LCC QL696.S83 (ebook) | DDC 598.9/7–
dc23/eng/20221219
LC record available at https://lccn.loc.gov/2022056296
LC ebook record available at https://lccn.loc.gov/2022056297
ISBN 978-1-009-10015-1 Hardback
Additional resources for this publication at www.cambridge.org/littleowl
Cambridge University Press and Assessment has no responsibility for the persistence
or accuracy of URLs for external or third-party internet websites referred to in this
publication and does not guarantee that any content on such websites is, or will remain,
accurate or appropriate.

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 Contents

Foreword ix
Jean-Claude Génot

Preface xi

Acknowledgements xv

Executive Summary 1

1 Introduction: Framework 5

2 History and Traditions 9

3 Fossil Evidence, Taxonomy and Genetics 20

Chapter Summary 20
3.1 Fossil Evidence 21
3.2 Discovery of the Species 22
3.3 Discovery of the Subspecies 27
3.4 Morphology and Coloration 45
3.5 Morphological Descriptions 48
3.6 Geographical Distribution 73
3.7 Genetic Approach 80
3.8 Vocalizations 87
3.9 Taxonomic Discussions 88

4 Morphology and Body Characteristics 89

Chapter Summary 89
4.1 Embryonic Development 89
4.2 Owlet Development 92
4.3 Plumage 101
4.4 Molt 107
4.5 Eyes 114
4.6 Ears 118

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 Contents

4.7 Bill 119

4.8 Measurements 122
4.9 Voice 131
4.10 Flight 144
4.11 Digestive Organs 145

5 Distribution, Population Estimates and Trends 146

Chapter Summary 146
5.1 Distribution 148
5.2 Population Numbers 152
5.3 Densities 155
5.4 Trends 163
5.5 Overview per Country 168

6 Habitat 239
Chapter Summary 239
6.1 Natural Habitats 240
6.2 Anthropogenic Habitats 241
6.3 Habitat Preferences 244
6.4 Habitat Suitability Factors 250
6.5 Assessments at Multiple Spatial Scales 264
6.6 Multilevel Resource Selection 269

7 Diet 271
Chapter Summary 271
7.1 Hunting 271
7.2 Capturing Prey 275
7.3 Prey Caching 276
7.4 Prey Species 278
7.5 Large Prey 279
7.6 Seasonal Variation 279
7.7 Geographical Variation 281
7.8 Different Methods 283

8 Breeding Season 324

Chapter Summary 324
8.1 Pair Formation and Pair Bond 324
8.2 Life Expectancy 327
8.3 Copulations 329
8.4 Nest Sites 331
8.5 The Nest 337

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 Contents

8.6 Eggs 337

8.7 Laying of Eggs 342
8.8 Laying Interval 344
8.9 Clutch Size 345
8.10 Replacement Clutches 354
8.11 Incubation 356
8.12 Hatching 358
8.13 Egg Failure 360
8.14 Prolonged Incubation 363
8.15 Number of Nestlings 364
8.16 Nest Success 368
8.17 Egg Success 371
8.18 Condition 373
8.19 Fledging Success 374
8.20 Mortality 380
8.21 Cainism 382
8.22 Lifetime Reproductive Success 382
8.23 Extra-Pair Copulation 385

9 Behavior 386
Chapter Summary 386
9.1 Body Movements 386
9.2 Locomotion 388
9.3 Roosting and Sleeping 388
9.4 Preening 390
9.5 Nest-Site Competitors 392
9.6 Enemies 395
9.7 Social Behavior 396
9.8 Territorial Behavior 397
9.9 Division of Tasks 398
9.10 Parents and Young 398
9.11 Handling Prey 400
9.12 Two Females Nesting Together 403
9.13 Behavior Towards Humans 404

10 Population Regulation 405

Chapter Summary 405
10.1 Introduction to Population Regulation 407
10.2 Environmental Limiting Factors 407
10.3 Demographic Limiting Factors 435

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11 Management and Conservation 480

Chapter Summary 480
11.1 Conservation Status and Threats 480
11.2 Conservation Strategy for the Little Owl 481

12 Research Priorities 553

12.1 Research Priorities 553
12.2 Recent Insights 553
12.3 Research Topics 555
12.4 Conclusion 562

References 563

Index 608

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 Foreword

When the first edition of the English monograph on the Little Owl was published in
2008, I was one of the authors since I was involved in the study of this owl at the time,
which I studied in France from 1984 to 2006. I was very happy to work with Dries and
David. A monograph represents an important work of collecting data and information,
as well as exchanges with specialists of the species in a large number of countries,
reading and writing in order to give the reader the most synthetic view possible on each
topic covered. For me, this work was very rewarding because I learned a lot while
writing this monograph. Today it is with pleasure that I write this foreword of the only
English updated monograph on the Little Owl, published by Cambridge University
Press.
Since the writing of the first edition, publications devoted to the species have
doubled, which made its updating essential. Techniques have advanced in many topics
allowing scientific studies to shed new light on the biology and ecology of this very
endearing owl. Hence, this monograph includes more data on aspects such as genetics,
morphology and biogeography of the different subspecies.
The drawings of Joris De Raedt are of beauty and of great precision, especially those
which illustrate the taxonomy. I am sure they will make this monograph enjoyable to
read for conservationists and scientists alike.
The Little Owl saw its numbers collapse in many European countries during the
twentieth century and the global causes are identified, linked in particular to agricul-
tural practices which have profoundly modified the habitats of the species. In France,
the National Museum of Natural History sounded the alarm in 2018 on the fall in bird
populations linked to agricultural areas: 33% decrease in 15 years, the decline having
accelerated between 2016 and 2017. Reasons for this decline are related to intensive
agriculture and pesticides. However, the species, being more generalist in terms of diet
and more eclectic in the choice of its habitat, seems to resist in a certain number of
landscapes, even increasing its population locally. The status of the species varies
according to the situation of the meta populations. It will be interesting to see what
impact global warming will have on this species, native to the Mediterranean basin.
The Little Owl is an interesting species because studying it illustrates our ecological
footprint and our impact on rural landscapes. It was the emblem of Athena, goddess of
wisdom and science among the Greeks, which is why its decline or its maintenance
sends back to us the symbolic image of our own “wisdom.”

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 Foreword

This book is the result of a huge amount of work, I hope it will be of use to all owl
enthusiasts. It is undoubtedly a reference book on the Little Owl. Many thanks to Dries,
Ronald and David for completely revising the first edition and making it an
indispensable work.
Jean-Claude Génot
Ecologist

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 Preface

Little Owls have always meant a great deal to me since my childhood. It was one of the
first bird species that I learned to know and appreciate. My favorite bird lived in an old
pollard willow nearby. More than ever little things are felt to be more important in life
than big things. The Covid pandemic showed the relativeness of all things in life and
forced us to go back to the basics. This was not an evident period for many people that
suffered from illness, lost loved ones or were exposed to extreme stress due to solitude
and psychological tension. This negative spiral led to sorrow and intense need for
consolation. No wonder that compassionate communities emerged, bringing people
and local organizations together at intimate places of consolation to find peace of mind
and get back on the road.
I dedicate this book to all people, especially youngsters, that had some mental
difficulties in this hard period, seeking comfort and balance in life. May this poem offer
solace and ease of mind like my Little Owl is still offering to me through each glimpse,
each hoot, each facial expression of this wonderful bird with the golden eyes.

Pollard Willow
I bury myself with soaking feet
in the grooves of my skin I preserve
the touches of grandchildren and grandparents
sometimes I break out of myself, my sap flows too fast
my hollows silence the birds
that have chosen me
pollarded I show my wounds shamelessly,
my growth is curtailed
I must germinate once more
capture light in my branches
find my voice again when the wind contradicts me

Steven Van Der Heyden

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 Preface

Special thanks to Trui, my wife and Juul and Siel, my children who helped me through
tough times when the combination between a family life, a busy stressful job, rebuild-
ing our house and writing a book were extremely energy-consuming. Thanks for the
support, the belief that this book would succeed and the comprehension that writing
this book simply had to be done.
This book is the result of a joint effort of the whole Little Owl community, the
multitude of volunteers and Little Owl enthusiasts throughout its distribution range. It
was very comforting as author to have access to all those data and insights that were
collected over decades with the utmost dedication. Hopefully these volunteers, profes-
sionals and not least the professional volunteers enjoy reading this book and realizing
that this book would never have been possible without their continuous efforts and
willingness to contribute their findings for the greater interest. It was an honor to sense
the belief that this book would act as a lever for all their work and dedication.
Last but not least I wish to thank Ronald, David and Joris for their remarkable effort
in making this book a success. Jean-Claude Génot, who retired to have more time to
enjoy playing with his grandchildren nowadays, was so kind to write the Foreword and
stimulated to make the update of this volume a real success. This book would never
have been possible without our very intense co-operation and the willingness of all of
us to deliver a piece of art that the Little Owl deserves.

Dries Van Nieuwenhuyse

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 Preface

This book holds plenty of insights that were obtained from the Zuidoost-Achterhoek
research area in the Netherlands. This body of knowledge is the result of a unique co-
operation with Pascal Stroeken who should be mentioned explicitly. As a team we
managed to collect this unique wealth of data and insights that this research has
yielded. Without our 35-year-old close friendship and the co-operation between us,
this information would never have been collected in this way. The number of hours
spent together in the field are innumerable. The same applies to the inspiring
exchanges of views on almost all aspects that are discussed in this book. Without
Pascal’s meticulous “accounting,” many results – especially in the Breeding chapter –
could not have been presented in this way.

Ronald van Harxen

The Little Owl is truly an international species whose conservation and cultural impacts
have reached far beyond its actual distribution. What an honor and wonderful oppor-
tunity it has been to work with Dries, Ronald and Joris on this book, and to bring
together our collective insights and skill sets about owls. I sincerely appreciate the
caring and thoughtful efforts of all of the people that have worked with the Little
Owl, sharing their publications and perspectives, assessments and analysis, and
amazing societal insights. I would also like to dedicate this book to the members of
the Global Owl Project. My life has become richer through our shared communications,
explorations, addiction to owls and the friendship I have had with you over the years. It
is upon this collective foundation that our book was possible. Thank you so very much.

David H. Johnson

Seeing my subjects in their natural habitat is often the very first step in the making of
my illustrations. Keeping a nature journal with field sketches helps me get familiar with
the character of the species and makes it easier to capture its essence in the final
illustration. Looking through a spotting scope I make (visual) notes of the animals
I’m studying. Travelling was a bit hard during the pandemic, but I was able to observe
Little Owls close to home in Belgium, Cyprus (definitively not Athene noctua lilith!) and
Spain in the preparation of the book. Luckily, Dries was also able to provide additional
pictures of all the subspecies, including study skins. He was even able to get footage of
the mysterious spilogastra and somaliensis.
As individual differences are often bigger than the differences between subspecies it
took lots of research and studying to represent the discriminative plumage of each
subspecies. More than any previous assignment I did, the habitat of these plates was
very important, visualizing the environment in which the owls thrive.
I would like to thank Dries, Ronald and David for their excellent guidance and
feedback during the making of the illustrations. I would also like to thank my partner
Linde, and my family for their patience and critique. Special thanks to Raf Beyers, Eddie
Schild, Niels Schild and Anders Gray for sharing their knowledge.

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 Preface

Figure P.1 Fieldwork by the artist in Edegem, Belgium.

The illustrations in this book are a combination of graphite sketches and digital
illustrations. Color and details are added on the computer using a graphic tablet. To see
more on my workflow, visit my website, jorisderaedt.com or follow me on Instagram;
@jorisderaedt.

Joris De Raedt

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 Acknowledgements

This book has many mothers and fathers, it shows our global Little Owl community at
its best. The idea of this update ermerged during the Nieuwvliet symposium in 2015 that
was held at the Belgian–Dutch border. Crossing borders has always been easy for people
in the Low Countries due to their very tiny size. This did not prevent their inhabitants
from playing a uniting role across Europe and even beyond. Our Dutch language that
we share also forces us to speak a broad range of foreign languages, which is also crucial
to capture plenty of gray literature and sometimes very valuable insights that never end
up in peer-reviewed journals.
So it is no surprise that we are endebted to many people, hopefully we are excused
for all those that were forgotten in this overview. In the first place we want to thank
several thousands of Little Owl enthusiasts that are active in conservation, education,
study, data organization and standardization, publication, . . . It is only through this
unique co-operation that projects such as Little Owl 2.0 can be successful.
A significant part of new insights in this book stems from camera observations
ranging from old-fashioned video recordings to online webcams that are followed by
armies of volunteers that monitor each movement of breeding Little Owls, leading to
unprecedented volumes of data, standardized and digitized, ready for analysis. Special
thanks go to the webcam detectives of Beleef de Lente (Vogelbescherming Nederland)
Marja Braams, Hansje Hardenberg, Anna Heijblok, Anjo Schouten, Ellen Verheul, Nel
van de Vin, Trudy de Wit and Geert Sterringa (co-ordination) who unravelled and
digitized hours of behavior that were never seen before.
The advantage of a second edition of this volume was the trustworthiness and the
unconditional co-operation of crucial multinational and standardized initiatives of
Little Owl data collection for ringing, distribution, population numbers and voice
recognition. We are highly endebted to Mark Balman (BirdLife International), Dorian
Moss (EURING), Verena Keller (EBCC) and Sergi Herrando (EBCC) for putting all recent
available data at our disposal for advanced large-scale analysis. We were lucky in our
search for updated high-quality data by different simultaneous initiatives that were
converging, e.g., Article 12 of the Bird Directive of the European Union (European
Environment Agency) yielded detailed distribution data at the 1010 km scale for all
EU member states (luckily still including the UK). The European Breeding Bird Atlas
(EBBA2) of the European Bird Census Council covered continental Europe up to the
Ural at the 5050 km level. More detailed information was obtained by EURING
revealing unique dispersion, fecundity and mortality insights. Zooming out to the

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 Acknowledgements

global range of the species was enabled by the Macaulay Library – The Cornell Lab of
Ornithology – at Cornell University that opened Pandora’s Chapter 4, Morphology and
Body Characteristics, outside Europe, allowing virtual travel and the opportunity to see
the Little Owl in all its diversity and very accurately behind a computer screen. Xeno-
canto allowed us to use hours of geocoded sound recordings. Geocoded occurrence data
for Little Owls were available from the GBIF – Global Biodiversity Information
Facility. We believe that by using all the available data from those organizations, we
were not only realizing our objectives, but also theirs, i.e., collect, standardize, analyze
and dissiminate insights and knowledge of species for the purpose of their long-term
conservation. We hope we have made our contribution through the publication of this
volume and are very grateful for the trust they had in us.
Literature is crucial in such a huge project. We are specially endebted to Alan
Sieradzki, Bruce G. Marcot, Al Vrezec, Anna Heijblok, Lex la Crois, Line Holm
Andersen, Lars Bo Jacobens, Gianpiero Calvi, Jevgeni Shergalin, Grzegorz
Grzywaczewski and Frank Schram for helping us out with a variety of aspects that are
related to tracing, collecting, scanning and organizing literature data. Al Vrezec shared
his much appreciated views on taxonomic and bibliographic challenges related to
Scopoli and Little Owls.
We are highly endebted to Irene Pellegrino who was so kind as to make all pictures
of Little Owl skins available to us as the raw material for the illustrations. Koen Aeyels
was kind enough to help us out with the maps.
We are delighted with the poem by Steven Van Der Heyden who also created an
English version for this project. The real-life weathered or Corten Steel version of the
poem is 170 cm tall and can be enjoyed in Herzele, Flanders.
Special thanks to Norbert Lefranc and Martin Grüebler for their advice and support
for this second edition and to Alan Sieradzki for going through the final manuscript and
turning it into real English. The next steps in the creation of this book were in the
hands of the production team at Cambridge University Press. We are grateful for the
continuous effort and belief in this project by Megan Keirnan, Aleksandra Serocka and
our content manager, Jenny van de Meijden and copy-editor Jo Tyszka alike.
Crucial input of Little Owl insights were obtained from Anita van Dooren, Christien
Hermsen, Geco Visscher, Henk Wanders, Ruud Leblanc, Philippe Smets, Ricardo Tomé,
Simon Birrer, Guido Desmarets, Magnus Robb, Michael Exo, Koen Van den Berghe,
Reuven Yosef, Luis Reino, Boris Nikolov, Grzegorz Grzywaczewski, Rottraut Ille,
Hubertus Illner and the many volunteers of Natuurpunt Herzele and beyond.
We are greatly endebted to the contributing experts from each country: Albania –
Taulant Bino; Algeria – Paul Isenmann; Arabian Peninsula – Michael Jennings;
Armenia – Karen Aghababyan; Austria – F. Grinschgl, F. Kloibhofer, H. Steiner,
H. Wurm, Ille Rottraut, K. Huber, L. Engel-Klien, Lugmair A., Paar J., R. Zink, Stefan J.,
W. Geringer, W. Pegler; Azerbaijan – Elchin Sultanov; Belgium – Didier Van Gheluwe;
Brussels – Jean-Paul Jacob; Bosnia and Herzegovina – Kotrosan Drazen; Bulgaria – Assen
Ignatov, Boris Nikolov, Ignatov Assen, Petar Iankov, Stoycho Stoychev; Croatia – Sven
Kapelj, Tibor Kopacki, Vesna Tutis, Vlatka Dumbovicmazal; Cyprus – Christina
Ieronymidou; Czech Republic – Alexandra Pruchova, Karel Stastny, Libor Opustil,

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 Acknowledgements

Martin Dobrý, Martin Šálek; Denmark – Lars Bo Jacobsen, Egon Østergaard, M. B. Grell,
Pelle Andersen-Harild, Timme Nyegaard; Egypt – Mohamed Habib; Flanders – Guido
Desmarets, Isabelle Binnemans, Philippe Smets; Former USSR – Jevegni Shergalin;
France – Dominique Bersuder, Laurent Couzi, Olivier Hameau, Sebastien Dalloyau;
Georgia – Alexander Abuladze; Germany – Bettina Gerlach, Christoph Sudfeldt, Doris
Siehoff, Hubertus Illner, Kai Gedeon, Michael Joebges; Hungary – Daniel Hamori,
Károly Nagy; Italy – Gianpiero Calvi, Marco Mastrorilli; Israel – Reuven Yosef, Yoav
Perlman; Jordan – Fares Khoury; Khazakhstan – Askarisa Bekov; Luxembourg – Marc
Junio, Mikis Bastian, Patric Lorgé; Moldova – Silvia Ursul; Morocco – Michel Thévenot;
Netherlands – Alex De Smet, Pascal Stroeken, SOVON; Pakistan – Filza Ghafoor, Pankaj
Koparde; Poland – Grzegorz Grzywaczewski, Ignacy Kitowski, Tomasz Chodkiewicz,
Tomasz Wilk; Portugal – Lourecq Luis, Luis Reino, Ricardo Tomé, Rui Lourenço;
Serbia – Dimitrije Radisic; Slovakia – Chrenkova Martina, Jozef Ridzoň, Monika
Chrenková; Slovenia – Al Vrezec, Tomaž Mihelič; Spain – Blas Molina, Hugo Framis,
Juan Carlos Del Moral; Switzerland – Beat Naef-Daenzer, Christian Meisser, Françoise
Schmit, Klaus Robin, Paul Schmid, Peter Knaus; Tunisia – Paul Isenmann; Turkey – Ali
Kayahan, Kerem Boyla; Ukraine – Andriy-Taras Bashta, Tatiana Kuzmenko; United
Kingdom – Gary Clewey, Neil Calbrade BTO, Roy Leigh; Wallonia – Jean-Yves Paquet.
This project was unique in its form due to the fact that crowd funding was organized
to finance the illustrations. Two centerfolds, 22 unique plates and 32 chapter logos by
Joris De Raedt could be sponsored and magnificent gift cards purchased (Figure 8.7).
Theo Boudewijn co-ordinated fundraising in the Netherlands, Trui Mortier in Flanders
succeeding in a joint effort to close the financial gap. Obviously, we want to specially
thank our sponsors for their generous support and belief in this project.
Double Plate: Vogelbescherming Nederland, Uilenwerkgroep Waasland.
Plate: Anja Menck, Astrid de Roos, Erwin de Visser, Frederik Dierickx, Hein Bloem
(International Owl Center, Huston, MN, USA), Jouke Altenburg, STONE
Steenuilenoverleg Nederland, Van Der Haegen Dirk BVBA.
Chapter Logo: Jules Tolboom, René De Boom, Piet Vanhaverbeke, Peter De
Ruytter, Koenraad Bracke, J. Ter Stege, Henk and Wilma Lammers, H. Lammers,
Guido Desmarets, Geert van Dijk, Francis Bathy, Eddy Vandenbouhede, Dorita
Schattorie, Bert Van der Krieken, Bart Theunis, Arjan Berben, Natuurwerkgroep De
Kerkuil vzw, Marietta Schramm, F. and H. Van den Broeck, Erwin De Meyer,
Bernard Mortier.
Gift cards: Werner and Marinelle Mory-De Paepe, Rita Saerens, Marleen
Achtergaele, Lucrèce Maeckelbergh, Katty Van Severen, Kaatje Toye, Eddy Bruggeman,
Dries Mertens, Helen Mortier, Lode Mortier, Jeroen Mortier, Jean-Marie Vande Waetere,
Céline Gendron, Marc Van Iseghem, Koenraad De Wolf, Nadine and Patrick
Rouckhout, Dirk and Dorien De Man DMD DESIGN BVBA, Bart Van de Vel, TMTS
BV, Renaat Polfliet, Raes Marnix, Steven and Christel Rowaert-Stalpaert, Karel
Schotsmans, Jozef and Mie Van Nieuwenhuyse, Maarten and Joline Bekaert-
Snauwaert, Jan Tanghe, Jan Lenaerts, Dimitri and Mieke De Beul-Dennequin, Danny
and Rita Schockaert, Verschuuren, Friedel and Lieve Nollet-Wemaere, Michiels
Bernadette, De Zutter-Geerts, Elke Borms, Rita Saerens, Lieve De Vuyst, De Middeleer

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 Acknowledgements

Lore-Mertens, CODE-Projects CV, Michel Vangheluwe, Marc Van Brantegem, Mia

Vande Casteele, Etienne Vande Casteele, Versteels Etienne, Sigrid Balduck, Katty and
Frank Westerlinck, Silvie Geerts, Marijke Van Nieuwenhuyse, Hilde Van
Nieuwenhuyse, Bart Van Nieuwenhuyse, Geert Vanacker, De Middeleer Wim and
Marianne, Filip and Veerle Renaer-De Clercq, Johny Huvaere, Geert Jackers, Ward De
Groote, Marie-Claire Pierreu, Lieve De Middeleer, Karel Schotsmans, Mieke Casaert,
Griet Van Den Bosch, Marleen Verhamme, Heidi Jansegers, Ingrid Diependaele, Koen
and Lieve Dhoore-Jansegers, Saskia Collage, Michelle Mortier, Jef Melckebeke, Bjorn
Crabs, Mireille Diependaele, Pierre Vidts, Hilde Snauwaert and Wim Duthoit.
The authors are highly endebted to all people and organizations that supported
them in this project and apologize to all those that were forgotten in this overview.

xviii

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 Executive Summary

In this book we synthesize the substantial literature and knowledge base on the Little
Owl Athene noctua and detail the current understanding of its range-wide ecology,
genetics and subspecies, its population status by country, and offer a conservation
management strategy and outline a monitoring program for its conservation.
The Little Owl (Plate 1) is a small (19–25 cm, 160–250 g), relatively long-legged
nocturnal “chunky” owl with a short tail and a round head. The updated world
distribution of the Little Owl and its subspecies is substantial, from the Atlantic coast
of Britain and Iberia, north to Denmark, Sweden and Latvia, east to Asia Minor, Levant,
Arabia, Asia to China and Manchuria, south to northern Africa and the Red Sea coast to
Somalia and Eritrea. It has been introduced in New Zealand and England. It is con-
sidered fairly common to common in 45 countries, and uncommon, rare or a vagrant in
29 (78 in total) other countries. The Little Owl occupies open country with groups of
trees and bushes, rocky country, deserts and semi-deserts with rocks, ruins, oases,
pastureland with scattered trees, old orchards, along rivers and creeks with pollarded
willow and other trees, parkland and edges of semi-open woodland, and farmsteads and
urban areas with surrounding cultivated lands. Home-range sizes for a breeding pair of
Little Owls tend to be small, averaging 0.5 km². Owls tend to be resident, with most
young dispersing relatively short distances (between 600 m and 4000 m). The diet of
the owl includes relatively small-bodied prey: invertebrates, rodents, reptiles, frogs and
small birds. The current population status of this owl differs by country, with declines
apparent or percieved in many countries. As it serves as an important ambassador for
the small-scale, half-open landscape (ranging from pastoral landscapes with scattered
trees to stony steppe deserts), we detail a conservation strategy and consistent survey
methodology to further the significant and long-term international efforts being con-
ducted by scientists, fieldworkers and volunteers.
The update of this volume after two decades of international co-operation, aided by
standardized methods for data collection, management and analysis, and dissemin-
ation led to the almost doubling of Little Owl publications, almost exclusively in
English across the entirety of Europe and gradually expanding over most of its distribu-
tion range. Technological advances like PDF, GIS, advanced statistics, international
standardized databases of geocoded and time-stamped pictures and sound recordings
led to real-time exchange of papers, data, advice, images, sound recordings and maps,
facilitating analysis and editing. The major difference was definitely the digital artwork

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 Executive Summary

FACT SHEET Athene noctua

LENGTH NEST CAVITY

19–25 cm Natural cavity of nestbox
approximately 3 m above the
ground. Breeds in buildings,
e.g., barns, sheds.

WINGSPAN DISTRIBUTION
54–58 cm (Figure 0.1) Resident throughout the global
distribution area.

WEIGHT TREND
160–250 g Breeding populations decline
because of loss of suitable habitat
and mechanization of agriculture.

CHARACTERISTICS MAXIMUM AGE

Small, compact, not much bigger than Average life expectancy is 2.3 to 2.8
Common Blackbird. Brown with white years.
spots. Yellow eyes.

FLIGHT BREEDING SEASON

Fast, low, undulating flight. Broad, rounded April–May. Mostly 3–5 eggs,
wings. sometimes 6 or 7. 28–33 days
incubation. After 32–34 days the
young fledge.

CALL
Mostly in spring, repetitive “kie-ew” or
“miaw.” Territorial call elongated
“kuuuup.” “kjie-kjie-kjie.” Juveniles
“hssss.”

HABITAT
Diverse and near short-grazed grassy
vegetation with hedges and older trees.

Figure 0.1 Silhouette wingspan of Little Owl.

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 Executive Summary

by Joris De Raedt who worked with photographic input of owls and landscapes,
collected by travelling from Iberia to China, to all the places where subspecies were
discovered, with Google Maps. Drawings were custom made, as ordered by the authors,
adapted, standardized and optimized, leading to excellent, tailor-made plates and
illustrations. This state-of-the-art contribution aims at perfecting the knowledge and
understanding of the species, study methods, proven conservation and management
approaches that will undoubtedly lead to international co-operation between conser-
vationists, well-trained academics and field workers across the entire distribution range
of the Little Owl. The ambition of this book is to serve as yardstick for advanced citizen
science and conservation management.

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 Chapter 1: Introduction: Framework

For centuries, all across the world, humans have had a continuous and strong cultural
relationship with owls (Marcot and Johnson 2003), traceable back 30 000 years to caves
in France. Some cultures view owls as omens of back luck, sickness and death, while
others view them as creator beings or helping spirits, having profound wisdom, oracular
powers or the ability to avert evil. Depending on where you are within the range of the
Little Owl, both of these divergent (bad or good omens) are still held for owls.
In the mid twentieth century, drastic declines in several owl species attracted the
attention of ornithologists, conservationists and researchers. Throughout Europe the
decline of the species is mainly being caused by habitat destruction, especially due to
the intensification and mechanization of agriculture. In order to counter this negative
situation, scientists and conservationists have realized the need for international co-
operation through a multidisciplinary approach.
Early literature on the Little Owl reflected general studies on biology, distribution,
diet, nesting and habitat. In 2008 our first edition of this book (Van Nieuwenhuyse
et al. 2008) was published. The publication gave a review of most of the literature that
was available at the time. Since then, substantial new findings on the Little Owl have
been produced from a growing number of countries within the range of the owl. Since
our first edition, much has changed. Literature is now always digitally available as pdf
files, primarily in English or at least containing English abstracts. Contacting authors
has become simple, no matter which country they originate from. High-quality digital
pictures are now available which are time-stamped and georeferenced as input for
solving the taxonomic challenges within the species, yielding sample sizes that are
10- to 100-fold times larger than in the early days of ornithology. The georeferencing
has allowed virtual travel to landscapes by means of maps and local landscape pictures
enabling landscapes across the entire range of the species to be discovered, even in the
most remote and hostile countries. Last but not least, the evolution of the artwork
proved crucial for this book, having high-quality illustrations available digitally created
by Joris De Raedt. This technological advancement in merely one decade has allowed
us to take a giant leap in relation to our first edition.
The aim of this current book is to synthesize the substantial literature and knowledge
base on this species and provide a more complete understanding of its range-wide
ecology and conservation status. While rooted in science, this book has the principal
aim of making the insights accessible to the wider international public. Recognition
and appreciation of volunteer work is also of crucial importance for nature conser-
vancy. Virtually unlimited input of volunteers not only helps to construct huge,

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 Chapter 1: Introduction: Framework

standardized and reliable databases, it also helps to broaden the social network of
nature conservation. In this way, nature conservancy is brought closer to the general
public. The best guarantee for long-term conservation is to obtain the critical scientific
insights resulting from co-operation between scientists and conservationists. The over-
all importance of detailed large-scale and long-term data collection by volunteers aided
by scientists, and their contributions to applied as well as to basic scientific research,
becomes increasingly important. Hence, a perfect blend of fieldworkers, data analysts,
photographers and publishers is needed.
Finally, the Little Owl is a useful ambassador for small-scale half-open landscapes
(ranging from pastoral landscapes with scattered trees to stony steppe deserts) and as
such deserves special attention. The species has a multitude of strategic features that
make it a flagship of the rural environment: the species is very well known among the
public, it is still present in reasonable numbers in most European countries and is
readily observed, and the Little Owl can offer us insights into methods to implement
nature restoration. Due to its “high cuddle factor” it is a perfect vehicle to transfer
nature conservation values to the broader public.
The framework of this book (Figure 1.1) reflects the complexity of the situation of
the species at different scales. To position the Little Owl in a cultural context we look at
the history and cultural traditions connected to the species and interpret this in the
light of current knowledge (Chapter 2). Results of a large-scale research project dealing
with owl myths and culture spanning over 6000 interviews collected in 26 countries are
reported. Then we start with the bird itself and describe the taxonomy, the races to
settle some taxonomic discussions (Chapter 3) and the morphology of the species
(Chapter 4). Major progress has been made on genetic, morphological and biogeo-
graphical insights into the subspecies of the Little Owl. More data were collected on
size and coloration of stuffed birds from various historical collections. Skins were
measured and photographed, opening up new standardized, historical data. Genetics
allowed for more clarification on the postglacial colonization of the different European
subspecies. The distribution of the different subspecies and recent population estimates
for the Western Palearctic are given in Chapter 5 to illustrate the diversity and local
peculiarities. We then zoom in on both biotic and abiotic factors that influence Little
Owl numbers. The habitat is described and the relationships between the landscape and
the species are characterized in Chapter 6. A principal biotic factor inside the habitat
is the food delivering the crucial energy input for the birds. Abiotic factors such as
breeding cavities and perches show their importance for breeding and foraging
efficiency to minimize the energetic cost for the birds (Chapter 7). Chapter 8 focuses
on the breeding season and the genetic offspring of the birds, including clutch size,
hatching and fledging success in relation the age of the birds. We then describe the
behavior of Little Owls (Chapter 9), mainly based upon captive breeding data and
complemented by almost two decades of webcam observations. The next section zooms
in on the limiting factors for population regulation that influence the number of owls
in a given geographic environment (Chapter 10). Input into the local population from
outside is through immigration, re-introduction or supplementation. Local offspring
feed back new owls into the local population too. Population outflow is observed

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 Chapter 1: Introduction: Framework

HUMAN ACTIVITIES
Conservation and
Management (11)
Research History
Priorities (12) and Traditions (2)

POPULATION
Breeding Biology (8)
Behavior (9)
Population Regulation (10)

HABITAT OF THE SPECIES

Habitat (6)
Diet (7)

(7)
SPECIES
Fossils, Genetics and Taxonomy (3)
Morphology and Body Characteristics (4)
Distribution and
Population Estimates (5)

POPULATION

HABITAT

SPECIES

HUMAN ACTIVITIES

Figure 1.1 Framework of the book.

through emigration and local mortality. Both external input and outflow, and local
offspring and mortality influence the number of owls in a given location. Mechanisms
that interact between local populations such as migration, metapopulation functions
and the occurrence of sinks and sources place the individual parameters in a wider

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 Chapter 1: Introduction: Framework

context. The insights obtained through population studies have proven to be crucial in
conservation. After describing the main causes for declines in the species, we summar-
ize the knowledge presented so far in the form of an international Little Owl conser-
vation and management strategy and best practices (Chapter 11). This chapter will also
discuss specific methods involved in reintroducing and translocating owls, captive
breeding and releasing Little Owls into the wild to establish or improve local popula-
tions. Finally, we conclude this book with an overview of the key points raised and the
most important open questions and advice for future studies (Chapter 12).
Readers will find supplementary resources online at www.cambridge.org/littleowl.
These include appendices as well as tables which previously appeared in the first edition
(Van Nieuwenhuyse et al. 2008).

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 Chapter 2: History and Traditions

Throughout human history, owls have variously symbolized dread, knowledge,

wisdom, creator beings, death, witchcraft and religious beliefs in a powerful spirit
world. A small but important body of literature is beginning to document the important
aspects of owls and culture both across civilizations and through time (e.g., Medlin
1967, Simmons 1971, Stuart 1977, Holmgren 1988, Gimbutas 1989, Weinstein 1989,
Enriquez and Mikkola 1997, Marcot and Johnson 2003).
The most ancient representations of owls date from the Upper Paleolithic (13 000
BC) and are seen in two caves in France. The first site reflects three owls (generally
considered Snowy Owls) in the “Gallery of the Owls,” in the French cave of Les Trois
Frères in Ariège. A second site, in the Cave Chauvet represents a single “eared” owl that
looks to be a Long-eared Owl (or Eagle Owl). Other prehistoric sites with owl art and
symbolism are found in an array of locations around the world, such as the Victoria
River region of northern Australia, the Columbia River region of northwestern USA and
a Mayan cultural site in what is now Guatemala (Marcot and Johnson 2003). In the east
Mediterranean, archaeological digs in Syria and Jordan have recovered anthropo-
morphized owls in the stone or clay as early as the prepottery Neolithic period
(8000–6500 BC)(Gimbutas 1989: 190–195). Representations of Little Owls were found
associated with the Xian culture (a society of agriculturists) in Inner Mongolia, dating
from 8000–7500 BC (Schönn et al. 1991).
Owls have been viewed by human societies in many different ways. Some viewed owls as
a single group of animals (e.g., species were not differentiated and any owl was a bad omen),
and others viewed large owls as bad (harbingers of bad luck, illness, death), whereas small
owls were not viewed in this way. Still others were attentive to specific owl species for
medicine, religious or hunting applications (and of course for current-day conservation
purposes). Societies and societal values change, and whereas owls may have been viewed in
one way or another in the past, current cultural views may differ. For example, during the
Middle Ages, the owl was linked to witches and bad spirits, but nowadays in Turkmenia, the
Little Owl is a sacred species and to kill one is a great sin (Shukurov in Khokhlov 1995).
Another example of this change may be seen (in general) in Western culture, where newer
perspectives pertain more to scientific understanding and conservation needs of owls; a
specific example is how Spotted Owls have become symbols of old-growth forest protection.
Included in her book on the religion of the Old European Great Goddess, Gimbutas
(1989) presents a pictorial “script” consisting of signs, symbols and images of divinities.
The main theme of goddess symbolism is the mystery of birth and death and the
renewal of life, not only human but all life on Earth. Symbols and images cluster around

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 Chapter 2: History and Traditions

the parthenogenetic (self-generating) Goddess and her basic functions as Giver of Life,
Wielder of Death and as Regeneratrix. Overall, owls were deeply feared, and viewed as
the harbingers of death. The Goddess in the guise of an owl was prominent in stone,
pottery (e.g., terracotta figurines, burial urns), engravings on statues, schist plaques,
bone phalanges laid in graves, drawings, amber figurines, wooden posts and gold
sculptures in this prehistoric religion which extended from the Neolithic to the Early
Bronze Age (general dates of 6500 BC to 2500 BC). Archaeological artifacts and images
of owls in this religious context were generic or abstract in shape and pattern, and
specific images distinguishable as Little Owls are not apparent. The features that char-
acterize owls (round eyes and beak) can be seen on statue menhirs of southern France
and the Iberian Peninsula, and in reliefs and charcoal drawings in the hypogea of the
Parisian basin. A series of stelae and drawings of the Owl Goddess from Brittany and the
Paris basin are depicted with breasts and one or more necklaces. Beautiful examples of
owl-shaped burial urns dating from ca. 3000 BC come from the Baden culture in
Hungary, from Poliochni on the island of Lemnos and from Troy. They have wings,
the characteristic owl beak connecting arched brows, and sometimes a human vulva or
a snakelike umbilical cord, symbols of regeneration. In continental Greece, gold
sculptures have been found in the tholos tombe (Kakovatos, Pylos) and from shaft
graves in Peristeria of the fifteenth century BC (Marinatos 1968: pl. 58). In spite of
the gloomy aura which surrounds it, the owl has also been endowed with certain
positive qualities, such as profound wisdom, oracular powers, and the ability to avert
evil. However, this latter ambivalent image is a dim reflection, diffused through time, of
the owl as an incarnate manifestation of the fearsome Goddess of Death. Perhaps she
was respected for her grim but necessary part in the cycle of existence, as the agony of
death which we take so much for granted was nowhere perceptible in this symbolism
(Gimbutas 1989).
With its large distributional range across Europe, the Middle East and Asia, and an
ability to co-exist as a commensal with many human habitations, not surprisingly, the
Little Owl has figured prominently in many cultural beliefs, and in a variety of ways.
The common names given to this species across the countries are linked to its activity
(Nightbird), to its voice (Kliwitt), to its morphology (Little Owl), to its food (Lark Owl or
Sparrow Owl), to beliefs (Death Bird), to its habitat (House Owl, Willow Owl, Stone
Owl) and to mythology (Bird of Minerve [Roman]).
The Little Owl was first formally described to science by Giovanni Antonio Scopoli in
1769 (Scopoli 1769). The origin of its scientific name, Athene noctua, combines the
genus Athene, derived from the goddess of wisdom Athena in Greek mythology, and the
species noctua, as derived from the nocturnal characteristics of this bird of prey. An early
name for the Little Owl “Strix passerina” was noted on Thomas Bewick’s 1797 drawing.
This scientific name was not used as the name Athena noctua more accurately placed the
Little Owl within the taxonomic naming convention of owls. While the origin of the
English name Little Owl is probably linked to the size of the owl, the German and Dutch
names – Steinkauz and Steenuil, respectively – refer to the open habitat of this species
that it uses in many countries, where it breeds in piles of stones. Common names for
the Little Owl in other countries include:

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 Chapter 2: History and Traditions

Italy: Civetta
Spain: Mochuelo Comun
Portugal: Mocho
France, Wallonia, Switzerland: Chevêche d’Athéna
Germany, Austria, Switzerland: Steinkauz
Greece: Κουκουβάγια (Koukouvaya)
Denmark: Kirkeugle
Netherlands, Flanders: Steenuil
Turkey: Kukumav
Russia: Domovogo sycha
Georgia: Tchoti
Hungary: Kuvik
Mongolia: Chotny bugeechej
Somalia: E’yu

The relationship between the Little Owl and Greece is very long indeed, and is one of the
most well-known affiliations between a particular owl species and a prominent society.
This affiliation is again alive and prospering, with the new Greek 1 Euro coin showing the
same Little Owl design as the former tetradrachm coin from 2500 years ago (Figure 2.1).
In Greek mythology, Athena was the daughter of Zeus and originally a Mycenaean
palace goddess. Her function later expanded to include the role of guardian of cities,
war goddess, patroness of arts and crafts, and promoter of wisdom. Always shown
modestly clothed and often armed, the Little Owl was her special bird.
An emblem refers to some distinctive and characteristic or activity for which the issuing
city is known. The Attica tetradrachm coin bearing an obverse head of Athena and reverse

Figure 2.1 Greek 1 Euro coin and former drachm silver coin showing the image of the Little Owl.

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 Chapter 2: History and Traditions

owl with olive branch is perhaps the best known example of a city emblem. The Greek
goddess Athena is a reference to the city (in Greek, Athenai) that honored her as its chief
protective diety. The owl as Athena’s favorite bird, and the olive, which was one of the
city’s most lucrative exports, in time came to stand for Athens throughout the
Mediterranean world. With rich silver mines at Laurium at the southern tip of Attica, the
Athenians were able to export bullion for foreign exchange at a time when most Greek
states restricted coinage to home use. The necessity to create a standardized Greek currency
that would be widely acceptable demanded a rigid uniformity in metallic purity, type and
style seldom seen elsewhere in the Greek world. What resulted from ca. 525 BC onward was
the famous series of tetradrachms that carried an obverse image of Athena’s head on one
side, and a Little Owl with an olive branch and a cresent moon on the reverse. This type
was kept in circulation for the next two centuries. The coins were made of silver, and their
nickname “owls” become synonymous with Athen’s commercial power. The Obal, Stater,
Drachm, Didrachm, Tetradrachm, Octodrachm, and Decadrachm are terms for common
denominations based on these weight systems.
About 77 BC, the Roman scholar Pliny the Elder assembled more information about
the owl in a few chapters of Book X in his Historia Naturalis. He specified the Little Owl,
the Eagle Owl and the Screech Owl. His observations were laced with beliefs not entirely
sound by today’s zoological standards, but were studied as gospel during the Middle
Ages (Medlin 1967: 20).
Little Owls are also of historical importance for the Italian Peninsula since they
appear in some classic fables (Mastrorilli 2005). Fedro’s classic fable “The Cicada and
the Little Owl” describes the capture of a cicada by a Little Owl during its diurnal hunt.
The fable describes the owl’s hunting strategy as a sign of its intelligence. It also
demonstrates that the Little Owl was a loved species in the Roman period.

XVI. Cicada et Noctua

Humanitati qui se non accommodat
plerumque poenas oppetit superbiae.
Cicada acerbum noctuae conuicium
faciebat, solitae uictum in tenebris quaerere
cauoque ramo capere somnum interdiu.
Rogata est ut taceret. Multo ualidius
clamare occepit. Rursus admota prece
accensa magis est. Noctua, ut uidit sibi
nullum esse auxilium et uerba contemni sua,
hac est adgressa garrulam fallacia:
“Dormire quia me non sinunt cantus tui,
sonare citharam quos putes Apollinis,
potare est animus nectar, quod Pallas mihi
nuper donauit; si non fastidis, ueni;
una bibamus.” Illa, quae arebat siti, simul gaudebat uocem laudari suam,
cupide aduolauit. Noctua, obsepto cauo, trepidantem consectata est et leto dedit.
Sic, uiua quod negarat, tribuit mortua.
Fedro’s classic fable (Latin version)

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 Chapter 2: History and Traditions

Leonardo da Vinci (1452–1519) wrote one fable on Little Owls and Thrushes.
He described the use of the Little Owl as a “decoy” during the hunting of Thrushes.
This is an important source of evidence that Little Owls were used for hunting after
the Middle Ages.
Illustrations became as important as word descriptions to naturalists and others.
Conrad Gessner (1516–1565), from Zurich, Switzerland, authored one of the first
printed books containing birds in the sixteenth century. His third volume, published
in 1555, was the largest in a series of five books on animals (i.e., Historia animalium), and
contained a nice image of a Little Owl (“Night Owl”) (Figure 2.2); [“Von der Eul/oder
Nacht-Eul/und erstlich von ihrer Gestalt Ulula”].

Figure 2.2 Image of the Little Owl from Conrad Gessner’s “Vogel-Buch” (bird book) published
in 1555.

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 Chapter 2: History and Traditions

Thomas Bewick published his authoritative History of British Birds in England in

1797 and 1804. Bewick rejected the copper-engraving medium that was common to
book illustration at that time, in favor of incizing the end grain of a hardwood block
with a graver, thus achieving detail in picture and durability in printing that was
previously impossible in wood. Bewick’s first edition of the History illustrated the
Long-eared Owl, Short-eared Owl, a female Horned Owl, the Snowy Owl, Tawny Owl,
and Little Owl (Figure 2.3). In the book Birds of Europe, Nilsson in Gould et al. (1837)
named the Little Owl Strix nudipes (Figure 2.4). Refinements were made in the drawings,

Figure 2.3 Image of the Little Owl from Thomas Bewick’s 1797 History of British Birds.

Figure 2.4 Little Owl image from the Birds of Europe Vol. 1 (Raptors), 1837 [Strix nudipes (Nilsson),
Noctua nudipes (Mihi) – illustrated by Edward Lear.

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 Chapter 2: History and Traditions

Figure 2.5 Engraving of a Little Owl from Howard Saunders’s Manual of British Birds (1889).

and Howard Saunders published his significant An Illustrated Manual of British Birds in
1889 (Figure 2.5).
Until the beginning of the twentieth century, hunters in France enjoyed shooting
the Little Owl (Delamain 1938). In the ninteenth century, despite the prejudices against
this species, the Little Owl was also appreciated by people and held as a captive bird
(von Risenthal 1879) and sold by fowlers in Germany (Wemer 1910). In France
(Crespon 1840) and in Belgium (Dupond 1943), the Little Owl was used to hunt little
birds such as larks, because the mobbing behavior of passerines towards the Little
Owls was a good technique to attract them. In Italy, Little Owls were domesticated
and kept (with their wings clipped) in houses or gardens, where they caught rodents,
slugs and insects (Goddard 1917).
A particularly interesting example of the cultural use of Little Owls comes from
Crespina, Italy. Crespina is located 25 km from Pisa in the Toscana region of Italy.
The community has a specific monument dedicated to the Little Owl called the “Place
of the Little Owl Fair” (Schaaf 2005). This little town was a center for the rearing of owls
in captivity to be used in a Little Owl competition. Around a century ago (ca. 1900),
Little Owls were taken out of their nests, reared and put in special boxes in farms of this
Italian region to be used as bait for the hunting of larks. They were then sold on the
Little Owl market while tied up on a roost (for an example of this market, see Figures
2.6–2.8). The nobility (upper-class people) commonly hunted in the countryside using

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 Chapter 2: History and Traditions

Figure 2.6 Cover of the journal La Domenica del Corriere in 1903 represented the Little Owl market.

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 Chapter 2: History and Traditions

Figure 2.7 Drawing from 1896 showing a seller of Little Owls and interested women.

the Little Owls as bait. People pulled the tethers to stimulate the Little Owl to fly;
larks were attracted by the owl and were themselves caught in nets. There was a
“civettaio” (“Owl man” or “Owl-keeper”) charged with tending to the Little Owls.
The owl market of Crespina took place annually on September 29. At these occasions,
it was possible to listen to a range of bird imitators and attend the Little Owl competi-
tion. This hunt reflects an ancient tradition that started in 350 BC and lasted until
the twentieth century in Italy, and from the seventeenth to the twentieth century in
Germany.
A national policy in Italy (L157/1992) and a regional policy approved in Toscana
(L3/1994) have recently banned the annual owl market. The laws forbid the killing,
catching or keeping of any owl species in aviaries, or to possess dead owls or parts
of them. The use of owls as bait is also forbidden. Because of the traditional aspect
of the hunting method, the policy was modified to allow some rearing stations

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 Chapter 2: History and Traditions

Figure 2.8 Photo of a hunter and his wife at Crespina market.

for wild animals. A Little Owl rearing station in Crespina covers 2 ha and was
sponsored by hunters.
Little Owls were introduced in England and in New Zealand at the end of the
nineteenth century and beginning of the twentieth century. After its release, the species
was considered a pest by many gamekeepers and farmers in England (Dawson-Smith
1913).
The history and traditions of the Little Owl are truly long, rich and varied, and grow
with additional recoveries of artifacts from archaeological sites, as well as evolving
cultural views. In closing this chapter, we urge reviewers of owl myths, traditions and
lore to closely scrutinize the information they assemble, to determine whether the ideas
and symbolism described in text and artifacts still apply in contemporary societies, or
whether they are part of a colorful but quaint past. Whatever the outcome, no doubt
the lure and fascination of the Little Owl will continue to make a significant impression
on peoples of the world.

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 Chapter 2: History and Traditions

Figure 2.9 Belgian Little Owl beer “Steenuilke” brewed by Brouwerij De Ryck as part of a European
Leader project dedicated to the conservation of the species in Flanders.

Many countries use the image of Little Owls on a diverse range of postage stamps.
Human appreciation for the Little Owl continues to grow, and the image of the owl has
recently been used for the marketing of a Belgian Little Owl beer (Figure 2.9).

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 Chapter 3: Fossil Evidence, Taxonomy
and Genetics

Chapter Summary
Different owl species and subspecies used to be defined in the early days of taxonomy
uniquely based upon morphological features like size and color in specific geographical
regions across the world. Recently, more differentiators are taken into account to define
new species to avoid upgrading them from a subspecies based upon only one or two
aspects. We consider the following concepts in the discussion of the subspecies of the
Little Owl, letting the old taxonomy prevail if there is a lack of alternative multi-
disciplinary evidence. The biological species concept (Mayr 1940) takes biological
characteristics into account, the morphological concept studies especially morpho-
metrics and coloration of the bird. Recently, a third principle was fully adopted, the
so-called phylogenetic concept (Cracraft 1983, Dickinson and Remsen 2013). Through
mitochondrial DNA studies, a lineage can be established that eventually leads to a
common ancestor. Different species thus have different ancestors, up to some point
in the past. This research is complemented with fossil evidence for the evolution of
species. Vocalizations of Little Owls across the range are increasingly documented and
taken into account. The last approach considers the geographical distribution and the
use of validated, geocoded, high-quality photographic input.
In this chapter we present the current fossil evidence for ancestors of the Little Owl.
We describe the historical context in which subspecies have been defined. For a while it
was a kind of international sport to distinguish as many subspecies as possible. Hence,
there was quite some evolution in the insights on the subspecies. We illustrate the way
the subspecies were described for which we now have evidence for their relevance. We
consider 14 subspecies of Little Owl where we have found substantial evidence. This
means that we have one more subspecies compared to the 13 subspecies that were
retained in Van Nieuwenhuyse et al. (2008). Since then Athene noctua impasta has been
studied in more detail and is found to deserve the status of real subspecies which we will
justify here. We cite the original descriptions of the subspecies as found in historical
publications in the original language and translated into English. We combined the
textual descriptions of Hartert (1913) and Vaurie (1960) based upon 216 skins in four
museums with those of Pellegrino et al. (2020) based on 282 skins from five other
museums and validated the descriptions by studying over 5000 georeferenced high-
quality pictures (macaulaylibrary.org) across most of the distribution range, including
Somaliland (somaliensis) and Eritrea (spilogastra). We were able to validate and

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 3.1 Fossil Evidence

complement all textual descriptions through pictures of typical plumage for the sub-
species, but we equally found aberrant plumage and clinal variations across the distri-
bution areas of most subspecies during the photographic validation. Futher, we
validated morphological trends and differences in geographical areas related to each
individual subspecies in detail by studying high-quality standardized pictures of the
skins of some of the type specimens (where available) from five major museums
(Pellegrino et al. 2020).
Each subspecies illustrated by Joris De Raedt was based upon pictures that were taken
at or near the type locations of the subspecies. Typical landscape elements were
obtained by studying Google Street View or available characteristic landscape pictures
presented by Google at or near the locations of the owl pictures.
Extraordinary specimens are illustrated separately in dedicated plates that are only
georeferenced without clean subspecies classification.

3.1 Fossil Evidence

Little Owls are small strigiform birds commonly present in Mediterranean islands
during the Pleistocene and Holocene with four fossil species: Athene angelis (Mourer-
Chauviré et al. 1997) from the Middle and Late Pleistocene of Corsica, Athene cretensis
(Weesie 1982) from the Late Pleistocene of Crete and Armathia, Athene trinacriae from
the Middle Pleistocene of Sicily, and Athene vallgornerensis (Guerra et al. 2012) from
the Early Pleistocene of the island of Mallorca (Balearic Islands).
Athene trinacriae (Pavia and Mourer-Chauviré 2002) is described within the
fossil assemblage of some Middle Pleistocene localities in Sicily. The species is
similar in size to the extant A. noctua, but differs by its longer legs and some
morphological characteristics, probably as a result of a mainly terrestrial lifestyle.
Athene trincacriae is smaller than A. angelis from the Pleistocene of Corsica, and has
a longer and thinner tibiotarsus and tarsometatarsus; it also differs by its smaller
size and by its relatively longer legs from the endemic form from the Pleistocene
of Crete and Armathia, Athene cretensis. Athene vallgornerensis (Guerra et al. 2012)
was discovered in a fossil assemblage from the Early Pleistocene of the island
of Mallorca (Balearic Islands) in Cova des Pas de Vallgornera (Llucmajor, south of
Mallorca). The presence of this new species in Mallorca improves the knowledge of
the evolution of Little Owls on the Mediterranean islands, where two main types
of insular Little Owls seem to have evolved, one displaying a short and robust
tarsometatarsus (A. angelis and A. vallgornerensis) and the other with a slender and
longer tarsometatarsus (A. cretensis and A. trinacriae), associated with relatively
shorter wings, which have been interpreted as indicative of more terrestrial life-
style than Athene noctua (Weesie 1982, Pavia and Mourer-Chauviré 2002, Louchart
2005). Long legs are also characteristic of A. cunicularia that suggests terrestrial
habits, while A. angelis would have had a less terrestrial lifestyle, according to its
body shape and proportions (Louchart 2005). The lifestyle of A. vallgornerensis
remains unkown.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

The presence of endemic species of Athene on Mediterranean islands during the

Pleistocene is related to particular environmental and ecological conditions. The long
geographical isolation of the islands from the mainland, the presence of abundant
terrestrial prey and the absence of terrestrial carnivores, together with the sedentary
habitus of the genus, favored genetic isolation and the formation of endemic forms, as
was also suggested by Weesie (1982). The extinction of the endemic forms is probably
related to other ecological factors, such as paleoenvironmental changes or competition
with terrestrial predators, e.g., foxes and hedgehogs, which arrived from the mainland.
Their arrival is due either to natural colonization after the new connection of the
mainland with the island, as is the case of Sicily, or to human introduction, as is the
case for Crete (Weesie 1982), or both. Such colonization would be a cause of concern
for the extinction of Athene trinicriae; this species in fact probably mainly fed on small
endemic rodents and its extinction is related to the arrival of the new faunistic element
of the Elephas mnaidriensis Faunal Complex.
The present distribution of Athene noctua on the major Mediterranean islands is
probably due to recent colonization during the Late Pleistocene. During the Early and
Middle Pleistocene, in fact, most of the Mediterranean islands were much more isolated
because of the higher sea level. During the Late Pleistocene the lower sea level, as a
result of eustatic changes and tectonic activity, made it possible for some terrestrial
animals, or sedentary birds like Athene noctua, to spread on many Mediterranean
islands.
Fossil Little Owls Athene noctua have been described from the Pleistocene and
prehistoric period from 14 sites in 11 countries across the species’ range (Brodkorp
1971, Andrews 1990).

3.2 Discovery of the Species

Since the Little Owl is one of the most synanthropic owl species in Europe, it is not
surprising that the species was already known to ancient authors such as Pliny the
Elder, since the Little Owl also inhabited Roman cities (Jashemski and Meyer 2002). The
Little Owl was later described or mentioned in many early ornithological works in
Europe under the names Noctua (Magnus 1495), De Noctua (Gesner 1555,
Aldrovandus 1646, Jonston 1650) and Noctua minor (Willughby and Ray 1676). Many
of these works contain detailed descriptions and even illustrations (Figure 3.1).
However, official scientific naming began with the publication of the 10th edition of
Systema Naturae by Carl Linnaeus (1707–1778) in 1758. Linnaeus (1758) described most
European owl species with descriptions that are still valid today, but also created some
major confusions, especially with species that do not live in Sweden and with which
Linnaeus therefore had no real experience. This includes the Little Owl, which occurs in
most parts of Europe, but not in Sweden.
Under the name Strix passerina, Linnaeus (1758) described an owl resembling the
Little Owl, citing several references to older European authors who described the bird
extensively. Linnaeus (1758) even referred to illustrations that clearly depicted the

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 3.2 Discovery of the Species

Figure 3.1 Little Owl illustration from Ornithologiae (Aldrovandus 1646) named De Noctua.

Little Owl, such as Frisch (1734–1763; Figure 3.2). The description by Linnaeus (1758)
was later taken up by many authors, introducing Strix passerina as the accepted scientific
name for the Little Owl (e.g., Latham 1790, Naumann 1803, Temminck 1815, Boie
1822, Freyer 1842). The first to revise Linnaeus’s owl descriptions was the Swedish
ornithologist Peter Gustaf Tengmalm (1754–1803). He checked several references and
illustrations such as Aldrovandus (1646), Jonston (1650) and Frisch (1734–1763) and
found that this small Italian owl does not live in Sweden and that Linnaeus (1758)
certainly made a mistake (Tengmalm 1793). There are two other small owl species
similar to the Little Owl that live in Sweden. Tengmalm (1793) referred to the illustra-
tions of Olof Rudbeck Jr. (1660–1740), professor at the University of Uppsala, whose
student was Carl Linnaeus (Walters 2003). Rudbeck’s excellent hand drawings were

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

Figure 3.2 Little Owl illustration by Johann Leonhard Frisch (1734–1763) cited by Linnaeus (1758) in
his description of Strix passerina.

admired by Linnaeus and became his main references in describing bird species
(Linnaeus 1746), although they were not mentioned in Systema Naturae (Linnaeus
1758). From these illustrations, Tengmalm (1793) recognized that Linnaeus (1758)
described the Boreal Owl under the name Strix funerea (Figure 3.3) and the Pygmy
Owl under the name Strix passerina (Figure 3.4). Subsequently, Tengmalm (1793) con-
cluded that the Little Owl was actually described by Scopoli (1769) from Carniola (now
Slovenia). Low accessibility to references largely contributed to misinterpretations of
the original species descriptions in the eighteenth and nineteenth centuries, and even
Tengmalm’s (1793) correct designation of Strix noctua for the Little Owl was later
incorrectly attributed to the Boreal Owl by Retzius (1800).

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 3.2 Discovery of the Species

Figure 3.3 Illustration of the Boreal Owl (Aegolius funereus) by Olof Rudbeck Jr., which matches the
description by Linnaeus (1758) Strix funerea.

Figure 3.4 Illustration of the Pygmy Owl (Glaucidium passerinum) by Olof Rudbeck Jr., which matches
the description by Linnaeus (1758) Strix passerina.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

Figure 3.5 Joannes Antonius Scopoli (1723–1788) – the author of the first scientific decription of the
Little Owl (Athene noctua) according to binominal Linnean nomenclature.

However, even Joannes Antonius Scopoli (1723–1788; Figure 3.5), the author of the
first scientific description of the Little Owl according to Linnean binominal nomenclat-
ure, was misled by the vague descriptions from Linnaeus (1758). In Annus I. Historico
Naturalis, Descriptiones Avium, Scopoli (1769) actually described the Little Owl in great
detail under the name Strix passerina and supplemented the description by Linnaeus
(1758) with the remark that the species is used by hunters to catch passerines and breeds
in chimneys. The Italian (Civetta) and German names (Tschiavitl, Hauseule, Stockeule,
Käuzlein) also clearly designate the Little Owl he found in Italy. The description by
Linnaeus (1758) was apparently so clear to Scopoli that he had no doubt about it and
did not refer to it, even in his correspondence with Linnaeus, in which they discussed, for
example, the species identity of the Scops Owl (Otus scops) (Soban 2004). Nevertheless,
the name Strix passerina was already used by Linnaeus (1758) for the Pygmy Owl, as
clarified by Tengmalm (1793). Therefore, the second description of the Little Owl by
Scopoli (1769) as Strix noctua must be taken into account (Tengmalm 1793, Hartert 1913):

Strix noctua
Diagn. Pallide rufa, fuscisque maculis longitudinaliter variegate. Irides flava.
In M.p. (Musei Proprii) Carniola indigena, et in sylvis circa Labacum copiosa.
Statura Columbae.

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 3.3 Discovery of the Subspecies

Strix noctua
Diagn. Pale red, with brown spots variegated longitudinally. Iris yellow.
In M.p. (Scopoli’s own collection) from Carniola where it is abundant in the woods
around Ljubljana.
The size of Columba [probably the Stock Dove (Columba oenas) is meant here,
described under the number 177 in Annus I. Historico Naturalis].

Hartert (1913) rated the description as poor and even uncertain, but it should be
given as benefit of the doubt. J. A. Scopoli apparently knew the bird well, because he
had the type in his own collection (Scopoli 1769), which was later destroyed, and also
the longitudinal spots and yellow eyes agree with the Little Owl, although the size of a
pigeon, which probably refers to the Stock Dove (length 28–32 cm), is slightly larger
than the Little Owl (21–23 cm). J. A. Scopoli conducted research mainly between
1754 and 1769 in the former Duchy of Carniola, which existed in the former Holy
Roman Empire and includes the territory of the western half of Slovenia (Vrezec et al.
2017). Scopoli (1769) referred to the city of Ljubljana as the type locality for Strix noctua,
where the species bred until recently (Tomé et al. 2005, Berce 2019).

3.3 Discovery of the Subspecies

3.3.1 Historical Rationale
For a while it was a kind of international competition to distinguish as many subspecies
as possible. Subsequently, of course, extensive discussion arose about how many sub-
species should or should not be distinguished and on the basis of what. Table 3.1 gives
an overview of the subspecies distinguished over the years by various authors.
Dickinson and Remsen (2013) state that many subspecies were distinguished at a
time when even the simplest statistical tests were not yet available. Subspecies were
mainly distinguished using differences in structural sizes or subtle nuances in the
plumage (morphological concept). However, many differences are arbitrary or based
on extremes and for that reason might be artificial. Too small samples, insufficient
comparisons with recognized species and too little control for wear and tear of the
seasonal plumage or discoloration increase the problem (Dickinson and Remsen 2013).
The division into subspecies is still ongoing, e.g., Hart et al. (in Del Hoyo et al. 2016)
note that many races or subspecies should probably be considered intermediate popu-
lations or reflect individual variation with unclear geographic boundaries. For example,
the subspecies vidalii and indigena meet in a wide area in northwestern Russia, noctua
and vidalli ranges border from southern France to the Czech Republic and Slovakia, and
noctua and indigena meet in the Western Balkans.
Many differences were not detected in the field, but were described by skins. Hartert
(1924) describes in a letter how he compared 29 saharae with 15 lilith to establish that they
differed too much from each other to be counted as belonging to the same subspecies.
A first overview of the different subspecies of the Little Owl was given by Hartert
(1913). It was not yet complete, as new subspecies were discovered in subsequent years

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28

Table 3.1 Overview of the subspecies distinguished over the years by various authors
Van
König Van Howard Nieuwenhuyse,
and Nieuwenhuyse and Johnson and Van
Hartert Peters Vaurie Schönn Weick Wink Weick et al. Mikkola Moore Clements IOC Harxen

Subspecies Year Author Country 1913 1940 1960 1991 2006 2008 2008 2008 2012 2013 2015 2016 2023

A. n. noctua 1769 Scopoli Italy 1 1 1 1 1 1 1 1 1 1 1 1 1

A. n. glaux 1809 Savigny France 1 1 1 1 1 1 1 1 1 1 1 1 1
A. n. bactriana 1847 Blyth UK 1 1 1 1 1 1 1 1 1 1 1 1 1
A. n. indigena 1855 Brehm Germany 1 1 1 1 1 1 1 1 1 1 1 1 1
A. n. vidalii 1857 Brehm Germany 1 1 1 1 1 1 1 1 1 1 1 1 1
A. n. spilogastra 1863 Heuglin Germany 1 1 1 1 1 1 1 1 1 1
A. n. plumipes 1870 Swinhoe UK 1 1 1 1 1 1 1 1 1 1 1
A. n. orientalis 1873 Severetzov Russia 1 1 1 1 1 1 1 1 1 1 1 1
A. n. somaliensis 1905 Reichenow Germany 1 1 1 1 1 1 1 1 1 1
A. n. sarda 1907 Kleinschmidt Germany 1 1 1 1 1
A. n. saharae 1909 Kleinschmidt Germany 1 1 1 1 1 1 1 1 1
A. n. lilith 1913 Hartert Germany 1 1 1 1 1 1 1 1 1 1 1
A. n. solitudinis 1924 Hartert Germany 1
A. n. ludlowi 1926 Baker UK 1 1 1 1 1 1 1 1 1 1 1 1
A. n. impasta 1928 Bangs and UK 1 1 1 1 1 1 1
Peters

9 15 13 13 10 12 8 13 7 13 13 13 14
 3.3 Discovery of the Subspecies

and he disregarded spilogastra and somaliensis as non-Palearctic. A first, complete over-

view with 17 subspecies was presented by Peters (1940). Vaurie (1960) retained only 13
subspecies, something that most current lists have followed so far (Van Nieuwenhuyse
et al. 2008, Dickinson and Remsen 2013, Clements et al. 2015, Gill et al. 2021). The
subspecies impasta was discussed by Vaurie (1960) but not retained due to a very limited
sample size of two specimens available from Bangs and Peters (1928) and seven in his
own collection plus a possible bias in coloration due to very fresh plumage that could
have explained the dark coloration rather than being a conclusive differentiation.
However, Vaurie could identify a clear difference in the size of seven newly collected
skins from impasta being smaller in wing length than ludlowi and larger than plumipes.
Furthermore, recent morphological insights (Lei et al. 1997) and genetic analysis of the
Chinese subspecies (Qu et al. 2002) confirm the relevance of A. n. impasta.

3.3.2 Discovery
All subspecies of the Little Owl were described in the nineteenth century and the first
quarter of the twentieth century with noctua (1769) as the exception and impasta (1928)
as the last. Discovering new species was relatively easy at that time since most were still
unknown. People travelled in all directions, i.e., the French to Egypt, the Germans first
into Europe, later to northeast Africa and the Arab countries, the English to India and the
surrounding area, and the Russians to the Caucasus (Walters 2003). They were adven-
turers, looking for the thrill of the unknown. They were scientists, yearning for new
knowledge. Above all, they were pioneers, going where others had not yet gone. They
shot and gathered birds; full shiploads were sent to museums (Mearns and Mearns 1998).
It was not until 1822 that Ferdinand Boie – a German lawyer, entomologist, herpe-
tologist and ornithologist – placed Little Owls in the Athene genus. The discovery of the
different subspecies is presented in chronological order.

1. Athene noctua noctua (Scopoli 1769) Italy to Slovakia, NW Romania and Croatia.
Type locality: Ljubljana (Slovenia)
See species discovery above

2. Athene noctua glaux (Savigny 1809) Coastal areas North Africa, Nile Valley, SW
Israel, Arabian Peninsula.
Type locality: Egypt

The first subspecies described is Athene noctua glaux. In the years 1798 to 1801,
Napoleon Bonaparte made a fierce attempt to conquer Egypt. In addition to increasing
the French sphere of influence in this part of the world, the campaign also served a
scientific purpose. In the wake of the Armée d’Orient, a small army of scientists of
various backgrounds went along, with the aim among other things of describing the
rich and hitherto virtually unknown flora and fauna of the Orient. Among them was
the zoologist Marie Jules César Lelorgne de Savigny. Although initially responsible for
the study of invertebrates, he had a special interest in ornithology. He probably derived
the name glaux from ancient Greek, where glaux (γλαύξ) means owl and is derived from

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

glaucos, which indicates something as sparkling or shiny (in relation to the eyes). The
Greek goddess of wisdom, Pallas Athene, was also referred to as glaukopis: “she with the
beautiful owl eyes” (Schönn et al. 1991). Savigny used the name Noctua glaux.

3. Athene noctua bactriana (Blyth 1847) Lower regions of Urals, Transcarpathia and
Iran to south Kazakhstan, West Pakistan
Type locality: Old Candahar (Afghanistan)

The subspecies bactriana was first described in a periodical edited by Edward Blyth, who
is referred to as the father of Indian ornithology. From 1841 to 1863 he curated the
Asiatic Society of Bengal Museum in Calcutta. In that capacity he was regularly more
concerned with ornithology than with museum matters. He started writing articles, and
trading and shipping wild animals from India to wealthy collectors in England. He was
a contemporary of Charles Darwin and corresponded with him. He provided Darwin
with a lot of information about birds and other animals in India.
Blyth got his information on bactriana from Thomas Hutton, Captain in the 37th
Bengal Native Infantry. Hutton served in the First Soviet–Afghan War (1839–1840)
between England, Russia and Persia. After his retirement he developed into an all-round
naturalist. In 1847 he described the new subspecies bactriana as common among the rocks
and ruins of old Candahar. The article was published in the Journal of the Asiatic Society,
edited by Blyth, who claimed ownership while honoring Hutton in the English name for
bactriana as “Hutton’s Little Owl.” The name bactriana (or bactrian as Hutton originally
calls it) refers to the region of northern Afghanistan around Bactra (now Balch) that was
called Bactria in ancient times and in 328 BC was conquered by Alexander the Great.

4. Athene noctua indigena (Brehm, C.L. 1855) Western Balkan peninsula to southern
Russia, in the south to Greece, Cyprus, Turkey, North Levant and Transcaucasia
Type locality: Attika (Greece)

Christian Ludwig Brehm was a German priest and ornithologist and is considered one of
the greatest bird connoisseurs of his time. In 1824 he was the publisher of Ornis, the
world’s first ornithology magazine. Brehm owned a collection of more than 15 000 skins
that proved to be of great importance for a better understanding of species and subspe-
cies. He called indigena “Der Griechische Steinkauz” and wrote not only that it lived in
Greece, but also that it migrated to Egypt, which was only speculation. The name indigena
comes from Latin and means “out of the land” or indigenous. Brehm here probably
referred to the cradle of the family, the Greek city-state of Athens. On Greek tetra- and
decadrachm coins from 500 BC the Little Owl is already prominently depicted.
Brehm also mentioned A. n. meridionalis (southern Little Owl) as a synonym of
A. n. glaux.

5. Athene noctua vidalii (Brehm A.E. 1857) Western and Central Europe, east to the
Baltic states and Belarus
Type locality: Murcia (Spain)

Alfred Eduard Brehm, son of Christian Ludwig Brehm, is famous for his monumental
work Brehms Tierleben, published in 1863–1869.

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 3.3 Discovery of the Subspecies

Brehm gained much of his knowledge during his travels abroad. He took part in a five-
year expedition to Northeast Africa and travelled to Scandinavia and Siberia. On his journey
to Sinai in the 1850s he was accompanied by Theodor von Heuglin who described the
subspecies spilogastra a few years later. In 1856 Alfred Brehm left with his brother Reinhard
for a two-year trip to Spain. There he saw in rock walls everywhere – what he presumed
was – a new Athene species, which he affectionately called “Vidals Käuzchen.” In the
Allgemeine Deutsche Naturhistorische Zeitung he described it as a new species of Athene vidalii.
Dedicated (nobis), on behalf of father and sons Brehm, to his “most loving friend” Ignatio
Vidali, director of the zoological museum in Valencia. In June of that year they had visited
the university museum in Valencia and there they met Ignacio Vidali y Cros. Together they
went on a few collection excursions. Ignacio was considered an excellent ornithologist and
helped both brothers financially. Alfred returned to Germany to become director of the
Zoologischer Garten in Hamburg and the Berliner Aquarium Unter den Linden.

6. Athene noctua spilogastra (von Heuglin 1863) Northeast Sudan, Eritrea and
Northeast Ethiopia
Type locality: coast of Abyssinia near Massawa (Eritrea)

Theodor von Heuglin was a German explorer and ornithologist. He made several trips
to Africa, in particular to countries along the Red Sea. In 1855 he published his opus
magnum Übersicht der Vögel Nordostafrikas in which he described 754 species. During
one of his travels in 1862 he discovered along the coast of Abyssinia (now Ethiopia/
Eritrea) the “African Höhlenkauz,” which he named Noctua spilogastra (von Heuglin,
1863). He found nests in burrows with a diameter of 10–15 cm in the high, almost
perpendicular walls of water channels. The name spilogastra is composed of the Greek
words for stain (spilos) and gaster (belly) and means (white) spotted belly (Heuglin:
abdomine albidis).
The illustration of the subspecies (Figure 3.6) gives the impression that the artist had
never seen the animal nor had a skin available and had to draw based upon textual
descriptions.

7. Athene noctua plumipes (Swinhoe 1870) Southern Altai Mountains and Mongolia to
Transbaikalia and north (central) and northeast China
Type locality: Shuato in Sichuan Province (China)

Robert Swinhoe – the first Western consul on Formosa, Taiwan – was an avid amateur
ornithologist. His diplomatic status gave him unique access to China and Taiwan at a
time when the door to parts of the empire was still firmly locked to Western researchers.
He published more than 120 articles on the region in which he gave a first description
of 231 (sub)species of birds and 26 mammals. He was a contemporary of Darwin and
admirer of his ideas.
On September 28, 1870, near Shato (Shuato in Sichuan Province), he saw a small owl
basking on the ruins of a brick oven. They could easily reach it. The animal was notable
for its feathered feet: “feet covered with shorter hair-like feathers, just showing scaled of
end of toes.” His proposal to recognize it as the new subspecies plumipes (with feathers
on the foot) was adopted by the professional ornithologists of his day.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

Figure 3.6 Athene noctua spilogastra (von Heuglin 1863).

8. Athene noctua orientalis (Severtzov 1873) Pamir and the Tan Sien Mountains to
Qinghai in western China
Type locality: Turkestan

The orientalis subspecies was first described by the Russian paleontologist and zoologist
Nikolai Alexeyevich Severtzov, son of Alexei Petrovich Severtzov who, like Savigny (ssp.
glaux), went to Egypt under Napoleon’s command. Nikolai Alexeyevich also travelled a
lot. While trekking through the steppes of Kyrgyzstan, he was seriously injured in an
attack by Turkestan bandits and held prisoner by them for a month. He collected a lot
of material on the taxonomy of birds in Russia and the former Turkestan. He also had a
large collection of no fewer than 12 000 specimens.
He published his findings in Messages of the Imperial Society of Lovers of Natural
Sciences, Anthropology and Ethnography in Russian.

9. Athene noctua somaliensis (Reichenow 1904) Somaliland.

Type locality: northern Somaliland

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 3.3 Discovery of the Subspecies

Anton Reichenow was a German ornithologist specializing in African birds. He was

married to the daughter of Jean Cabanis, founder of the Journal für Ornithologie in 1853,
one of the oldest ornithological journals. In 1894, Reichenow succeeded his father-in-
law as editor, a position he would remain in for more than 25 years. He undertook
expeditions to West Africa in 1872 and 1873. From 1874 to 1921 he was associated
with the Humboldt Museum in Berlin. He recorded his findings in Die Vögel Afrikas,
which appeared in three volumes in the period 1901–1905.
The East African subspecies somaliensis comes off very scantily in its first description
in Die Vögel Afrikas. In part 1, in which the owls are discussed, of the Little Owls only
the subspecies spilogaster, previously described by Heuglin, is mentioned. Only in the
addendum to the third part does Reichenow mention somaliensis incidentally as a
subspecies of spilogaster which was then regarded as an independent species
(Reichenow 1904). The author assumed certain statements, indicating that no real hard
evidence had been collected, nor did he indicate the name of the observer. Somaliensis
obviously refers to Somalia.

10. Athene noctua sarda (Kleinschmidt 1907)

Type locality: Sardinia (Italy)

Konrad Ernst Adolf Otto Kleinschmidt for decades combined the profession of an
evangelical clergyman with that of an ornithologist. He had great significance for the
animal system through the introduction of the term “Artenkreis.” Kleinschmidt recog-
nized that there are species that, while morphologically different, can mate and repro-
duce successfully with each other and must be classified as one and the same species.
The renowned German–American evolutionary biologist Ernst Mayr built on his ideas
with the name superspecies, nowadays polytypical species, a species that consists of
several subspecies and of which the Little Owl could be considered an excellent
example.
In Falco 3 (1907) Kleinschmidt describes the geographic variation of 96 Little Owls
(which he still classifies in the genus Strix) on the basis of skins from his own and
Erlanger’s collections, supplemented with some from a few other sources. Kleinschmidt
mentioned for the first time the subspecies sarda occurring in Sardinia and distin-
guished by the very small spots on the shoulder feathers that are divided in two by a
dark shaft stripe along the length of the feather. Currently the taxon sarda is not
accepted as a valid subspecies (Gill et al. 2021) although there is strong genetic support
for it (Pellegrino et al. 2015).

11. Athene noctua saharae (Kleinschmidt 1909) Sahara, east to the Red Sea (excluding
the Nile Valley), and Arabian Peninsula
Type locality: Mouleina by Biskra (Algeria)

Although Kleinschmidt already included saharae in his overview in 1907, it was removed
again in the correction, partly because its description was based on only two skins from
the collection of Professor Flückiger and the evidence was therefore very thin. Later he
received a new series of skins from Flückiger who regularly visited the north of Africa on
collection trips. The skins came from the Biskra region in Morocco, and to the south of it.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

On that basis, he decided that this saharae deserved the status of a separate subspecies.
The name saharae undoubtedly refers to the geographic distribution area.

12. Athene noctua lilith (Hartert 1913) Sinai, north of the Arabian Peninsula, Jordan,
Central Syria to southeast Turkey and Iraq
Type locality: Deir ez-Zor by Euphrat (Syria)

Athene noctua lilith was first described by the German ornithologist Ernst Hartert. For
almost 40 years Hartert was curator of the Zoological Museum in Tring. The museum
belonged to Lord Lionel Walter Rothschild, of the famous and wealthy Rothschild
banking family. With the help of the family capital he was able to realize his childhood
dream on the family estate and set up his own museum. The museum contains the
largest collection of stuffed animals in the world. Ernst Hartert will have regularly sat
next to him, discussing the content of the museum magazine Novitates Zoologicae that
they published together for 35 years.
In the years 1910–1922 he wrote the three volumes of Die Vögel der Paläarktischen
Fauna: Systematisch Übersicht der in Europa, Nord-Asien und der Mittelmeerregion forkom-
menden Vögel, including the Little Owl. He distinguished nine subspecies, including
lilith as a new subspecies (novum). Unfortunately, he did not explain the reason why he
chose the name lilith.
Given the region in which the subspecies occurs, however, we may assume that it
refers to the female figure Lilith who appears in both (pre-) Babylonian and Hebrew
mythology and writings. In the former, Lilith was a storm or night demon. The name
would mean “night” in proto-Semitic (Carvalho 2009). She was the wandering spirit of
those who died prematurely who had not yet been able to fulfill their destiny in life
(Broekema 2014). Lilith was part of a group of four like-minded people who haunted at
night and took pleasure in assaulting sleeping men. However, in some Hebrew sources,
she was Adam’s first wife. Because she refused to submit to him, she left him. She is
depicted, together with two lions and two owls, in the famous Burney relief, which is
dated between 1800 and 1750 BC (Broekema 2014).

13. Athene noctua ludlowi (Baker 1926) south and east of the Tibetan Plateau (Ladakh,
Nepal, Western Sichuan)
Type locality: Rhamtso Lake in Tibet (China)

Edward Charles Stuart Baker was an English police officer who spent most of his career
in India. In his spare time he studied and collected birds. His estate also contained some
50 000 eggs from Indian birds. His most important contributions to ornithology were
the eight volumes of The Fauna of British India, including Ceylon and Burma. Birds. The
1926 Bulletin of the British Ornithologists Club mentions his submission of the descrip-
tions of subspecies of four oriental owls, including a Little Owl.
The subspecies ludlowi is named after Frank Ludlow, an English officer who was
stationed as a naturalist in India and for a time in the Tibetan capital Lhasa. At the time,
he collected about 7000 birds, all of which are now kept in the British Museum,
including a specimen of the newly (novum) described subspecies ludlowi, 10-12-1923
shot at Rhamtso Lake in Tibet. Baker notes that although the subspecies closely resembles

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 3.3 Discovery of the Subspecies

plumipes (which also occurs in China), it is distinctively larger (wing 160–165 mm for
plumipes and 169–173 mm for ludlowi).

14. Athene noctua impasta (Bangs and Peters 1928) northeast Qinghai,
southwest Gansu.
Type locality: Lake Kokonor (China)

Outram Bangs was an American zoologist. Together with James Lee Peters, curator of
Harvard University’s Museum of Comparative Zoology and president of The American
Ornithologists’ Union (Wetmore 1957), he described in the museum bulletin the
collection of John F. Rock during his journey westward to Kansu (China) and eastern
Tibet. Peter’s most important ornithological achievement was the publication of the
Check-list of Birds of the World, better known as Peters’s check-list. In part 4, among
other things, owls are discussed and the Little Owl with 17 subspecies is described.
Born in Vienna, later naturalized as an American, Joseph F. Rock was a widely qualified
scientist. He was proficient in anthropology, botany and linguistics. He also practiced the
crafts of filmmaker, photographer, explorer and author. At 16 it seems he already spoke
Arabic and Mandarin. From 1924 to 1927, he led an expedition through western China
and Tibet on behalf of Harvard University to collect plants and animals for the Arnold
Arboretum and Museum of Comparative Zoology. During that voyage – in September
1925 – he caught an adult female Little Owl at Lake Kokonor (the Mongolian name for
Lake Qinghai in today’s Qinghai province), at an altitude of 3261 m. It looked a bit like
plumipes, but deviated from it because of its dark color. He collected two more, a female and
a doubtful case that he considered somewhat intermediate between plumipes and impasta
(Bangs and Peters 1928). While Vaurie (1960) had some doubts on the representativeness
of the small sample to describe the subspecies, some more evidence for its relevance has
been obtained recently. Lei et al. (1997) studied the distribution, habitat and clinal
variations of the subspecies of the Little Owl in China, confirming impasta as being
distinctively intermediate between ludlowi and plumipes in body coloration, as well as in
body size and claw length. The morphological arguments for considering it as a subspecies
were confirmed genetically by Qu et al. (2002). The results confirmed A. n. impasta and
A. n. plumipes as two distinctive subspecies rather than synonymous.
The name impasta comes from the Latin impastus, which means not fed, hungry
(Jobling in Del Hoyo 2016).

3.3.3 Original Descriptions of Subspecies

The original descriptions of the 14 subspecies are presented in their original language
and translated into English. The description by Hartert (1913) can be considered as the
first formal morphological description of the nominate form noctua.

[ATHENE NOCTUA NOCTUA] SCOPOLI (1769) [LATIN]

15. STRIX NOCTUA.
DIAGN. Pallide rufa, fuscisque maculis longitudinaliter variegata.
Irides flavae. In M. p. Carnoliae indigena, et in sylvis circa Labacum copiosa.
Statura Colombae.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

17. STRIX PASSERINA.

Strix capite laevi, remigibus albis : maculis quinque ordinum, Linn. l.c.n.12 Kram.
L.c.n. 6.
Ital. Civetta
Germ. Tschiavitl, Hauseule, Stockeule, Käuzlein.
DIAGN. Inservit aucupio inprimis Sylviarum.
Nidificat in caminis.

[ATHENE NOCTUA NOCTUA] SCOPOLI (1769) [ENGLISH]

[15. Pale red spots, facial disk variegated lengthwise. Iris yellow. Native to M. p.
Carnolia and woods around Ljubljana. Pigeon-like stature.
17. Used for bird catching, particularly passerines? Breeds in chimneys.]

[ATHENE NOCTUA NOCTUA] HARTERT (1913) [GERMAN]

♂ ♀ ad. Oberseite erdbraun, wie Sepia oder Ridgways ‘bistre’, Oberkopf mit längeren
oder mehr rundlicheren, kürzeren Schaftflecken, die die Spitze nicht erreichen und
weiß mit einem rahmfarbenen Anfluge. Federn an Nacken und Hinterhals rahmweiß
mit grauen Wurzeln und breiten, erdigbraunen Rändern, so dass ein deutliches, breites,
weißes Flankenband entsteht angelegt, auf der übrigen Oberseite sind die weißlichen
Flares kleiner und oft doppelt, mehr oder minder durch die breiten Säume verdeckt,
aber an den Skapularen und größeren Flügeldecken aber treten sie bedeutend mehr
hervor; sie sind dort groß und die Flecke der beiden Fahnen bilden zusammen einen
großen Fleck. Schwingen tiefbraun, Außenfahnen mit rahmfarbenen Flecken,
Innenfahnen mit weißlichen, den Schaft nicht erreichenden Querbinden, außerdem
ein an den Armschwingen deutlicherer, beziehungsweise breiterer, weißlichen
Spitzensäum. Steuerfedern wie der Rücken, mit schmalem Endsaum und 4, selten 5 hell
rahmfarbenen Querbinden, die mitunter am Schaft nicht gansz susammenhëngend
sind. Kinn und Vorderhals weiß, durch ein braunes Fleckenban voneinander getrennt;
übrige Unterseite weiß mit rahmfarbenem Anflug und dunkel- oder erdbraunen
Flecken, meist in Gestalt von in der Mitte etwas schmaleren Längsflecken, an den
Brustseiten rundlicher werdend. Analgegend und Unterschwanzdecken ungefleckt,
letztere oft mit braunen Schäften oder ebensolchen aber nur sehr schmalen Streifen.
Laufbefiederung rahmweiß, Unterflügeldecken ebenso, aber mit einigen braunen
Flecken. Iris gelb. Schnabel wachsgelb, an der Wurzel grünlich.

[ATHENE NOCTUA NOCTUA] HARTERT (1913) [ENGLISH]

[Upper side earth brown, like sepia or Ridgways “bistre,” upper head with longer or more
rounded, shorter shaft spots that do not reach the tip and are white with a cream-colored
tint. Feathers on the neck and back of the neck creamy white, with gray roots and broad,
earthy brown edges, so that a distinct, wide, white flank band is created. On the rest of the
upper side the whitish bands are smaller and often double, covering the broad edges more
or less thickly, but they stand out much more on the scapula and larger elytra; they are big
there and the spots of the two bands together form a big spot. Wings deep brown, external
flags with cream-colored flecks, inner flags with whitish cross-bands that do not reach the
shaft, and a more distinct or brighter, whitish lace border on the wings. Back feathers with

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 3.3 Discovery of the Subspecies

a narrow bottom edge and four, rarely five light cream-colored transverse bands, which are
sometimes not completely connected on the shaft. The chin and front neck are white,
separated from one another by a brown band, the rest of the underside is white with cream-
colored patches and dark or earth-brown patches, mostly in the form of slightly pale
longitudinal patches in the middle, becoming more rounded on the chest. Anal area and
under the tail unspotted, the latter often with brown shafts or similar but only very narrow
stripes. Fletching creamy white, underwing coverts as well, but with a few brown spots. Iris
yellow. Bill wax yellow, greenish at the root.]

[ATHENE NOCTUA BACTRIANA] BLYTH (1847) [ENGLISH]

Athene bactrianus, mihi, n.s. ? [Strix persica (?) Nouv. Dict. d’Hist. Nat., VII, 26.]
Length about 9 inches; of wing 6.25 inches, and tail 3.5 inches: tarse 1.25 inch.
Plumage of the upper parts somewhat rufescent clay-brown, with large white spots on
the features, more or less concealed, and wholly so on those of the middle of the back:
coronal feathers with whitish streaks: face white; some of the radiating feathers on the
sides of the beak terminating in black vibrissae: chin, throat, lower tail-coverts, and the
tibial and tarsal plumes, white also the fore-part of the under-surface of the wing: a
longitudinal broad streak on each feather of the breast and abdomen: on the hindneck,
the white so predominates upon the feathers as to give the appearance of a half-collar:
the great wing-feathers have broad incomplete pale bands, disposed alternately on their
two webs; and the middle tail feathers have a double row of semi-alternating pale spots,
passing into dull bands on the outer tail feathers: beak (in the dry specimen) whitish;
and claws pale horn-color. Common upon the rocks and ruins of old Candahar.

[ATHENE NOCTUA GLAUX] SAVIGNY (1809) [FRENCH]

ORDRE 1er, FAMILLE 3, LES CHOUETTES, 287

Jambes à plumes courtes. Les plumes de tout le corps molles et douces au toucher.

Genre XIII. NOCTUA.

Caractères principaux.
Bec épais, très court, brusquement incliné, peu comprimé, convexe dessous; cire très-
renflée sur les narines et comme gibbeuse de chaque côté; narines écartées, très
petites, parfaitement rondes, tournées en devant; mandibule inférieure ayant
échancrures marginales vers le bout.
Langue ovale, épaisse, pourvue de deux côtes en dessous, très obtuse.
Tarses laineux de toutes parts.
Doigts velus jusqu’à la base des dernières phalanges.
Ongle intermédiaire sans crénelures.

Caractères accessoires.
Ailes peux pointues, dépassée par la queue.
Quatre à cinq rémiges légèrement échancrées; la première assez courte, la troisième la
plus longue.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

Queue égale.
Cercles périopthalmiques médiocres et peu réguliers.
Oreilles externes petites, rondes et dénuées d’opercules.
Tête sans aigrettes.

Espèce
22. NOCTUA GLAUX, LA CHEVÊCHE. (v)
NOCTUA iridibus pallidé flavis; abdomine maculato; gula alba

[ATHENE NOCTUA GLAUX] SAVIGNY (1809) [ENGLISH]

[Main Characters

Beak thick, very short, sharply inclined, slightly compressed, convex underneath; wax
very swollen on the nostrils and as a gibbous on each side; nostrils wide apart, very
small, perfectly round, turned forward; lower mandible with marginal notches
towards the end. Tongue oval, thick, with two ribs underneath, very obtuse. Woolly
tarsi on all sides.
Toes feathered to the base of the last knuckles.
Intermediate nail without crenellations.

Accessory Characters
Wings little pointed, exceeded by the tail. Four to five slightly indented flight feathers;
the first rather short, the third the longest. Equal tail.
Poor and irregular periopthalmic circles. Outer ears small, round and without
operculum. Head without crags.
NOCTUA Iris pale yellow; spotted belly; white throat]

[ATHENE NOCTUA IMPASTA] BANGS AND PETERS (1928) [ENGLISH]

ATHENE NOCTUA IMPASTA subsp. Nov.

Type. – Adult female, No. 239416Mus. Comp. zool.; grass country south of Lake
Kokonor, 10 700 feet; collected September 1925 by Joseph F. Rock (orig. no. 408).
Characters. – Similar to A. n. plumipes (Athene plumipes Swinh., P.Z.S. 1870, p.448 Shato,
north China) in having feathered toes, but much darker; pale markings on the top of
the head linear rather than guttate; light tail bands interrupted; markings
underneath darker and more extensive.

Number Sex Wing Tail Tarsus

239416 ♀ (type) 167 108 38

239417 ♀ 168 102 34
239418 ♀ (♂?) 157 91 34

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 3.3 Discovery of the Subspecies

Athene noctua plumipes From Shansi

87567 ♀ 166 92 34.5

87568 ♀ (♂?) 155 85 35

Of the specimens of our new form listed, in addition to the type, one was secured on
the grasslands at the heat of the Serchen valley (13 300 feet) three days east of Radja,
26 July 1926 and the third was taken in May 1925 in the forests covering the slopes of
the mountains of Choni (9000 feet). The latter specimen as might be suspected on
geographic grounds, is somewhat intermediate between impasta and plumipes. It
approaches the latter in having the light tail spots almost forming bands, but tends
toward the former in the extent of the streaking underneath and in having linear pale
crowns-stripes – on the whole it is best placed with impasta.
We have not seen Athene noctua ludlowi Stuart Baker of southern Tibet, but that form is
said to be about the color of A. n. plumipes. Our new form is much darker, in fact little paler
than A. noctua noctua of Europe. Our bird also comes from a region that is quite different
faunally, and although we hesitated at first, we now see no course open but to name it.

[ATHENE NOCTUA INDIGENA] BREHM C.L. (1855) [GERMAN]

Fang der Kaüze

3) Der Griechische Steinkauz. Athene indigena, Brm.

Er steht in der Zeichnung swischen Nr. 2 und 4 mitten inne, hat auf dem Ünterkörper
sehr verwachsene Flecken, lebt in Griechenland und wandert nach Aegypten.

4) Der Südlichen Steinkauz. Athene meridionalis, Brm. (Strix meridionalis, auct.

Noctua nilotica, Paul de Wrttbg.)

Er ähnelt der vorhergehenden; allein sein Oberkörpfer und die Flecken des
Unterkörpfers ziehen sharf ins Rostbraune. Er lebt in Aegypten und Südeuropa.

[ATHENE NOCTUA INDIGENA] BREHM C.L. (1855) [ENGLISH]

[3) The Greek little owl. Athene indigena, Brm.

It is in the middle of the drawing between nos. 2 and 4, has very overlarge spots on the
lower body, lives in Greece and migrates to Egypt.

4) The southern little owl. Athene meridionalis, Brm. (Strix meridionalis, auct. Noctua
nilotica, Paul de Wrttbg.)

It resembles the previous one, but his upper body and the spots on the lower body turn
sharply rusty brown. He lives in Egypt and southern Europe.]

[ATHENE NOCTUA LILITH] HARTERT (1913) [GERMAN]

1461. Athene (oder Carine) noctua lilith subsp. Nov.
Noch bleicher, mehr sandfarben, als A. n. bactriana, auch die Flecke der Unterseite
blasser und mehr röthlichgelblich, ausserdem die Zehen nur mit Borsten bedeckt, nicht

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

dicht befiedert wie bei A. n. bactriana. Flügel von 12 Exemplaren 154-168 mm. – Typus:
♀ Der-ez-Zor amd Euphrat, 15. V. 1911, gesammelt von J.Aharoni, im Tring-Museum.
Palästina und Mesopotamien (oberer Euphrat), und das südwestliche Persien.
Untersucht wurden exemplare in den Museen von Tring und Berlin und aus
Witherbys Sammlung.

[ATHENE NOCTUA LILITH] HARTERT (1913) [ENGLISH]

[Even paler, more sand-colored than A. n. bactriana, the spots on the underside are also
paler and more reddish-yellow, moreover the toes are only covered with bristles, not
densely feathered as in A. n. bactriana. Wings of 12 specimens 154–168 mm. - Type: ♀
Der-ez-Zor amd Euphrates, 15th V. 1911, collected by J. Aharoni, in the Tring Museum.
Palestine and Mesopotamia (Upper Euphrates), and southwestern Persia. Examples
were examined in the museums of Tring and Berlin and from Witherby’s collection.]

[ATHENE NOCTUA LUDLOWI] BAKER (1926) [ENGLISH]

In general color intermediate between A. n. noctua and A. n. bactriana – in fact, very
similar to A. n. plumipes from Sheni, China, but decidedly bigger than that bird, which
has a wing between 160 and 165 mm. The amount of feathering on the toes varies
considerably, both individually and seasonally, but in winter most birds have the
plumelets extending down the toes almost to the base of the claws.

Colors of soft parts. Iris yellow; bill bright yellow; legs gray, soles yellow (F. M. Bailey).
Measurements: 4 ♂, 2 ♀. Wing 169 to 173 mm; tail 88 to 96 mm; tarsus 31 to 32 mm;
culmen 18 to 20 mm.
Distribution: Tibet. A bird from the Mishmi Hills is nearest to the present race, but is
smaller (wing 164 mm) and rather darker.
Type in British Museaum. ♂. Dochen, Rhamtso Lake, Tibet, alt. 15 000 ft, 10.12.23.
Collected by F. Ludlow, No. 40. Brit. Mus. Reg. No. 1926.11.11.1.

[ATHENE NOCTUA ORIENTALIS] SEVERETZOV (1873) [RUSSIAN]

38. Athene orientalis от Европейской A. noctua отличается тем, что бурый цвет последней
A.orientalis гораздо бледнее, переходит в серо-рыжеватый с желтоватым оттенком как
A. noctua meridionalis, от которой A. orientalis отличается густым оперением пальцев,
кроме ногтевого сустава. Пальцы оперены также густо, как у A. (Nyctale) Tengmalmi, но
кроме своего темного цвета, A. Tengmalmi отличается ушными отверстиями,
полулунными и гирокими, во всю вышину черепа, между тем как у A.orientalis ушное
отверстие овально и гораздо меньше, в пол-черепа, как у A.noctua. По Киргизски
A.orientalis ай-хай, по крику: тоэе бай-кыль (богатая девица).
По соединению этих признаков, светлого цвета с пушистыми пальцами, A. orientalis я
считаю не видоизменением, а особым видом.

[ATHENE NOCTUA ORIENTALIS] SEVERETZOV (1873) [ENGLISH]

[38. Athene orientalis is distinct from the European A. noctua by the fact that the brown
color in the latter is darker. A. Tengmalmi and A. orientalis are more pale, transforming

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 3.3 Discovery of the Subspecies

into gray-russet with a yellowish tint as A. noctua meridionalis, from which A. orientalis is
distinguished by the dense feathering of the toes, excluding the nail joint. The toes are
feathered more densely, like A. (Nyctale) Tengmalmi, apart from the dark color. A.
Tengmalmi differs by the ear hole, semi-moon and wide, the entire height (upper part)
of the skull, whereas in A. orientalis the ear hole is oval and much smaller, half of the
skull, like A. noctua. In the Kyrghyz language A. orientalis is ai-khai, from its call; also
bai-kyz (rich young lady).
By combining these observations, the light color with feathered toes, I consider A.
orientalis not as a species variation, but as a separate species.]

[ATHENE NOCTUA PLUMIPES] SWINHOE (1870) [ENGLISH]

A. bactriana (Blyth) of Thibet, by its short tarse covered with long down, and by its well-
clothed toes. I propose to distinguish it as the
ATHENE PLUMIPES, sp. Nov.
Throat white, the white extending in crescent-form up each cheek in rear of ear-
coverts (the lower white neck-ring of A. noctua is wanting); lores, round eye, and middle
of belly also pure white; upper parts light reddish brown, with drops of reddish white
on the head, and marked and spotted much as in A. noctua; underparts, leg- and feet-
feathers cream-color, on the breast and flanks broadly streaked with reddish brown, like
A. glaux (A. noctua being spotted with white on a dark ground and wanting the white on
the center of the belly); bill yellow tinged with green; irides light yellow; claws
blackish brown.
Length about 8.5 inches; wing 6.2, of similar-proportioned quills to those of A.
noctua; tail 3.6, of 12 equal feathers; tarse to base of hind toe 0.85, densely clothed
with down-like feathers, 0.65 long; feet covered with shorter hair-like feathers, just
showing scales at the end of toes; soles bare and yellow.

[ATHENE NOCTUA SARDA] KLEINSCHMIDT (1907) [GERMAN]

9. Sardinien. Strix Athene sarda form. Nov.
3 Stück augenblicklich in meinen Händen. Ich have aber mehr gesehen. Dem
mitteldeutschen Vogel ähnlich, im Herbst bisweilen dunker, im Sommer bisweilen
lichter als Mitteldeutschland und Rhein. Unterscheidet sich durch die engere
Zeichnung. Der Schwanze eng und schmal weisslicht gebändert. Am deutlichsten wird
der Unterschied an den Schulterfedern. Die weissen Querflacken sind hier sehr klein
und werden von einem dunkeln Schaftstrich längs der Federmitte geteilt. Das sardi-
nische Käuzchen bildet eine hübsche Parallel zu Falco Peregrinus brookei.

[ATHENE NOCTUA SARDA] KLEINSCHMIDT (1907) [ENGLISH]

[Three pieces currently in my hands. But I’ve seen more. Similar to the Central German
bird, sometimes darker in autumn, sometimes lighter in summer than Central Germany
and the Rhine. Differs in the narrower markings. The tail is narrowly banded with
white. The difference is most evident on the shoulder feathers. The white transverse
bands are very small here and are divided by a dark line along the middle of the feather.
The Sardinian owl forms a nice parallel to Falco Peregrinus brookei.]

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

[ATHENE NOCTUA SAHARAE] KLEINSCHMIDT (1909) [GERMAN]

2. Strix saharae
Formenkreis Strix Athene. Terra typica Algerische Sahara. In Falco 1907 p. 66 No. 15 war
obiger Name schon gedruckt. Er wurde bei der Korrekty wieder gestrichen, weil dat
Vorhandensein von nur zwei Stücken einerseits und die rote Abbildung der Strix
numida (Lavaillant) die Benennung bedenklich machten. Nun hat mir aber Herr
Flückiger noch eine ganze Serie zugesandt, die er auf seiner ersten Reise bei Biskra und
weiter südlich gesammelt hat. Das Flügelmaximum erhöht sich auf 16,3 cm. Sonst
bleibt es bei der an der citierten Stelle gegebenen Charakteristik. Typus von Moulaina.
Verbreitung von Biskra südlich (vielleicht bis ins südliche Tunesien). Strix numida ist
eine rote Phase der Mittle-Algerische Form, wenn man eine genaue Deutung dieser
Abbildung überhaupt wagen will.

[ATHENE NOCTUA SAHARAE] KLEINSCHMIDT (1909) [ENGLISH]

[Form circle Strix Athene. Terra typica Algerian Sahara. In Falco 1907 p. 66 No. 15 the
above name was already printed. It was deleted from the Korrekty because of the
presence of only two pieces on the one hand and the red image of the Strix numida
(Lavaillant) made the name questionable. But now Mr. Flückiger has sent me a whole
series that he collected on his first trip to Biskra and further south. The wing maximum
increases to 16.3 cm. Otherwise the characteristics given in the quoted passage remain.
Type of Moulaina. Distribution of Biskra south (perhaps as far as southern Tunisia). Strix
numida is a red morph of the Middle Algerian form, if one dares to interpret this figure
at all.]

[ATHENE NOCTUA SOMALIENSIS] REICHNOW (1904) [GERMAN]

Bd. I Seite 671:
553a. Athene spilogaster somaliensis Rchw.
Athene noctua spilogaster Erl. J. O. 1904, 238.
Im Somalilande scheint eine von der abessinischen A. Spilogaster abweichende Form
vorzukommen: Flügel und Schwanz kürzer; Braun der Oberseite dunkler; Oberkoph
einfarbig oder weis gefleckt, aber nicht schmal gesrichelt; braune Strichelng der
Unteseite dunkler. Lg. etwa 160-180, Fl. 130-140, Schw. 60-70, Schn. v. d. Wachsh.
12-13, L. 26-28 mm.

[ATHENE NOCTUA SOMALIENSIS] REICHNOW (1904) [ENGLISH]

[In Somaliland there seems to be a form deviating from the Abyssinian A. Spilogaster:
wings and tail shorter; top darker brown; top of head monochrome or spotted white,
but not narrowly dashed; brown dashes on the underside darker.]

[ATHENE NOCTUA SPILOGASTRA] VON HEUGLIN (1869) [LATIN]

Nr. 84. Noctua spilogastra. (Tab. IV.)
Athene spilogastra, Heugl. In Cab. Journ. 1863. P. 15. – A. troglodytica, Hartl. M.S.
♀ Supra pallide et dilute rufescente-fumosa, ex albido guttatim et fasciatim varia,
colore albido fulvescente-lavato; rectricibus albidis, fasciis 6-7. Rufescente-fumosis;
facie, pectore postico et abdomine albidis, vix fulvescente-lavatis et striolis obsoletes

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 3.3 Discovery of the Subspecies

rarioribus subrufescentibus; pectore superior eodem colore vario; tarsis antice plumosis
albis; pilis digitos supra obtegentibus, setoris, albidis; subalaribus omnino albidis; rostro
flavido; ceromate et unquibus brevissimis nigricante-corneis; iride flava; orbitis nudis
nigricantibus; digitis incarnate-plumbeis; long. tot. 7”. 6’’’. – rostr. a fr. 7.5-8’’’. – al. 5”-
5”.1’’’. – caud. 2”.9’’’-3”. tars. 1”-1”.2.5’’’. – dig. med. c. u. vix 12’’’. – halluces
c. u. 5.75’’’. – caudae apice alas superante 4-5’’’.

[ATHENE NOCTUA SPILOGASTRA] VON HEUGLIN (1869) [ENGLISH]

[♀ Above pale and faintly rufescent-smoky, variable whitish spots and streaks of a
yellowish-washed white color; whitish tail feathers with six to seven bands. Rufous-
ashy; the face, the chest, and the abdomen are whitish, scarcely washed with a yellow-
ish shade, with worn-out and faint subrufous streaks; the upper part of the breast is of
the same variable color; the tarsi are white feathered in front; the toes have bristly,
whitish hairs above; feathers under the wing are completely white; beak yellow; cere
with few short blackish bristles; iris yellow; orbital ring blackish; incarnate-lead fingers;
Measurements: total length 7”. 6’’’. – beak to front 7.5-8’’’. - wing 5’’’-5”.1’’’. – tail 2’’.9’’’-
3’’. tarsi 1’’-1’’.2.5’’’. – middle toe <12’’’ – thumb (first toe or toes in general)
c. u. 5.75’’’. – the tip of the tail exceeding the wing tips 4-5’’’.]

[ATHENE NOCTUA SPILOGASTRA] VON HEUGLIN (1869) [GERMAN]

Die vierte Schwinge die längste, die dritte kaum kürzer, die erste ungefähr gleich der
siebenten. – Die erste Schwinge auf der aussenfahne deutlich gesägt, auch auf der
zweiten eine Andeutung hievon.
In Bezug aud Flügelschnitt steht diest gute neue Art der Athene perlatat am
nächsten, nur erscheint die Flügelsitze unbedeutend länger. Die Zehenrücken sind sehr
schwach mit borstenähnlichen Federchen besetzt; die Tarsen erscheinen auf der
Hinterseite nackt, doch ist es nicht unmöglich, dass dies Folge von Nachlässigheit des
Ausstopfens ist. Auffallend kurz und stumpf sind die Krallen dieser Eule.
Wir fanden den afrikanischen Höhlenkauz nur an einer einzigen Lokalität, nämlich
im abessinischen Küstenland unfern Umkulu und Harkiko und zwar im Monat Juli. Ob
derselbe Standvogel sei, kann ich nicht angeben. Er lebt meist paarweise längs der tief in
Geröllmassen eingerissenen Regenbetten, in Steinhaufen un auf Stapelien-Gebüsch.
Wie Athene meridionalis udn A. Perlata fliest der Vogel auch bei Tag aus und nährt
sich von Mäusen, kleinen Vögeln, Eidechsen, Käfern und Heuschrecken. Verfolgt ver-
steckt er sich in Erdlöcher und unter Geröllmassen und er dürfte ein wirklicher
Höhlenbewohner sein, worauf auch die kurzen Krallen hindeuten; an hohen, fast
senkrechten Wandungen der Wasserrinnen trafen wir öfter horizontale Gänge von
4-6 Zoll Durchmesser, ohne Zweifel die Nesthöhlen dieser Eule, da wir sie hier ab-
und zufliegen sahen.

[ATHENE NOCTUA SPILOGASTRA] VON HEUGLIN (1869) [ENGLISH]

[The fourth primary is the longest, the third barely shorter, the first about the same as
the seventh. The first primary clearly sawn on the outer vane, also a hint of this on
the second.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

With regard to the wing cut, this good new species is closest to Athene perlatat, only
the wing seats appear slightly longer. The backs of the toes are very sparsely covered
with bristle-like feathers; the tarsi appear bare on the back, but it is not impossible that
this is the result of careless stuffing. The claws of this owl are noticeably short
and blunt.
We only found the African cave owl in one small location, namely in the Abyssinian
coastal region near Umkulu and Harkiko, in the month of July. I cannot say whether it
is the same resident bird. It usually lives in pairs along rain channels that are cut deeply
into rubble, in piles of stones and on stapelia bushes. Like Athene meridionalis and
A. Perlata, the bird also flies during the day and feeds on mice, small birds, lizards,
beetles and grasshoppers. If chased, it hides in holes in the ground and under debris and
might actually be a cave dweller, which is also indicated by the short claws; on the high,
almost vertical walls of the gullies, we often met horizontal tunnels 4-6 inches in
diameter, no doubt the nest holes of this owl, since we saw it flying to and fro here.]

[ATHENE NOCTUA VIDALII] BREHM A.E. (1857) [LATIN/GERMAN]

Zweite Zunft. Nachtraubvögel. Raptatores nocturne.

I. Familie. Eulen. Striginae.

Erste Sippe. Käutzchen. Athene, Boje.
30. Ath. Vidalii, nobis *). Vidals Käutzchen.
Spanish : Mochuelo. Galizianisch: Mucho.

Diagn. Nostrae Athene noctuae similis, sed paulo minor, obscurior in corpore super-
iori, maculis et striis nigris distinctionribus angustioribus et longioribus.
Die genaue Beschreibung wird folgen. Paarweis oft häufig in ganz Spanien in
Felsenwänden der Gebirge. Standvogel.
*) Ignatio Vidali, Professori academico ordinario, Musei zoologici Valenciae
Directori, viro doctissimo et amico maxime amabili hanc avem, sapientii symbolum
dedicant pater et filii Brehm.

[ATHENE NOCTUA VIDALII] BREHM A.E. (1857) [ENGLISH]

[Second Guild. Nocturnal Raptors

I. Family. Owls. Striginae.

First Kin. Owlet. Athene, Boje.
30. Ath. Vidalii, nobis *). Vidals Owlet.
Spanish : Mochuelo. Galician: Mucho.

Our Athene-like owl is a bit smaller, darker in the body, to the higher, narrower and
longer than spots and streaks with black distinction.
The detailed description will follow. In pairs often common throughout Spain in
rock faces of the mountains. Standing bird.]
[*) Ignatius Vidali, professor of academic ordinary Museum Zoologicum Valenciae
Director, learned friend and most loving way in which this bird is a symbol of wisdom
and a devoted father and son Brehm.]

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 3.4 Morphology and Coloration

3.4 Morphology and Coloration

3.4.1 Geographic Variation in Coloration
The Little Owl Athene noctua is a species with a wide range in the Palearctic. Several
subspecies have been described based on coloration or size differences. A reference work
by Charles Vaurie (Vaurie 1960) included a taxonomic revision where he accepted the
presence of 13 subspecies which we complemented by A. n. impasta which has recently
been confirmed with larger samples for morphological comparison (Lei et al. 1997) and
through genetic research (Qu et al. 2002).
The geographical variation in Little Owl is conspicuous and involves size, coloration
and vocalizations, but also varies individually and tends also to vary locally in some
regions. The individual and local variations are more evident in the southern part of the
range, and the local variations may represent adaptations to climatic or other factors
that are of critical importance in a sedentary and diurnal species which inhabits deserts
or very arid open regions. However, in the equally arid and open, but colder, parts of
Asia the birds are much more consistant in coloration. Possibly they may be forced to
wander more throughout the year because of the more severe climate and, as a result,
are less closely adapted to local conditions. The plumage of A. noctua also bleaches more
quickly in arid and sunny regions. The individual and local variations, or those due to
wear and bleaching, are not unusual, but should be taken into account as they are
possible sources of confusion in the case of the southern subspecies, glaux, saharae, lilith
and bactriana.
Vaurie reached his conclusion based on a comparison of about 200 specimens of
these four races. At that time it was customary to shoot the birds and prepare the
specimens to be sent to different museums. We have analyzed and validated the
descriptions by Vaurie through over 5000 geocoded high-quality pictures of Little
Owl subspecies across the global range (macaulaylibrary.org). In order to obtain pictures
that are geographically the nearest to the subspecies definitions we selected those that
were closest to the type location of each subspecies.
Vaurie outlined the existence of a geographic trend towards paler colors in arid
lands, but this tendency was questioned by the author himself, because the trend was
true only for a few subspecies, while some populations in desert regions are dark.
The climatic conditions in southern Spain and northern Morocco are similar, but
vidalii is very much darker than glaux. On the other hand, vidalii grades into the paler
noctua in Hungary, and a cline of decreasing color saturation runs eastward to bactriana
of Iran and Transcaspia, via indigena of southeastern Europe and Asia Minor, accom-
panied by a slight cline of increasing size from noctua to bactriana. However, although
the body plumage of indigena is intermediate in coloration between that of noctua and
that of bactriana, its tail is darker than that of either, the pale buff markings on the top
of the tail of indigena being more reduced in size and number and being less regular
than in either noctua or bactriana, in which the pattern is similar.
Owls from the more humid ranges of the Tian Shan (orientalis) are paler than those of
the arid lowlands (bactriana), although one would expect the reverse (Vaurie 1960). The
population of the Tibetan Plateau (ludlowi) is relatively dark, distinctly darker than

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

orientalis. The population of western China (impasta) is intermediate between ludlowi

and plumipes from northern China, and in northwestern Mongolia the birds appear to
be intermediate between plumipes and orientalis.
Pellegrino et al. (2020) utilized observations on 282 museum specimens collected
over a large area spreading through Europe, North Africa and the middle-east of Asia,
aimed at examining of the variation in structural size, the coloration of the feathers,
with particular attention to the pattern of light and dark pigmentation on the tail, and
to spectrophotometry reflectance characteristics to test whether or not size and
plumage show clear-cut differences between the subspecies.
Little attention has been paid to the intensity and distribution of light or dark
pigmentations over tail feathers, despite the fact that this is a character that varies
noticeably in this species (Keve and Kohl 1961). Pellegrino et al. (2020) present a new
tail score system (as illustrated in Figure 3.7) that highlights a difference between
Balkan individuals with respect to all other geographic areas, except for Italy, where
an intermediate pattern was observed. Tail score had significantly higher values in the
Balkans and in southern, central and northern Italy, i.e., these birds had wider dark

Tail score

4
Tail score

1
Iberia
France
England
Central Europe
N-Italy
C-Italy
S-Italy
Sardinia
Balkans
Tunisia Algeria
Egypt
Cyprus
Middle East
Arabic Peninsula

Afghanistan Pakistan
China
Ethiopia Somalia
Iraq

Figure 3.7 Tail score taken in different geographic areas corresponding to distinct described
subspecies (after Pellegrino et al. 2020).

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 3.4 Morphology and Coloration

pigmentation, with reduced light spots in the tail. Little Owls located outside the
Balkans and Italy had lower score values, i.e., their tails had a less dark and whiter
spotted pattern. Pellegrino et al. (2020) did not find any statistically significant differ-
ences when comparing the dark parts of Little Owl plumage in different geographic
areas. Light or dark individuals can be commonly found everywhere.
This result contrasts with old qualitative descriptions of the three European subspe-
cies, i.e., vidalii, noctua and indigena. According to older descriptions, the vidalii Little
Owls are darker with brown and some olive, but not russet-brown feathers, while
indigena are paler and more russet, with the plumage of noctua intermediate (Vaurie
1960, Mikkola 1983, Cramp 1985). Our photographic validation, however, was able to
clearly identify specimens near the type locations that corresponded very well to the
respective textual descriptions.
Plumage color does not differ between females and males. This is in accordance with
the observation that most owl species lack distinctive sexual color dimorphism
(Pellegrino et al. 2020). Plumage is not considered reliable for distinguishing sex.
Natural selection appears to have favored monomorphic plumage coloration for most
owl species (Holt et al. 1999), with the noticeable exception of Tytonidae (Roulin 2016).
However, this argument does not really hold for crepuscular or diurnal owls like the
Little Owl because there are increasing insights into the role of color in Little Owl adults
and juveniles, showing a possible informative role of brightness for colored bare parts
and the iris of Little Owls, as female little owls with brighter bills were larger in size and
produced larger owlets (Avilés and Parejo 2012), nestlings with more yellow bills were
positively discriminated by their feeding parents (Avilés and Parejo 2013) and iris
yellowness was positively associated with nest success (Passarotto et al. 2020).
Individuals found in the same geographical area can differ strongly in color. The
high variation between Little Owls should not be surprising since in raptors individual
differences in plumage pigmentation have been shown to be widespread and can be
influenced by various parameters (Pellegrino et al. 2020). Pigmentation may be related
to behavioral traits (Gomes et al. 2018, Wails et al. 2018, López-Idiáquez et al. 2019),
body conditions (Avilés and Parejo 2013, Wails et al. 2018), parasite resistance (Roulin
et al. 2019) and can have a genetic basis (Roulin and Ducrest 2013, Roulin et al. 2018),
which involves the regulation of genes involved in the melanogenesis pathway (Corti
et al. 2018).
The results of Pellegrino et al. (2020) support the hypothesis that evolutionary
pressure constrains plumage coloration in owls, while other aspects (size, genetics,
vocalizations) might have more discriminative power to show evolution and
differentiation.
The Little Owl plumage also varies individually and tends to vary locally (Cramp
1985, Holt et al. 2019). Furthermore, even Vaurie (1960) recognized from older museum
descriptions the presence of problems related to the comparison of individuals that
were collected when freshly molted and darker with respect to birds collected far from
the molting period with a worn and lighter plumage. Moreover, another difficulty arises
when comparing birds conserved for several decades in museum expositions, which
tend to be paler due to the bleaching effect of light, with stuffed specimens preserved in

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

dark cabinets (Martin et al. 2018). This fact can increase difficulties when attempting to
find reliable differences between areas when comparing birds of different putative
subspecies if birds have been collected in different seasons, the skins were stored in
different manners and the effects of these differences have not been taken into account.
The individual diversity in coloration is very important based upon our study of
georeferenced pictures. Despite exceptional individuals (see Plate 2 for aberrant color-
ations) all subspecies considered and formally described could be recognized, confirmed
and validated based upon the geographical boundaries of the subspecies.

3.4.2 Geographic Variation in Size

The morphological characteristics that were usually utilized in past centuries to describe
Little Owl subspecies were those linked to structural size, i.e., bill, tarsus, tail and wing
lengths, or to characteristics of plumage, i.e., the general pattern of feather coloration.
A comparison of biometrical parameters taken in different geographic areas
(Pellegrino et al. 2020) is shown in Figures 3.8 and 3.9 (tarsus length, wing/tarsus ratio).
Individuals from different areas differed significantly. Pellegrino et al. (2020) found a
noticeable morphometric variability in the Western Palearctic portion of its distribu-
tional range. Size showed two different trends. The first involved the wing length,
which was smaller westward in Iberia, Sardinia, Cyprus, Tunisia, Egypt and southern
Italy, and longer in the Balkans, Iraq, Afghanistan and China. The second trend
concerned the length of the tarsus, which was smaller in Cyprus, Tunisia, Egypt,
Afghanistan, Arabia, China and Iraq, and longer in Iberia, France, England and central
Europe. The samples from Ethiopia had very small wings, tarsus, bills and tails. This
result is in line with the descriptions of Vaurie (1960), which have been previously
reported to pertain to the somaliensis (Somaliland) and spilogastra (Eritrea) subspecies,
both characterized by small sizes. Georeferenced pictures of both subspecies showed a
clear distinction between the two smallest subspecies based on plumage and general
form. Pellegrino et al. (2020) did not find a consistent pattern in the studied areas for
tail and bill lengths.
Pellegrino et al. (2020) give evidence for the presence of a clinal variation in size
from western to eastern Europe, and a particular tail pattern in the Balkan area.
However, discriminative classification of single individuals to the conventional
(Vaurie 1960) subspecies, or to the recently described genetic clades (Pellegrino et al.
2015), has a limited classification accuracy of 70% for the four European subspecies
(vidalii, noctua, sarda, indigena) and 94% for the two furthest apart (vidalii and indigena).
The lack of statistical differentiating power between 18 geographical regions clearly
illustrates the clinal gradient that is typical for this species.

3.5 Morphological Descriptions

The morphological descriptions are based upon the original descriptions of the 14 sub-
species complemented by the descriptions of Hartert (1913) and Vaurie (1960) and
validated by georeferenced pictures of Little Owls at or near the type locations.

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Tarsus length
3

saharae/glaux
2

indigena

lilith/saharae
indigena
Tarsus length (STD)

plumipes/ludlowi
1

bactriana
lilith

spilogastra/somaliensis
0

-1

-2
sarda
vidalii
noctua

-3
Iberia
France
England
Central Europe
N-Italy
C-Italy
S-Italy
Sardinia
Balkans
Tunisia Algeria
Egypt
Cyprus
Middle East
Arabic Peninsula

Afghanistan Pakistan
China
Ethiopia Somalia
Iraq

Figure 3.8 Tarsus length taken in different geographic areas corresponding to distinct described
subspecies (after Pellegrino et al. 2020). Green: western Europe; blue: Italy; yellow: Sardinia; red:
Balkans. Labels in brown indicate the subspecies as defined by Vaurie (1960).

The descriptions of the subspecies are presented in a geographical order from west to
east.

1. Athene noctua noctua Scopoli (1769). This race (Plate 3) is rufous brown and paler
than vidalii, the white spots above and the dark streaks underneath somewhat
less contrasting.

Hartert (1913) describes this subspecies as follows. Upper side earth brown, like
sepia, upper head with longer or more rounded, shorter shaft spots that do not reach
the tip and are white with a cream-colored touch. Feathers on the neck and back of the
neck creamy white with a cream-colored touch with gray roots and broad, earthy brown
edges, so that a distinct, wide, white flank band is created, on the rest of the upper side
the whitish flares are smaller and often double, more or less thick the broad hems are
covered, but they stand out much more on the scapular and larger elytra; they are big
there and the flecks of the two flags together form a big spot. Wings deep brown,
external flags with cream-colored flecks, inner flags with whitish cross-ties that do
not reach the shaft, and a more distinct or brighter, whitish lace border on the
wings. Control feathers like the back, with a narrow end hem and 4, rarely 5 light

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 3.5 Morphological Descriptions

Wing/Tarsus ratio

Wing/Tarsus ratio 5.5

5.0

4.5

4.0
Iberia
France
England
Central Europe
N-Italy
C-Italy
S-Italy
Sardinia
Balkans
Tunisia Algeria
Egypt
Cyprus
Middle East
Arabic Peninsula
Iraq
Afghanistan Pakistan
China
Figure 3.9 Wing/tarsus ratio taken in different geographic areas corresponding to distinct described Ethiopia Somalia
subspecies (after Pellegrino et al. 2020).

cream-colored transverse bands, which are sometimes not completely connected on the
shaft. The chin and front neck are white, separated from one another by a brown patch
band, the rest of the underside is white with cream-colored patches and dark or earth-
brown patches, mostly in the form of slightly pale longitudinal patches in the middle,
becoming more rounded on the chest. Anal area and lower ceilings unspotted, the latter
often with brown shafts or similar but only very narrow stripes. Running fletching
creamy white, lower wing covers as well, but with a few brown spots. Iris yellow. Bill
wax yellow, greenish at the root.
Pellegrino et al. (2020) was not able to statistically distinguish between noctua and
vidalii based upon size and coloration. The majority of pictures taken in Western Europe
however are considerably darker than pictures taken in Italy (noctua) and the Balkans
(indigena).

2. Athene noctua vidalii Brehm A.E. (1857)

This race (Plates 4–6) differs by being very much darker, dark umber brown, tinged
faintly with rufous brown in some individuals. It is the darkest race of the species;
darker in the body, with higher, narrower and longer spots and black streaks. It is also
more regularly barred on the tail and more conspicuously spotted with purer white on

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

the upper parts. New feathers are always very distinctly darker than the old ones in birds
that are molting or in very fresh plumage.

3. Athene noctua sarda Kleinschmidt (1907)

This race (Plate 7) is poorly differentiated from noctua, but its general coloration is
somewhat paler, with smaller white spots on the scapulas. It differs in the narrower
markings. The tail is narrowly banded with white. The difference is most evident in the
shoulder feathers. The white transverse bands are very small here and are divided by a
dark line along the middle of the feather.

4. Athene noctua indigena Brehm C.L. (1855)

This race (Plate 8) differs from noctua by averaging somewhat paler, and more rufous, in
general coloration and by averaging a little larger, but essentially by being less exten-
sively and sharply patterned with buff on the tail. It has very overgrown spots on the
lower body. In indigena, the pale and buff markings are reduced in number and size and
form rounded or elongated spots rather than bars and are irregularly arranged in two
rows, less often three, as against three or four rather regular bands in noctua. It
approaches bactriana in size, but differs clearly from it in coloration and by having
the toes less thickly feathered; in bactriana the tail is well and regularly barred as
in noctua, the general coloration is considerably paler, more grayish, than in indigena,
and the crown, nape, wings and scapulas are more profusely marked with larger
white spots. Some authors consider that indigena is intermediate between
nominate noctua and bactriana, but indigena is not truly intermediate other than in
the length of the wing. It is much more similar to noctua than it is to bactriana, with the
exception of the reduced and irregular pattern on the tail, and in this respect differs
from both.

5. Athene noctua bactriana Blyth (1847)

Plumage of the upper parts is somewhat rufescent clay-brown, with large white spots
on the features, more or less concealed, and wholly so for those in the middle of the
back (Plate 9). Coronal feathers with whitish streaks. Face white, some of the radiating
feathers on the sides of the beak terminating in black vibrissae. Chin, throat, lower tail-
coverts, and the tibial and tarsal plumes, are white as is the fore-part of the under-
surface of the wing. There is a longitudinal broad streak on each feather of the breast
and abdomen. On the hindneck, the white predominates upon the feathers so as to give
the appearance of a half-collar. The great wing feathers have broad incomplete pale
bands, positioned alternately on their two webs and the middle tail feathers have a
double row of semi-alternating pale spots, changing into faint bands on the outer tail-
feathers. Beak whitish and claws pale horn-color.
On average larger than the preceding races with the toes more thickly feathered.
Paler than glaux but darker than saharae and lilith. Less sandy, more greyish brown, less
spotted with white than lilith. More greyish, less rufous brown above than saharae, and
more heavily streaked underneath with darker and browner streaks.

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

6. Athene noctua lilith Hartert (1913)

Even paler, more sand-colored (Plate 10) than A. n. bactriana, the spots on the underside
are also paler and more reddish-yellow. Moreover the toes are only covered with
bristles, not densely feathered as in bactriana. Whiter on the crown, nape, back
and wings. Clearly less streaked underneath and the tail is more regularly barred
with buff.
This race differs from saharae by being distinctly paler throughout, very pale “sandy”
in coloration, whiter on the head, back and wings and, especially, on the nape, where
the white spots form a large patch. It is the palest race of the species and also differs
from saharae by being very distinctly, less heavily, streaked underneath with paler,
more reddish, less brownish streaks; its tail is also more regularly barred with buff
on average.
Vaurie (1960) spread out 29 saharae side by side with 15 lilith from Arabia, finding
that the latter averaged much paler, with especially the heads being lighter; only 3 of 15
could be mistaken for saharae, while only 2 of our saharae would be called lilith if their
origin was unknown.

7. Athene noctua saharae Kleinschmidt (1909)

This race (Plate 11) is similar to glaux but differs from it by being paler throughout and
by being less heavily and darkly streaked underneath, but more profusely spotted with
white on the crown and the entire upper parts. It varies more, individually and locally,
than glaux.
Vaurie had specimens from Arabia, the population of which is said to be very
variable individually and locally, some individuals being similar to saharae, others to
the paler lilith, while the rest are intermediate to a varying degree between these
two races.
Our findings by studying georeferenced pictures identified clear saharae and a vari-
ation with rusty glaux-like lilith features without large reddish patches on the belly
underneath the wings from the Arabian Peninsula (Plate 12).

8. Athene noctua glaux Savigny (1809)

This race (Plate 13) is relatively dark on the upper parts and the streaks underneath,
which are heavy, being rufous brown or pale chocolate brown; the tail is irregularly
spotted or barred with brownish buff. The wings are slightly pointed and shorter than
the tail. There are four to five slightly indented flight feathers; the first rather short, the
third the longest. The tail is equal. There are poor and irregular periopthalmic circles.
The outer ears small, round and without operculum. The beak is thick, very short,
sharply inclined, slightly compressed and convex underneath; wax very swollen on
the nostrils and as a gibbous on each side; nostrils wide apart, very small, perfectly
round, turned forward; lower mandible with marginal notches towards the end. The
tongue is oval and thick, with two ribs below, very obtuse. There are woolly tarsi on all
sides. The toes are hairy to the base of the last knuckles. The nail is intermediate
without crenellations. Iris pale yellow; spotted belly; white throat.

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 3.5 Morphological Descriptions

9. Athene noctua spilogastra von Heuglin (1863)

This race (Plate 14) is similar to saharae in coloration but smaller. Above pale and faintly
rufescent-smoky, variable whitish spots and streaks of a yellowish-washed white color;
whitish tail feathers with six to seven bands. Rufous-ashy; the face, the chest, and the
abdomen are whitish, scarcely washed with a yellowish shade, with worn-out and faint
subrufous streaks; the upper part of the breast is of the same variable color; the tarsi are
white feathered in front; the toes are with bristly, whitish hairs above; feathers under
the wing are completely white; beak yellow; cere with few short blackish bristles; iris
yellow; orbital ring blackish; incarnate-lead fingers.The fourth primary feather is the
longest, the third slightly shorter, the first about the same as the seventh. The first
primary is clearly sawn on the outer vane; there is also a hint of this on the second. The
backs of the toes are very sparsely covered with bristle-like feathers; the tarsi appear bare
on the back. The claws are noticeably short and blunt. This race lives in pairs along the
rain beds that are deeply torn in debris, in piles of stones and on stacks of bushes. Birds
hide during the day and feed on mice, small birds, lizards, beetles and grasshoppers.
When chased it hides in holes in the ground and under debris.

10. Athene noctua somaliensis Reichenow (1904)

In Somaliland there is a form (Plate 15) deviating from the Abyssinian (Eritrea) spilogas-
tra, with wings and tail shorter, top darker brown, top of head monochrome or spotted
white, but not narrowly dashed, brown dashes on the underside darker. This race differs
from spilogastra by being smaller and distinctly darker. It is darker brown on top, more
heavily streaked with darker brown underneath, and less spotted or streaked with white
on the crown, these markings being reduced in number and extent than in somaliensis
or absent altogether.

11. Athene noctua orientalis Severtzov (1873)

This race (Plate 16) differs from noctua by the fact that the brown color in the latter is
darker, orientalis is paler, changing to gray-russet with a yellowish tint as in glaux, from
which orientalis is distinct by dense feathering of the toes, excluding the nail joint. The
ear hole is oval, and half the skull. The general coloration of the upper side of body is
usually lighter than in bactriana (sometimes there are birds indistinct from bactriana in
this respect), clayish-pale-gray, less brown. The dark pattern on the under-side of body
is somewhat more contrasting than in bactriana. The extent of the white spots on the
upper side of body is more developed than in the other races.
This race is similar to bactriana in size and in having thickly feathered toes, but is
paler above, more grayish, less brownish, more conspicuously and abundantly spotted
with larger white spots on the whole of the upper parts and wings, and somewhat less
heavily and darkly streaked below. It is much paler than indigena, but the base color-
ation of its upper parts is considerably darker than that of lilith, more grayish and
“cooler” in tone, less “sandy” and yellowish, although almost as well spotted with
white. It also differs from lilith by having darker, more brownish, less reddish, streaks
underneath, a longer wing and thickly feathered toes.

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 3.5 Morphological Descriptions

According to Dementiev (1931) orientalis differs from bactriana by being darker and
larger, and by having more thickly feathered toes and less spotted with white on top
than bactriana and more heavily and darkly streaked underneath. The difference in
coloration between the two races is therefore remarkably constant for such a species in
which individual variation is often great in some regions.

12. Athene noctua plumipes Swinhoe (1870)

Throat white, the white extending in crescent-form up each cheek behind the
ear-coverts; the lores, around the eye and middle of the belly are also pure white;
the upper parts are light reddish brown, with spots of reddish white on the head,
and marked and spotted as much as noctua. The underparts, leg- and feet-feathers are
cream-colored, with the breast and flanks broadly streaked with reddish brown, like
glaux. The beak is yellow tinged with green. The iris is light yellow and claws blackish
brown.
The general coloration of the upper side of body (Plate 17) is close to bactriana and
slightly darker, more brown-gray, than in orientalis. The white spots on the upper side of
body are similar to orientalis. This race differs from ludlowi by being a little darker, more
rufous brown, on top and by being considerably smaller; it is similar to glaux from
Africa in general coloration.

13. Athene noctua ludlowi Baker (1926)

In general color intermediate between noctua and bactriana (Plate 18) – in fact, very
similar to plumipes, but decidedly bigger. The amount of feathering on the toes varies
considerably, both individually and seasonally, but in winter most birds have the
plumelets extending down the toes almost to the base of the claws. Iris yellow; bill
bright yellow; legs gray, soles yellow.
A relatively dark race, darker than orientalis, similar to bactriana in general coloration,
but more spotted with white. This race differs from orientalis by being darker and
considerably larger, being the largest race of the species. It is more rufous brown on
top than orientalis, less grayish, and is less spotted on average, the spots being also more
buff and less white; the under parts are also slightly more buff and less white, and are
distinctly more heavily and darkly streaked.

14. Athene noctua impasta Bangs and Peters (1928)

Similar to plumipes in having feathered toes, but much darker; pale markings on the top
of the head (crowns-stripes) are linear rather than guttate; light tail bands are inter-
rupted; markings underneath are darker and more extensive. Impasta is much darker
than plumipes and ludlowi, a little paler than noctua.
Impasta (Plate 19) is intermediate between ludlowi and plumipes in body size and claw
length. In higher latitudes, the wings and claws of races become shorter. High-quality
georeferenced pictures illustrate a clear distinct brown color of all individuals near the
type location.
Plate 20 gives an overview of all 14 subspecies.

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 3.6 Geographical Distribution

3.6 Geographical Distribution

3.6.1 Global Range
The global distribution of the Little Owl ranges from the Atlantic shore of Britain into
Eurasia, and eastward to northeastern China and the Korean Peninsula. It extends
northward through Belgium, the Netherlands, Denmark, Latvia, regions of Pskov and
Moscow, to the middle part of Meshchera, north to the 56th north latitude. It is found
in the region of the Ural ridge to the 54th parallel, to the Ilek River mouth, in the north-
central part of Kazakhstan and in eastern Kazakhstan to the 49th parallel, to south-
eastern Altai, to the Tannu-Ola ridge, the southwestern Trans-Baikalia and the
Kheiluntszyan Province of northeastern Mongolia.
The species is found southward to the Mediterranean Sea, to the northern shore of
the Arabian Sea, northeastern Pakistan, southern Tibet, and in central and eastern
China, approximately to the 35th parallel. It is found on the islands of the
Mediterranean Sea, on the Arabian Peninsula, northern and northeastern Africa,
in western Africa south to the 22nd parallel, to Sudan and Somaliland (northern
Somalia).
The southern boundaries of the distribution in the Sahara were either confirmed or
highly supposed to be in higher altitude areas with availability of water, such as around
Zemmour, northern Mauretania (22), the mountain massifs of Ahaggar Tamanrasset,
southern Algeria (23) and Aïr (18) in northern Niger, Tibesti, northern Tchad (20),
Ennedi and Am-Djarass, and northeastern Tchad (17).
The owl was introduced in England with Dutch specimens (Sharrock 1976) and in
New Zealand (Kinsky 1973) with German specimens.
The global distribution of the Little Owl is shown in Figure 3.10. This map offers a
compilation of the maps of Vaurie (1960), Cramp (1985), Del Hoyo et al. (1999) and
König et al. (1999), taking into account the most recent distribution from EBBA2 (Keller
et al. 2020) at 5050 km UTM squares, and the status and trends of EU bird populations
datasets from Article 12, Birds Directive 2009/147/EC reporting at the 1010 km UTM
squares level. In addition, the insights that were obtained from our study of the
georeferenced pictures are used to settle subspecies discussions.

3.6.2 Subspecies
ATHENE NOCTUA VIDALII BREHM A.E. (1857)
Range: Western Europe, Iberia (Spain and Portugal) north to France, Belgium,
Netherlands, Germany, Denmark, east through Germany and Poland to Baltic
States, to the Ural River. Introduced in Britain; intergrades with indigena in
Croatia, Bosnia-Herzegovina, southeastern Austria, Hungary and northern
Rumania and west of Carpathian mountains, Ukraine north of Kiev, Voronezh, and
Orenburg. Introduced to Britain with Dutch birds and New Zealand with
German birds.
Type locality: southeastern Spain.

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 Figure 3.10 Global distribution of all 14 subspecies of Athene noctua.
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 3.6 Geographical Distribution

ATHENE NOCTUA NOCTUA SCOPOLI (1769)

Range: mainland Italy, Sicily, southeast Austria, Slovenia, Slovakia and Hungary,
intergrading locally with vidalii in southern France, with indigena in Croatia, Bosnia-
Herzegovina, southeastern Austria, Moldova, intergrading with vidalii in southern
Germany, northern Switzerland and western Austria is unlikely due to the
geographical barrier of the Alps during the postglacial re-colonization of Europe and
has not been genetically confirmed (Pellegrino et al. 2020).
Type locality: Carnolia, Ljubliajana, Slovenia.

ATHENE NOCTUA SARDA KLEINSCHMIDT (1907)

Range: Sardinia throughout the island, south of Corsica.
Type locality: Sardinia, Italy.

ATHENE NOCTUA INDIGENA BREHM C.L. (1855)

Range: Albania, Bosnia, southern and eastern Rumania, south Moldova and Georgia,
and further east to the Caspian Sea, south to Crete, Cyprus, Rhodes and Turkey (except
southeast), Transcaucasia, intergrading with bactriana in southern Trans-Uralia.
Type locality: Attica, southern Greece.

ATHENE NOCTUA GLAUX SAVIGNY (1809)

Range: coastal North Africa, along the Nile, separated from saharae in Morocco and
Atlantic Sahara by the Draa River south of the Atlas Mountain range.
Type locality: Egypt.

ATHENE NOCTUA SAHARAE KLEINSCHMIDT (1909)

Range: northern and central Saharan Desert (Africa), south of glaux, on the Arabian
Peninsula overlapping with lilith.
Type locality: Biskra, Algeria.

ATHENE NOCTUA LILITH HARTERT (1913)

Range: inland Levant from Sinai to southeast Turkey, intergrading with bactriana in
Iraq and overlapping with saharae in Saudi Arabia.
Type locality: Deir ez-Zor, eastern Syria.

ATHENE NOCTUA BACTRIANA BLYTH (1847)

Range: from the eastern shore of the Caspian Sea, southeast Azerbaydzhan and eastern
Iran, and the Ural River valley to east to eastern edge of Balkhash-Alakol hollow, the
foothills of Tarbagatai, Dzhungarskoe (Jungarian) Alatau, the foothills of Tien Shan
and the foothills of the Alai system. In the central and eastern part of Kazakhstan,
the northern border is the 49th parallel, extending southward to the southern border
of the country. Within these boundaries it inhabits lowlands, and does not occupy
the upland mountains. Near the western border of its distribution (e.g., the western
bank of the Ural River) it intergrades with indigena, and in the western and northern

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

foothill region of Tarbagatai, Jungarian Alatau, Tien Shan and the Alai system, it
supposedly integrades with orientalis.
Type locality: Candahar, Afghanistan.

ATHENE NOCTUA LUDLOWI BAKER (1926)

Range: from northern Kashmir (the foothills of the Karakoram), eastward over the
Tibetan Plateau to the region of Litang in the east-central region of Sichuan and
western Qinghai in southwestern China. North to the upper basin of the Yangtse
and Yalung in Qinghai. In Kashmir found in Baltistan and Ladakh, to the Kunlun
Mountains in Xinjiang.
Type locality: Dochen, Lhasa, Namtso Lake, southern Tibet.

ATHENE NOCTUA ORIENTALIS SEVERTZOV (1873)

Range: Pamiro-Alai Mountains country, Tien Shan, Jungarian Alatau, Tarbagatai up to
Lake Balkhash. In the western and northern foothill region of this territory it
supposedly intergrades with bactriana, in the region east of Tarbagatai supposedly
with impasta.
Type locality: Issyl Kul Lake, Tjensjan, Kirghizstan.

ATHENE NOCTUA IMPASTA BANGS AND PETERS (1928)

Range: southeastern Altai (Kazakhstan – Russian Federation), Tannu-Ola (Russian
Federation – Mongolia), southwestern, southern and southeastern Trans-Baikalia
(Russian Federation), ranging in China to Qinghai, Gansu and southern Shaanxi. In
the region east of Tarbagatai (Kazakhstan) it supposedly intergrades with orientalis.
Type locality: Qinghai Lake, eastern Qinghai, western China.

ATHENE NOCTUA PLUMIPES SWINHOE (1870)

Range: most eastern subspecies ranging in China from Shaanxi to Shandong and Hebei
to the Yellow Sea, and bordered by the Jangtsekiang River in the south. The eastern
extension of the range into North Korea might be restricted to a straggler or winter
visitor, according to Austin (1948).
Type locality: Juyong Guan, Changping District, China, northwest of Peking, Hebei.

ATHENE NOCTUA SPILOGASTRA VON HEUGLIN (1863)

Range: on the African coast of the Red Sea from Port Sudan in the north to at least as far
south as Buri Peninsula in Eritrea and possibly Ethiopia.
Type locality: Massawa, Eritrea.

ATHENE NOCTUA SOMALIENSIS REICHENOW (1904)

Range: on the African coast of the Red Sea in Somaliland, supposedly into Ethiopia
along the Somalian border.
Type locality: northern Somaliland.

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 3.6 Geographical Distribution

3.6.3 Intergrading of Subspecies

Intergrading of subspecies is important in certain countries, such as Kazakhstan, Israel,
China and Russia.
Based on morphology, Kazakhstan has five subspecies (Gavrilov 1999) ranging as follows:

vidalii: northern part of Volga–Ural inter-river area (south of the 49th parallel) and the
Ural River valley eastward to the lower Ilek River valley.
indigena: southern part of Volga–Ural inter-river area northward to the 49th parallel,
eastward to the lower Ural River valley.
bactriana: from the eastern shore of the Caspian Sea and Ural River valley to eastern
border of Balkhash-Alakol hollow, Tarbagatai foothills, Jungar Alatau and Tien Shan.
orientalis: Tien-Shan, Jungarian Alatau, Tarbagatai, probably foothills of Altai and Kalba.
impasta: Altai; in the region of Tarbagatai it supposedly intergrades with orientalis.

Based on morphological data, Russia has five subspecies (Stepanyan 1990) with distri-
butions as follows:

vidalii: from the western border of the country eastward to the Uralian ridge and Lower
Ilek valley. The northern border of the species’ breeding range is in this region; the
southern border is the northwestern shore of the Black Sea, Lower Dnieper River,
northern shore of the Azov Sea, in the region of the Volga–Ural inter-river area to the
49th parallel. The southern border of its distribution is in the region of the Crimean
Isthmus, the southeastern part of Ukraine, north Caucasia; it integrades with
indigena at the region of the 49th parallel in the Volga–Ural inter-river area.
indigena: Crimea, Great Caucasia, Trans-Caucasia, southern part of the Volga–Ural inter-
river area, eastward to the Nizhniy (Lower) Ural River area. Northward to the
Crimean Isthmus, the northern shore of the Azov Sea, in north Caucasia and in the
Volga–Ural inter-river area north to the 49th parallel. It integrades with noctua near
its northern border of distribution, in the region of the Crimean Isthmus,
southeastern Ukraine and eastwards in the region of the 49th parallel. It integrades
with bactriana near the eastern borders of its distribution, in the region of the
western bank of the Ural River.
bactriana: from the eastern shore of the Caspian Sea and Ural River valley eastward to
the eastern edge of the Balkhash-Alakol hollow, foothills of Tarbagatai,
Dzhungarskoe (Jungarian) Alatau, foothills of Tien Shan and foothills of the Alai
system. Near the western borders of its distribution, in the region of the western
bank of the Ural River, it intergrades with indigena. It supposedly integrades with
orientalis in the western and northern foothills of Tarbagatai, Jungarian Alatau, Tien
Shan and the Alai system.
orientalis: occupies the Pamiro-Alai Mountain country, Tien Shan, Jungarian Alatau,
and Tarbagatai. It supposedly intergrades with bactriana in the western and northern

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

foothills of the Tarbagatai, Jungarian Alatau and Tien Shan region. It supposedly
intergrades with impasta in the Tarbagatai region.
impasta: southeastern Altai, Tannu-Ola, southwestern, southern and southeastern
Trans-Baikalia. In the region of Tarbagatai it supposedly intergrades with orientalis.

The distribution of Athene noctua bactriana (Hutton’s Owlet) covers the area from Iran
and Iraq, east-northeastwards through Afganistan and north Pakistan, Kashmir to Syr
Darya. In addition, it is not uncommonly encountered in suitable habitat in western
Ladakh, in the Indus Valley. The range of Athene noctua ludlowi (Tibetan Owlet) is much
more restricted, being limited to the area from eastern Ladahk eastwards through Tibet,
north Sikkim and north Bhutan. Ladakh can therefore be considered as the zone where
the two subspecies meet (Pfister 2001).
China has four subspecies (Lei et al. 1997, Qu et al. 2002) ranging as follows: ludlowi
in Tibet, orientalis in Xinjiang, plumipes in Shanxi and impasta in Qinghai and Gansu.
Serbia-Montenegro features two subspecies: Athene noctua noctua in the northern
part and A. n. indigena in the southern part (Matvejev 1950).
Israel is a meeting point of four subspecies but only lilith occurs in its typical form.
The three others (indigena, glaux and saharae) represent only intergrading populations
with lilith (Shirihai 1996). The border between the coastal races indigena and glaux
appears to be situated around Haifa. Our analysis of georeferenced photographs did
not reveal any pictures of glaux nor saharae for Israel. The glaux reference might be
referring to the Arabian variant of the glaux-like lilith without the large reddish patches
underneath the wings.

3.7 Genetic Approach

An increasing body of knowledge is becoming available through genetic analysis. Most
studies focus on genetic variability within local populations or across different subspe-
cies. Although sample sizes remain modest and the number of subspecies incomplete,
they offer a better view on the genetic diversity and on the evolution of the species and
its subspecies. Table 3.2 offers an overview of all genetic studies so far and the subspe-
cies that they are dealing with. Unfortunately, an overall genetic study covering all
14 subspecies is still lacking.

3.7.1 Genetic Variability

The genetic variability was studied with multilocus enzyme electrophoresis in Italy
(Randi et al. 1991). The caryotype of the species is 2n = 82 with 18 macrochromosomes
and 64 microchromosomes (Capanna et al. 1987). The polymorphism (P) of the Little
Owl is 0.13 and heterozygosity (Ho) 0.026, which is similar to the Barn Owl, but less
than the Long-eared Owl (P = 0.23; Ho = 0.024) and Tawny Owl (P = 0.16; Ho = 0.041).
Because of the low heterozygote frequency, the Little Owl could be subdivided in
different and more or less isolated geographic populations, with different allele frequen-
cies at some polymorphic loci. However, the studied samples were not large enough to
allow intraspecies population analysis (Randi et al. 1991).

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Table 3.2 Overview of genetic studies on Athene noctua and the subspecies accounted for

Wink
and
Génot Müller Qu Pellegrino Sauer-
Randi et al. et al. et al. Bouchy Wink et al. Gürth
Genetic
Subspecies Year Author Country 1991 2000 2001 2002 2004 2008 2014 2015 2021 studies

Athene noctua 1769 Scopoli Italy 1 1 1 1 1 5

noctua
Athene noctua 1809 Savigny France 1 2
glaux
Athene noctua 1847 Blyth UK 0
bactriana
Athene noctua 1855 Brehm Germany 1 1 1 1 4
indigena
Athene noctua 1857 Brehm Germany 1 1 1 1 1 1 1 7
vidalii
Athene noctua 1863 Heuglin Germany 0
spilogastra
Athene noctua 1870 Swinhoe UK 1 1 1 3
plumipes

continues
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82

Table 3.2 (cont.)

Wink
and
Génot Müller Qu Pellegrino Sauer-
Randi et al. et al. et al. Bouchy Wink et al. Gürth
Genetic
Subspecies Year Author Country 1991 2000 2001 2002 2004 2008 2014 2015 2021 studies

Athene noctua 1873 Severetzov Russia 1 1

orientalis
Athene noctua 1905 Reichenow Germany 0
somaliensis
Athene noctua 1907 Kleinschmidt Germany 1 1 2
sarda
Athene noctua 1909 Kleinschmidt Germany 0
saharae
Athene noctua 1913 Hartert Germany 1 1 2
lilith
Athene noctua 1926 Baker UK 1 1
ludlowi
Athene noctua 1928 Bangs and UK 1 1
impasta Peters

Number of 1 1 1 4 1 6 4 4 5
subspecies
per study
 3.7 Genetic Approach

3.7.2 Genetic Fingerprinting

In France, genetic fingerprinting was carried out to examine the basis for a
supplementation experiment of a Little Owl population (Génot et al. 2000). Here,
genetic work was conducted to establish the polymorphism within birds from different
regions of the country and from different localities in the same region. The DNA bands
were analyzed to examine the degree of genetic similarity. Technically speaking, X =
2Nab/Na + Nb, where Na and Nb are the numbers of bands in individuals a and b, and
Nab is the number of bands shared by a and b. If X is near 1, the birds are closely related
genetically, if X is near 0, they are distant. The analysis of owls from different geo-
graphic origins revealed that genetic variability is not unique to each region. Two
unrelated birds from the same region may be more distant genetically (X = 0.36) than
two birds from different regions (X = 0.52). However, the sample size of nine individuals
was too restricted to establish with any certainty the degree of genetic diversity between
two unrelated birds. As far as owls from different localities were concerned, the marked
similarity coefficients indicated the existence of an important genetic proximity
between the different individuals studied (X = 0.8). Such values thus reflect both the
absence of great genetic diversity locally and the homogeneity of the similarity
coefficients obtained.
DNA fingerprint analyses were also carried out in Germany to calculate band-sharing
coefficients (0.580.09; n = 53) between putative fathers and nestlings to check a high
rate of paternal care and no extra-pair young in a Little Owl population (Müller et al.
2001).
In France, a study (Bouchy 2004) used five polymorphic microsatellites (DNA
markers), isolated de novo, for three aims. First, to assess the genetic structure of five
French populations of Little Owls, three from the Vosges, one from the Drôme and one
from the Oléron Island. Second, to confirm or invalidate demographic observations
obtained from mark–re-capture programs, as well as to give reliable estimations of
dispersion rates that could not be accurately evaluated with classic mark–re-capture
methods in these cases. Third, to determine if some populations, genetically close to
those from the Vosges, could be used as sources of wild birds for the re-inforcement
program in the Vosges (instead of the captive birds used, which was a failure). The
populations exhibited quite high internal genetic diversity, which means that the
authors did not find any genetic impact of fragmentation. All analyzed populations
appeared substructured. Results indicated that the dynamics of the populations were
not correctly described and that further sampling efforts should be organized in light of
this study. Although this genetic study on Little Owls revealed a significant genotypic
differentiation between the populations studied, geographically distant populations
were genetically close. Thus, instead of using local birds from captivity in this re-
inforcement program, one could use birds from other wild and healthy populations.
However, the real utility of this conservation action should be studied carefully.
Indeed, some populations appeared endangered, according to their supposed dynamics.
This study showed that sampling methods were not appropriate in the Drôme case, and
that birth and death rates were probably not correctly estimated, especially in the case

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

of the Northern Vosges. Hence, other sampling efforts, given all the remarks made and
evidence found, should be organized to improve our knowledge of these populations,
prior to any conservation decision.
A mixed demographic genetic model was constructed for the subpopulations of the
Northern Vosges. Deleterious mutations have an impact not only on population
dynamics, but also on the rate of effective exchanges. Under a pessimistic demographic
scenario, all three local Northern Vosges subpopulations are sustained by a high
immigration rate from a large German population located nearby (2–3 km) in Sarre.
Inbreeding depression has therefore little impact on the vital rates and mutation
accumulation does not occur. Under an optimistic scenario, local demographic rates
are sufficient to give population sustainability, and deleterious mutations may pose a
serious threat to persistence within a 100-year period, especially if the number of
German migrants is low (Bouchy 2004).

3.7.3 Phylogenetic and Phylogeographic Relationships

Since owls are nocturnal, their morphological traits are often very similar (König et al.
1999) and a systematic differentiation or speciation event can easily be overlooked.
Together with morphological and vocal characters, nucleotide sequences of marker
genes help to define phylogenetic relationships, systematics and evolution of owls.
The mitochondrial cytochrome b gene is regularly amplified and sequenced. Because
this gene shows a good resolution at the genus level and has large samples for this gene
(Wink and Heidrich 1999, Wink et al. 2004), it is used as a platform to cover all taxa of owls.
About 109 taxa of the Strigidae and 21 of Tytonidae have been studied and a phylogeny based
on cytochrome b data has been published. In addition to Athene, a selection of representa-
tive owls of the genera Glaucidium, Surnia and Aegolius are usually combined (Wink 2008),
since these genera form a monophyletic group together with Athene. A sister group to this
assemblage consists of members of the genera Bubo, Strix, Otus, Megascops, Asio and Ptilopsis.
Both groups form the subfamily Strigidae that clusters as a sister group to the Tytonidae
(Wink and Heidrich 1999). The phylogeny methods maximum parsimony (MP) and
maximum likelihood (ML) were employed by Wink (2008) to reconstruct phylogenetic
relationships in the present dataset (Figure 3.11). Athene forms a monophyletic genus
Athene clustering basal to Aegolius, Surnia and Glaucidium as part of monophyletic subfamily
Strigidae comprising all non-Tyto owl genera. These relationships could be confirmed using
nucleotide sequences of a nuclear marker gene (LDH b intron) (Wink et al. 2004).
Nucleotide sequences of the cytochrome b gene were used to infer phylogenetic and
phylogeographic relationships in members of the genus Athene. Surprisingly, Ninox super-
ciliaris turns out to be a member of the monophyletic Athene clade and is regarded as Athene
superciliaris by Wink (2008). Within Athene noctua several well-defined clades are apparent
which agree with recognized subspecies. A substantial differentiation might be found in
the Little Owl as more data become available. Because some of the genetic distances
between the subspecies are in the typical range for established species, it might be suggested
that Athene noctua comprises a monophyletic species complex that could be subdivided
into several “good” species (according to Wink in: Van Nieuwenhuyse et al. 2008).

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 3.7 Genetic Approach

MP
Bootstrap
Gallus
97 Phodilus. badius.6222
Tyto a.guttata LZ 17
98 Aegolius a. acadicus.6242
100 Aegolius.harrisii.6315
Aegolius funereus FL 6332
100 Surnia ulula FINL 6305
52 Surnia ulula FINL 6306
89 Glaucidium.perlatum.S Africa.6074
Glaucidium passerinum1 AU 6067
59 69 Glaucidium brasilianum2 ARG.6053
98
Glaucidium gnoma CR 6149
60 Glaucidium.cuculoides.CHINA.9576
Glaucidium.capense.KENYA 6288
Athene.barma.6212
A.noctua.lilith3680 TAU Israel
91 98 A.noctua.lilith TUR9.6307
A.noctua.lilith.CYP.20883
99 100 Athene.n.indigena.GR2878
99 A.noctua.glaux 3683 gz Israel
93
A.noctua.glaux 3684 gz Israel
100 Athene.noctua.plumipes.MONG.23811
100
Athene.noctua.plumipes.MONG.23812
91
Athene.noctua.vidalii.8228
100
68 87 Athene n.noctua8 DE
Athene n.noctua AU4
Athene.superciliaris.MADAG.10419
94 83 Athene cunicularia punensis AF448257 300
51 Athene cunicularia nanodes AF448255 300
100 Athene c.cunicularia NEArgent
93 Athene cunicularia floridana AF448254 300
Athene cunicularia.hypugaea BAJA 9027
Bubo b. buboSW5
90
95 Bubo virginianus. Utah 23142
Bubo [Nyctea] scandiacus 4
88 Otus megalotis 1
100 Otus bakkamoena 1
Otus.scops.13 77.6251
87 82 Megascops choliba4
Megascops flammeolus
96 Asio.flammeus.465
71 Asio.o.otus.NOR.2478
100 Ptilopsis.leucotis.22328
Ptilopsis.granti.6171
99 Strix a.aluco DE6
Strix butleri2

Figure 3.11 Molecular phylogeny of Athene and other owls inferred from nucleotide sequences of the
cytochrome b gene; MP-boot-strap cladogram with boot-strap values (from 1000 replications) at
branches (after Wink in Van Nieuwenhuyse et al. 2008).

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

The analysis of the Athene complex by Wink (2008) so far only covers the subspecies
vidalii, noctua, indigena, lilith, glaux and plumipes. Some of the African and Asian taxa are
not represented, thus leaving the analysis incomplete at present. The present dataset
reveals three phylogenetic lineages that are supported by boot-strap values between
93 and 100%.
Athene n. noctua and A. n. vidalii represent sister taxa that appear to be well recog-
nized (genetic distance 4.1%) and clearly separated from A. n. indigena in the Balkans.
Athene n. plumipes clusters at the base of the clade leading to Little Owls of the Balkans,
and the Near and Middle East. Its genetic distance from the other Little Owl subspecies
is between 6 and 9%. A. n. lilith, A. n. indigena and A. n. glaux are closely related, but
differentiated. A. n. lilith from Cyprus shows an identical haplotype to Little Owls from
Turkey and Israel. Own findings by studying georeferenced pictures and local observa-
tions (J. De Raedt personal communication) did not lead to confirmed observations of
lilith in Cyprus. The genetic distances between lilith and indigena are 2.2–2.6%, whereas
lilith and glaux differ by 1.1% only. However, this group differs from European Little
Owls by 6.8–7.6%.
Genetic distances of the cytochrome b gene above 1.5–2% have been regarded as
indicative of species level in owls (Wink and Heidrich 1999) and show that a gene flow
has not occurred between sister taxa within the last one million years (assuming that
2% distance equals one million year divergence; Wilson et al. 1987, Tarr and Fleischer
1993). Within defined populations, genetic distances are usually below 0.5% (as also
found by Wink In: Van Nieuwenhuyse et al. 2008). According to this argumentation,
noctua, vidalii, indigena, lilith and plumipes represent not only well-recognized subspe-
cies, but might even be considered as distinct species under the phylogenetic species
concept (Wink 2008).
The present data could indicate that Little Owls represent a species complex, similar
to the situation of the Barn Owl (Tyto alba) (Wink and Sauer-Gürth 2021). Also in the
Barn Owl, many geographically defined subtypes have been recognized, of which many
have attained species status due to morphological and genetic differences (König et al.
1999).
The morphological arguments by Lei et al. (1997) for considering A. n. impasta as a
subspecies were genetically confirmed by Qu et al. (2002) using PCR to study orientalis,
plumipes and impasta, with three repeat sequence primers and five random amplified
primers over 70 polymorphic DNA bands that showed differences in intraspecific individ-
uals. The cluster analysis separated seven intraspecific individuals of Athene noctua into
three branches, which followed the three recognized subspecies. The results confirmed
A. n. impasta and A. n. plumipes as two distinctive subspecies rather than synonymous.
Based on mitochondrial DNA research on 326 samples from 22 locations, Pellegrino
et al. (2014) found that the Little Owls in Europe belong to two different monophyletic
clades (group with a common ancestor): a western haplogroup (female parentage
group) in western and northwestern Europe, and an eastern haplogroup in central
and southeastern Europe. The eastern group is divided into three subgroups: Balkans,
southern Italy and Sardinia. Successive ice ages caused the European range of the Little
Owl in prehistoric times to be sometimes larger, sometimes smaller. Re-colonization

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 3.8 Vocalizations

probably took place several times from three refugia (Iberian Peninsula, Italy and the
Balkans) as much of Europe had become unsuitable habitat during the preceding Ice
Age. A natural barrier in the shape of the Alps not only created a western line and an
eastern line, but also made it difficult for the eastern line to advance to the north. As a
result, the western line was able to penetrate into central Europe and make contact with
the eastern group in northern Italy and Hungary (Pellegrino et al. 2015). In contrast to
Wink (2008), Pellegrino et al. (2015) found samples from Cyprus to group in the Balkan
cluster, suggesting indigena as the subspecies rather than lillith.
Pellegrino et al. (2015) state that although their genetic findings are in line with the
four subspecies described for Europe (vidalii, noctua, indigena and sarda), the similarity
between genetic and morphological patterns is often weak. Future studies considering
the whole distributional range of the species and both mtDNA and other molecular
markers (e.g., microsatellite DNA) should be able to clarify the distribution boundaries
at the subspecies level. Pellegrino et al. (2015) showed a strong genetic structure and no
evidence of genetic depletion in European Little Owl populations. The genetic insights
are crucial to re-stocking programs for endangered populations. The strong genetic
structure and diversity of European Little Owls suggest that their management units
should encompass areas of limited extension, and prospective population
supplementation should be set up from nearby stable populations. Re-introduction
programs based on birds bred in captivity should be based upon birds of local origin.

3.8 Vocalizations
Natural selection appears to have favored monomorphic plumage coloration for most
Strigidae (Holt et al. 1999). Hence, Magnus Robb and The Sound Approach (2015) give
more importance to vocalizations as taxonomic arguments and use the wide mitochon-
drial cytochrome c oxidase I (COI)-divergence between the western (vidalii) and eastern
(noctua, indigena, sarda) clades found by Pellegrino et al. (2014) as supportive evidence
for splitting Athene noctua vidalii from the other subspecies into Athene vidalii based
upon a distinct vocal difference in the “aggressive song.” They distinguish two separate
species based upon the aggressive song (“the most important long-distance signals”):
Athene vidalii in Western Europe and an Athene noctua group named Cucumiau in
southeastern Europe and northern Africa. Vidalii produces an aggressive song that
consists of a single type of sound: MIAU, MIAW, MIAU while this call by all noctua
sampled owls is preceded by one or two low chugging sounds: cucuMIAU, cucuMIAU,
cucuMIAU etc. The name Cucumiau is used in many Mediterranean languages,
including Turkish: kukumav, Bulgarian: kukumyavka, Greek: koukouvaya and
Serbian: kukumayka. However, sampling might be biased since our study of all Little
Owl recordings from xeno-canto.org yielded similar vocalizations for vidalii too
(See Table 4.9).
The distribution map of the two suggested Athene species by vocalization only
seems purely based upon the insights from Pellegrino et al. (2014), despite the fact that
these authors only analyzed genetic samples from the four European subspecies and

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 Chapter 3: Fossil Evidence, Taxonomy and Genetics

promoted further studies considering the whole distributional range of the species, and
both mitochondrial DNA and other molecular markers.
The limited sample sizes of recordings from Portugal, Spain, Bulgaria, Turkey, Oman
and Morocco, however, remain anecdotal and lacking statistical grounds to split the
vidalii subspecies into a separate species. Our conservative opinion on this split is
furthermore supported by a warning by Pellegrino et al. (2014) that the divergence
time estimates based upon the traditional 2% rate should be not be applied due to large
variations across lineages and loci but rather replaced by the more reliable fourfold
neutral rate. Hence, so far there is not enough evidence for splitting vidalii and noctua
into two separate species.

3.9 Taxonomic Discussions

Besides using vocalizations for taxonomic purposes suggesting Athene vidalii and Athene
noctua by Robb and The Sound Approach (2015), König et al. (2008) suggest keeping
both subspecies, but splitting Athene lilith and Athene spilogastra (with subspecies spilo-
gastra and somaliensis). Despite the fact that König et al. (2008) indicate that studies of
ecology, vocalizations, behavior and genetics are required to clarify the taxonomy of
the A. noctua complex, which we highly support, they split these two subspecies into
species, which we do not support. The isolated geography of spilogastra is the only
argument that might suggest a further analysis to study a split. However, the excep-
tional distribution of lilith by König et al. (2008) in the Middle East bordered by glaux in
northern Africa and indigena in Turkey and bacteriana in Afghanistan appears to be
artificial and against biogeographical logic of a clinal species like Athene noctua. Hence,
so far there is not enough evidence for splitting lilith and spilogastra into two
separate species.

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 Chapter 4: Morphology and
Body Characteristics

Chapter Summary
This chapter gives an overview of the embryonic development and morphological
characteristics of the Little Owl. We first look at how the egg develops, with special
attention on the temporary asymmetric ears, then zoom in on owlet development as
they grow and the plumage of adult birds and how molt takes place. The eyes are special
and the species has retinal cells similar to diurnal birds of prey. While Little Owls can
differentiate several colors, the species does not see infrared light. It has an auditory
sensitivity to locate small rodents with an accuracy of up to 1%. The bill is yellowish
and its color functions as a signal for the fitness of both juveniles and adults. The Little
Owl has differential biometrical measures (such as length of wing, tail, tarsus) according
to the subspecies or according to the sex (e.g., weight – females are heavier close to the
breeding time). We finally examine the voice. The species has a large vocal repertoire
including 40 acoustic signals and combinations with regional specificities. The chapter
concludes with specific characteristics of flight and anatomy.

4.1 Embryonic Development

The developmental stages of Little Owl embryos from 23 to 40 were described by Krings
et al. (2020). They found a correlation between the weight of the eggs (normalized by
volume) and the developmental stage. Eggs containing embryos close to hatching were
significantly lighter than eggs containing embryos at an earlier stage of development
(Pearson correlation coefficient r = –0.441, p = 0.026, t-test, n = 20). The average weight
of the Little Owl eggs was 12.04 g (n = 76); their average size was 34.89
29.12 mm (n = 20).
Many of the embryos were classified as stage 40 which is the last stage before external
pipping (where the eggshell is pecked from within). Collectors reported dead stage
40 embryos in many unhatched eggs of bigger clutches with successfully hatched siblings.
The forebrain of the Little Owl grows disproportionately from stage 28 on. The eyes
are oriented laterally up to stage 32. At stage 33, the lens of the eye shows a forward
orientation. The eyelids start to cover the eye at stage 34. Conjunctival papillae and
scleral ossicles are hard to detect in the embryos.

Supplementary resources are available online at www.cambridge.org/littleowl. These include

appendices as well as tables which previously appeared in the first edition (prefixed with ‘S’).

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 Chapter 4: Morphology and Body Characteristics

The Little Owl shows a pronounced cere (unfeathered skin surrounding the nostrils).
Due to this swollen cere, the line connecting the neurocranium and beak is almost straight
(Figure 4.1). The rhamphotheca develops at stages 36/37 and starts to turn gray at stage 40.
The claws on the toes of the Little Owl appear at stage 35 and start to turn white at stage 38.
The claws are completely white at stage 39 and have even turned gray by stage 40.
The first feather germs appear at stage 34. The plumage is long and dense from stage
38 on. The secondary wing feathers measure more than 7 mm at stage 40.
The weight and limb length of embryos increase exponentially, while head length
and width, as well as beak length, increase linearly. The maximum limb length of Little
Owl embryos that died at stage 40 is 22.8 mm. The average weight of the Little Owl at
stage 40 is 7.7 g.
Little Owl embryos at advanced developmental stages are more densely feathered
and the feathers are longer than in Tengmalm’s Owl or Barn Owl (Köppl et al. 2005).
Mebs and Scherzinger (2000) described the neoptil of Little Owl hatchlings as plushy
and dense. Barn Owl and Tengmalm’s Owl hatchlings show several unfeathered areas
(Mebs and Scherzinger 2000).
Glutz von Blotzheim and Bauer (1980) report that the hen of the Little Owl inten-
sively warms the hatchlings under her plumage just for a week, while intensive
warming in Tengmalm’s Owl lasts at least 2 weeks and may even be partly done by
the male. Despite slight differences in warming, nestling time and clutch size of Little
and Tengmalm’s Owls are similar (Glutz von Blotzheim and Bauer 1980, Mebs and
Scherzinger 2000). The differences in the neoptil between these two species can be
explained by their breeding strategies and cavities: tree crevices are easier to keep warm
and dry (Mebs and Scherzinger 2000) than rocky crevices and rodent burrows. Hence,
the chicks of Little Owls are adapted to the adverse conditions of open breeding sites
(Mebs and Scherzinger 2000).
Barn Owl embryos hatch in a less developed state compared to Little Owls with
respect to size and weight (Köppl et al. 2005). In the more advanced Little Owl,
pigmentation of the claws is accelerated, the claws even turned gray before hatching.
In the Barn Owl, feather germs appear at stage 32, whereas in the Little Owl appearance
of the feather germs is slightly postponed until stage 34 when plumage development is
accelerated, leading to densely feathered hatchlings. Hatchlings of the Barn Owl and
Tengmalm’s Owl were barely covered with down.
Krings et al. (2018) showed that a small, but significant ear asymmetry occurs in the
embryonic development of Little Owls, despite the presence of symmetrical ears in adults.
Asymmetries were not observed in adult Athene noctua. In European birds from the
subspecies A. n. vidalii, the left ear measured 37.06 mm² while the right ear measured
37.61 mm2 (t = 0.332, n = 7, SD = 5.33 mm2). Three specimens had a bigger left ear
opening, four specimens showed a bigger right ear opening. Specimens from northern
Africa belonging to the subspecies A. n. glaux were symmetrical too (left ear: 35.95 mm²,
right ear: 35.96 mm², t = 0.006, n = 17, SD = 7.54 mm2).
In the stages 35–40 they observed a dorsal movement of the ear openings. The move-
ment occurred in parallel on the left and right side of the head. They did not observe an
asymmetry in the positions of the ears in any stage. General growth of the head was
indicated by all marker points, including the ear openings, moving away from the center.

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eggtooth

St. 23 St. 25 St. 27 St. 29 St. 31 St. 35

eyelid closing ramphotheca swollen cere claws white

St. 36 St. 37 St. 38 St. 39 St. 40

feathers growing 1 cm

Figure 4.1 Embryonic development of the Little Owl (after Krings et al. 2020).
 Chapter 4: Morphology and Body Characteristics

In the asymmetric period, the left ear opening is bigger in area than its counterpart on the
right. Clear asymmetry was found at stage 37; the left ear opening averaged 3.36 mm2 and
the right ear opening averaged 2.87 mm2 (n = 3). Afterwards asymmetry increased substan-
tially at stage 38 (left ear: 4.25 mm²; right ear: 3.23 mm²; n = 13). At stage 39 less asymmetry
was found (left ear: 4.27 mm²; right ear: 3.91 mm²; n = 13). At stage 40 the difference in the
size of the ears was small again (left ear: 2.80 mm²; right ear: 3.09 mm²; n = 3).
The asymmetry in the Little Owl occurs at the same stages at which the asymmetry
in the American Barn Owl (Tyto furcata) develops, but in the former the asymmetry
vanishes shortly before hatching. The disapperance of the asymmetry is interpreted as
an indication of a secondarily evolved diurnal activity in Little Owls.
Krings et al. (2018) considered it likely that the symmetrical development of ear
position in Athene was due to growth rather than migration of the ear openings.

4.2 Owlet Development

Little Owl nestlings display a rather systematic evolution in weight and wing length
according to their age in days while in the nest (day 10–25), allowing the age to be
estimated (Juillard 1984, Stroeken and van Harxen, 2003, 2019) and the relative condition
of each nestling to be determined (Stroeken et al. 2010, Hallmann et al. 2011). It is possible
to determine the age of the juveniles by measuring their growing primaries (Juillard 1979).

4.2.1 Age Determination

Stroeken and van Harxen (2003) calibrated a linear growth model to calculate the age
based upon the wing length of the largest nestling of 19 nests sampled in 1999 and 2000.
They observed a linear relationship between age in days and wing length between days
10 and 25, assuming that the largest nestling was representative for the age of all nestlings
as hatching of all eggs largely took place within 15 hours, exceptionally up to 24–26 hours
(van Harxen et al. 2018). The largest nestling was assumed to have had the best food
conditions, removing the variance of insufficient delivery of food to the nonrepresentative
smaller nestlings (Groen et al. 2001). The linear model of age versus wing length of
the largest nestling yielded a regression model of y = 5.1121
age in days – 16.039, which
allowed prediction of the age using the wing length of largest nestling (age = (wing length +
16.039)/5.1121). The model was re-calibrated by 68 largest nestlings, of which the exact
hatching data were observed as fresh hatchlings that were still wet, noisy eggs about to
hatch, broods with hatchlings all weighing 10–12 g or broods that were monitored by
video or webcam observed from 2002 to 2018 (n = 950 broods) (Stroeken and van Harxen
2019). The linear regression yielded a daily growth of 5.2515 mm (y = 5.2516
age
in days – 18.321; age = (wing length + 18.321)/5.2516). To study whether there was a
gender effect in the largest nestling, the sex of 253 largest nestlings was determined yielding
50.3% females and 49.7% males, indicating no predominance of either sex as largest
nestling. Both linear models yielded very similar results.
Table 4.1 shows the expected wing length per day for both Stroeken and van Harxen
(2003, 2019).

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 4.2 Owlet Development

Table 4.1a Expected wing length per day of age of Little Owl largest nestling per nest
Stroeken and van Harxen
(2003) Stroeken and van Harxen (2019)

Wing length
Predicted Predicted
Age Observations wing length Observations wing length Mean Median Min Max

10 ? 33–37 0 32–36
11 ? 38–42 0 37–42
12 0 43–47 0 43–47
13 ? 48–52 5 48–52 51 51 48 54
14 ? 53–57 31 53–57 55 55 44 66
15 ? 58–62 0 58–63
16 ? 63–68 1 64–68 65 65 65 65
17 ? 69–73 1 69–73 60 60 60 60
18 ? 74–78 1 74–78 75 75 75 75
19 0 79–83 2 79–84 86 86 81 95
20 ? 84–88 5 85–89 87 86 84 102
21 ? 89–93 14 90–94 94 94 88 102
22 ? 94–98 4 95–99 95 96 90 98
23 ? 99–103 0 100–105
24 ? 104–108 2 106–110 101 101 98 104
25 ? 109–114 2 111–115 114 114 112 116

Table 4.1b Expected weight per day of age of Little Owl largest nestling per nest
Age Expected weight Age Expected weight Age Expected weight

0 10.1 15 113.9 30 146.6

1 14.7 16 118.5 31 147.2
2 20.4 17 122.6 32 147.7
3 27 18 126.2 33 148.2
4 34.3 19 129.4 34 148.5
5 42.2 20 132.2 35 148.9
6 50.5 21 134.7 36 149.2
7 58.9 22 136.9 37 149.4
8 67.2 23 138.8 38 149.6
9 75.2 24 140.4 39 149.8
10 82.9 25 141.8 40 150
11 90.2 26 143.1 41 150.1
12 96.9 27 144.1 42 150.2
13 103.1 28 145.1 43 150.3
14 108.8 29 145.9 44 150.4

Stroeken and van Harxen 2003, 2019

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 Chapter 4: Morphology and Body Characteristics

4.2.2 Relative Nestling Condition

Weight is often used as a measure of fitness in birds. An index for the condition of a bird
can be expressed as ratio:
Condition ¼ ðobserved weight=expected weightÞ

where the expected weight is estimated from a series of weight measurements or

resulting from statistical models. A fitness index value >1 indicates that the individual
has above-average fitness, while a fitness index value <1 indicates that the individual
has below average condition. Weight development is S-shaped in relation to age
(Figure 4.2a), and in relation to wing length is a flattening, saturating, nonlinear curve
(Figure 4.2b). This implies that in order to arrive at a meaningful fitness index, the age
at which the young are weighed, or by the wing length at the time of weighing must be
taken into account. As a result, the condition is expressed as:
Condition ¼ ðobserved weight at age x=expected weight at age x

or
Condition ¼ ðobserved weight for wing length x=expected weight for wing lengthÞ

To calculate a fitness index it is therefore necessary to enter an age-specific or wing

length-specific expected weight.
Hallmann et al. (2011) collected 2049 nestling weight measurements of 295 broods
found in 94 territories, excluding 115 measurements of nestlings that did not fledge,
from 2002 to 2010 in order to cover both good and bad years. For 1140 measurements,
the wing length of hatchlings was also measured at 10 days old. This large dataset
served as the basis for the calibration of a sigmoid Gompertz-curve (Gompertz 1825),
which resulted in a standardized weight–growth curve for Little Owl nestlings based
upon the age of the nestlings in days. A von Bertalanffy model (von Bertalanffy 1934)
performed best to predict weight based upon the wing length.
The weight and wing length measurements of nestlings cannot be considered to be
independent of each other. After all, there are multiple measurements per individual,
multiple individuals per brood and multiple broods per territory. It can be expected that
there is a correlation in repeated weight measurements within the same brood (sibling
owlets share basically the same microclimate and the same prey supply) or within
broods in the same neighborhood. Because of such effects, the best model is refined
with random effects, which correct for this dependence, and thus it is essentially a
mixed-effects nonlinear model, making the estimated weight more representative of
the population.
Due to the fact that all nestlings are born on the same day and to the fact that
measuring wing length in owlets younger than 8 days is error-prone, the weight
approach is preferred (P. Stroeken personal communication).
Hallmann et al. (2011) also examined whether brood size and breeding season (year)
have an effect on the estimated parameters and thus the growth curve of young Little
Owls. Brood size can be expected to influence the growth of young Little Owls, e.g.,

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 4.2 Owlet Development

(a)

200
Asymptote = 150.94 grams
150
Weight (grams)
100
50

0 10 20 30 40
Age (days)

(b)
200
180
160

Asymptote = 146.11 grams

Weight (grams)
140
120
100
80
60

20 40 60 80 100 120 140 160

Wing length (mm)

Figure 4.2. Fitted weight gain profile of Little Owl (Athene noctua) nestlings based upon the age and
the wing length (Hallmann et al. 2011): (a) age = Gompertz curve, (b) wing length = von
Bertalanffy curve.

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 Chapter 4: Morphology and Body Characteristics

limiting available food per young. Annual variations in weather conditions and/or food
availability can also have an effect on the weight of nestlings.
The Gompertz model was found to be the best fit for the weight–age ratio. The
weight to wing length ratio was best approximated by the von Bertalanffy model.
The Gompertz model approximated the growth curve better than a logistic model.
This has, in addition to a practical consequence, a biological interpretation. The logistic
model assumes that the inflection point of the curve occurs at half (50%) of the
asymptote, while the Gompertz model shows this inflection point at 36.7% of the
asymptote. For Little Owls this implies that maximum growth is reached in the first
half of the growth phase. Similar findings have also been made in other species,
suggesting a trade-off between number of nestlings and their condition (Hotchachka
and Smith 1991). There is no significant effect of the year factor on the estimated
parameters of the Gompertz model, suggesting that the growth of young Little Owls
is not affected by annual variations in food and weather conditions, or that food or
climate conditions differ more between territories than between years. It seems that
parent pairs successfully cope with variation in annual food or climate conditions.
However, the relation between weight and wing length is less accurate, especially in
the early stages, implying less strong conclusions can be drawn with regard to the
condition determination of the Little Owls. In addition, the relationship between
weight and wing length is only valid for a wing length measurement from 20 mm,
i.e., only from an age of 8–9 days, which is the basis for using these results for condition
determination for very young Little Owls. The age model should be preferred when
determining the relative condition of Little Owl nestlings. The weight gain of Little Owl
nestlings using age in days is shown in Figure 4.2a.

4.2.3 Expected Weight Based on Age

The expected weight for the given age is shown in Figure 4.2a. Pursuant to the data
pertaining to pulli of 0–44 days, the weight asymptote or the weight represented by the
saturated or flattening curve has been estimated to be 151 g. The maximum growth rate
has been estimated at 8.38 g per day, with the inflection point at an age of 6.6 days.
Brood size has a clear negative effect on the asymptote parameter (LRT test = 34.08,
degrees of freedom = 1644, p < 0.0001). The relationship between brood size and mean
expected weight can be expressed based on the current model as: asymptote = 168.104 –
5.0183
brood size. The nest average asymptote thus decreases by about 5 g per young,
which may generally imply that from larger broods, lighter young fledge, or that the
owlets from larger broods take longer to reach their asymptote. Fitting in the factor of
years as a predictor of the asymptote does not lead to a significant improvement of
the model.

4.2.4 Expected Weight Based on Wing Length

The expected weight for given age is shown in Figure 4.2b. The asymptote or saturation
point according to this model is estimated at 146 g, which is clearly lower than

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 4.2 Owlet Development

according to the age model. The lower number of weight measurements that may have
been paired with wing length measurements for later ages (>30 days) caused the
difference between the estimates from the two models. Because the model for wing
length measurements <30 mm, the relationship between weight and wing length is less
accurately described (too high) and condition index numbers based on young chicks
should be applied sparingly.
From the curve, expected or reference values can be deduced for a nestling’s weight
in relation to its age, offering a standardized method for determining the relative
physical condition of nestlings. Dividing the measured weights by the reference value
yields the condition index, which expresses the relative condition of nestlings as a
possible approximation of the relative habitat quality.
Chick development is illustrated in Plate 21, feather development in Figure 4.3.

4.2.5 Days 1–5

When the owls hatch they are covered in a short white down, their eyes are closed and
they rely on their mother and each other for warmth in a “warmth pyramid” (Schönn
et al. 1991). The average weight is 10 g (Ataev 1977), 11 g (Bakaev in Sagitov 1990) or
12 g (Juillard 1979). The diamond-shaped egg tooth on the bill to break the egg is visible
until days 13–15. The owlets stay on their abdomen or breast with their angular legs
under or near the body; the head is supported by the bill or the chin, the wings are
feeble. When they stand on their talons, they can trot a few steps (Schönn et al. 1991).

4.2.6 Week 1
The down on the owlets changes from a white color to gray and has a more fluffy
texture; the primary feather pins are just beginning to show; the legs, feet and talons are
developing. Their cries sound like whistled begging (“chsij”) or a shrill defense (“uin,”
“wuin”) (Schönn et al. 1991). Between days 1 and 10, the weight increases from 12–75 g.
In captivity, they weigh between 20 and 45 g at 5 days, between 60 and 90 g at 10 days
and up to 150 g between days 20 and 25 (Génot and Sturm 2001). Festetics (1952) noted
the following weight evolution: 20 g (day 1), 56 g (day 5), 112 g (day 10), 120 g (day 15)
and 120 g (day 20).

4.2.7 Week 2
The eyes usually open between days 8 and 10 (Schönn et al. 1991), but sometimes this
can occur at days 6 or 7 (Bakaev in Sagitov 1990). The owlets develop the ability to tear
prey provided for them. The down has become more sparse, and has changed color to
light gray. They are now able to regulate their own temperature, allowing both parents
to forage for food. Some sibling skirmishes begin to occur at this age with the oldest
becoming the most dominant in the nest, therefore getting priority over the other
siblings when parents bring in food. Short facial feathers begin to develop, the beak
seems to have grown faster than other parts of the body and dominates the face (see bill

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 Chapter 4: Morphology and Body Characteristics

Figure 4.3 Feather development at day 14, day 18, day 23 and day 29.

coloration and brightness as informative chromatic signal for parents revealing aspects
of quality of nestlings later). The legs and feet have grown faster and proportionally
larger than the other body parts. The owlets can move forwards and backwards on their
talons with the wings stretching out to balance. They can also climb. They sleep lying
or squatting by lowering one wing or leaning against a wall (Schönn et al. 1991). At
days 10–12 the wing length of the largest nestling is 33–42 mm. At days 12–14 the
primaries start to grow as blue-gray shafts without plumes, wing length 43.52 mm, at
days 14–21 the plumes leave the shafts, wing length 53–88 mm. At day 14 the owlets
begin to be able to focus their eyes on something. The main feathers of the wing and
tail appear at 9 days and 12–16 days respectively.

4.2.8 Weeks 3–5

An accelerated transformation occurs between weeks 2 and 3, with the head, breast,
flanks and nape feathers all developing in this week. The eyes are clearer, the legs and
feet have lost the fatty appearance, the talons have grown in line with the bone
structure and the face pattern is now characteristic of the Little Owl. The owlets are
alone in the nest site because the female left them at day 16. The behavior has also
developed, the owls show more mobility in the nest chamber than before, sometimes
running, and therefore predator evasion is greater. It is the beginning of handling prey,
which they can catch at day 28 and are able to kill at day 34 (Ille 1983). At days 21–28
the wings of the largest nestling measure 89–116 mm, the shafts are mainly blue-gray
and the tail feathers start growing; at days 28–30 the wing length is 117–126 mm, the

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 4.3 Plumage

tips of the feathers are pale, the tail is clearly visible and nestlings are ready to fledge.
The owlets strengthen their begging behavior, getting up to their legs (Schönn et al.
1991). Within the nest the sibling rivalry tends to be dominated by the oldest owlet.
The owlets will become more adventurous, and will climb out of the nest onto the tree
trunk, climbing up and along the branches using their bill. This provides good exercise,
and again will minimize the effect of predators on the young in the nest. The third
primary feather (P8) measures on average 75 mm (Juillard 1979).

4.3 Plumage
This description applies to the best-studied subspecies, Athene noctua vidalii.

4.3.1 Juveniles/First Year Adults

The first down (neoptile) of the young is short, dense and white. It becomes mottled
gray after one week, when the mesoptile starts to grow. The second down (mesoptile) of
the juvenile is rather feather-like. The feathers are still distinctly softer and shorter than
in first year and adult owls. The structure of the flight feathers, tail and greater primary
coverts are as in the adult. The neoptile clings to the tips of the mesoptile for up to 3–4
weeks (retained longest on crown, flanks and thighs). The mesoptile is similar to the
adult, but with a paler base color and more grayish. The crown, back, rump and upper
tail coverts are almost uniform gray-brown or with faint buff spots. The face is the same
as in the adult, but grayer and less contrastingly marked.

4.3.2 Adult
The main color of the upper parts and upper wings of the adult is dark chocolate-brown,
usually with a slight olive-brown tinge. The forehead and crown are closely marked
with cream-buff streaks, each streak often widens lightly towards the feather-tips, and
the crown shows elongated spots. A distinct white V-mark (Figure 4.4) is noticeable on
the hind neck from behind the ear down to the center of the neck in a “face-like”
pattern (Scherzinger 1986).
The mantle and scapulas are covered with large white or pale cream rounded
subterminal spots, often joined to form a single wide subterminal bar. The back, rump
and upper tail have large rounded subterminal spots of cream-buff. The facial disc is
poorly developed, and is not surrounded by a distinct ruff. The ring round the eye is
white and is widest between the eye and the base of the bill.
The chin is white, bordered below by a buff-and-gray spotty band across the throat to
the sides of the neck. The upper chest is also white and the lower chest, the breast and
the upper flanks are dark fuscous-brown to olive-brown. Each feather presents a paired
subterminal pink-buff or white spot or short bar. The base color of the tail is similar to
or slightly paler olive than the upper parts of the owl; the terminal bars or spots on the
tail can be lighter due to bleaching from exposure to sunlight.

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Figure 4.4 Distinct white V-mark on hind neck of the Little Owl.

The pattern of the pale bars on the tail is somewhat variable. Starting at the base of
the tail feather and working towards the tip: one straight bar is just visible on the tail
feathers at the tips of the longest upper tail coverts. A similar second and third (and
sometimes fourth or fifth) bar are found across the middle part of the tail. At the feather
tips, traces of a terminal bar are often present in the form of the a spot, sometimes
contiguous with a white border. Thus, while there are usually four, sometimes four bars
and traces of a fifth bar are visible across the tail feathers. Pellegrino et al. (2020)
developed a classification of tail bar patterns (Figure 4.5) that was used in Chapter 3
on the subspecies to study regional differences.

Figure 4.5 Classification of the tail patterns of Little Owls after Pellegrino et al. (2020).

The flight feathers are dark olive-brown or fuscous-brown, darkest along the shafts and
outer primaries. These primaries have three to four well-spaced pale spots along the outer
webs, and much larger spots or short bars are seen on the inner webs. The secondaries
are similar to the primaries, but have only two to three rows of spots or bars; the tips of

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 4.3 Plumage

Figure 4.6 Box and whisker plots of the distribution of male and female head height versus width
ratios (n = 20 males, n = 20 females).

the secondaries have large or small, white, paired spots; sometimes these spots or bars
are completely absent. The upper wing coverts, like the mantle and scapulars, have white
spots smaller than those that are found on most of the primary coverts, and are often
restricted to one web only. The under-wing coverts and axillaries are cream-white, slightly
spotted and mottled gray. The sexes are similar, but the male tends to have a whiter face
and a rounder head. A t-test analysis of the oval shape of random 20 adult males and
20 females (van Harxen and Stroeken personal communication) yielded a significant
difference in height versus width ratios, with males having rounder heads (mean ratio male
heads = 0.819, mean ratio female heads = 0.764, t-value = 4.085, DF = 38, p = 0.000219);
Figure 4.6 shows two box and whisker plots of the distributions of both samples.
Figure 4.7 illustrates the clear difference in head shape for males (rounder) and
females (flatter). Figure 4.8 illustrates some further variation in head shape depending
on the mood and the sex of the bird.
Primaries are quite long and not very wide. The longest is P7 at up to 143 mm.
Rectrices are relatively short. Where whole sets of rectrices are considered, any except
the members of the central pair may be found to be longest. Hansen et al. (1988) found
the longest rectrices ranged from 77–95 mm (n = 96). The fingers on the outer primaries
are quite long. There is shallow emargination of both vanes on the five outer primaries,
though this is only marginal on P6. The inner emargination angle is shallow and
slightly dilated. Secondaries and rectrices are quite broad. The transitional secondary

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 Chapter 4: Morphology and Body Characteristics

Figure 4.7 Male and female heads with rounder heads for males, flatter for females.

is S12 or S13. The oval shape of the shafts of the primaries is clearly perceptible as
feathers are turned between a person’s fingers. Juvenile remiges are only slightly
narrower than those of adults, thus precluding confident identification of an age
category on the basis of single feathers (Cieślak 2017).
The first year adult is very similar to the adult (J. Bultot personal communication,
Etienne 2012, Cieślak 2017). They retain juvenile flight feathers and tail into adult-
hood, with the pattern variable at all ages. The main differences include:

• the shape of the tips of the primaries – rather pointed in first year; tips are almost
square in adult owls (Figure 4.9). The size of the spots on the tips of the primaries are
larger in first year birds than in adults.

• the plumage of the adult is fresh until the onset of the breeding season; the flight
feathers, tail and in particular the tertials are slightly worn in the first autumn,
distinctly worn in spring and heavily worn in summer.

• the texture and shape of the tertials is rather soft, narrow and tapered towards the tip
in the first year bird; they are broader and the tips are squarer in the full adult.

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 4.3 Plumage

Figure 4.8 Diversity of different head shapes of Little Owls.

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Figure 4.9 Difference in the shape of the tips of the primaries in (a) first year and (b) adult Little Owls.

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 4.4 Molt

Adult and first year owls have a lemon-yellow iris. The eyelids are a dark slate-blue.
The bill is lemon-yellow or greenish-yellow. The cere is slate-gray or olive-black. The
tarsus is grayish or grayish-yellow and the toes are gray-brown or olive-black. The claws
are dark brown to brownish-black. At hatching, the skin under the down of the young,
the cere, and the toes are pink; the bill and claws are whitish- or grayish-pink. The eyes
open around days 8–10 and the iris is pale yellow at first. The skin partly darkens
to gray after a few days, beginning on the toes and the cere. At fledging, the iris and
eyelids are the same as the adult. The bill is olive-yellow with the tip of the culmen a
pale yellow. The cere becomes slate-blue or blackish violet-green. The tarsus is yellow
or yellow-gray, the toes flesh-gray or dusky gray and claws black with a gray-blue base.
Some anomalies can occur in the plumage, as in a case of partial albinism (Bisseling
1933), (partial) leucism (Brinzal personal communication, Framis et al. 2020) or a russet
form (Paris 1909). König et al. (2008) mention the occurrence of gray and rufous
morphs (see Chapter 3 on Taxonomy).

4.4 Molt
The molt of adults starts after breeding, at the end of June or beginning of July, when
the young fledge or are independent (Haverschmidt 1946, Dementiev and Gladkov
1951, Stresemann and Stresemann 1966). However, late breeders, especially those with
a replacement clutch, may start molting at about the time of hatching (Ullrich 1970),
although some breeders can be found without any molt (Exo in Schönn et al. 1991).
The onset date of the molt depends on the age of the owls and is earlier in older birds
(Piechocki 1968, Hartmann-Müller 1973, 1974).
To illustrate the molt we will first illustrate the topology of a Little Owl wing
(Figure 4.10) and present the different stages of the molt according to age
(Figures 4.11–4.17).
The molt begins with the loss of the first primary (P1) in mid-May to mid-July, and is
completed with P10 in early September to early November (Stresemann and
Stresemann 1966, Piechocki 1968). A different view of the molt sequence was offered
by Dementiev and Gladkov (1951) who considered that replacement of the primaries
occurs centrifugally. There is little geographical variation in the onset of the molt, but
northern birds starting from mid-May may be nonbreeders and eastern birds start
earlier, as in Turkmenistan (Dementiev and Gladkov 1951). It is probably due to a
breeding cycle that is more extended in the year. Normally, the complete molt starts
after P1 is shed (Glutz Von Blotzheim and Bauer 1980) but in other cases after a
secondary or tail feather is shed (Exo in Schönn et al. 1991).
In two captive owls, P10 was shed 98–99 days after P1 (Piechocki 1968) and three
months for a male in Turkmenistan (Bel’skaya 1992).
The secondaries (S) molt from three centers: S10 starts with the shedding of P3–P4,
S5 starts with P6–P7, and S1 starts with P5–P7 (Cramp 1985). S4, S7 and S8 molt last,
shed at about the same time as P10 (Piechocki 1968). Most secondaries are replaced
during the last stages of the primary molt (early August to mid-October).

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Figure 4.10 Topology of wing and tail of the Little Owl. Primaries are counted from the middle of the wing outwards to the tip and the secondaries from the
middle of the wing inward to the body. When the wing length is measured, the wing has to be closed and the distance taken from the wrist or bend to the longest
primary feather. The tail length is the distance from the base of the central tail feather to the tip. Tail feathers are counted from the center outwards. Each feather has
a shaft and an outer and inner web.
 4.4 Molt

Figure 4.11 Comparison of the juvenile and neighboring remiges of adult plumage of the same
individual Little Owl.

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Figure 4.12 Juvenile (a) and adult (b) primaries.

Both sexes undergo a complete postbreeding molt, normally the male molts before
the female (Martinez et al. 2002). The primaries commence from P1 on to P10. The
secondaries are shed in three groups S1–S4, S5–S7 ascending, the third group of second-
aries are molted descending from S10–S8. Tail molt occurs simultaneously, usually
coinciding with the period when P4 and P5 are growing. Juvenile Little Owls tend to
undergo a partial body molt and will molt some wing coverts, normally between August
and November. The primaries are molted for the first time in the owl’s second summer.

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 4.4 Molt

Figure 4.13 Juvenile (a) and adult (b) inner secondaries.

The tail feathers are shed sometimes simultaneously, but in general within a few
days up to 2 weeks (Piechocki 1968, Exo in Schönn et al. 1991) with an irregular
sequence. The tail feathers are shed 20–30 days after P1 (Piechocki 1968) or when P4–
P5 are growing, and the molt is completed during the growth of P7 (Stresemann and
Stresemann 1966).
The scheme of molt is different according to sex among Little Owls in Turkmenistan
(Bel’skaya 1992). Complete replacement for females took seven months

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Figure 4.14 1Y spring (3). During partial molt, all greater coverts are still juvenile.

Figure 4.15 2Y spring (5). Contrast between molted 1–9 greater coverts and nonmolted inner greater
coverts and primary coverts.

(May–November) and eight months (April–November) for males. Replacement of small

feathers in females took four months (June–September) and three months (July–
September) for males. Replacement of the tail feathers in females took three months
(June–August) while it was two months (July–August) in males.

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 4.4 Molt

Figure 4.16 3Y+ spring (6). Molting. Two generations of adult feathers. S3–S6 not molted. Rest of
remiges molted.

Figure 4.17 3Y+ spring (6). All greater coverts are adult. There is no contrast between them and
primary coverts or remiges.

The body molt starts with the lesser and median upper wing coverts, soon followed
by the mantle and scapulars (Cramp 1985). The skin comes out in shreds in May–June
(Bel’skaya 1992).
For fledged juveniles the molt begins in July at the age of 70–75 days, when they are
separated from their parents (Bel’skaya 1992). The face feathers and lesser and median

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upper wing coverts are molted first (early July to mid-August) and this is largely
completed 6–7 weeks later (September or early October). The molt of the crown, neck
and feathering of the legs and toes occurs up to late November (Cramp 1985). The molt
is partial for head, body and wing coverts.
The general molt pattern in Spain is given by Martinez et al. (2002) (EURING codes
between square brackets):

1Y autumn (3)/2Y spring (5) [Bird in first calendar year to second calendar year]: Partial
molt affecting head, body and some wing coverts. Inner greater coverts retained.
Rather worn plumage before first molt of remiges, especially noticeable in tertials.
2Y+ autumn (4)/3Y+ spring (6) [Bird in second calendar year or older to third calendar
year or older]: Most birds undergo complete molt, but some retain a few secondaries.
Adult-like tertials. If some secondaries have been retained, the contrast between new
and old secondaries should be checked.
2Y autumn (5)/3Y spring (7) [Bird in second calendar year to third calendar year]: Strong
contrast between retained feathers (old juvenile) and the fresh ones.
3Y+ autumn (6)/4Y+ spring (8) [Bird in third calendar year or older to fourth calendar
year or older]: Less contrast.

4.5 Eyes
Owls have always fascinated mankind for their highly sensitive vision. Potier et al.
(2020) summarize what is presently known about the eyes as well as the visual abilities
of owls and raptors, and point out knowledge gaps. They discuss visual fields, eye
movements, accommodation, ocular media transmittance, spectral sensitivity, retinal
anatomy and what is known about visual pathways. The specific adaptations of owls to
dimlight vision include large corneal diameters compared to axial (and focal) length, a
rod-dominated retina and low spatial and temporal resolution of vision.
Little Owl eyes have been poorly studied compared to Tawny Owls (see references in
Potier et al. 2020). We give an overview of the most relevant adaptations of owls and
detail the aspects that have already been published for Little Owls.
Ille (1983) showed that the perception of prey is linked to sight. The big eye with a
large cornea has rods in the retina. The length and number of rods explain the good
sight of the species during twilight (Rochon-Duvigneaud 1934). Further investigations
revealed that the night vision of the owls is linked to the organization of the outer
plexiform layer of the retina, specifically to the stratification of the synoptic bodies of
the single and oblique cones. The adaptation to nocturnal light could be due to the lack
of oblique cones (Gallego et al. 1975). The neuronal cells of the Little Owl’s retina are
divided into 20 types of simple and complex synapses (Yew 1980). Yew speculates that
the contact between certain cells, in particular the amacrine-ganglion contact, are
related to vision in dim light. Morphological differences in the retina cells of the Barn
Owl (Tyto alba) and Little Owl show that the latter has retina cells more similar to
diurnal birds (Tarres et al. 1986). But the visual acuity of the Little Owl is the lowest in

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 4.5 Eyes

comparison with other birds of prey (Gaffney and Hodos 2003). Binocularity is an
advantage for the Little Owl (Porciatti et al. 1990), especially under low
light conditions.
The highest activity of the Little Owl in an aviary was recorded with a luminescence
of about 150 lux (Erkert 1967). But in nature, the main activity throughout the year
occurs when the light intensity is under 1 lux, and it is exceptional when an owl leaves
its day roost when there is a light intensity as much as 50 lux (Exo 1989).
In the Little Owl, the impact of scotopic vision (rods) in electroretinographic flicker
experiments is larger than the impact of photopic vision (cones) (Porciatti et al. 1989).
In this respect, the retinal function of the Little Owl resembles nocturnal rather than
diurnal animals (Porciatti et al. 1989).

4.5.1 Absolute Sensitivity

The low flicker fusion frequency of 50 Hz, as measured using electroretinogram record-
ings of Little Owls (Porciatti et al. 1989), together with their large corneal diameters,
long rod outer segments and rod-dominated retinae, indicate that their vision is opti-
mized for high sensitivity with sufficient acuity in dim light. Unfortunately, absolute
sensitivity has only been determined in the Tawny Owl, whose eyes are only 2.5 times
more sensitive than those of humans. Some species of owls that are diurnally active use
cone-based vision and can discriminate colors in such bright light intensities, as dem-
onstrated for the Little Owl (Meyknecht 1941). Through experiments, Little Owls can
see yellow, green, blue, red (although less well) and also differentiate gray colors
(Meyknecht 1941). Kopystynska (1962) showed that the Little Owl does not see infra-
red rays. The presence of a greater number of rods than cones in the retina indicates an
increased ability to discriminate shades of gray which would allow orientation under
low light levels at night (Kopystynska 1962).

4.5.2 Visual Wulst

The signals from photoreceptors are sent to the retinal ganglion cells via bipolar cells,
with horizontal and amacrine cells providing intra-retinal interactions (Potier et al.
2020). Birds have complete optic nerve crossing and retinal ganglion cells project to the
brain via the tectofugal pathway to the tectum opticum, the thalamofugal pathway to
thalamic nuclei and then the visual wulst, the putative homolog to the mammalian
primary visual cortex and the accessory optic system (Potier et al. 2020). Important
work has been published on the anatomical and functional organization of the visual
wulst in the Little Owl (Casini et al. 1988) and its role in the binocularity of the species
(Porciatti et al. 1989, Bagnoli et al. 1990, Porciatti et al. 1990, Casini et al. 1992). Cells
in the visual wulst represent the binocular visual field, but lack representation of the
central retina, including the central fovea. Soms cells in the visual wulst of the Little
Owl respond to sine wave gratings of contrast as low as 1% (Porciatti et al. 1989), which
is remarkable, as most birds, including the Barn Owl, do not respond to contrasts lower
than 10% in behavioral tests (Potier et al. 2020). The binocular representation of the

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Figure 4.18 Colobome as a malformation of Little Owl eyes.

frontal visual field might be related to the need for precise object fixation in the last
moments of prey capture (Güntürkün 2000).

4.5.3 Malformations
The eyes of the Little Owl have a diameter of 10 mm. The juvenile iris is white-yellow, at
hatching pale yellow (Glutz Von Blotzheim and Bauer 1980). The yellowness increases
at 20–35 days towards yellow lemon then sulfur yellow. Old owls can have the “chia-
radiae” aberration in the form of a dark brown iris (Kleinschmidt 1906). Schönn et al.
(1991) illustrate an example of a grainy marble-patterned iris.
Malformations in the Little Owl iris have been reported as tear-like and downward
iris lesions (colobome by Juillard 1981; Figure 4.18) and are observed regularly as a
consequence of accidents (Siehoff 2011). Van Harxen and Stroeken (personal commu-
nication) observed in 2017 out of 99 pairs, one female with loss of the right eye and one
male with a colobome, in 2018 out of 101 pairs, the same male with colobome was
paired with a female that had lost the right eye, in 2019 out of 100 pairs the same male
with a colobome and one with the right eye lost were observed.
Van Harxen and Stroeken (2005) documented a breeding pair in which both the
male and the female had only one good eye. The pair was together for four years and
bred every year with on average only 2.5 eggs/year without any young ever fledging.
This bad breeding efficiency is probably related to obvious problems with food supply
due to the loss of the eyes despite its availability. Barn Owls bred successfully in the
vicinity during the same period. Furthermore this pair changed nest site every year. The
female drowned at the age of 5 years, 3 months and 28 days.

4.5.4 Iris Yellowness

Among colored phenotypic traits, eyes remain largely overlooked, with only a few
studies suggesting a potential signaling function or a role in mate recognition and
crypsis. However, iris color is a candidate trait for pigment-based signaling in owls.

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 4.5 Eyes

The brightness of avian irises is due to structural elements (i.e., stromal purine crystals
in reflecting organelles or superficial blood vessels) as well as pigments (Ferris and
Bagnara 1972, Oliphant 1987, Oliphant and Hudon 1993). Chromatographic analyses
have revealed that pteridines (also called pterines) and purines are the most widely
distributed pigments in the irises of bird species with brightly colored eyes, although
carotenoids are also present in families such as Strigidae, Ardeidae and Anatidae
(Oliphant 1987).
Iris color is a remarkably striking feature in the wholly cryptic pattern of many owls,
and may potentially play a signaling function. Passarotto et al. (2020) studied variation
and potential signaling of iris yellowness as an indicator of quality in parent–offspring
communication and other social contexts in the Little Owl. Yellowness did not differ
between sexes in either of the species, however adults had more vividly yellow irises
than owlets. Owlet yellowness varied more between than within nests in Little Owls,
and the total variation in yellowness within nests was 17.0%, indicating that owlets
raised in the same nests tended to share similar levels of yellowness. Yellowness
variation was not related to owlet body mass within nests. Little Owl owlet yellowness
was not significantly related to clutch size, egg volume or nest success (percentage of
fledglings relative to number of hatched eggs). The average iris yellowness of Little Owl
owlets was higher in later broods, possibly because owlets in later nests received more
food deliveries per hour than those in earlier nests. A significant negative relationship
between owlet iris yellowness and breeding success (percentage of fledglings relative to
number of laid eggs) was found, indicating that owlets in nests with fewer fledglings
had high average yellowness. Yellowness variation was not related to adult condition
(i.e., body mass). Although the iris yellowness of adult Little Owl females was positively
associated with nest success, there was no significant relation with clutch size, egg
volume or hatching date.
Passarotto et al. (2020) provide support for a potential role of iris color in parent–
offspring communication in owls. However, in Little Owls, owlets raised in the same
nest showed low variation in iris yellowness, and variation in owlet iris color was not
related to body condition. Therefore, it seems unlikely that adult owls might use iris
yellowness as an indicator of the young’s condition or rely on it to adjust their parental
effort, as sometimes happens with other color traits of owlets, such as bill and cere
(Parejo et al. 2010, Aviles and Parejo 2013).
Interestingly, rather than indicating the owlets’ own quality, owlet iris color may be
an indicator of parent quality in Little Owls, as Passarotto et al. (2020) found owlets
with more-yellow irises later in the breeding season. This apparent finding contradicts a
large body of empirical evidence suggesting that individuals of better quality are usually
those raised in early broods (Verboven and Verhulst 1996, Arnold et al. 2006, Saino
et al. 2012), which would predict high yellowness levels in early Little Owl clutches.
However, Passarotto et al. (2020) also reported that nest-associated yellowness was
inversely related to breeding success, and that breeding success tended to decrease as
the season progressed, which in combination suggest that high values of iris yellowness
in the late Little Owl nests might result from brood reduction in those nests. Therefore,
one possibility is that negative selection against owlets in the poorest condition (i.e.,

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Figure 4.19 Skull of the Little Owl.

those with low yellowness) at the end of the breeding season results in an increase in
average nest-associated yellowness. A mutually nonexclusive possibility is that the
pattern results from differences in parental food allocation strategies in later clutches,
as adults could distribute food among fewer owlets in these cases, which may promote
an increase in yellowness due to a greater amount of food.

4.6 Ears
Despite the fact that owls are well adapted to nocturnal hunting by means of vision
tuned to low-level light conditions, silent flight and asymmetrical ears, the Little Owl,
with its crepuscular or diurnal lifestyle, is more eye focused and has symmetrical ears as
an adult (Schönn et al. 1991) (Figure 4.19).

4.6.1 Symmetrical Ears

The Little Owl is not strictly nocturnal like many other owl species, but also hunts at
dusk and dawn, and even during daylight (Negro et al. 1990). Hence, the species is not

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 4.7 Bill

as adapted to hunting in darkness as other owl species. This is confirmed by the fact
that its leading-edge wing serrations that are one of the adaptations of owls to silent
flight are less well developed than those in Barn Owls (Weger and Wagner 2016). Little
Owls mainly use optical cues for catching prey (Ille 1983, Schönn et al. 1991). This
species possesses color vision, as do other owl species (Porciatti et al. 1989, Schönn et al.
1991). The facial ruff, which acts as a sound collecting and amplifying system, and thus
improves hunting success through sound localization, is rather flat in Athene (Schönn
et al. 1991, König et al. 2008). Furthermore, adult Little Owls show no ear asymmetry
(Schönn et al. 1991), an anatomical adaptation that facilitates sound localization in
many owl species.
Kaup (1859) described the ears of Athene noctua noctua, A. n. glaux and Athene brama
as asymmetrical and presented a schematic drawing of the ear openings, without clearly
stating which ear was right and which left. Krings et al. (2018) suggest Kaup (1859)
might have examined an exceptional case or probably examined a specimen from a
different species.
Based on their findings Krings et al. (2018) proposed that crepuscular and diurnal
activity in Little Owls evolved secondarily from the strictly nocturnal lifestyle common
in most owl species. This is consistent with the results of Weger and Wagner (2016).
Serrations of the primary flight feathers improve silent flight and are thought to have
evolved as an adaptation to acoustically guided hunting. Consequently, serrations are
more developed in nocturnal species (Weger and Wagner, 2016). The typical serrations
are also observed in Little Owls, but they are short and may be secondarily reduced
(Weger and Wagner 2016). If Athene evolved from a strictly nocturnal to a more
crepuscular or even diurnal lifestyle, ear asymmetry might be an ancestral state in
Athene that still occurs in its embryonic development but disappears a few days before
hatching (Krings et al. 2018).
The frequency sensitivity of hearing in the Little Owl is 3–4 kc/s. It is correlated with
the morphology of the hearing apparatus and the way of life of the birds. The Little Owl
can locate small rodents with an accuracy of up to 1% thanks to their rustling and
squeaking (Golubeva et al. 1970).

4.7 Bill
Many owl species are sexually and socially active at night, when there is probably not
enough light for efficient color vision (Kelber et al. 2003). Plumage designs in nocturnal
species are dominated by white plumage patches intermingled with high-contrasting
dark backgrounds (Cramp 1985) producing achromatic (intensity-related but color-
blind) designs enhancing conspicuousness and allowing achromatic-based visual com-
munication (Kelber et al. 2003). Recent findings support the relevance of achromatic
signals for visual communication in nocturnal owls (Penteriani et al. 2007). Male eagle
owls Bubo bubo behaved more aggressively toward intruder males displaying low-
brightness white badges (Penteriani et al. 2007). Color cues are also used in parent–
offspring communication in nocturnal Scops Owls, Otus scops (Parejo et al. 2010).

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Little Owls are essentially considered crepuscular and nocturnal regarding their
hunting and vocal activities (Hardouin et al. 2008). However, adults can be found to
be active at any time during the day, both before and during reproduction (Negro et al.
1990), suggesting that important aspects of their life cycle can be completed in a wide
range of luminal conditions, from dim light around dawn and dusk to daylight condi-
tions. Hence, bill coloration and brightness might be relevant for both females and
their offspring.

4.7.1 Bill Coloration of Female Adults

Adult Little Owls show colorful yellow bills in both sexes (Cramp 1998) with unknown
function. Previous studies have found that the yellow-orange beak of diurnal raptors
may function as a carotenoid-based signal of adult quality, playing a role in mate choice
(Bortolotti et al. 2000, Mougeot and Arroyo 2006). Carotenoids cannot be synthesized
de novo and must thus be ingested in the diet, therefore carotenoid-based colorations
may inform on an individual’s capacity to acquire food rich in carotenoids and
to assimilate and process those nutrients (Hill 1991). Carotenoids also have important
anti-oxidant and immunostimulant properties (Møller et al. 2000), which may also
confer honesty on color signals through the competition for carotenoids to be
deposited as ornamentation versus serving physiological functions (von Schantz et al.
1999).
Avilés and Parejo (2012) studied color variation and potential signaling of the yellow
bill – a trait functioning as a sexual signal in diurnal raptors – during the breeding
season in female Little Owls. They found that the yellow-red chroma of the bill was
highly variable between individuals and positively correlated with female fitness pro-
spects (i.e., brood size at fledging). In addition, they found that females with brighter
bills were larger in size and produced owlets with a higher mass at fledging. This study
suggests that yellow bill coloration in female Little Owls may potentially play a role in
sexual signaling and may constitute the first evidence of chromatic color signaling in a
nocturnal bird.
Avilés and Parejo (2012) studied whether yellow bill coloration in breeding female
Little Owls might potentially function as a visual signal by investigating its covariation
with female quality (i.e., size and condition) and female fitness prospects (brood size at
fledging and fledgling body mass). To investigate the possible mechanism behind bill
color variation in Little Owls, they explored whether yellow bill coloration was related
to plasma carotenoid concentration. They reported a huge variation in the coloration of
the bill between different females. The reflectance of the bill of all females showed a
main peak in the yellow-red region (550–700 nm) and the spectral shape of a typical
yellow-orange object to the human eye. Descriptive statistics of bill color variables
(mean  SE) were as follows: total brightness: 517.12  22.49; hue: 511.66  0.90 nm;
yellow-red chroma: 61.72  0.05%. A greater yellow-red chroma in the bill was associ-
ated with a higher hue (Rp = 0.60, p = 0.008, n = 18), but no significant relationship
was found between total brightness and the other color variables. The yellow-red
chroma of the bill of wild females was not associated with body size and/or condition

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 4.7 Bill

once controlled for potential seasonal effects. However, yellow-red chroma of the bill
was associated with female fitness prospects. Females with bills with a greater yellow-
red chroma during incubation reared more fledglings. Additionally, they found that
females with brighter bills were larger in size, as estimated by tarsus length, and
produced owlets with a higher mass at fledging. The study by Avilés and Parejo
(2012) was the first showing a possible informative role of brightness for colored bare
parts of nocturnal birds.
Although disentangling the exact mechanism promoting the link between yellow
bill coloration and female fitness prospects clearly deserves further experimental
research effort, Avilés and Parejo (2012) confirmed the existence of such a link, which
is a prerequisite for the use of bill coloration as indicator of individual quality in mate
choice and/or rival assessment in intra-sexual contests.

4.7.2 Bill Coloration of Nestlings

Experimental manipulations of different color attributes of the bill of Little Owl nest-
lings were done by Avilés and Parejo (2013) by painting randomly chosen nestling bills
a brighter yellow, respectively less yellow and duller. Chromatic signals of offspring
quality have been shown to play a role in parent–offspring communication in diurnal
birds, but are assumed to be useless in dim light conditions because color-based dis-
crimination probably requires more light. A major ecological and evolutionary conun-
drum in this scenario is why the nestlings of some nocturnal owls display colorful
beaks. They tested the hypothesis that yellow bill coloration of owlets may function as a
chromatic signal revealing to parents aspects of quality of their offspring. They first
examined physical variation in bill coloration and its covariation with owlet quality.
Next they studied parental provisioning in relation to an experimental manipulation of
bill coloration of owlets.
The dark bill of Little Owl owlets adopts an adult-like colorful olive-yellow of
unknown function appearance a few days before fledging (Cramp 1998). Little Owl
owlets can climb up to the nest entrance at 3 weeks, and can even receive food deliver-
ies from their parents outside the nests while branching for a few weeks before being
fully fledged (Del Hoyo et al. 1999). Bill color can change over a short time period when
it relies on deposition of pigments (e.g., Faivre et al. 2003, Navarro et al. 2010, Karubian
et al. 2011), and so might communicate information to parents (e.g., Dugas 2009,
Thorogood et al. 2011).
Despite the owlet bill being perceived by humans as yellow, reflectance spectra
revealed that, besides a prominent peak around 550–625 nm (i.e., the yellow part of
the spectrum), Little Owl bills had a second marked peak in the UV (350–400 nm) part
of the spectrum.
Avilés and Parejo (2013) studied parental food allocation in relation to owlet bill
coloration by exaggerating a presumed carotenoid-rich bill (i.e., enhanced) of randomly
chosen owlets within each brood and comparing their body mass gain with that of
siblings in which brightness of the bill was slightly reduced (i.e., dulled) to compensate
for the possible achromatic effect. To enhance bill coloration, the bill was painted with

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a nontoxic yellow marker; to dull bill color, a gray marker was applied. After the
manipulation, color spectra of enhanced and dulled bills were within the range of
variation of bills in our population. Visualization of the significant color effects showed
that heavier Little Owl owlets, both between and within nests, displayed more-yellow
bills than lighter owlets.
Owlets had an average mass gain (mean  SE of 4.55  0.93 g) per night, which
suggests that parents were actively feeding nestlings during the experiment.
The effect of bill coloration on parental favoritism changed with brood size (inter-
action brood size treatment: F1, 22 = 5.75, p = 0.027; treatment effect: F1, 40 = 1.64, p =
0.2; brood size effect: F1, 22 = 1.02, p = 0.32) once controlled for the random effects.
Parents raising large broods preferentially fed enhanced over reduced bill-colored
owlets, whereas those raising small broods did not significantly bias feeding in relation
to owlet bill coloration.
Given that in birds, color vision functions only in bright light (Kelber and Lind
2010), a quality signaling function of chromatic aspects of the bill in adult (Avilés and
Parejo 2012) and young (Avilés and Parejo 2013) Little Owls was found, possibly
representing an overlooked but general pattern of signaling among nocturnal birds
inhabiting a wide range of luminal conditions.

4.8 Measurements
This section gives an overview of the measures of the Little Owl. The sizes of the Little
Owl are given in relation to the other related Athene species before focusing on the
differences between the subspecies. In Tables 4.2–4.11 we have directly brought forward
detailed body measurements from a number of publications and from Pellegrino et al.
(2020) for different subspecies. Included in these data are references to subspecies
designations of the Little Owl that we adapted in light of the most recent taxonomic
and biogeographical insights.

4.8.1 Interspecies Differences

According to the tarsus (t) length and the wing (w) length (shown as t; w, respectively),
the categorization between the different species of Athene is the following (Cramp
1985): Athene blewetti (26; 148), Athene brama (28; 152), Athene noctua (32–35;
156–162), Athene cunicularia (44; 164) (Schönn et al. 1991). There appears to be a
relationship between the habitat and the size of the species. The more open the habitat,
the taller the species, thus Athene cunicularia, the biggest Athene, has the longest legs
because it lives on the ground in areas of steppe vegetation (Scherzinger in Schönn et al.
1991).

4.8.2 Body Characteristics

The following measurements are most commonly used: weight (g), wing length (mm),
tail length (mm), bill size (mm) and tarsus length (mm). A general overview of these

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Table 4.2 General body characteristics of male and female Little Owls

Sex Wing (mm) Tail (mm) Bill (mm) Tarsus (mm) Weight (g) Author Subspecies

Male 161.3 (n = 25) 82 (n = 18) 29.7 (n = 15) 143.7 (n = 32) Simeonov et al. 1989 indigena
Female 164.5 (n = 21) 83.2 (n = 15) 30 (n = 11) 151.3 (n = 27)
Male 169 (n = 4) 86 (n = 4) 13 (n = 4) 31 (n = 4) 182 (n = 4) Khokhlov 1992 indigena
Female 170 (n = 4) 89 (n = 4) 13 (n = 4) 30 (n = 4) 220 (n = 4)
Male 162.2 (n = 8) 153.1 (n = 10) Brichetti personal vidalii
Female 164.8 (n = 15) 165.8 (n = 17) communication
Male 162.9 (n = 20) 77.8 (n = 22) 13.5 (n = 6) 34.5 (n = 8) 160.1 (n = 14) Mlikovsky and vidalii
Female 166.5 (n = 26) 79 (n = 27) 13.4 (n = 13) 37.2 (n = 9) 153 (n = 13) Piechocki 1983
Male 165.07 (n = 87) 195.07 (n = 243) Van Harxen and vidalii
Female 168.39 (n = 84) 208.60 (n = 223) Stroeken personal
communication
Male 161.8 (n = 26) 82.1 (n = 26) 39 (n = 9) 155.3 (n = 9) Martinez et al. 2002 vidalii
Female 163.6 (n = 26) 82 (n = 25) 41.1 (n = 7) 185.7 (n = 7)
Male 157.8 (n = 25) 34.59 (n = 27) 159.37 (n = 27) Mastrorilli 2005 noctua
Female 160.5 (n = 6) 35.39 (n = 10) 176.8 (n = 9)
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 Chapter 4: Morphology and Body Characteristics

210

200 male
female
Weight (grams)

190

180

170

160

150

Figure 4.20 Seasonal changes in weight of males and females in captivity during the annual cycle in
1995–1999 (26 pairs weighed; Génot and Sturm 2001).

measurements is given in Table 4.2. Martinez et al. (2002) also reported mouth size
(mm), back claw (mm), front claw (mm), talon length (mm), P8 length (mm), wingspan
(mm) and total length (mm). The terms used are explained in detail in the
Supplementary Glossary section.

4.8.3 Weight
As with the majority of owl species, female Little Owls are heavier than their male
counterparts at the onset of the breeding season. According to the mean weight over
the year, the maximum difference in weights occurs during the breeding season, and is
the least during the end of summer and autumn. Data recorded during a ringing study
in northeastern France (from March to July) gave the average weights for males at 164 g
(n = 36) and 181 g for females (n = 64) (Génot personal communication). The weight
was very variable according to the annual cycle, as shown in England, with an average
weight of 164 g in winter (n = 61) and 178 g in spring (n = 61) (Schönn et al. 1991).
These data were also reported from owls in captivity (Figure 4.20) where the most
important weight difference between males and females occurred in the breeding
season (March–June), with a peak in April and May (28 g), the laying and incubation
time. The smallest weight difference (1–2 g) occurred at the end of summer and in
autumn (Génot and Sturm 2001).
The average weights of 223 males and 243 females in northeast Netherlands were
195.07 and 208.60 g respectively; the difference between the averages was significantly
different (p <0.001) (Figure 4.21). During the breeding season in northeastern
Netherlands the weights of both male and female owls actively engaged in nesting
followed a pattern of decline until the young dispersed (Table 4.3, R. van Harxen and
P. Stroeken personal communication).
Maximum average weights were 200–240 g for males and 220–250 g for females
(Schönn et al. 1991). Minimum average weights were 160–170 g for males and

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 4.8 Measurements

Table 4.3 Variation in male and female Little Owl weights during the breeding season

March–April Incubation Young 0–10 days Young >11 days

Male 198.2 (n = 10) 170.9 (n = 15) 138 (n = 1)

Female 220.1 (n = 8) 209.2 (n = 136) 190.6 (n = 52) 179.6 (n = 37)

Van Harxen and Stroeken personal communication

–
ou

Figure 4.21 Box and whisker plot of male and female Little Owl weights, Achterhoek, the Netherlands,
2002–2021 (Van Harxen and Stroeken personal communication).

170–180 g for females (Schönn et al. 1991). In midwinter, the adults were 20–40%
heavier than at the end of the breeding season (Schönn et al. 1991).
Some loss of weight can occur between the beginning and the end of winter, even
when prey is abundant. This was shown in Belgium where owls were weighed on two
occasions. One male went from 225 g in December 2002 to 185 g in March 2003; a
female went from 260 g in December 2002 to 205 g in March 2003; a male went from
240 g in November 2001 to 205 g in February 2002; a male went from 230 g in
December 2002 to 165 g in February 2003 (J. Bultot personal communication). There
was also a difference of weight for males and females between the prebreeding
seasons (November–March (males 186.2 g (n = 45); females 204.9 g (n = 59)) and the
time during which nestlings are present (May–June) (males 159.3 g (n = 7); females
171.1 g (n = 56)).

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 Chapter 4: Morphology and Body Characteristics

25

males (n = 44)
20
females (n = 43)

15
Percentage

10

0
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
Wing length (mm)

Figure 4.22 Wing lengths of male and female Little Owls (Kämpfer and Lederer in Schönn et al. 1991).

The lowest weight record from a wild owl in nature was 98 g (Morbach in Schönn
et al. 1991). Génot (personal observation) found a recently dead 110 g adult during
a snowy winter in eastern France. Glutz Von Blotzheim and Bauer (1980) consider
that a weight of 120–125 g is the lower limit when an owl is hungry – death is likely
imminent.
The heaviest weights recorded were 270–280 g in nature and 300–350 g in an aviary
(Schönn et al. 1991).

4.8.4 Wing Length

The data shown in Table S4.1 show that females have wings that are on average 3–5 mm
longer than males. Females have tails on average 1–2 mm longer and tarsi on average
2.7 mm longer than males. In Spain, the wing length of 24 males and 25 females and
the weights of nine males and seven females were found to be statistically significantly
different. Other measures of males and females were not found to be significantly
different (Martinez et al. 2002). In other cases, the average wingspan does not differ
significantly between males and females (Mittel Westphalia, Germany, Kämpfer and
Lederer in Schönn et al. 1991) (Figure 4.22). In general, both sexes are not statistically
distinguished by wingspan for western European birds. The subspecies lilith, glaux and
saharae with relatively short wings do not feature distinguishable differences, while the
larger orientalis and plumipes show more significant differences (females on average
5–10 mm longer wings).

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 4.8 Measurements

180

–
o

Figure 4.23 Box and whisker plot of male and female Little Owl wing length, Achterhoek, the
Netherlands, 2002–2021 (van Harxen and Stroeken personal communication).

The average wing length of 87 males and 84 females in northeast Netherlands were
165.07 and 168.39 mm, respectively; the difference between the averages was signifi-
cant (F = 36.915; p <0.0001) (Figure 4.23) (van Harxen and Stroeken, personal
communication).
ANOVA tests comparing male and female wing lengths of stuffed birds per
subspecies from Pellegrino et al. (2020) (Table 4.4) only yielded sufficiently large
samples sizes (n >30) and significant differences for A. n. noctua (females 3.155 mm
longer wings than males; p = 0.0253) and A. n. vidalii (females 1.922 mm longer
wings than males; p = 0.038), but no statistical differences for A. n. sarda or
A. n. indigena.
According to the wing length of adults, the largest Little Owls come from Tibet
and the Kashmir Mountains. The following subspecies have a mean wing length of
180 mm: ludlowi, orientalis and plumipes. The smallest come from northeast Africa,
with the subspecies spilogastra and somaliensis having a mean wing length of 136 mm.
In Europe, the wing length of the species ranges between 151 and 178 mm; one-year-
old owls have an average wing length that is 3 mm shorter than the older adults.
Glutz Von Blotzheim and Bauer (1980) saw a geographical gradient for the wing
length from the shortest in western Europe to the longest in western Asia and the
Middle East. The geographic trend, however, is masked by local variation in wing

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 Chapter 4: Morphology and Body Characteristics

Table 4.4 Comparison of wing lengths of female and male Little Owls

Female

Subspecies Mean n Min Max SD Q25 Median Q75

A. n. bactriana 165.00 3 161.00 168.00 3.61 161.00 166.00 168.00

A. n. glaux 155.43 7 152.00 159.00 2.76 153.00 156.00 158.00
A. n. indigena 159.83 12 153.00 168.00 4.67 155.50 161.00 163.00
A. n. lilith 160.50 2 158.00 163.00 3.54 158.00 160.50 163.00
A. n. noctua 159.37 27 153.00 173.00 5.41 155.00 157.00 163.00
A. n. plumipes – – – – – – – –
A. n. saharae 153.50 2 153.00 154.00 0.71 153.00 153.50 154.00
A. n. sarda 157.74 19 149.00 165.00 4.38 154.00 158.00 162.00
A. n. somaliensis 140.50 2 139.00 142.00 2.12 139.00 140.50 142.00
A. n. spilogastra – – – – – – – –
A. n. vidalii 160.67 30 150.00 169.00 4.63 158.00 161.50 164.00
104

Male

Subspecies Mean n Min Max SD Q25 Median Q75

A. n. bactriana 166.43 7 161.00 171.00 4.16 163.00 165.00 171.00

A. n. glaux 155.70 10 152.00 162.00 2.98 153.00 156.00 157.00
A. n. indigena 160.46 24 151.00 168.00 4.85 156.50 160.50 164.00
A. n. lilith 159.20 5 153.00 163.00 4.44 156.00 162.00 162.00
A. n. noctua 156.19 27 145.00 166.00 4.72 153.00 156.00 159.00
A. n. plumipes 163.50 2 160.00 167.00 4.95 160.00 163.50 167.00
A. n. saharae 154.60 5 152.00 158.00 2.70 153.00 153.00 157.00
A. n. sarda 157.46 13 153.00 163.00 3.41 155.00 158.00 160.00
A. n. somaliensis 141.00 2 139.00 143.00 2.83 139.00 141.00 143.00
A. n. spilogastra 145.00 1 145.00 145.00 – 145.00 145.00 145.00
A. n. vidalii 158.74 47 151.00 165.00 3.35 156.00 159.00 162.00
143

ANOVA

Subspecies Difference of Averages n F p

A. n. indigena –0.625 36 0.14 0.71

A. n. noctua 3.155 54 5.31 0.0253
A. n. sarda 0.275 32 0.04 0.85
A. n. vidalii 1.922 77 4.5 0.0380

After Pellegrino et al. 2020

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 4.8 Measurements

length (Schönn et al. 1991). Males and females of German Little Owls in Mittel
Westphalia do not significantly differ from the subspecies indigena and bactriana.
In Kazakhstan, where five subspecies are found, the wing lengths, given as ranges
and means (Gavrilov 1999) are the following:

vidalii: males 152–169 (160), females 158–177 (168) mm

indigena: males 163–170 (164), females 166–173 (168) mm
bactriana: males 159–174 (166), females 159–177 (169) mm
orientalis: males 165–172 (168), females 170–181 (176) mm
plumipes: males 158–170 (163), females 167–178 (173) mm

4.8.5 Tail Length

The tail length of adult male and female Little Owls ranges between 69 and 89 mm
(Table S4.2). Measurements of Dutch stuffed birds showed, for first year birds, signifi-
cantly (3–5 mm) shorter tails than for adults. Females of noctua, vidalii, indigena, glaux
and saharae had, on average, 1–2 mm shorter tails than males. While some data sets
show no sex difference in tail length, some Dutch data show a considerable average
difference of 3.7 mm longer tails for females (Cramp 1985).
ANOVA tests comparing male and female tail lengths of stuffed birds for subspecies
from Pellegrino et al. (2020) (Table 4.5) only yielded sufficiently large samples sizes
(n >30) but no significant differences for A. n. sarda, A. n. indigena, A. n. noctua nor
A. n. vidalii.

4.8.6 Bill Size

The size of the bill measured between the tip of the bill and the start of the cere (culmen
length) (Figure 4.24) varies between 11.9 and 12.4 mm. Table S4.3 illustrates the
comparison of bill lengths of male and female Little Owls measured between the tip
of the bill and the start of the cere.
ANOVA tests comparing male and female bill size of stuffed birds from Pellegrino
et al. (2020) (Table 4.6) only yield sufficiently large sample sizes (n > 30) and significant
differences for A. n. vidalii (females 0.523 mm smaller bill than males; p = 0.03) and
A. n. noctua (females 0.527 mm smaller bill than males; p = 0.013) but not for A. n. sarda
or A. n. indigena. All bill sizes were measured by the same person with a calliper to the
nearest 0.1 mm (Pellegrino et al. 2020).

4.8.7 Tarsus Length

The length of the tarsus (Figure 4.25) shows significant differences between the subspe-
cies (Table S4.4). The smallest measures were observed in birds from Cyprus (males
30.0 mm; females 30.3 mm), which were, on average, 20% smaller than the birds
measured in Nordrhein-Westphalia (Germany) (males 36.7 mm; females 39.4 mm)

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 Chapter 4: Morphology and Body Characteristics

Table 4.5 Comparison of tail lengths of female and male Little Owls

Female

Tail Subspecies Mean n Min Max SD Q25 Median Q75

1 A. n. bactriana 79.33 3 76.00 83.00 3.51 76.00 79.00 83.00

2 A. n. glaux 73.71 7 70.00 76.00 2.75 70.00 75.00 76.00
3 A. n. indigena 75.42 12 72.00 79.00 2.07 73.50 76.00 77.00
4 A. n. lilith 75.00 2 72.00 78.00 4.24 72.00 75.00 78.00
5 A. n. noctua 76.04 26 69.00 85.00 3.75 73.00 75.00 78.00
6 A. n. plumipes – – – – – – – –
7 A. n. saharae 79.00 2 73.00 85.00 8.49 73.00 79.00 85.00
8 A. n. sarda 74.58 19 68.00 78.00 2.97 72.00 75.00 77.00
9 A. n. somaliensis 63.50 2 62.00 65.00 2.12 62.00 63.50 65.00
10 A. n. spilogastra – – – – – – – –
11 A. n. vidalii 72.77 30 68.00 79.00 2.98 71.00 73.00 74.00
103

Male

Tail Subspecies Mean n Min Max SD Q25 Median Q75

1 A. n. bactriana 78.14 7 74.00 84.00 3.34 75.00 78.00 80.00

2 A. n. glaux 73.00 10 70.00 76.00 2.36 71.00 73.50 75.00
3 A. n. indigena 76.00 24 71.00 83.00 3.15 74.00 76.00 79.00
4 A. n. lilith 76.00 5 72.00 80.00 3.08 74.00 77.00 77.00
5 A. n. noctua 75.54 28 68.00 90.00 3.89 73.00 75.00 77.00
6 A. n. plumipes 82.00 2 82.00 82.00 0.00 82.00 82.00 82.00
7 A. n. saharae 73.60 5 71.00 75.00 1.67 73.00 74.00 75.00
8 A. n. sarda 75.38 13 72.00 78.00 1.76 74.00 75.00 77.00
9 A. n. somaliensis 63.50 2 61.00 66.00 3.54 61.00 63.50 66.00
10 A. n. spilogastra 72.00 1 72.00 72.00 – 72.00 72.00 72.00
11 A. n. vidalii 72.61 46 68.00 83.00 2.98 70.00 72.00 74.00
143

ANOVA

Tail Subspecies Difference of Averages n F p

3 A. n. indigena 0.583 32 0.34 0.566

5 A. n. noctua 0.503 50 0.23 0.631
8 A. n. sarda –0.806 28 0.77 0.388

After Pellegrino et al. 2020

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 4.9 Voice

Figure 4.24 Different bill measurements showing the (A) culmen length with cere, (B) beak depth and
(C) culmen length.

(Schönn et al. 1991). The Mediterranean populations, except for the Iberian birds, have
relatively small tarsi. The significant spread in average measures within the noctua
subspecies shows the important local variation. The length of the tarsus was found to
differ between the sexes in two German studies (Schönn et al. 1991), with females
having, on average, 2.7 mm larger tarsi than males. All other studies showed a differ-
ence in length below 1 mm.
ANOVA tests comparing male and female tail lengths of stuffed birds for subspecies
from Pellegrino et al. (2020) (Table 4.7) only yield sufficiently large samples sizes
(n >30) but no significant differences for A. n. vidalii, A. n. indigena, A. n. noctua nor
A. n. sarda.

4.8.8 Other Measures

Besides the most common body measures given above, Martinez et al. (2002) also
reported mouth size (males: 18.33  1.6 mm; females: 19.60  0.55 mm), back claw
(males: 9.67  1.7 mm; females: 9.31  1.04 mm), front claw (males: 10.72  1.10 mm;
females: 11.64  0.85 mm), talon length (males: 43.56  1.40 mm; females: 43.57 
1.90 mm), P8 length (males: 117.00  3.20 mm; females: 119.12  3.88 mm), wingspan
(males: 557.67  11.90 mm; females: 565.64  13.15 mm) and total length (males:
234.44  11.00 mm; females: 243.57  10.29 mm).

4.9 Voice
Most studies are carried out through the recording of owl vocalization in the field, with
results offered as descriptions of sound spectrograms or sonagrams (Glutz Von
Blotzheim and Bauer 1980, Exo 1984, 1990, Scherzinger 1988, Exo and Scherzinger

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 Chapter 4: Morphology and Body Characteristics

Table 4.6 Comparison of bill lengths of female and male Little Owls

Female

Bill Subspecies Mean n Min Max SD Q25 Median Q75

1 A. n. bactriana 14.33 3 13.70 15.40 0.93 13.70 13.90 15.40

2 A. n. glaux 13.88 6 13.40 14.90 0.58 13.40 13.75 14.10
3 A. n. indigena 14.45 11 12.40 15.50 0.91 13.90 14.70 15.10
4 A. n. lilith 14.55 2 14.50 14.60 0.07 14.50 14.55 14.60
5 A. n. noctua 13.77 26 12.80 16.40 0.79 13.20 13.60 14.20
6 A. n. plumipes – – – – – – – –
7 A. n. saharae 13.55 2 13.00 14.10 0.78 13.00 13.55 14.10
8 A. n. sarda 14.47 19 13.40 14.90 0.42 14.10 14.60 14.80
9 A. n. somaliensis 12.45 2 12.00 12.90 0.64 12.00 12.45 12.90
10 A. n. spilogastra – – – – – – – –
11 A. n. vidalii 13.24 30 11.20 14.70 0.75 12.80 13.30 13.50
101

Bill Subspecies Mean n Min Max SD Q25 Median Q75

1 A. n. bactriana 13.96 7 13.20 14.60 0.52 13.50 14.00 14.60

2 A. n. glaux 14.40 10 13.10 15.80 0.85 13.90 14.60 14.90
3 A. n. indigena 14.23 23 12.40 15.90 0.90 13.70 14.10 14.80
4 A. n. lilith 14.10 4 13.70 14.60 0.37 13.85 14.05 14.35
5 A. n. noctua 14.30 24 13.30 16.00 0.65 13.80 14.10 14.55
6 A. n. plumipes 14.00 2 13.50 14.50 0.71 13.50 14.00 14.50
7 A. n. saharae 13.96 5 13.30 15.30 0.85 13.40 13.50 14.30
8 A. n. sarda 14.28 13 13.70 15.40 0.53 13.80 14.30 14.50
9 A. n. somaliensis 12.25 2 11.60 12.90 0.92 11.60 12.25 12.90
10 A. n. spilogastra 12.90 1 12.90 12.90 – 12.90 12.90 12.90
11 A. n. vidalii 13.76 46 12.30 15.40 0.69 13.20 13.80 14.30
137

Bill Subspecies Difference of Averages n F p

3 A. n. indigena 0.224 30 0.46 0.504

5 A. n. noctua –0.527 46 6.59 0.013
8 A. n. sarda 0.191 28 1.31 0.262
11 A. n. vidalii –0.523 72 9.782 0.003

After Pellegrino et al. 2020

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 4.9 Voice

Figure 4.25 Skeleton of Little Owl. The foot consists of toes with claws and the tarsus or
tarsometatarsus. The expression “foot” is used when no specific part is referred to. Owls have four toes,
three in front and one hind toe. Of the front toes the outer one may be turned backwards (as owls often
do when they are roosting).

1989, Schönn et al. 1991). Exo and Scherzinger (1989) recorded 22 distinct call notes
(23 including nonvocal bill snapping) (Table 4.8). Two are specific to young owls and
disappear during ontogenetic development. The juvenile repertoire contains 12 defined
call notes which can be summarized by three or four basic call notes. Males and females

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 Chapter 4: Morphology and Body Characteristics

Table 4.7 Comparison of tarsus lengths of female and male Little Owls for different
subspecies

Female

Tarsus Subspecies Mean n Min Max SD Q25 Median Q75

1 A. n. bactriana 34.87 3 33.30 37.70 2.46 33.30 33.60 37.70

2 A. n. glaux 32.00 7 30.50 34.20 1.32 30.90 32.30 32.80
3 A. n. indigena 33.28 12 30.60 34.70 1.24 32.40 33.60 34.30
4 A. n. lilith 31.85 2 31.50 32.20 0.49 31.50 31.85 32.20
5 A. n. noctua 34.70 26 31.80 38.50 1.51 33.40 34.80 35.90
6 A. n. plumipes – – – – – – – –
7 A. n. saharae 32.40 2 31.70 33.10 0.99 31.70 32.40 33.10
8 A. n. sarda 34.39 17 30.60 37.50 1.99 33.30 34.30 35.60
9 A. n. somaliensis 28.05 2 27.90 28.20 0.21 27.90 28.05 28.20
10 A. n. spilogastra – – – – – – – –
11 A. n. vidalii 36.36 30 33.50 39.30 1.52 35.50 36.55 37.40
101

Male

Tarsus Subspecies Mean n Min Max SD Q25 Median Q75

1 A. n. bactriana 31.81 7 30.10 33.60 1.25 30.70 31.70 32.80

2 A. n. glaux 33.48 10 31.30 37.10 1.93 31.70 33.20 34.20
3 A. n. indigena 33.68 24 30.30 35.50 1.43 32.45 34.10 34.85
4 A. n. lilith 34.06 5 31.80 39.10 3.03 31.90 32.90 34.60
5 A. n. noctua 35.04 26 31.70 40.70 1.85 34.10 34.50 35.80
6 A. n. plumipes 32.25 2 31.80 32.70 0.64 31.80 32.25 32.70
7 A. n. saharae 33.55 4 31.20 36.40 2.18 32.00 33.30 35.10
8 A. n. sarda 34.44 13 31.90 38.80 1.79 33.90 34.50 34.70
9 A. n. somaliensis 28.65 2 27.80 29.50 1.20 27.80 28.65 29.50
10 A. n. spilogastra 28.20 1 28.20 28.20 – 28.20 28.20 28.20
11 A. n. vidalii 35.76 45 31.50 39.70 1.76 34.80 35.60 36.70

ANOVA

Tarsus Subspecies Difference of Averages n F p

3 A. n. indigena –0.399 32 0.68 0.416

5 A. n. noctua –0.338 48 0.52 0.474
8 A. n. sarda –0.050 26 0.01 0.944
11 A. n. vidalii 0.608 71 2.391 0.126

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 4.9 Voice

Table 4.8 Different calls of Little Owls according to age and sex

Relative
volume Sound Juveniles Females Males

Contact calls 1 Contact call x x x

Feeding call 5 Begging-calls x
juvenile
Feeding call 5 Begging snoring x
Feeding call 5 Feeding call x (x)
Feeding call 5 Luring/tempting (x) x x
Feeding call 5 Feeding X
twittering
Advertising calls 7 Begging-calls x x
adult
Advertising calls 7 Begging snoring X
Advertising calls 7 Luring/tempting x x
Advertising calls 7 Chugging ? x
Advertising calls 7 Nest showing X
Advertising calls 7 Yelping x ?
Advertising calls 7 Copulation call X
Enemy call and defense 6 Alarm call x x x
Enemy call and defense 6 Chattering/ x x x
cackling
Enemy call and defense 6 Peeping/ x
cheeping
juvenile
Enemy call and defense 6 Defence x x ?
screeching
Enemy call and defense 6 Chirping x x x
Enemy call and defense 6 Hissing/blowing X ? ?
Enemy call and defense 6 Bill snapping (x x x)
Intra-specific aggression 6 Trembling X
Intra-specific aggression 6 Peeping x x x
Intra-specific aggression 6 Excitement calls x x x
Intra-specific aggression 6 Aggressive calls ? x
Intra-specific aggression 6 Whispering calls ? x x
Intra-specific aggression 6 Copulation call x
Song 2 Song, ordinary x x
Song 2 Glissando song X

Total 12 15 15

Brackets indicate uncertain assignment. X indicates gender- or juvenile-specific call.

After Exo and Scherzinger 1989

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 Chapter 4: Morphology and Body Characteristics

(a) Hooting call

Frequency (kHz)

4.5

1.5

0
1.5 3
Time (s)

(b) Chewing call

Frequency (kHz)

12

0 1 4
Time (s)

Figure 4.26 (a) Hooting or “guhk” call of male Little Owl; (b) Chewing or “miaw” call of male Little
Owl (after Hardouin 2006).

each have 15 call notes out of a total of 20 altogether. Only three for the male and two
for the female are sex-specific. The final repertoire includes 40 acoustic signals with
mixtures and combinations. The most common are indicated here and are based
mainly on Exo and Scherzinger (1989). Most typical calls are the territorial hooting or
“guhk” call and the intra-specific excitement chewing or “miaw” call (Figure 4.26).
A comparative study of the vocalization repertoires within the genus Athene
(Scherzinger 1988) showed a strong similarity of calls among the Little Owl Athene
noctua, Spotted Owlet Athene brama and the Burrowing Owl Athene cunicularia. This
study supported the placement of the Burrowing Owl in the genus Athene.

4.9.1 Calls of Adults

Song (Cramp 1985, Exo and Scherzinger 1989), Guhk (Exo 1984) or Hoot (Hardouin
2006). This is the primary call used in pair formation and territorial defense. It is
produced by both male and female adult owls (Exo 1984). The use of this call for
census work, aiming at exact population estimates (Exo and Hennes 1978a) rather than

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 4.9 Voice

Figure 4.27 Territorial song of Little Owls: (a) female song, (b) male song I and II short calls; III
Glissando Song of up to one second with increasing frequency (after Exo and Scherzinger 1989).

determining Little Owl presence, should pay special attention to “guhk”-like female
calls (Exo 1984).
The hooting call sounds like “goooek,” a loud, questioning “huui” (Haverschmidt
1946) or “ghu(k),” sometimes given in a crescendo series; towards the end of these series
it changes to an excited “guiau,” or “kwiau” and ends abruptly with a shrill “hoo-ee,” or
“miju” (Haverschmidt 1946, Glutz Von Blotzneim and Bauer 1980, Exo and Scherzinger
1989).
The female’s call is shorter than the male’s and the tone is generally higher in pitch
(Figure 4.27). Most calls are louder and clearer when given by the male and repeated
monotonously in a varied group of notes, while the notes occur singly when uttered by
the female (Exo 1984). During the main time of courtship the male repeats the “guhk”
call, often for several minutes, whereas for females spacing between the notes is irregu-
lar and the number of calls given in a session variable between three and five, or even
one (Exo 1984). Typical for the male calls is the steep frequency increase at the
beginning. The basic tone of the call is mostly between 0.4 and 0.5 kHz at the start,
rather constant or slightly increasing in the middle between 0.7 and 1.3 kHz. In
contrast to the opening and middle sections, the end of the call is often less pure in
terms of sound with both lower and higher frequencies up to 2.2 kHz, but mostly
limited to 2 kHz. Female hoots are mostly above 0.5 kHz, ranging from 0.6 to 1.1 kHz in
the middle part. The hoots have an average length of 0.58  0.10 s and 0.36  0.07 s for
males and females, respectively (Exo 1984).
The basic monosyllabic form of the territorial song can be rather variable and is
described in different ways as a melanchonic “kiiu” (Mikkola 1983), a nasal increasing
“uuhg” (Kehrer 1972), a clear “üü” (Stadler 1945), a rifled “ghuk” (Kleinschmidt 1934),
an interrogative “guhg” (Bergmann and Helb 1982), an interrogative “ui,””huu” or
“huui” as a mating call (Haverschmidt 1939, 1946). The number of calls by males varies
according to the excitement of the bird between 12 and 20 per minute (Bergmann and
Helb 1982). Males can sing for up to 20 minutes, while females utter a maximum of five
calls (Exo 1984). During the courtship, duets occur. Simultaneous hooting of both
partners or neighboring owls can lead to aggressive calls.

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 Chapter 4: Morphology and Body Characteristics

Figure 4.28 Comparison of the six most abundant “Ghuk” types from (a) northwest German and
(b) eastern English male Little Owls. I. Basic form with characteristic roughly pressed final element.
II. Basic form with a constant frequency final element. III. Basic shape with high-pitched final
element. IV. Song with frequency jump in the 1st element part. V. Singing with frequency jump in
the 2nd element part. VI. Transition to “aggressive singing.”

The glissando song sounds more like a whisle like “ghuui” or “gluui” and “üüüü”
(Stadler 1945) or up to three syllabic calls like “goojhüi” (Exo in Schönn et al. 1991).
Despite all the variability, the basic structure of the typical courtship call is always
retained. A male can utter both simple “uuu” or ““gjuu” syllables, as well as nasal high-
drawn “gjuüi” or long glissando syllables whose continual rise in frequency sometimes
breaks off with an almost clicking element. Females lack the glissando song completely.
Sonagrams of territorial songs of male Little Owls were studied by Exo (1990) in
eastern England and compared with songs of birds from northwest Germany. The basic
repertoire is almost the same in both populations, with the same syllable types and
characteristics (Figure 4.28); the different syllable types occur in similar frequencies of
occurrence (Chi-square test of frequencies nonsignificant). Population-specific syllables
were not observed. A detailed analysis of the syllable structure of the different call notes
showed that most notes of Little Owls from eastern England last significantly longer
than of birds from northwest Germany. Furthermore, the pitch was higher in English
Little Owls than in German conspecifics. The differences might indicate an actual
genetic evolution due to a limited population density and the entire isolation from
the continental populations supporting the local evolution of vocalizations.
The Excitement Call is variable and described as “jau,” “mija,” “kwiau,” “miji,”
“iwidd” and “kuwitt”(Exo and Scherzinger 1989). This “kiew” (Cramp 1985), “miau”
(Exo in Schönn et al. 1991) or chewing call (Hardouin 2006) is the most commonly
given. It is a clear “(k)weew,” “huu” or “gwauu”; a sharp, complaining, “kee-ew”;
repeated irregularly. It is used in many cases of social contact, such as mating, feeding
young, sometimes at the end of the ordinary song during courtship behavior, during
copulation and nest-showing, or during disturbance at the nest site, occurrence of
predators (e.g., feral cats) and territoriality. The final shrill call “kuwitt”can be confused
with the call of a female Tawny Owl, Strix aluco (Bettmann 1951, Haller 1951, Runte
1951, Ammersbach 1952). This is a fast call that has been recorded at 28–36 calls/min

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 4.9 Voice

Figure 4.29 Excitement calls of Little Owls: I single “jau,” II strong “jau,” III “mija,” IV “mlijau,”
V “mijau,” VI “mlji.jau” (after Exo and Scherzinger 1989).

(Glutz Von Blotzheim and Bauer 1980). It is mainly given in association with copula-
tion and other aspects of heterosexual behavior, but also is given in fights
between rivals.
The basic frequency of excitement calls ranges between 0.8 and 1.8 kHz. The loudest
sounds are mostly produced for the heighest frequencies. Repeats can be as frequent as
20–30 calls per minute. The variation in excitement call is substantial (Figure 4.29),
with the shrillness going up with increasing arousal. When the excitement goes up, the
syllables and intervals shorten, while frequencies rise and sounds become clearer and
sharper (Exo 1987). The bill is mostly wide open during the calls. These calls are
regularly combined with other calls, yielding multisyllabic sounds like “guiau(u)” in
combination with hooting, “go-go jau” in combination with chugging, one to four
syllables “gho-cho-cho-guau” in combination with begging or in combination with
cackling/luring such as “gwju,” “guau” or “guoau.”
The variations yield many different descriptions and spelling, e.g., “giou” (Stadler
1932), “whirrou” (Mikkola 1983), Lapwing-like “iwidd” (Stadler 1945), “kiut” or
“kuwitt” (Kleinschmidt 1934). Excitement calls are mostly single or in longer series
broadcasted from exposed perches or in flight.
A special sequence of excitement calls leads to aggressive calls, while territorial song
has crescendo to shrill calls. Such compilations sum up to 60 calls and are uttered with
breaks of several minutes and repeated up to 50 times (Illner in Glutz Von Blotzheim
and Bauer 1980). Excitement calls always precede the aggressive song and increase in
speed, volume and aggressive arousal.
Excitement and aggressive calls are very variable and frequently combined with
other calls. Robb and The Sound Approach (2015) illustrate the “cu-cu-miau” sequence
as a combination of two alarm calls followed by an excitement call. Similar to this, we
found many different combinations in the xeno-canto database of recordings, includ-
ing the reverse order of “miau-cu-cu,” amongst other combinations. A check of 300
vidalii, noctua and indigena recordings at www.xeno-canto.org showed that the number
of alarm calls varies between none and three before and after the “miau”-call
(Table 4.9). A verification by Exo (personal communication) of the most unusual
combinations indicated that amongst the different subspecies, no structural differences
could be observed and all the calls sounded very familiar, even the “cu-cu-miau.” Such
variations were observed by Exo during his entire research of Little Owl vocalizations

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 Chapter 4: Morphology and Body Characteristics

Table 4.9 Little Owl recordings downloadable from www.xeno-canto.org indicating a

large variation of sequences of calls across all subspecies.

Xeno-canto multisyllabic excitement calls

Subspecies Recording Country Sequence url

vidalii XC584228 Belgium miau-cu-cu www.xeno-canto.org/

584228
vidalii XC584232 Belgium miau-cu-cu www.xeno-canto.org/
58432
lilith XC586495 Cyprus cu-cu-miau www.xeno-canto.org/
586495
vidalii XC543093 France miau-cu-cu-cu www.xeno-canto.org/
543093
vidalii XC581158 France cu-cu-cu-miau www.xeno-canto.org/
581158
vidalii XC632858 France cu-miau www.xeno-canto.org/
632858
vidalii XC640751 France miau www.xeno-canto.org/
(same 640751
bird as
XC640752)
vidalii XC640752 France cu-miau www.xeno-canto.org/
640752
vidalii XC666245 France miau-cu www.xeno-canto.org/
666245
vidalii XC582853 Germany miau-cu www.xeno-canto.org/
582853
noctua XC104903 Italy cu-cu-miau www.xeno-canto.org/
104903
noctua XC130646 Italy cu-miau www.xeno-canto.org/
130646
noctua XC280410 Italy cu-cu-cu-cu- www.xeno-canto.org/
cu-cu-miau 280410
noctua XC303918 Italy cu-cu-miau www.xeno-canto.org/
303918
noctua XC76846 Italy cu-miau www.xeno-canto.org/
76846
indigena XC289312 Turkey cu-cu-cu-cu- www.xeno-canto.org/
cu-cu-miau 289312
indigena XC386897 Azerbaijan hoot www.xeno-canto.org/
386897
lilith XC329983 Cyprus miau www.xeno-canto.org/
329983

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 4.9 Voice

Table 4.9 (cont.)

Xeno-canto multisyllabic excitement calls

Subspecies Recording Country Sequence url

vidalii XC402735 Denmark very long hoot www.xeno-canto.org/

402735
vidalii XC542901 France miau-miau www.xeno-canto.org/
542901
vidalii XC563549 France arousal www.xeno-canto.org/
563549
vidalii XC579256 France weird www.xeno-canto.org/
579256
vidalii XC625972 France hissing www.xeno-canto.org/
625972
vidalii XC632855 France miljau www.xeno-canto.org/
632855
vidalii XC637312 France alarm www.xeno-canto.org/
637312
vidalii XC637939 France male hoot www.xeno-canto.org/
637939
indigena XC383820 Greece miau www.xeno-canto.org/
383820
indigena XC384037 Greece miau www.xeno-canto.org/
384037
noctua XC608765 Hungary queck-queck www.xeno-canto.org/
608765
noctua XC431705 Italy screech www.xeno-canto.org/
431705
noctua XC551298 Italy very long miau www.xeno-canto.org/
551298
noctua XC585163 Italy begging www.xeno-canto.org/
585163
lilith XC167696 Kuwait hoot www.xeno-canto.org/
167696
indigena XC379810 Greece miau www.xeno-canto.org/
379810
orientalis XC498062 Mongolia miau www.xeno-canto.org/
498062
vidalii XC565134 Netherlands nestlings www.xeno-canto.org/
565134
vidalii XC635668 Netherlands female hoot www.xeno-canto.org/
635668

continues

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 Chapter 4: Morphology and Body Characteristics

Table 4.9 (cont.)

Xeno-canto multisyllabic excitement calls

Subspecies Recording Country Sequence url

vidalii XC574057 Poland male hoot www.xeno-canto.org/

574057
vidalii XC574060 Poland kieuw www.xeno-canto.org/
574060
vidalii XC200789 Portugal miau www.xeno-canto.org/
200789
vidalii XC594439 Portugal miau www.xeno-canto.org/
594439
vidalii XC628295 Portugal kiwit www.xeno-canto.org/
628295
lilith XC215633 Qatar hoot www.xeno-canto.org/
215633
vidalii XC663140 Spain hissing www.xeno-canto.org/
663140
lilith XC209241 UAE hoot www.xeno-canto.org/
209241
vidalii XC572506 UK male hoot www.xeno-canto.org/
572506
vidalii XC592428 UK nestlings www.xeno-canto.org/
592428
vidalii XC592429 UK female as male www.xeno-canto.org/
enters with 592429
food
vidalii XC592431 UK pair interaction www.xeno-canto.org/
592431
vidalii XC592434 UK begging www.xeno-canto.org/
592434
vidalii XC592435 UK begging www.xeno-canto.org/
592435
vidalii XC592436 UK bill snapping www.xeno-canto.org/
592436
vidalii XC592437 UK screech www.xeno-canto.org/
592437
vidalii XC615449 UK duet www.xeno-canto.org/
615449
indigena XC401499 Ukraine hoot www.xeno-canto.org/
401499

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 4.9 Voice

with a number of variations in which the “cu-cu” can preceed or follow the “miau,” or
the “cu-cu” is not heard. A first analysis of a limited sample from different countries
gives the idea that one will detect these variations in all countries. So-called differences
could simply be artifacts of a limited number of sequences analyzed before assuming
geographical differences. Our limited unrepresentative sample size gives the idea that
the concentration of “cu-cu” in front seems to be higher in Italy, but certainly not
exclusively, and currently it has not been found in the samples from the Balkans
(indigena) and only once in a recording from Turkey and Cyprus (indigena).
So far there is no evidence to support Athene noctua vidalli being a different species
compared to the other subspecies. A differentiation based upon specific variations in
vocalizations alone is not conclusive and statistically not sound. Hence we see no
reason why there would be the need to split A. n. vidalii from A. n. noctua, indigena,
glaux, sahare and lilith based upon a three-syllabic “cu-cu-miau” (Robb and Sound
Approach 2015) simply because the sample of vidalii individuals from Portugal might
be too small and statistically biased to not produce this variation.
The Whispering Call, similar to “shrie,” is used as a contact call. It is prominently
given by the female during courtship (Glutz Von Blotzheim and Bauer 1980).
Begging Calls of the female are extentions of those from juveniles, “tsiech,”
“schräää” and “siej” sounds (Glutz Von Blotzheim and Bauer 1980, Exo and
Scherzinger 1989).
The Feeding Trill from females is used when feeding young. It is a rapid, nasal “gek-
gok-gok-gok,” or hoarse cackling, sometimes interrupted by “uuh” as with the “kiew”
call (Glutz Von Blotzheim and Bauer 1980, Exo and Scherzinger 1989).
The Nest-Showing Call of the male is similar to the “zick-zick” of the Kestrel Falco
tinnunculus, or “tjuck-tjuck” of the domestic cockerel given during nest-showing; it is
also occasionally given during mating (Glutz Von Blotzheim and Bauer 1980, Exo and
Scherzinger 1989).
The Copulation Call is a soft “oo-oo”given by birds sitting close together, before or
after copulation. The female shrieks during copulation. The male often sings “goooek”
before mating and sometimes during mounting (Haverschmidt 1946, Exo and
Scherzinger 1989). The male also uses the “kiew” call, the excitement call and the nest
showing nest-showing call.
The Alarm Call is used as an alarm and given during times of anxiety and for warning. It
is a loud, chattering “kek-kek” when an owl is disturbed at its nest (Haverschmidt 1946,
Exo and Scherzinger 1989). Calls based on “queb”and “keck” sounds express fear; short
“kja” or “kju” are given as a warning to a rival or predator; “quip” expresses anxiety;
“quijep” is given as warning to young at fledging age. Snoring or screeching sounds are
given in displeasure or fear (Glutz Von Blotzheim and Bauer 1980). Bill-snapping is a
typical alarm noise of the Strigiformes family (Schönn et al. 1991). Little Owls can produce
this sound from 10 days of age (Exo in Schönn et al. 1991). Some authors (Gooch 1940,
Haverschmidt 1946) have mistakenly interpreted this sound as tongue-clicking.
The alarm call (Figure 4.30) is sometimes combined with excitement calls or aggres-
sive calls including “cu-cu-miau” or “miau-cu-cu” in most subspecies, including vidalii.
An under-representation of samples from the Balkans might be the principal reason for

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 Chapter 4: Morphology and Body Characteristics

Figure 4.30 Alarm call and bill-snapping sound of Little Owls: (a) Alarm call – I “kek,” II “kju.kek,”
III “kau,” IV “kju.kau,” V “kju.i.pep.pep” (I, II and V: adult birds; III and IV fledged birds).
(b) Bill-snapping (30-day-old juvenile birds).

the lack of such combinations. Only about half of Italian samples (noctua) produce “cu-
cu-miau” combinations.
The Hissing Call is given in threat and is often linked to the alarm call (Schönn
et al. 1991).
Other Calls: A squeaky “uik” is given during allopreening, which is a rare behavior
(Glue and Scott 1980). A hissing or rasping sound is produced by males and females, probably
in a begging or contact calling context (Glutz Von Blotzheim and Bauer 1980). A curious
snoring is sometimes given by day in spring, sounding like the exhalation of person in
deep sleep (Witherby et al. 1938). Owls also give various shrieking, yelping, grumbling
and rasping calls (Glutz Von Blotzheim and Bauer 1980, Exo and Scherzinger 1989).

4.9.2 Calls of the Young

A food-begging call is given from unhatched (still within the egg) and small young. It is
a monosyllabic “psiep,” “szip,” or “srie” and a disyllabic “uiiet.” In the second week
after hatching it becomes a harsh, “chsij,” “chriie” or hissing “schwo”; in the fourth
week a rasping snoring like a Barn Owl Tyto alba is given. Several adult calls, e.g., “gjuu”
as the “kiew” call, “kek,” snoring (alarm call) and hissing develop while the young are
still in the nest or shortly afterwards. The typical territorial song is given during the first
autumn (Ullrich 1973, Glutz Von Blotzheim and Bauer 1980).

4.10 Flight
The flight of the Little Owl is direct, swift, easy and wave-like, very similar to a
woodpecker’s flight. The owl’s wing-beats are as silent at high frequencies as they are
at low ones, and have no ultrasonic noise (Thorpe and Griffin 1962) despite the fact
that its leading-edge wing serrations that are one of the adaptations of owls to silent
flight are less well developed than those in Barn Owls (Weger and Wagner 2016).
In certain cases, in particular when hunting, Little Owls display hovering flight like
Kestrels (Martin and Rollinat 1914, Malmstigen 1970, Wahlstedt 1971). The hovering takes
place 2–3 m or sometimes as much as 20 m above the ground (Gyllin 1968), with the owl
finally closing its wings and dropping like a stone on the prey (Gregory 1944, Tayler 1944).

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 4.11 Digestive Organs

4.11 Digestive Organs

In the context of a taxonomical study, the chemical composition of the uropygial gland
waxes of the Little Owl was compared to those of diurnal birds of prey and confirmed
that owls seem to be an isolated order (Jacob and Hoerschelmann 1984).
Some data on the digestive organs were collected in Turkmenistan (Broun 1986).
Therelative weight of the glandular stomach is 0.54% of the overall body weight. The
relative weight of the stomach muscle is 2% and it has comparatively thin walls. The
bowel is comparatively short, being 198% of the body length. There are blind appendi-
ces between the thin and thick parts of the bowel, representing 14.5% of the total
length of the bowels. Histomorphological observations of the digestive apparatus
showed that the third blind intestine of the Little Owl is absent and developed glands
are plentiful. The muscular tunics and the glands of the duodenum and the organ of
Meckel are well developed compared to those of granivorous birds (Richetti et al. 1980).
A comparitive neurohistological study of the esophagus in granivorous, omnivorous
and carnivorous birds showed that carnivorous birds have one remarkable development
in the innervation of the esophagus, advanced compared to granivorous and omnivor-
ous birds. The nervous plexus of the esophagus is supplied with many nervous cells and
the muscularis mucosae is well developed (Costaglia et al. 1981).

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 Chapter 5: Distribution, Population
Estimates and Trends

Chapter Summary
In this chapter, we examine aspects of distribution and population for the Little Owl over
its global range which covers 78 countries. Three recent publications give an overview of
population numbers and short- and long-term breeding population and breeding distri-
bution trends for the EU28 (European Environment Agency 2021) and for continental
Europe (BirdLife International 2015, Keller et al. 2020). The European Union (EU28) Red
List assessments were based principally on the official data reported by EU Member States
to the European Commission under Article 12 of the Birds Directive. All Member States are
requested by the Birds Directive to monitor bird species and send a report every six years.
This information includes population sizes and trends (short and long term) for breeding
and wintering populations, as well as pressures and threats for Special Protection Area
trigger species. In addition, population status and trends are assessed at the EU level. The
reporting period 2006–2012 data were used by BirdLife International (2015) and Keller et al.
(2020). The reporting period 2013–2018 data were used by the European Environment
Agency (2021). Detailed information about the reporting under Article 12 of the Habitats
Directive can be obtained from www.eionet.europa.eu/etcs/etc-bd/activities/reporting/art
icle-12/. The reported data as well as the assessments of status and trends at the EU
level can be visualized in the web tool http://bd.eionet.europa.eu/article12/.
For the European Red List assessments outside the EU28, similar data were sourced
from BirdLife Partners and other collaborating experts in other European countries and
territories (BirdLife International 2015, Keller et al. 2020). Some further declining
populations were observed in central European (e.g., Poland, Austria, Czech Republic,
Slovakia, Slovenia) and northern European (e.g., Denmark, Estonia, Latvia) countries,
stable or fluctuating populations in Mediterranean countries (e.g., Portugal, Spain, Italy,
Greece, Cyprus). Positive trends, either long term or short term are observed in western
European countries (e.g., Germany, Belgium, Netherlands, France, Luxembourg,
Switzerland), where conservation initiatives have yielded or are starting to yield positive
results. Balkan countries were not monitored in detail. Short-term trends were more
positive than long-term trends.

Supplementary resources are available online at www.cambridge.org/littleowl. These include

appendices as well as tables which previously appeared in the first edition (prefixed with ‘S’).

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Table 5.1 General short-/long-term trends in population numbers and distribution at different spatial scales

Range Scope Reference Years + – = F X Total

Europe Short-term population BirdLife International 2015 1998–2012 2 5 13 4 13 37

trends
EU28 Short-term distribution European Environment Agency 1996–2018 2 6 12 4 24
trends 2021
EU28 Short-term population European Environment Agency 2001–2018 2 8 9 5 24
trends 2021
Global Long-term population 12 10 23 33 78
Distribution trends
Europe Long-term population BirdLife International 2015 1973–2013 3 9 5 1 19 37
trends
EU28 Long-term population European Environment Agency 1973–2018 2 13 4 5 24
trends 2021
Global Long-term distribution 12 10 25 31 78
Distribution trends

–: Decrease; +: Increase; =: Stable; F: Fluctuating; U: Uncertain; X: Unknown

147
 Chapter 5: Distribution, Population Estimates and Trends

Data outside the European continent were obtained from individual publications.
In very general terms, all cumulative data suggest that the global distribution of the
Little Owl increased in 12 countries, decreased in 10, remained unchanged in 25 and
was insufficient for determination in 31 countries (Table 5.1).
For population numbers, the data suggests that the number of owls increased in 12
countries, decreased in 10, remained unchanged in 23 and was insufficient for deter-
mination in 33 countries.
This chapter gives a description of the global distribution and some limiting factors
such as latitude and elevation, and presents maps of European population numbers and
short- and long-term trends. For each country we focus on the population estimates for
currently existing populations and present distribution maps when available.
In Table S5.1 we give details per country on the evolution of the distribution and the
population estimates. Except the countries of western and central Europe, where the
Little Owl has decreased for the last 40 years due to land-use changes, the species
remains common or relatively widespread in all the Mediterranean countries from
southern Europe and the Middle East to North Africa and in many Asiatic republics of
the former Soviet Union, where natural habitats of the Little Owl still occur. Changes in
Little Owl numbers reflect the effects of regulatory factors acting on the populations
(see Chapter 10). Some western European countries were able to counter the downward
progressions through a professional conservation and management strategy and local
volunteer actions. (For management best practices see Chapter 11.)

5.1 Distribution
A map showing the rough approximation of Little Owl global distribution is given in
Figure 5.1. The distribution is principally determined by the latitude, elevation, climate
and land use. We discuss latitude and elevation in this chapter, while land use is
discussed in relation to habitat selection in Chapter 6.

5.1.1 Latitude
The species occurs in middle and lower latitudes (mainly between the 22nd and 51st
parallels and at the extremes between the 9th and 56th parallels), both continental and
marginally oceanic, mainly temperate, steppe and Mediterranean biotopes, but extending
to boreal and tropical. While adapting to windy and rainy climates, it favors warm, even
semi-arid conditions and is vulnerable to severe frosts and snow cover (Glutz Von
Blotzheim Von Blotzheim and Bauer 1980). Hunting becomes very difficult, if not impos-
sible, for the owl with a snow cover over 10–15 cm (Juillard 1984, Schönn 1986), limiting
distribution in both latitude and elevation. Only one observational record exists for the
Opochka district (Pskov Region, north of Leningrad, Russia) and the species is extremely
rare in Estonia (Malchevskiy and Pukinskiy 1983) and probably exinct in Latvia too. Figures
5.2–5.3 illustrate the northern border of the global distribution at different spatial scales.
The Republic of Bashkortostan forms the northern limit around the Ural Mountains
(Karyakin and Kozlov 1999), where only a few breeding pairs have been recorded.

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(A)

Figure 5.1 Global distribution of Little Owl. (A) EBBA2 50
50 km accuracy for Continental Europe. (B) Article 12 of Birds Directive at 10
10 km accuracy
for EU28.
 Figure 5.1 (cont.)
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 5.1 Distribution

(A)

Breeding assessment
Possible (673)
Probable (449)
Confirmed (1220)
Not Available (8)

(B)
Abundance
1-9 (370)
10-99 (957)
100-999 (758)
1,000-9,999 (236)
Not reported (29)

Figure 5.2 European distribution of Little Owl in 50
50 km grid cells (after Keller et al. 2020).
(A) Breeding evidence. (B) Abundance.

The southern limit is northern Africa where the species is found north of the Sahara
Desert in the mountainous regions of Aïr (Niger), Tibesti (northwest Chad), Am Djarass
(Ennedi, northeast Chad), Murzuq (Lybia), Mellit (Sudan) and south through Sudan
and Ethiopia to Somalia. Further eastward the species is found in the Arabian Peninsula,
Kuwait, Iraq, Iran, Pakistan, Kashmir, the Tibetan Plateau and north Szetschwan to
the Yellow Sea south of Xinhailian, China.

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 Chapter 5: Distribution, Population Estimates and Trends

Figure 5.3 EU28 distribution in 10
10 km grid cells of Little Owl as obtained from data (2018)
reported by EU Member States to the European Commission under Article 12 of the Birds Directive
(after European Conservation Agency 2021).

5.1.2 Elevation
In the northern and middle parts of its range the Little Owl is a lowland species that is
rarely found above 600 m, even in mountains of central Europe (Glutz Von Blotzheim
and Bauer 1980). Further south, it has been found up to 1140 m in the Causse Méjean
(Lozère, France) (Juillard et al. 1992), 1200 m in the Spanish Pyrenees (Kostrzewa et al.
1986), 1230 m in the Orobie Alps (Lombardia, northern Italy) (Mastrorilli 2001),
1900–2000 m in Georgia and Armenia, up to 1260 m (Parashka town) in the Lviv
Region (Kijko and Yakubenya 1995), at 1600 m in Catalonia (northeast Spain), up to
2300 m in favorable territories in Sierra Nevada (southern Spain) (Olea 1997), up to
2300 m in Bulgaria (Simeonov and Delov 1989), up to 2000 m in Armenia (Lyaister and
Sosnin 1942), up to 2200–2300 m in Uzbekistan (Abdusalyamov 1971), up to 2300 m,
probably even higher, in Gissar-Karategin, Tajikistan (Popov 1959), up to 2000–2800 m
in Altai, up to 2800 m on the southern side of the Todra gorges, Morrocco (Thévenot
et al. 2003), 4200 m in Pamir (Dementiev and Gladkov 1951) and sometimes higher in
Tajikistan (Beik at 4260 m, Zor-Kul and Bash-Gumbez at 4200 m; Abdusalyamov 1971)
and 4400 m in Turkey (Kasparek 1992). In central Europe the population density tends
to decrease with elevation in Germany (Loske 1986).

5.2 Population Numbers

The species is a widespread resident across much of Europe (except the north), which
accounts for less than half of its global range. Its European breeding population is large

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 5.2 Population numbers

(618 000–1 170 000 pairs, BirdLife International 2015), but has undergone a moderate
decline between 1970 and 1990. Although the species was stable or increased across
parts of its European range during 1990–2000, several populations have declined and
the species has probably undergone a moderate decline overall in that timespan.
Consequently the species has been provisionally evaluated as Declining by BirdLife
International (2004). In several countries conservation activities have proven successful
from 2008–2012, resulting in the status Stable in Europe (BirdLife International 2015)
and Decreasing in the EU28 (European Conservation Agency 2021).
Little Owl populations are fluctuating, especially in the north of the range, where
marked decreases follow severe winters (Büchi 1952, Poulsen 1957, Dobinson and
Richards 1964, Kämpfer-Lauenstein and Lederer 1995). Some populations recover after
severe winters; however, in some countries such winters result in local extinction when
habitat deterioration comes into play. In northern Europe the distribution is shrinking,
i.e., Estonia and Latvia no longer have Little Owls. Denmark is suffering probably
terminal declines.
Recent decreases, often marked, over much of Europe have been observed. They were
ascribed mainly to habitat changes, including loss of suitable nest sites (offset in some
areas by provision of artificial sites – see, e.g., Juillard 1980, Ullrich 1980, Bultot et al.
2001). Estimation of population numbers are subject to possibly severe misinterpret-
ations (Génot and Lecomte 1998). This is partly due to the fact that most researchers
work in areas more densely populated by owls (K. M. Exo personal communication).
The following is an overview of the three main methods by which population estimates
are made.

5.2.1 Population Estimates Through Consolidation of Study

Results Across a Geographic Region
Consolidation of geographic results is the simple addition of locally based owl
numbers. Using geographic population numbers has several drawbacks, i.e., data
are never recorded in one single year, sometimes with a spread of more than 10
years, and during this period population numbers may fluctuate substantially.
Further, data for these estimates are generally obtained through the use of inconsistent
methods, i.e., some observers record singing males, while others record singing
individuals or confirmed breeding pairs. Finally, the habitats vary from one place to
another, and even within regions, making extrapolations error-prone. The main advan-
tage of this estimate is that a minimal population is obtained without risking over-
estimations.

5.2.2 Population Estimates Through Partial Densities in

Different Habitats
These estimates are derived through main habitat associations, whereby owl numbers
and habitats within known study areas are recorded, and then extrapolated to areas
without owl surveys, but in which the main habitats are mapped. These estimates

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 Chapter 5: Distribution, Population Estimates and Trends

assume that strong habitat preferences exist. Previous work on the Little Owl has
identified a potential correlation between owl numbers and the percentage of grassland
(Exo 1983, Loske 1986, Génot and Wilhelm 1993, Zuberogoitia 2002). On a larger scale,
similar observations were made (Van Nieuwenhuyse et al. 2001b). However, a recent
study in Belgium was not able to find any association of the species with grassland (Van
Nieuwenhuyse and Bekaert 2002). In France, no correlation between average popula-
tion densities and the area of grassland was found (Ferrus et al. 2002). Thus, researchers
are cautioned against using grasslands as an estimator for the population of Little Owls.
Using average densities of the species to calculate population numbers only makes
sense when habitat-specific calculations are made. Further, this is only reasonable if
an indication of the small-scale mosaic-like landscape structure is taken into account,
e.g., using the length of parcel perimeters instead of areas (Van Nieuwenhuyse et al.
2001b). An additional limitation to this method is that it does not take the clustered
distribution of the species into account, as seen when apparently unsuitable habitats
are occupied and apparently suitable habitats remain empty (Génot and Lecomte
1998).

5.2.3 Population Estimates Through Habitat Modeling

A final method to estimate population numbers is through statistical modeling using
habitat characteristics of grid cells that are, in turn, based on the area used by a breeding
pair (i.e., 25 ha) (Van Nieuwenhuyse et al. 2001c). After a complete survey of a region
covered by 25 ha grid cells, habitat models are estimated through logistic regression
analysis. These models predict the probability that, based on the habitat conditions
within the grid cell, a cell will harbor Little Owls. Following this, all of the cells
containing a probability of 50% or more are counted and summed for the region.
A range of population numbers is obtained by estimating using different models
(stratified or not). Estimates through habitat modeling require both a large number of
volunteers, a consistent survey method (see Johnson et al. 2009) and reliable digital
descriptions of the landscape. More and more countries are assembling national
working groups that organize volunteers, and more digitized geographic datasets are
becoming available.
Ilanloo et al. (2020) used MaxEnt to model habitat suitability in Iran. This maximum
entropy algorithm is based on a machine learning technique that evaluates the prob-
ability of the distribution of maximum entropy values affected by limitations from
environmental variables that are important for the species distribution (Elith et al.
2011). Modeling based on this method only needs presence data and has the ability
to consider both continuous and discrete variables in the model. In addition, the
method helps to identify the most important environmental variables affecting species
distribution. Ilanloo et al. (2020) used 70% of the presence data for model creation and
30% for model validation. The area under the curve (AUC) acquired from an ROC
(receiver operating characteristic) curve is used to evaluate the model. Distance to
forests (51.2%) and distance to human settlements (16.2%) were the most important
variables in habitat selection in Iran.

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 5.3 Densities

5.2.4 Population Estimates and Trends

Despite the shortcomings of all these methods, it is still possible to give a rough
overview of population numbers around the distribution range. It should be stressed
that the data quality that we present is very heterogeneous from one country to
another. For the European countries that were included in the EBCC Atlas (Van
Harxen in Keller et al. 2020), the data were collected using similar methods across
Europe. The historical and current population estimates are given in Table 5.2.
The recent observed trend in maximum estimates in Little Owl populations was
negative throughout Europe for countries that had details available in 1994 (Manez in
Tucker and Heath 1994, BirdLife International 2004, Van Nieuwenhuyse et al. 2008).
Recent trends are more positive, in some countries due to conservation activities.
Some central European countries have undergone a well-documented negative pro-
gression between 1994 and 2004, i.e., Czech Republic –9%, Germany –24%, Poland
–50%, probably due to a severe winter in 2001/2002 (G. Grzywaczewski personal
communication), which continued till 2020, except for Germany. Other countries
have undergone a decrease due to a reduction in reproductive offspring, i.e., the
Netherlands –46% (Willems et al. 2004) or Luxembourg –43%, UK –29%, Slovenia
–63%. This trend continued till 2021 in some central European countries, e.g., Czech
Republic –50%, Slovenia –70%, Slovakia –40%, Austria –50%. Moreover, recently Little
Owl populations have appeared to recover or become stable, e.g., Germany, short-term
growth of 60–106% and Belgium, long-term growth of 16%. Switzerland featured a
recent increase of 153% in its few small, well-protected and managed populations
(Meisser et al. 2016). For other countries some important ajustments to the maximum
population estimates were carried out compared to Manez in Tucker and Heath 1994,
either upwards (Belgium +115%, Hungary +25%, Portugal +50%, Spain +54%, Denmark
+33%) or downwards (France –30%, Greece –90%, Italy –40%). Population estimates
report stable populations in 2004 for the UK, Germany, Portugal, Belgium, Austria,
Greece and Italy. From 2004–2020 the UK showed a severe decline of –48%.
In Groningen (the Netherlands), increasing fragmentation of a decimated popula-
tion featured a negative population trend that more significantly affected territories
that were isolated compared to those that were clustered, despite their more favorable
habitat composition (van’t Hoff 2001).

5.3 Densities
Using European abundancy data on the numbers of owl pairs (Figure 5.4) and the total
area of each country in Europe, we calculated the density of the species for earlier and
current data using the surface area of each country and the breeding area as obtained
from European Conservation Agency (2021). We found a significant relationship
between the latitude and the maximum observed densities (European Bird Census
Council; Van Nieuwenhuyse et al. 2008) when removing Portugal from the analysis
(due to extremely high densities). The density drops to 0.09 pairs per km² for each
increase of 1000 km latitude (Figure 5.4).

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Table 5.2 Historical and current population estimates (1994–2004 and 2015–2021)
156

Manez (1994) BirdLife International Van Nieuwenhuyse BirdLife International European Environment Agency
(2004) et al. (2008) (2015) (2021)

Min Max Min Max Min Max Min Max Min Max Mean (Min,
pairs pairs pairs pairs pairs pairs pairs pairs pairs pairs Max) pairs

Albania 5000 10 000 4000 8000 4000 8000 4000 8000

Armenia 800 1500 800 1500 1000 1500
Austria 40 60 70 100 60 60 130 170 140 190 165
Azerbaijan 2000 10 000 2000 10000 2000 10 000
Belarus 2000 4000 400 1000 400 1000 400 1000
Belgium 4500 6600 12 500 14 000 9000 14 200 8000 13 000 7900 12 200 10 100
Bosnia and 2000 3500
Herzegovina
Bulgaria 4000 10 000 5000 8000 4000 10 000 10 000 14 000 5000 8000 6500
Croatia 6000 8000 500 1000 6000 8000 4000 6000 4000 6000 5000
Cyprus 2000 4000 5000 15 000 2000 4000 4000 10 000 4000 10 000 7000
Czech Republic 700 1100 200 400 500 1000 250 500 100 150 125
Denmark 150 150 100 200 175 200 43 10
France 10 000 50 000 20 000 60 000 10 330 35 000 21 000 50 000 25 000 50 000 37 500
Georgia Present 3500 4000 Present
Germany 5000 10 000 5800 6100 7000 7600 7500 9000 7500 8500 8000
Gibraltar 1 5 5
Greece 5000 10 000 5000 15 000 5000 1000 5000 15 000 5000 15 000 10 000
Hungary 1500 2000 1500 2500 2000 2500 1500 2500 3000 4000 3500
Italy 10 000 50 000 30 000 50 000 10 000 30 000 40 000 70 000 40000 70 000 55 000
Kosovo 4000 6000
Latvia 10 30 10 30 10 30 0 0
Lithuania 10 50 5 10 5 10 1 10 5 10 8
Luxembourg 80 150 40 80 45 85 12 25 20 30 25
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FYRO 1400 2400 1400 2400 3000 7000

Macedonia
Moldova 5000 7000 3200 4200 3200 4200 3200 4200
Montenegro 1000 2000
Netherlands 9000 12 000 5500 6500 5500 6500 7000 9000 7500 8500 8000
Poland 1000 3000 1000 2000 1000 1500 500 1000 500 1000 750
Portugal 10 000 100 000 50 000 150 000 50 000 150 000 58 000 137 000 40 000 60 000 50 000
Romania 20 000 40 000 40 000 60 000 20 000 40 000 15 000 40 000 15 000 40 000 27 500
Russia 10 000 100 000 10 000 100 000 11 500 103 300 50 000 70 000
Serbia 10 300 15 000
Serbia and 10 000 15 000 14 000 22 000
Montenegro
Slovakia 800 1000 800 1000 800 1000 400 700 300 600 450
Slovenia 500 800 150 200 200 300 100 150 100 150 125
Spain 50 000 65 000 20 000 100 000 36 643 100 000 39 433 39 433
Switzerland 30 40 60 70 91 101 80 110
Turkey 5000 50 000 300 000 600 000 5000 50 000 300 000 600 000
Ukraine 11 000 12 000 15 000 22 000 15 000 22 000 15 000 22 000
UK 6000 12 000 5800 11 600 4000 8500 2338 4888 3613
Total incl. 184 320 568 980 555 840 1 267 900 235 159 649 986 617 849 1 128 365
Turkey
Total excl. 179 320 518 980 255 840 667 900 230 159 599 986 317 849 528 365
Turkey
Total excl. 169 320 418 980 245 840 567 900 218 659 496 686 267 849 458 365
Turkey and
Russia
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 Chapter 5: Distribution, Population Estimates and Trends

Cyprus Belgium

y = –9E–08 meter latitude + 0.5859

R² = 0.1818
Albania Netherlands

Yugoslavia
Maximum density

Romania

Italy
Moldova, Republic
Spain Of
Croatia
Bulgaria France United Kingdom
Greece Luxembourg
Slovenia
Germany Latvia
Turkey Georgia Ukraine Belarus
Poland
Austria
Switzerland Russian Federation
Lithuania

Latitude

Figure 5.4 Relation of latitude and population densities (densities calculated as population estimates
divided by the area per country (data after Manez in Tucker and Heath 1994).

The local density of a wildlife population can have a strong “feedback” effect on the
demographic performance of that population. For example, the higher the local dens-
ity, the fiercer the competition (for food and territory) might be and the greater the
subsequent impact on breeding success will be (Newton 1998). With this in mind, we
focus this section on examining Little Owl densities that have been observed within the
breeding range of the species. Densities of owls are best determined through the use of
radio-telemetry, with detailed locations over a full year (or more) on both male and
female owls, yielding an accurate determination of the area (ha) being utilized by a pair
of owls. More frequently though, densities are measured as the number of observations
(e.g., pairs, calling males, calling individuals) per km². Also, densities are given by
quantifiying the distances between neighboring owls (called the “nearest-neighbor
distance”).
Amongst the publications we reviewed for this book, numbers of Little Owls have
been reported in several different ways, making comparisons difficult. Some authors
report owl numbers obtained by playback; however, these are often (incorrectly)

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 5.3 Densities

interpreted and reported as confirmed breeding pairs. In some other cases calling
males are reported, and in others all calling individuals are reported, no matter
the sex. Some authors denote the density of owls as the number of territories, or
the number of breeding pairs, for a given study-area size. While this type of density
estimate often reflects the number of territorial pairs within the studied area, it may
not reflect the actual area used by the pairs within it (e.g., owls used habitat outside
of the study area, or, conversely, there may have been substantial areas within
the study area that were not used by owls). In particular, while small study areas
(e.g., <2500 ha) might have very high apparent densities, in actuality these esti-
mates are biased and ecologically inappropriate for the species because substantial
habitat outside the study area was used by the owls. Density estimates determined
from large study areas are most appropriate, so long as the entire study area was
thoroughly and consistently surveyed for owls. As a common measure for our review
of owl densities, we use vocal observations as the basis for comparisons across
all locations.
Study areas are seldom chosen at random. Most researchers favor areas with
dense populations of owls; however, such areas yield biased views on owl numbers,
with serious over-estimations possible if extrapolating local densities to larger
areas. Local extremes in densities are reflected in the following examples: 41.7
territories/km² on 12 ha (Exo 1988), 11.6 territories/km² on 86 ha (Estoppey 1992)
and 15–20 territories/km² (Glutz Von Blotzheim 1962, Visser 1977). Owl densities
are also measured by quantifying the nearest-neighbor distances and by characteriz-
ing the home-range area (through radio-telemetry or detailed observations of
known individuals). These measures correlate rather well since the average nearest-
neighbor distance decreases fairly constantly with increasing number of observed
owls in a given area (Bultot et al. 2001). The higher the densities, the shorter the
nearest-neighbor distance, and the smaller the expected home range of the owls
would be. We illustrate the heterogeneity of the densities first by describing the
number of observed individuals per square kilometer (km²) at the country level, then
as the number of pairs per occupied km² and conclude with descriptive statistics
of nearest-neighbor distances. Since densities tend to be different for different
regions, we give an overview of observed densities per country where data are
available (Table 5.3).
In the country-specific accounts we do not offer vocal observations as a basis for
comparisons but rather offer a wide mix of numbers (calling individuals, pairs, calling
males), hence we are reporting the results as reported in the papers or through personal
communication by the observers (Table S5.2).
Most nearest-neighbor distances are calculated from relatively small samples (Table
S5.3), and tend to confirm the clustered nature of local populations, rather than overall
densities (Table 5.4). The closest distances between nests in Britain were 240 m and
320 m (Glue and Scott 1980); the closest recorded nests in Switzerland were 50 m (Glutz
Von Blotzheim and Bauer 1980). The closest calling males in Flanders, Belgium were:
Herzele – min, 42; average, 210; max, 862; n ¼ 210; Geraardsbergen – min, 157;

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Table 5.3 Historical relative densities per country in relation to the country surface and the breeding area (data after Manez in Tucker and Heath
160

1994)

Densities/1000 km² country area Densities/1000 km² breeding area

BirdLife Van BirdLife European

Manez International Nieuwenhuyse International Environment
(1994) (2004) et al. (2008) (2015) Agency (2021) European Environment Agency (2021)

Mean Breeding area Mean

Surface (Min, from gridded (Min,
Country (km²) Min Max Min Max Min Max Min Max Min Max Max) maps (km²) Min Max Max)

Albania 28 890 173 346 138 277 138 277 138 277
Armenia 29 743 27 50 27 50 34 50
Austria 83 790 1 1 1 1 1 2 2 2 2 2 4000 35 48 41
Azerbaijan 86 600 23 115 23 115 23 115
Belarus 199 900 10 20 2 5 2 5 2 5
Belgium 30 710 147 215 407 456 293 462 261 423 257 397 329 27 629 286 442 366
Bosnia and 51 209 39 68
Herzegovina
Bulgaria 111 400 36 90 45 72 36 90 90 126 45 72 58 106 085 47 75 61
Croatia 52 900 113 151 9 19 113 151 76 113 76 113 95 26 059 153 230 192
Cyprus 8531 234 469 586 1758 234 469 469 1172 469 1172 821 6500 615 1538 1077
Czech 78 260 9 14 3 5 6 13 3 6 1 2 2 5700 18 26 22
Republic
Denmark 42 800 4 4 2 5 4 5 1 2500
France 546 900 18 91 37 110 19 64 38 91 46 91 69 266 800 94 187 141
Georgia 85 010 41 47
Germany 355 300 14 28 16 17 20 21 21 25 21 24 23 64 833 116 131 123
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Gibraltar 7 3 333 1667 1667

Greece 131 500 38 76 38 114 38 8 38 114 38 114 76 191 100 26 78 52
Hungary 92 890 16 22 16 27 22 27 16 27 32 43 38 50 967 59 78 69
Italy 302 000 33 166 99 166 33 99 132 232 132 232 182 261 100 153 268 211
Kosovo 10 887 367 551
Latvia 63 840 0
Lithuania 77 810 1 100
Luxembourg 2668 30 56 15 30 17 32 4 9 7 11 9 303 66 99 83
FYRO 25 713 54 93 54 93 117 272
Macedonia
Moldova 49 440 101 142 65 85 65 85 65 85
Montenegro 13 812 72 145
Netherlands 34 760 259 345 158 187 158 187 201 259 216 245 230 29 500 254 288 271
Poland 308 600 3 10 3 6 3 5 2 3 2 3 2 N/A N/A N/A N/A
Portugal 88 830 113 1126 563 1689 563 1689 653 1542 450 675 563 39 800 1005 1508 1256
Romania 237 300 84 169 169 253 84 169 63 169 63 169 116 200 500 75 200 137
Russia 16 620 000 1 6 1 6 1 6 3 4
Serbia 88 361 117 170
Serbia and 102 400 98 146 137 215
Montenegro
Slovakia 49 030 16 20 16 20 16 20 8 14 6 12 9 7815 38 77 58
Slovenia 20 560 24 39 7 10 10 15 5 7 5 7 6 2500 40 60 50
Spain 505 600 99 129 40 198 72 198 78 78 142 271 277
Switzerland 40 830 1 1 1 2 2 2 2 3
Turkey 782 700 6 64 383 767 6 64 383 767
Ukraine 615 100 18 20 24 36 24 36 24 36
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 Chapter 5: Distribution, Population Estimates and Trends

Table 5.4 Nearest-neighbor distances for the Little Owl in western European
anthropogenic landscapes

Nearest-neighbor
distance
No. of
Reference Region (country) territories Year Mean Min Max

Own observations Lorraine (France) 8116

Loske 1986 Baden-Württemberg 10 1976–1984 3470
(Germany)
Own observations Montagne Reims 2834
(France)
Loske 1986 Rheinland-Pfalz 11 1976–1984 2781
(Germany)
Loske 1986 Niedersachsen 5 1976–1984 2070
(Germany)
Loske 1986 Münsterland 76 1976–1984 1950
(Germany)
Own observations Vosges du Nord 1776 300
(France)
Loske 1986 Schleswig-Holstein 23 1976–1984 1145
(Germany)
Own observations Causse Méjean 36 1075
(France)
Own observations Brotonne (France) 23 1045
Loske 1986 Saarland (Germany) 11 1976–1984 682
Loske 1986 Pays de Herve 40 1995 680 300 1700
(Belgium)
Loske 1986 Jülicher Börde 12 1976–1984 596
(Germany)
Loske 1986 Essen (Germany) 13 1976–1984 585
Müller 1999 Kreis Viersen 250 1998 556 250 925
(Germany)
Lucas 1996 Condroz (Belgium) 26 1995 460 225 950
Loske 1986 Niederrhein 26 1976–1984 456
(Germany)
Loske 1986 Mittelwestfalen 29 1976–1984 435
Own observations Scarpe-Escaut 14 403
(France)
Own observations Geraardsbergen 310 1998–2000 330 157 942
(Belgium)
Own observations Meulebeke 78 2000 388 173 811
(Belgium)
Own observations Herzele (Belgium) 210 1998–1999 210 42 862

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 5.4 Trends

average, 330; max, 942; n ¼ 310; Meulebeke – min, 173; average, 388; max, 811; n ¼ 78
(Van Nieuwenhuyse et al. 2008). Breeding sites of three pairs in Ankara (Turkey) were
situated at 120 m from each other around 32 apartment blocks (11 ha), with males
hunting within 20 m of each other when using electricity poles as perches (Vaassen
2000). In southern Portugal, the nearest-neighbor distances of breeding pairs in an area
of 16 km² of steppe-like habitat were: min, 97.5 m; average, 313.7 m; max, 555 m;
n ¼ 37 and in a woodland area of 6 km²: min, 67.5 m; average,447.5; max 1230 m;
n ¼ 39 (Tome unpublished data). The minimum distances between nests in the
Mediterranean climates of Israel were 200 m in Jerusalem, 300 m at Marj Sanur, in
the desert climates of Nizzana, and 500 m in northwestern Negev (Shirihai 1996). In the
United Arab Emirates three nests have been recorded at a spacing of 500 m, although
occupied nests were not less than 1000 m apart in Kuwait. At one farm in central Arabia,
which was rich in rodents but poor in Little Owl roosting and nesting sites, 12 adults
were seen on a line of rock heaps within 1 km (Génot and Van Nieuwenhuyse 2002). In
the lower Zarafshan area of Uzbekistan, six Little Owls and three nests were registered
over the breeding period during a road survey of 15 km. In the sandy desert of the lower
parts of Kashka Darya (Uzbekistan), on a road survey of 20 km (50–60 m along each side
of the road surveyed) seven pairs were found (Sagitov 1990). In Badkhyz, Turkmenistan
the highest number is recorded on clayish precipices of the Kyzyldzhar ravine, where
there were 1.6 individuals per km of precipice (Simakin 2000). In Tajikistan the species
can be considered as common everywhere, but nowhere in the Republic at such
considerable concentrations as the one precipice, stretching almost uninterrupted
along several kilometers in the “Tigrovaya Balka” Nature Reserve. Three pairs
nested at a distance of 1–1.5 km from each other. As a rule, the separate breeding
pairs are situated at 8–10 km from each other (Abdusalyamov 1971). In the Stavropol
region, Little Owl density was two individuals per km of road surveyed (Khokhlov et al.
1998).

5.4 Trends
Table 5.5 illustrates the short-term and long-term trends in distribution and population
numbers. Figure 5.5 illustrates the short-term trends in population sizes across Europe
and Figure 5.6 shows the long-tem trends. In the short term, populations are increasing
in Russia and Germany and declining in Denmark, Poland, Slovakia, Slovenia and
Italy, while in the remaining countries they are stable. In the long term, Belgium,
Germany and Switzerland show increases, while Spain, France, Netherlands, Austria,
Czech Republic, Slovakia, Slovenia, Russia, Albania, Greece and Turkey all show
declining populations. However, some of these countries might currently exhibit a
temporary downward direction as part of natural population fluctuations. Central
European countries might not be able to counter the decline through conservation
activities due to much lower initial densities than in those countries where activities
seem to be fruitful.

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164

Table 5.5 Short-term and long-term trends in population numbers and distribution

BirdLife International (2015)

Short-term population trend Long-term population trend

Country Direction Magnitude (%) Year(s) Quality Direction Magnitude (%) Year(s) Quality

Albania = 0 2002–2012 Medium – 0–5 1980–2012 Poor

Armenia X X
Austria = 0 2001–2012 Medium – 20–50 1980–2012 Poor
Azerbaijan X X
Belarus = 0 2000–2012 Medium = 0 1980–2012 Medium
Belgium = 0 2000–2012 Poor + 10–78 1973–2012 Poor
Bosnia and X X
Herzegovina
Bulgaria = 0–5 2000–2012 Medium = 0–5 1980–2012 Medium
Croatia X X
Cyprus = 0 2001–2012 Medium X
Czech Republic X – 50–64 1985–2012 Medium
Denmark – 50–100 1999–2011 Medium – 50–100 1980–2011 Medium
France = 0 2000–2012 Medium – 20–50 1975–2011 Medium
Georgia X X
Germany + 60–106 1998–2009 Good + 179–318 1988–2009 Good
Greece X X
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Hungary X X
Italy – 30-40 2000–2011 Poor X
Kosovo X X
Latvia – 100 1980–1994 Good
Lithuania = 0 2001–2012 Medium = 0 1980–2012 Medium
Luxembourg F 0–30 2000–2012 Medium – 50–80 1980–2012 Medium
FYRO X X
Macedonia
Moldova = 0 2000–2010 Medium = 0 1980–2010 Medium
Montenegro X X
Netherlands F 0 2002–2011 Good – 50–63 1980–2011 Good
Poland – 50 2000–2012 Medium – 50–70 1980–2012 Medium
Portugal F 0 2004–2011 Good X
Romania X X
Russia + 5–30 2000–2012 Poor – 5–30 1980–2012 Poor
Serbia = 0 2000–2012 Medium = 0 1980–2012 Medium
Slovakia – 10–20 2000–2012 Poor – 30–50 1980–2012 Poor
Slovenia – 20–30 2001–2012 Medium – 30–50 1980–2012 Medium
Spain = 0 2006–2012 Good – 1980–2012 Medium
Switzerland = 0 2001–2012 Medium + 19–73 1990–2012 Medium
Turkey = 0 2000–2012 Poor – 0–19 1990–2013 Poor
Ukraine F 10–20 2001–2012 Medium F 10–25 1980–2012 Medium
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Table 5.5 (cont.)

166

European Environment Agency (2021)

Breeding population trend Breeding distribution trend

Short term Long term Short term Long term

Magnitude Magnitude Magnitude Magnitude

% min | % min | % min | % min |
Country Period Direction max (best) Period Direction max (best) Period Direction max (best) Period Direction max (best)

Austria 2007–2018 = N/A | N/A 1981–2018 – –80 | –50 2007–2018 – N/A | N/A 1981–2018 – –60 | –30
(10) (N/A) (–17) (N/A)
Belgium 2008–2018 = –25 | 16 1973–2018 + 8 | 67 (38) 2008–2018 = N/A 1973–2018 = N/A
(–4)
Bulgaria 2000–2018 – –40 | –20 1980–2018 = 0 | 5 (N/A) 2000–2018 = 0 | 5 (N/A) 1980–2018 = 0 | 5 (N/A)
(N/A)
Croatia 2007–2018 X N/A 1980–2018 X N/A 2007–2018 X N/A 1980–2018 X N/A
Cyprus 2007–2018 = 0 | 0 (N/A) 1980–2018 X N/A 2007–2018 + N/A | N/A 1980–2018 X N/A
(18)
Czech 2007–2018 – N/A | N/A 1980–2018 – N/A | N/A 2002–2016 – N/A | N/A 1986–2016 – N/A | N/A
Republic (–50) (–90) (–67.85) (–87.38)
Denmark 2006–2017 – –97.28 | 1990–2017 – –98.36 | 1996–2017 – N/A | N/A 1974–2017 – N/A | N/A
–85.42 –89.42 (–67.94) (–88.63)
(–93.51) (–95.77)
France 2007–2018 = 0 | 0 (N/A) 1980–2018 – –50 | –20 2007–2018 = 0|0 1985–2017 – –10 | –2
(N/A) (4.1) (N/A)
Germany 2004–2016 = –8 | 5 (–1) 1980–2016 = N/A | N/A 2004–2016 = –10 | 10 1980–2016 – –65 | –26
(–1) (N/A) (N/A)
Gibraltar 2001–2018 – –5 | –1 (–1) 1980–2018 – –5 | –1 (–1) 2001–2018 = 0 | 0 (0) 1980–2018 = 1 | 5 (5)
Greece 2007–2018 = N/A | N/A 1980–2018 = N/A | N/A 2007–2018 X N/A 1980–2018 X N/A
(0) (0)
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Hungary 2007–2018 + 20 | 50 1980–2018 X N/A 2007–2018 + N/A | N/A 1980–2018 + N/A | N/A
(N/A) (10) (20)
Italy 2007–2018 X N/A 1993–2018 + 130 | 300 2007–2018 = N/A 1993–2018 + 20 | 25
(N/A) (N/A)
Latvia 2007–2018 = 0 | 0 (0) 1980–2018 – –100 | 2013–2017 = 0 | 0 (0) 1980–2017 – –100 |
–100 –100
(–100) (–100)
Lithuania 2013–2018 = 0 | 0 (N/A) 1980–2018 = 0 | 0 | (N/A) 2006–2018 = 0 | 0 | (N/A) 1980–2018 = 0 | 0 | (N/A)
Luxembourg 2007–2018 + 0 | 30 (N/A) 1980–2018 – –70 | –50 2007–2018 – –30 | –10 1980–2018 – –50 | –30
(N/A) (N/A) (N/A)
Netherlands 2006–2017 = –7 | 11 (2) 1980–2017 – –45 | –43 2000–2015 = N/A | N/A 1977–2015 – N/A | N/A
(–44) (–8.66) (–18.28)
Poland 2007–2018 X N/A 1980–2018 – –67 | –54 2007–2018 X N/A 1980–2018 X N/A
(N/A)
Portugal 2009–2017 – N/A 1980–2018 X N/A 2005–2018 – N/A | N/A 1980–2018 = N/A
(–43)
Romania 2014–2018 U –8 | 21 1980–2018 X N/A 2007–2018 = N/A 1980–2018 = N/A
(N/A)
Slovakia 2007–2018 – –30 | –20 1980–2018 – –70 | –40 2007–2018 – –30 | –20 1980–2018 – –70 | –50
(N/A) (N/A) (N/A) (N/A)
Slovenia 2002–2017 X N/A 1980–2017 – –87 | –70 2007–2018 X N/A 1980–2018 – N/A | N/A
(N/A) (–18)
Spain 2007–2018 – N/A | N/A 1980–2018 – –40 | –20 2007–2018 = N/A 1980–2018 – N/A
(–25) (N/A)
UK 2004–2016 – N/A | N/A 1980–2016 – N/A | N/A 1989–2009 = 0.81 | 0.81 1970–2009 – –10.35 |
(–48.03) (–69.77) (N/A) –10.35
(N/A)

– Decrease; + Increase; = Stable; F Fluctuating; U Uncertain; X Unknown

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 Chapter 5: Distribution, Population Estimates and Trends

Figure 5.5 Breeding population sizes and short-term trends across Europe (BirdLife International
2015).

5.5 Overview per Country

The aim of this section is to give an overview of the species distribution within
individual countries. We have assembled within-country maps of Little Owl distribu-
tions for 77 countries. These maps are shown under each of the respective countries as
they are discussed in this section. Factors that limit the local distribution of the species
are given for each location whenever available.

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 5.5 Overview per Country

Figure 5.6 Breeding population sizes and long-term trends across Europe (BirdLife International
2015).

5.5.1 Belgium
5.5.1.1 DISTRIBUTION
The species was found in 86% of the country except the snow-rich High-Belgium, and
extensively wooded areas like the Ardennes and Belgian Lorraine in the south of the
country (Delmée 1988). The highest densities were observed in southern West-Flanders,
East-Flanders and Limburg (Vercauteren 1989). The Leemstreek region holds 40% and
the Zand-Zandleemstreek region 46% of the total population, the Kempen and Polders
regions were reported to have limited numbers in 1998–2000 (Van Nieuwenhuyse
et al. 2001e). The Little Owl in Flanders (northern half of Belgium) can be characterized

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 Chapter 5: Distribution, Population Estimates and Trends

along three gradients: (1) the gradient between open and closed landscapes, (2) the
gradient between dry and wet soils and (3) the gradient between sand and loam soils.
The species avoids extremely open (polders, lakes) and extremely closed (cities, forests)
landscapes, avoids soils with “excessive” and “bad” drainage and prefers loam soil to
sand. A mosaic-like landscape structure was found to be more important than the actual
land-cover types.
In Wallonia, the species is still found in all types of habitat, even in the Ardennes
and Lorraine. Some sectors seem to have been unoccupied for a long period of time,
e.g., some empty grid cells were already present in 1973–1977 (Delmée 1988) around
Saint-Vith and part of the Luxembourg Ardennes. In the recent Walloon Atlas project a
significant proportion of the surveyed grid cells are no longer occupied, with
fragmentation and a reduction in the distribution area emerging. The Little Owl may
possibly not have recovered from the severe winter of 1996–1997. Nevertheless, disap-
pearing pairs continue to be registered from 2000, at least in Lorraine.
In Brussels around 10 pairs are reported on the outskirts of the city (Weiserbs and
Jacob 2007).

5.5.1.2 POPULATION ESTIMATES

The species has decreased markedly, probably due to habitat destruction and pesticides.
Estimates indicate around 12 000 pairs in 1950, around 4000 pairs in 1972 (Lippens and
Wille 1972), around 7300 pairs during 1973–1977 (Delmée 1988), and 4500–6600 pairs
during 1981–1990, but decreasing (Manez in Tucker and Heath 1994). In 2003, the total
Belgian population was estimated at 12 500–14 000 pairs (BirdLife International 2004).
BirdLife International (2015) estimated the total population at 8000–13 000 pairs in
2008–2012. In 2013–2018, the population was estimated at 7900–12 200 breeding pairs
(European Environment Agency 2021).

5.5.2 Flanders
Van Nieuwenhuyse et al. (2001c) developed recent population estimates for Flanders.
Here, they estimated the number of occupied grid cells (500
500 m) with a probabil-
ity of Little Owl presence of  50%. Their results indicated a range of 12 527 (Figure 5.7)
to 16 046 occupied grid cells for the models without regional stratification. When
summing all of the minimum and maximum estimates of stratified models respectively,
a range of 13 646–17 361 occupied cells was obtained. Both estimates indicated two
main regions in absolute numbers, i.e., 40% of the Flemish Little Owl population was
situated in the Leemstreek ecological region and 46% in the Zand-Zandleemstreek
region, while the Kempen and Polders regions had only limited owl numbers. The
highest average probabilities calculated were observed principally in the Leemstreek.
The total Flemish Little Owl population was estimated at 6000–10 000 pairs (Van
Nieuwenhuyse 2004).
From 2000–2018 the percentage of built-up areas (principally housing) increased by
3.9 percentage points from 24.4% to 28.3% and 7 percentage points in the last 30 years
(Binnemans et al. 2021). This increase was most observed around the larger cities,

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 5.5 Overview per Country

Figure 5.7 Little Owl distribution in Flanders (2001) (Van Nieuwenhuyse et al. 2001c).

within the Flemish Diamond (Antwerp, Leuven, Brussels, Ghent) and in coastal areas.
In 8241 500
500 m grid cells sampled, the overall occupation rate was 37%
(2018–2020) compared to 38.6% (1998–2000). Of 3053 re-visited cells (of 12 152 visited
cells in 2000) a statistically significant decline of 10 percentage points from 46.1%
(1998–2000) to 36.1% (2018–2020) was observed (Figure 5.8).
This evolution is different locally, high-density locations in 2000 (e.g., Lievegem
Figure 5.9) feature the highest losses, while suboptimal regions (e.g., Meulebeke)
showed important increases with the highest counts since 1988 (Figure 5.10).
The re-visits scores were worst, possibly because of preferred sampling in high-
density locations in 2000. The largest declines were observed in the Leemstreek (59%
occupation in 2000 to 35% occupation in 2020) and Zand-Zandleemstreek (45.8% in
2000 to 39.7% in 2020) which scored best in 2000. Possible reasons for this negative
progression are thought to be habitat loss, leading to losses of breeding cavities and
food shortage, the application of new pesticides, e.g., neonicotinoids, traffic mortality
and possibly the recent increase in Stone Marten and Tawny Owl populations
(Binnemans et al. 2021), of which the latter was observed to have a significant negative
impact on Little Owls in Switzerland (Michel et al. 2015). More research is needed to
unravel the relative importance of each of these factors (see Chapter 10, Population
Regulation).

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 Chapter 5: Distribution, Population Estimates and Trends

Occupation repeated visits 2000 2020

REOCCUPIED (634)
NEW OCCUPATION (469)
NEVER OCCUPIED (1177)
DISAPPEARED (773)

Figure 5.8 The evolution of the re-occupation of Little Owls in Flanders. Red circles indicate declines
in the occupation, green circles increases (after Binnemans et al. 2021).

45

40
40
37 y = –2.5357x + 39.429
35
R² = 0.5367

30 30
30
27

25
21
20
20

15

10

0
2014 2015 2016 2017 2018 2019 2020
Lievegem Noord Lievegem Oost Lievegem Zuid Linear (Lievegem Oost)

Figure 5.9 Local evolution of responses to standardized playback in three areas of Lievegem, Flanders,
2014–2020 (Lode Van de Velde personal communication).

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 5.5 Overview per Country

140

y = 13.457x + 34.733
120 R² = 0.7519

100

80

60

40

20

0
1988 1994 2000 2006 2012 2018

Figure 5.10 Local evolution of responses to standardized playback in Meulebeke, Flanders 1988–2018
(own observations).

5.5.3 Wallonia
Large differences are found between different sectors. The species is still rather common
in Middle Belgium, Condroz and in the Pays de Herve and Fagne-Famenne. The most
observed abundance classes are 11–20 and 21–40 pairs/40 km²; with two grid cells with
41–80 pairs. The species is rare along the Meuse valley with most grid cells having only
6–10 pairs/40 km²). It is still present, but very rare (between 1–5 and 6–10 pairs/40 km²),
around the large urban centers (Liège, Huy, Namur, Charleroi, La Louvière, Mons). The
urban centers include some meadows and gardens that still offer some habitat. In
Ardenne and Lorraine the species is very rare with densities around: 0 up to 1–5 pairs/
40 km² and a few grids with 6–10 pairs/40 km².
The total estimate for Middle Belgium for 2000–2003 was 1800–2400 pairs, for
Condroz, Fagne-Famenne-Pays de Herve 950–1300 pairs, Ardennes 200–400 pairs and
Lorraine 30–60 pairs. The total Walloon population is thus estimated at 3000–4200
pairs (J. P. Jacob personal communication) (Figure 5.11).
The most recent estimate for Middle Belgium is 2442 pairs (66%), Condroz 629
(17%), Fagne-Famenne 370 pairs (10%), Ardennes 259 pairs (7%), Lorraine 30–60 pairs.
The total Walloon population is estimated at 3700 pairs (Bultot and Van Nieuwenhuyse
in Jacob et al. 2010).

5.5.3.1 TRENDS
In Wallonia, 71% of grids cell were still occupied in 2001–2007 (Jacob et al. 2010) as in
1973–1977 (Delmée 1988). In Famenne, densities decreased between 1985 and 1989
(0.4 pairs/km²) and in 2001–2006 (0.2 pairs/km²). Most of the changes in distribution
(new and abandoned grid cells) are situated in low-density areas like Ardenne, Lorraine
and Condroz. Abandonment was found in the Pont region, Grune, Awenne and

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 Chapter 5: Distribution, Population Estimates and Trends

1 to 5 breeding pairs : 128 (36 %)

6 to 10 : 100 (28 %) 3700 breeding pairs
11 to 20 : 84 (24 %)
21 to 40 : 39 (11%)
41 to 80 : 2 (1%)
Total : 353 (100 %)

Figure 5.11 Little Owl distribution in Wallonia in 2001–2007 (Bultot and Van Nieuwenhuyse in: Jacob
et al. 2010).

Ochamps, re-occurrence in Waremme, Nassogne and Moyenne-Semois (Bultot and Van

Nieuwenhuyse in Jacob et al. 2010).

5.5.3.2 DENSITIES
High densities of Little Owls have been found in larger areas in or near Belgium: e.g., 3.4
calling individuals/km² in Geraardsbergen with 42 of 91 km² squares with at least four
individuals; four calling individuals/km² in Herzele with 24 of 45 km² squares with at
least four individuals; 2.17 calling individuals/km² in Meulebeke with 8 of 29 km²
squares with at least four individuals (own observations). Locally high concentrations
are also found in Condroz, with two of 33 km² squares with four calling males (Lucas
1996). North of Charleroi densities up to 6–8 pairs per village are observed in a region of
300 km², where local conservation initiatives were very successful (Bultot et al. 2001).
At Theux, 15 sites were occupied in high-stem orchards (0.5 calling males/km²). Near
Ath, 75 playback sessions in 40 km² yielded 35 calling males (Bultot and Van
Nieuwenhuyse in Jacob et al. 2010).

5.5.4 Netherlands
5.5.4.1 DISTRIBUTION
The Little Owl was observed in 52–54% of the breeding bird atlas-surveyed grid cells
between 2013 and 2015 (Van Harxen et al. in Sovon Vogelonderzoek Nederland 2018)
(Figure 5.12). In the two northern provinces, Groningen and Friesland, the species is
almost completely absent, but in the southwestern part of Drenthe a remarkable
recovery took place due to the activities of volunteers placing nestboxes. In the western
part of the country some population clusters remain, in the former fruit-growing areas

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 5.5 Overview per Country

Figure 5.12 Little Owl breeding evidence 2013–2015 in the Netherlands. Reproduced with
permission from Sovon Vogelonderzoek Nederland 2018.

of West-Friesland, and in the Beemster, the Green Heart and Hoekse Waard. In Zeeuws-
Vlaanderen, the part of Zeeland bordering Flanders, the species showed, though still
widely spread, a decline of almost 35% between 2007 and 2019, mainly due to habitat
loss (De Smet 2019). In the same years the well-conserved small population in Zuid-
Beveland (north of the Westerschelde) expanded its area and increased from less than
40 breeding pairs in 2007 to more than 70 pairs in 2020 (0.5 pairs/km2) and is about to
re-colonize Walcheren (P. Boelee personal communication). Riverine areas along the
Rhine and Waal and the former high-stem orchard areas in the Betuwe have become
less important. Nowadays the stronghold of the population is found in small-scale
landscapes of sandy soils in the eastern and southern parts of the country, especially
in the Achterhoek, Liemers and the adjacent parts of Twente and Salland. The species is
absent on the Wadden Isles, in the IJsselmeerpolders and in the vast woods of
the Veluwe.

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 Chapter 5: Distribution, Population Estimates and Trends

5.4.4.2 POPULATION ESTIMATES

The species has declined since around 1935, mainly due to habitat changes and loss of
breeding places. The first estimates of 6000–8000 pairs for 1973–1977 (Teixeira 1979)
were probably too low. In the second Dutch atlas (Sovon Vogelonderzoek Nederland
1987) a new estimate of 8000–12 000 pairs was given for 1978–1983. Osieck and
Hustings (1994) estimated 10 000–14 000 pairs for 1979–1985 and 9000–12 000 pairs
for 1989–1991. In 1998–2000 the population was estimated at 5550–6500 pairs (Sovon
Vogelonderzoek Nederland 2002, BirdLife International 2004). The latest estimates
(2012–2015) were published in the most recent breeding bird atlas: 7500–8500
(Sovon Vogelonderzoek Nederland 2018). The differences between the estimates in
the last 50 years are at least partly due to difficulties in correctly establishing numbers.
Recently, well-researched regions, e.g., Achterhoek, Liemers and parts of Twente,
Brabant and Limburg, feature the highest current densities. The northern parts of the
Netherlands (Groningen, Friesland) and the western coastal regions have the
lowest densities.
Between 1973 and 1977 (Sovon 1987) and 2013–2015 (Sovon Vogelonderzoek
Nederland 2018) the species lost 17% of its distribution area expressed in atlas grid
cells (1089 to 906 of 1685), which equates to a geographic range reduction of roughly
4.575 km2 (Figure 5.13).
The situation is striking in the northern provinces, such as in Drenthe, where the
species is still absent in the central and eastern parts of the province. The number of pairs
in Friesland ðn ¼ 5–10Þ and Groningen ðn ¼ 5–10Þ is only a fraction of what they once
were. In Groningen there were up to 300–400 pairs in the 1960s (van’t Hoff 2001). The
presence of the species in 111 new atlas squares all over the country in the latest atlas is
probably due to more intensive survey work rather than an expansion of the owl’s range.
It remains unclear what the exact causes of the decline in distribution area are, as
reproduction as well as survival showed no significant decrease over the last 50 years.
However, the biggest decrease took place in the northern and western parts, where almost
no connection to other populations (apart from the eastern part of the country) is possible
due to geographical borders (sea) or low densities elsewhere (northern Germany). So
population dynamics in combination to habitat loss may have played a major role.

5.5.4.3 DENSITIES
Within the Netherlands, owl densities differ markedly (Figure 5.14). In the northwes-
tern part of the country, densities vary from 0.04–0.4 pairs/km². In the half-open
cultivated areas in the southeast, densities can reach 1.0–1.5 pairs/km² over larger areas
(Stroeken et al. 2001). A density of 3.43 pairs/km2 was found in a plot of 6.7 km2 near
Winterswijk (Van Harxen 2021).

5.5.5 Luxembourg
5.5.5.1 DISTRIBUTION
The species is limited to a population in the western and central part of the country and
a smaller one in the southeastern part. Dramatic declines took place between 1990 and

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 5.5 Overview per Country

Figure 5.13 Relative abundance change in breeding 2013–2015 versus 1998–2000. Reproduced with
permission from Sovon Vogelonderzoek Nederland 2018.

2004 in the northern part of the country. The main population is nowadays found in
the western part of the country, where a species action plan was set up by Station
Biologique Sicona and partners (Sicona 2021).

5.5.5.2 POPULATION ESTIMATES

In 1960, the population was estimated at 3400–4200 pairs (Hulten and Wassenich
1961). The Atlas of Breeding Birds in Luxembourg (Melchior et al. 1987) estimated
the population at 300–500 pairs between 1976 and 1980. As of 2003 the population was
estimated at 45–85 pairs, with only 15 pairs left in the northern part of the country.
Dramatic declines took place up to 2005 in the western and northern parts of the
country. The best remaining populations are reported in the southeast. In 2004, several
conservation initiatives started with intensive monitoring and nestboxes, yielding only
10 confirmed territories. The population hence was estimated in 2004 at 10–20 pairs.
(P. Lorgé and E. Melchior personal communication). BirdLife International (2015)

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 Chapter 5: Distribution, Population Estimates and Trends

Figure 5.14 Relative abundance in breeding 2013–2015. Reproduced with permission from Sovon
Vogelonderzoek Nederland 2018.

estimated the total population at 12–25 pairs in 2008–2012. In 2013–2018, the popula-
tion was estimated at 20–30 breeding pairs (European Environment Agency 2021).
Using recent surveys together with new data on distribution and species action
plans, the population in 2021 was estimated at 40–45 pairs (Lorgé and Melchior 2010).

5.5.6 United Kingdom

5.5.6.1 DISTRIBUTION
Previously a rare vagrant to the British Isles from the continent, the Little Owl was
introduced into England in the 1870s and 1880s and established a feral breeding popula-
tion (Marchant et al. 1990). Initial releases were made in Kent and Northamptonshire,
and by the close of the 1800s there were two separate breeding groups, in southeast
England and the east Midlands. The spread was subsequently very rapid, aided by further
releases, and soon Little Owls were present west to the River Severn and north to the

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 5.5 Overview per Country

River Trent. By 1900, Little Owls were regular breeders in several English counties and a
period of rapid expansion followed, lasting into the 1930s (Parkin and Knox 2010). The
first Welsh breeding record came in 1916 and that for Scotland, where the species has
subsequently lost its foothold, came in 1958. There have been only four records from
Ireland (Dempsey and O’Cleary 1993), none during 2007–2011. By 1925 they had
colonized much of Wales and expanded northwards to the central parts of Lancashire
and Yorkshire. Durham and Northumberland were reached in the 1930s, the Lake
District in the 1940s, southeast Scotland (Borders and Lothian) from 1958, then
Dumfries and Galloway from the 1970s (Marchant et al. 1990).
The winter and breeding-season distributions are similar, reflecting the sedentary
nature of the species in Britain (Balmer et al. 2013) (Figure 5.15). The species is largely
restricted to England and the Welsh borders, with isolated populations in northwest
and coastal Wales. In parts of the Little Owl’s native range, low temperatures and lying
snow reduce survival and this may explain why the British uplands remain unoccupied.
Hence, the Little Owl remains extremely rare in Scotland and has not successfully
penetrated north of the Edinburgh–Glasgow line.

5.5.6.2 POPULATION ESTIMATES

Though peripheral expansion continues, the initial explosive phase after the introduction
had slowed by the mid-1930s and was followed by reports of decreases in density over large
parts of central and southern England. A run of severe winters in the 1940s might have
adversely affected the Little Owl. Some improvement occurred in the early 1950s, but there
was a further decline during 1955–1963, which coincided with the organochlorine pesti-
cide era and the cold winters of 1961–1962 and 1962–1963 (Parslow 1967, Sharrock 1976).
The Common Bird Census (Marchant et al. 1990) showed fairly regular spacing of peaks
and troughs between 1964 and 1988. The peak years have been at intervals of three to five
years and an average of four years. This has all the appearance of being cyclical. It seems
possible that reports of Little Owl declines after the mid-1930s included such cyclical
oscillations reflecting natural variations in population levels and may have contributed
to other low points mentioned in the literature. Superimposed onto the cyclical pattern
has been some longer-term variation in Little Owl densities, though it cannot yet be said
that an overall trend has been confirmed. After recovery in the early 1960s, the average
population level varied little until 1975, but an increase in the late 1970s and early 1980s
produced the highest-ever index figure in 1984. This overall increase occurred despite two
very cold winters, 1978–1979 and 1981–1982, which seemed to have had little effect;
1982 was a natural low point in the cycle. In 2000 the British Trust for Ornithology and the
Hawk and Owl Trust completed a three-year national survey of Barn Owls within the UK
(Toms et al. 2000). As part of this project they also gathered information on breeding Little
Owls, and have produced a national population estimate of pairs. The Little Owl popula-
tion estimates were calculated using a stratified random sampling approach based on
1100 tetrads (2
2 km squares) spread across the UK. A boot-strapping procedure was
then used to calculate 95% confidence intervals for the estimates produced from the
sampling procedure. Individual national estimates were calculated for each of the three
survey years to allow for any short-term climatic effects on breeding performance. The

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 Chapter 5: Distribution, Population Estimates and Trends

Breeding Distribution 2008–11

Non-breeding

Possible

Probable

Figure 5.15 Little Owl distribution in the UK in 2007–2011 after Balmer et al. (2013). Map reproduced
from Bird Atlas 2007–11, which is a joint project between, BTO, BirdWatch Ireland and the Scottish
Ornithologists’ Club. Map reproduced with permission from the British Trust for Ornithology.

estimates of Little Owl pairs were found to be similar across each of the three years
(numbers of pairs (range): 1995: 7103 (5136–9136); 1996: 6142 (4075–8301); 1997: 5865
(4337–8186)). The estimates of the models using landscape data were comparable: 1995:
7468; 1996: 6653; 1997: 6253. The estimates were also re-calculated for the known
breeding range of the species within the UK (rather than for the whole UK). These data
suggest a population estimate of 4000–8500 breeding pairs and represent the first reliable
and repeatable estimate for the UK. The previous estimate of 6000–12 000 breeding pairs
(Gibbons et al. 1996) was derived from an Atlas survey for all breeding species, rather than
specifically looking at Little Owls and may be based on less accurate mean breeding

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 5.5 Overview per Country

Breeding Relative Abundance 2008–11

High

Low

Figure 5.16 Little Owl breeding abundance in the UK in 2008–2011 after Balmer et al. (2013). Map
reproduced from Bird Atlas 2007–11, which is a joint project between, BTO, BirdWatch Ireland and the
Scottish Ornithologists’ Club. Map reproduced with permission from the British Trust for Ornithology.

densities. BirdLife International (2004) estimates the total population at 5800–11 600
pairs. In 2013–2018, the population was estimated at 2338–4888 (3613) breeding pairs
(European Environment Agency 2021). Abundance maps (Figure 5.16) show that the Little
Owl currently is more abundant within East Anglia, the Midlands and central northern
England than in the more westerly parts of its breeding range. Little Owls generally are
most abundant within areas of mixed habitat, where there are small parcels of farmland,
woodland edge and hedgerows, which has also been confirmed by Balmer et al. (2013) in
Britain.

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 Chapter 5: Distribution, Population Estimates and Trends

5.5.6.3 TRENDS
Whereas there is evidence of expansion within northeast England since 1988–1991, this
was outweighed by the marked range contraction throughout southwest England and
across much of Wales between 1968–1972 and 1988–1991. Since the 1980s, Little Owls
have undergone declines of 47% in Gwent and 81% in Pembrokeshire in the numbers of
occupied tetrads (Venables et al. 2008). The abundance change map (Figure 5.17) shows
similar losses throughout the western part of the range since the 1988–1991 Breeding

Breeding Distribution Changes all atlases

Figure 5.17 Little Owl breeding distribution change in the UK between 1968–1972 and 2008–2011
after Balmer et al. (2013). Map reproduced from Bird Atlas 2007–11, which is a joint project between
BTO, BirdWatch Ireland and the Scottish Ornithologists’ Club. Map reproduced with permission from
the British Trust for Ornithology.

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 5.5 Overview per Country

Atlas, in line with a population decline of 40% during 1995–2010 (BBS Report 2011). It is
thought that changing farming practices might have depressed juvenile survival by redu-
cing their food supply (Woodward et al. 2020). The decreased fledging success reported in
Denmark (Thorup et al. 2010) is not currently apparent in Britain (Woodward et al. 2020).
Balmer et al. (2013) report a 40-year change of –11% and a 20-year index change of –0.22.
The breeding evolution change between 1968–1972 and 2007–2011 shows mainly aban-
donment in Wales (205 grid cells) and gains in northern England.

5.5.7 Germany
In Germany, the Little Owl is a synanthropic bird that is found in lowlands with
grassland and old trees, close to villages and farms with traditional orchards and
pastures. Currently it occurs in three distinct regions (Figure 5.18): one region extends
from the Lower Rhine region to the Cologne lowlands and to the Münsterland region,
another hotspot is the Geest region in Schleswig-Holstein and the third region ranges
from the Upper Rhine valley to the uplands of western Hesse and the Neckar valley. The
continental northeast German lowland is sparsely populated with concentrations
limited to 8–20 territories per 11
11 km grid cell in the area of the Middle Elbe.
Smaller occurrences also exist in the Recknitz valley, the Lower Havelniedringen, the
Havelländisches Luch, the Fiener Bruch and the Belziger Landschafwiesen, as well as in
the northern Harzvorland and the western Sächsischen Hügelland (Gedeon et al. 2014).

5.5.7.1 POPULATION ESTIMATES

The Little Owl has shown a marked decline in Germany, probably due to habitat
changes. Numbers were severely affected by hard winters (Glutz Von Blotzheim and

Territories

German Grid TK25

Figure 5.18 Little Owl distribution 2005–2009 in Germany per 11 by 11 km grid cell (after Gedeon
et al. 2014) and 2011–2016) on the German Grid TK25. Reproduced with permission from
Dachverband Deutscher Avifaunisten (DDA) e.V.

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 Chapter 5: Distribution, Population Estimates and Trends

Bauer 1980, Bauer and Thielcke 1982). Estimates for the former East Germany were
250–300 pairs for 1985–1986 (Schönn 1986), 1035 pairs in North Rheinland for 1999
(M. Wink perssonal communication), 5000–10 000 pairs in 1985 in the entire country
(Manez in Tucker and Heath 1994). The total population around the year 2000 was
estimated at 5800–6100 for 1996–1999 with only a minor decrease for 1999–2002 to
6000 pairs (Bauer et al. 2002). A detailed meta-analysis using all possible data sources
yielded a population estimate of 7000–7600 pairs (Illner in Van Nieuwenhuyse et al.
2008). BirdLife International estimated the total population at 5800–6100 pairs in 2004
(BirdLife International 2004) and 8000–9500 territories in 2005–2009 (Gedeon et al.
2014), 7500–9000 pairs (BirdLife International 2015). In 2013–2018, the population
was estimated at 7500–8500 breeding pairs (European Environment Agency 2021). The
population stronghold is situated in Nordrhein-Westphalia, which harbors 75% of the
national population. Since the mid-twentieth century, harsh winters and habitat
changes have caused population declines accompanied by range contractions. The
long-term population trend is negative. In recent decades, the population size has
increased again.

5.5.7.2 TRENDS
The population is declining in the long term and increasing in the short term (1988–2009)
(Sudfeldt et al. 2013). In Germany as well as the other Central European countries there
have been strong reductions and land losses since the middle of the twentieth century
(Schönn 1998, Haase 2001, Jöbges and Franke 2006, Brandt et al. 2012). Causes of decline
are hard winters, habitat changes, loss of breeding cavities associated with a decrease in
food availability. Especially strong declines were recorded in Eastern Germany (Nicolai
1993, 2000, Mebs and Scherzinger 2000). The population decreased there between
1980 and 2004 from 470 to around 40 pairs (Jöbges 2004). Larger concentrations with a
large number of individuals were retained in parts of the northwest German lowlands and
the western low mountain range (Rheinwald 1993, Bauer and Berthold 1996).
Intensive promotional and protective measures resulted in population recoveries in
these regions from the 1980s onwards (Hölzinger 2001, Struwe-Juhl 2003, Hessische
Gesellschaft für Ornithologie und Naturschutz 2010), favored by the mild winters
towards the end of the 1980s (Bauer et al. 2005). In North Rhine-Westphalia a signifi-
cant recovery of the population occurred in the 1990s, but did not result in a full
recovery to 1960 levels (Jöbges and Franke 2006). Significant increases were also
recorded in Hesse, where the population was less than 250 territories in 1976, increased
to 1000 in 2008 (Hessische Gesellschaft für Ornithologie und Naturschutz 2010). Also
for Baden-Württemberg a strong increase of over 50% between 1980 and 2004 was
documented (Hölzinger et al. 2007) (Figure 5.19). For re-settlement as well as for
population conservation population re-inforcement initiatives took place in
Vorpommern and the northern Harzvorland; however, these were mostly unsuccessful
(Haase 2001, Behl 2006, Ryslavy and Mädlow 2008, Fischer and Dornbusch 2010).
In comparison with previous population estimates of 6000 pairs for the period
around 1985 (Rheinwald 1993), 4900–7300 pairs around 1995 (Witt et al. 1996) and
5800–6100 pairs around 2000 (Bauer et al. 2002) significantly higher population

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 5.5 Overview per Country

Population increase

Figure 5.19 Little Owl population increase 1985–2009 in Germany (after Gedeon et al. 2014).
Reproduced with permission from Dachverband Deutscher Avifaunisten (DDA) e.V.

estimates were reported for 2005, up to 8200–8400 territories (Südbeck et al. 2007). In
the most recent atlas (Gedeon et al. 2014) estimates are even higher. However, in some
areas there is still a decline in the population and loss of area (Jöbges and Franke 2006,
Brandt et al. 2012).
The historical highest rates of decline (–3.0 to –4.3% per year) were found in the
federal states north and east of the stronghold in Northrhein-Westphalia; moderate
rates of decline were found in the southern federal states (–1.2 to –1.9%). The central
state of Hessen, neighboring Northrhein-Westphalia, is the only area with an apparent
increase over the last decades. A common feature is also that federal states with yearly
decline rates of more than 1.7% all lost significant parts (>about 50%) of the breeding
range of the species, even Hessen (with a population increase asssumed) shows a loss of
about one-third of the breeding range occupied in the 1970s. Even in Northrhein-
Westphalia, some parts of the former breeding range were lost, e.g., the higher parts
in south Westphalia and parts of the industrial area Ruhrgebiet.

5.5.7.3 DENSITIES
In Germany densities recorded below 1 territory/km² were reported from Saxony, with
12 pairs on 20 km² (Augst and Manka 1997). While higher densities were once found in

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 Chapter 5: Distribution, Population Estimates and Trends

Northrrhine-Westphalia (Exo in Galbraith et al. 1992) the situation has changed in the
Aachen region where only 34 calling males were recorded on 160 km² (Toschki 1999).
In Bönningheim, Landkreis Ludwigsburg, densities up to 1 pair/km² were recorded
locally (Eick 2003), with an average of 0.17 pairs/km² for the entire region (120 pairs
across 690 km² (Keil 2001). In the orchards of southwest Germany, a density of 3.5
breeding pairs/km² was found, with a nestbox density of 1 box per 15 ha (Otto and
Ullrich 2000).
The population center in northwest Germany, which is dominated by the Atlantic
lowland ranges from the Niederrheinischen Tiefland and the Kölner Buch far into
Münsterland, can reach densities of 21–50 or 51–150 territories per grid cell (Gedeon
et al. 2014). This occurrence currently comprises around 70% of all breeding pairs in
Germany and, with adjacent stocks in the Netherlands and Belgium, forms one of the
most important density centers in Central Europe. The populations in the Oldenburger
Münsterland have up to 21–50 territories per grid cell, largely uninterrupted
towards the East Frisian Oldenburg Geest. More isolated occurrences can also be
found within the Ems and Wesermarschen, as well as along the Aller and Leine. The
concentration within the low mountain range extends from the Oberrheinischen
Tiefland to Westhessische Bergland and the central Neckar area. With 51–150 territories
per grid cell there are density centers in the Wetterau, in the Rhine-Main area and in
the Neckar valley near Stuttgart. Separate from this are smaller clusters in the area of
the Bliesgau, the Saargau, north of the Moselle and in the Grabfeld, east of the
Rhön. Isolated occurrences with 1–3 territories per grid cell are found in the
Weserbergland and the Fulda valley on the Vogelsberg, as well as areas along the
Tauber, in Steigerwald and in the Zittauer Becken. The only occurrence in the Alpine
foothills is located in the Bodenseebecken near Markdorf. A smaller concentration is
found on the Schleswig-Holstein Geest, in particular in the Hohe Geest and in the river
plains of Eider and Stör, where densities of 8–20 territories per grid cell were reported
(Gedeon et al. 2014).

5.5.8 Denmark
5.5.8.1 TRENDS
An extreme restriction in the distribution of the Little Owl has taken place during the
last four decades and the species was only found in 2% of the atlas squares of Atlas III
2014–2017, half of them confirmed as breeding. The total number of breeding records
has been reduced by 79% between the Atlas surveys in 1993–1996 and 2014–2017
(Vikstrøm and Moshøj 2020) (Figure 5.20).

5.5.8.2 DISTRIBUTION
The major part of the Danish Little Owl population breeds in north and west Jutland.
The most import areas are Himmerland (between Ålborg and Mariager), Salling (north
of Skive) and the central part of Ringkøbing County. Outside these areas, small popula-
tions are found in Vendsyssel (north of Ålborg, between Kattegat and Fjerritslev, north
to Hjørring), in the region between Randers Fjord and Mariager Fjord, on Djursland

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 5.5 Overview per Country
(a) Breeding Distribution
Confirmed/Probable
Possible

1971–1973

(b) Breeding Distribution

Confirmed/Probable
Possible

1993–1996

Figure 5.20 Historical distribution of the Little Owl in Denmark in (a) 1971–1973, (b) 1993–1996,
(c) 2003 and (d) 2014–2017 (after DOF, BirdLife Denmark and the Museum of Zoology, Copenhagen).
Large dots represent confirmed breeding, small dots possible breeding. Reproduced with permission
from Dansk Ornitologisk Forening/ BirdLife Denmark (DOF BirdLife).

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 Chapter 5: Distribution, Population Estimates and Trends

(c) Breeding Distribution

Confirmed/Probable
Possible

2003

(d) Breeding distribution

Confirmed
Probable
Possible

2014–2017

Figure 5.20 (cont.)

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 5.5 Overview per Country

(between Randers/Århus and Grenå) and in the western part of Ribe County. In the
counties of Sønderjylland and Vejle only a few scattered breeding records exist. In Fyn
only one confirmed breeding record and another possible one has been recorded. The
Little Owl was found breeding in Falster in 1937 but has probably not bred on the
islands east of Store Bælt (Great Belt) since the 1940s (Grell et al. 1998). In 2021 the
major part of the remaining Danish Little Owl population was breeding in northern
Jutland. The most important area was Himmerland (between Aalborg and Hobro).
Outside this area only a few scattered breeding records exist annually (Vendsyssel north
to Aalborg and in the central and southwestern part of Jutland close to Esbjerg
(Vikstrøm and Moshøj 2020, Sunde 2021).

5.5.8.3 POPULATION ESTIMATES

Fewer grazed pastures close to appropriate buildings for nesting, together with the
general decrease in biological diversity caused by agricultural intensification, possibly
pesticides and over-fertilizing in open country, have been proposed as some of the
main causes of decline (Grell et al. 1998). The species also seems to be quite vulnerable
to the increasing levels of faster motorized traffic over the years (Glue 1971a). Lack of
good nesting sites due to better maintenance of buildings and predation by Stone
Martens (Martes foina) are also possible problems. BirdLife International (2004)
estimated the total population at 100–200 pairs. BirdLife International (2015) estimated
the total population at 43 pairs in 2008–2012. In 2013–2018, the population was
estimated at 10 breeding pairs (European Environment Agency 2021). In
2019 and 2020 only 9–11 pairs were known with an additional seven or eight single
birds, so the Little Owl is very close to extinction in Denmark (Sunde 2021). This is a
very critical situation and most of the pairs are fed by food supplementation with small
dead chicks, some only in the breeding season and others all year around (Sunde et al.
2021).

5.5.8.4 TRENDS
An extreme reduction in the distribution of the Little Owl has taken place during the
last two decades of the twentieth century. The total number of breeding records has
been reduced by 65% during 1993–1996, compared with the results of the first atlas
project in 1971–1974. Approximately 300 atlas squares distributed throughout the
area of Jutland and Funen that reported breeding in 1971–1974 did not report any
breeding activity during 1993–1996. In particular, a high proportion of the deserted
breeding areas were observed in eastern and southern Jutland and Funen. But the
species has also apparently disappeared from Thy (north of Limfjorden and west of
Fjerritslev) and on Mors, in western Vendsyssel, and in middle and western Jutland
(Grell et al. 1998). Further reduction in distribution has taken place over the last four
decades and Little Owls were only found in 2% of the atlas squares in Atlas III
2014–2017, half of them confirmed as breeding. The total number of breeding records
has been reduced by 79% between the Atlas surveys in 1993–1996 and 2014–2017
(Vikstrøm and Moshøj 2020).

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 Chapter 5: Distribution, Population Estimates and Trends

5.5.9 France
5.5.9.1 DISTRIBUTION
In France, the Little Owl is a sedentary species, present across the whole country except
the mountains. Yeatman (1976) indicated that the mountains where the snow stays for
a long time are not suitable for the species. The species breeds up to 1080–1100 m on
the Causse Méjean in Lozère and up to 1155 m in Haute-Loire (Juillard et al. 1990). The
Little Owl is absent from the islands of Brittany (Guermeur and Monnat 1980), but is
very common on the island of Oléron (Bavoux and Burneleau 1983, Bretagnolle et al.
2001). The species is very rare in Corsica, where just a few cases of breeding have been
recorded in the last century, in the middle-east plain, near Alerià (Thibault 1983,
Thibault and Bonaccorsi 1999). The last breeding record for Corsica was in 2000
(Mastrorilli 2000). The species is believed to compete with the Scops Owl (Otus scops)
on Corsica, and is present on the island of Frioul, not far from Marseille (Cheylan 1986).
Population strongholds are found in the west between Normandy, Pays-de-la-Loire and
Midi-Pyrénées, northeast of Massif Central (Auvergne and Rhône-Alpes) and along the
Mediterranean Sea between Provence and Pyrénées-Orientales. Elsewhere densities
decline, especially in the northeast between Portarlier and Etretat (Figure 5.21)
(Hameau in Issa and Muller 2015). Most of the adults remain in their territories in
autumn and winter. Juveniles disperse from September to October, principally within a
radius of 10 km around their place of birth. The distribution in France reflects the
specific situation where Mediterranean populations are inter-connected to populations

Breeding evidence
Confirmed (1197)
Probable (1039)
Possible (716)

Figure 5.21 Little Owl distribution in France 2005–2012 (after Issa and Muller 2015).

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 5.5 Overview per Country

in the neighboring countries of Spain, Italy and Belgium. For most of France, the Little
Owl is distributed in population clusters, sometimes irrespective of the apparent quality
of the habitat (Génot and Lecomte 1998).

5.5.9.2 POPULATION ESTIMATES

The French population of the Little Owl was estimated at 5000–50 000 pairs at the
beginning of the 1990s (Génot 1994b) and 11 000–35 000 pairs in 1998 (Génot and
Lecomte 1998). A detailed analysis was done by Génot and Lecomte (1998) yielding
population estimates per administrative district (Table S5.4). The total minimal popu-
lation at the end of the 1990s was estimated at 10 330–35 000 pairs by summing partial
estimates (Génot and Lecomte 1998). BirdLife International (2004) estimated the total
population at 20 000–60 000 pairs. Issa and Muller (2015) estimated the population at
25 000–50 000 pairs in 2009–2012, BirdLife International (2015) estimated the total
population at 21 000–50 000 pairs in 2008–2012. In 2013–2018, the population was
estimated at 25 000–50 000 breeding pairs (European Environment Agency 2021).

5.5.9.3 DENSITIES
In 1997, Camuzat and Faucon (unpublished data) found 70 calling males in an area of
44 km² (1.6 males/km²). In 1998, Vivier and Telle (unpublished data) recorded
115 calling males in an area of 95 km² (1.2 males/km²) in meadows with pollard
willows in northern France.
In 1993, Geslin (unpublished data) recorded 24 calling males in an area of 12 km²
(2 males/km²) in the Loire valley and Gailliez (unpublished data) noted 136 calling
males/133 km² (1.02 males/km²) in orchards and hedges of northern France. One
territory/km² was found in Bouches-du-Rhône (Barthelemy and Bertrand 1997). In
1999–2000, 157 calling males/320 km² were recorded in western France, with the
highest average density in the edge of the “bocage” (0.62/km²) and the lowest average
density in the sandy dunes area (0.05/km²) (Signoret 2002). Crespon and Chretienne
(unpublished data) recorded 88 calling males/1175 km² in the natural regional park of
Perche, with average densties of 0.07–0.83/km². Locally high densities of 6.7 calling
males/km² were found on Île d’Oléron (18 km²) (Bretagnolle et al. 2001). Monitoring in
10 regional parks revealed densities ranging from 57 calling males/44 km² in Livradois-
Forez (1.3 males/km²) to 7 calling males/200 km² in Lorraine (0.035 males/km²) (Génot
in Van Nieuwenhuyse et al. 2008).

5.5.9.4 TRENDS
Issa and Muller (2015) reported an evolution in distribution patterns between
1985–1989 and 2005–2012 of 524 versus 1197 confirmed, 272 versus 1039 probable
and 90 versus 716 possible occupied grid cells. A limited decline was reported in
1975–2012 and a stable situation in 2000–2012. In 1999 an overview of regional atlases
showed a population decline of 20–50% since 1970 (Rocamora and Yeatman-Berthelot,
1999). Between 1970 and 1985, 22% of grid cells in Bretagne were abandoned (Clec’h
1993). In Isle-de-France, 1000 pairs were estimated in 1950 dropping to 300-400 pairs in

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 Chapter 5: Distribution, Population Estimates and Trends

1990 (Lecomte, 1995). The population of Vaucluse dropped from 700 in 1983 to 350 in
1993 (Olioso 1996). In the Vosges du Nord, 19 pairs in 1984 dropped to 9 in 1991 and
recovered to 19 in 1996 (Génot 1992c). Since the monitoring project Observatoire
National Inter-Parcs unfortunately stopped in 2008, a representative view on national
trends is no longer available (Hameau in Issa and Muller 2015).

5.5.10 Spain
5.5.10.1 DISTRIBUTION
The Little Owl nests almost everywhere in the Iberian Peninsula, but is absent from the
Balearic and Canary archipelagos, where it has only been recorded in winter in Palma
de Mallorca (Olea 1997), suggesting migration of the species to that regioin. The species
avoids excessively wet zones, dense forests and high mountain habitats (Muntaner et al.
1983, Olea 1997). The Little Owl is less common in the Euro-Siberian Region than in
the Mediterranean Region of the Iberian Peninsula. This could be indirectly related to
the weather, e.g., the oceanic weather favors forests that are avoided by the species
(Zuberogoitia and Martínez-Climent 2001). However, there are high-density popula-
tions of Little Owls in places with traditional agriculture and poultry breeding, and their
associated field and grasslands. They are not common in mountainous areas, although
there are populations established in more open areas at higher elevations in the
Pyrenees (1600 m) and Sierra Nevada (2300 m).
Muntaner et al. (1983) and Urios et al. (1991) observed that the winter temperature
was also a limiting factor. In fact, Muntaner et al. (1983) established the distributional
limit as the 1C isotherm for January in Cataluña and Andorra, and Urios et al. (1991)
identified the 3C isotherm as the distributional limit in Valencia. This can be correl-
ated with other variables, like the persistence of snow cover during winter that makes it
difficult for the owls to find adequate prey.

5.5.10.2 POPULATION ESTIMATES

According to Purroy (1997) there were 50 000–65 000 estimated breeding pairs in Spain
for the period 1975–1995. The Atlas de las Aves reproductoras de España (Marti and Del
Moral 2003), suggests the Spanish Little Owl population to be at least 36 000 pairs,
although it is necessary to point out there is no abundance data for almost 25% of the
squares (each square being 625 km²) where the species was recorded (Garcia and Muñoz
2003). The map of the III Atlas de las Aves en Época de Reproducción en España
(Figure 5.22) (Molina et al. 2021) only reports confirmed breeding in 1988 grid cells
for 2020 compared to 1675 grid cells in 1998–2002 (Marti and Del Moral 2003). This
might indicate an increase, but is probably due to less detailed data gathering in the
most recent survey.
A long-term reduction of more than 20% of the population has been observed over
the last 30 years, following the European trend. However, the short-term population
tendency for the species was considered to be positive during 1998–2001. Furthermore,
the Little Owl is reported as the most abundant nocturnal raptor in Spain (SEO/BirdLife
2002). Changes in agricultural practices and land use are the main factors responsible

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Figure 5.22 Little Owl distribution (a) and probability of occurrence (b) in 2020 in Spain (after Molina
et al. 2021).

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for the decrease of the species. Urban development plans are a real threat, especially in
the eastern coastal semi-arid habitats and in the Basque countryside on the Atlantic
coast (northern Spain) (Zuberogoitia 2002, Martínez and Zuberogoitia 2004a). The
reduction of dry-farmed tree crops, in particular the carob tree (Ceratonia siliqua), has
caused severe declines on the eastern coast. Pesticides diminish prey availability, land
re-allocation schemes reduce suitable sites for nesting and hunting, and the reduction
of hedges and trees along roads increases the risk of collisions with vehicles (Olea 1997,
Garcia and Munoz 2003). Likewise, the enlargement of forested areas, especially in the
Pre-Pyrenees has led to a reduction in suitable habitats (Martinez and Zuberogoitia
2003, J. Estrada personal communication). BirdLife International (2004) estimated
the total population at 20 000–100 000 pairs. BirdLife International (2015) estimated
the total population at 39 433 pairs in 1998–2002. In 2013–2018, the population was
estimated at 39 433 breeding pairs (European Environment Agency 2021).

5.5.10.3 DENSITIES
There is limited data on Little Owl abundance in the Iberian Peninsula: 0.2 individuals/
km² in Granada (southern Spain), up to 1 pair/km² in Catalonia, and 0.03–0.11 pairs/
km² in Galicia (northwest Spain) (Olea 1997). Fajardo et al. (1998) recorded Little Owl
numbers during road surveys in the province of Seville (southern Spain). During the
breeding season, they recorded 17 individual owls/km² in olive groves and seven
individuals/km² in sunflower fields, respectively. During July, they recorded up to
86.8 individuals/km². Zuberogoitia and Campos (1997) found 1.3 territories/km² in
Biscay; Olea (1997) reported 1 pair/km² in olive plantations and Pinus halepensis forests
in Cataluna.

5.5.11 Portugal
5.5.11.1 DISTRIBUTION
The Little Owl is probably the most common and widespread owl in Portugal. Iberian
breeding populations are sedentary, and young birds undertake local movement. There
is evidence that some peninsular birds could be wintering in Iberia (Díaz et al. 1996).
The species is more common south of the River Tagus, mainly in the provinces of Alto
Alentejo and Baixo Alentejo (Rufino 1989), in southern Portugal. It is scarce or absent in
the extensive forest areas of central and northern Portugal and occurs in a great variety
of habitat types including regions from sea level to over 1000 m. Little Owls are
frequently found in parks and gardens in cities, small towns and villages. It is a
common species in Lisbon, frequenting the great urban parks and some gardens.
Although uncommon, they can be found using the upper parts of buildings (Reino in
Van Nieuwenhuyse et al. 2008). Figure 5.23 shows the detailed distribution in
2005–2014 (Lourenço et al. 2015) compared to 1999–2005 (Equipa Atlas 2008).

5.5.11.2 POPULATION ESTIMATES

As of 1989, the estimate for the Portuguese Little Owl population was between
10 000 and 100 000 pairs with populations considered stable (Manez in Tucker and

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 5.5 Overview per Country

Figure 5.23 Detailed distribution of the Little Owl (Athene noctua) in 2005–2014 (large circles) versus
1999–2005 (small circles Equipa Atlas 2008) (after Lourenço et al., 2015).

Heath 1994). The latest estimate for the Little Owl population in Portugal was 50 000–
150 000 breeding pairs, and probably closer to 90 000 pairs (Tomé in Van
Nieuwenhuyse et al. 2008). BirdLife International (2004) estimated the total population
at 50 000–150 000 pairs. BirdLife International (2015) estimated the total population at
58 000–137 000 pairs in 2008–2012. In 2013–2018, the population was estimated at
40 000–60 000 breeding pairs (European Environment Agency 2021).

5.5.11.3 DENSITIES
In southern Portugal (Bixo Alentejo) densities of up to 2.36 pairs/km² were recorded in
16 km² of steppe-like habitat, and 6.95 pairs/km² in an area of 6 km² of open Holm-oak
Quercus rotundifolia (Tomé in Van Nieuwenhuyse et al. 2008).
During 2010–2017 the Working Group on Nocturnal Birds of SPEA (GTAN-SPEA)
carried out a volunteer monitoring program (NOCTUA-Portugal) to determine the
population trends of seven owl species at a national scale (Lourenço et al. 2021). They
used generalized estimating equation models to assess trends. In addition, they used the
distribution data from breeding bird atlases, observations sent by collaborators and data
from the public online database PortugalAves/eBird to detect changes in distribution
over a four-decade period (1978–2017). Four periods were defined to study potential
changes in distribution of owls in Portugal: 1978–1984 (first breeding bird atlas),
1999–2005 (second breeding bird atlas), 2006–2014 (interval between second and third
breeding bird atlases) and 2015–2017 (ongoing third breeding bird atlas). The Little Owl
had negative population and distribution trends. Figure 5.24 shows the evolution in the
four sampling periods since 1978.

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 Chapter 5: Distribution, Population Estimates and Trends

Figure 5.24 Distribution of the Little Owl (Athene noctua) in the sampling periods: first breeding
bird atlas BBA1 (1978–1984), second breeding bird atlas BBA2 (1999–2005), period between
second and third breeding bird atlases – inter-atlas (2006–2014), third breeding bird atlas BBA3
(2015–2017) (after Lourenço et al. 2021).

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 5.5 Overview per Country

4.0
3.5
Breeding pairs/10 x 10 km square
3.0
2.5
2.0
1.5
1.0

2010 2011 2012 2013 2014 2015 2016 2017

Year

Figure 5.25 Annual abundance estimates (number of breeding pairs per sampled 10
10 km square
with 95% confidence intervals) for the Little Owl (Athene noctua) in the period 2010–2020 resulting
from the NOCTUA-Portugal monitoring program (after GTAN-SPEA 2021).

5.5.11.4 TRENDS
Figure 5.25 shows the evolution of the abundance estimates per 10
10 km square
(GTAN-SPEA 2021). The distribution trend estimates for 1978–2017 (Lourenço et al.
2021) gives a percentage of the sampling units with breeding/presence information for
four periods, i.e., 1978–1984, BBA1 first breeding atlas, 92% occupation; 1999–2005,
BBA2 second breeding atlas, 74% occupation; 2006–2014, inter-atlas period between
second and third breeding atlas, 64% occupation; 2015–2017 BBA3 third breeding atlas,
65% occupation. The eight-year population trend for 2010–2017 from a GEE-GLM
model yields an estimated index decline of –0.24.

5.5.12 Italy
5.5.12.1 DISTRIBUTION
In Italy the Little Owl is a sedentary species, being widespread across the mainland, but
can also be found on the isles of Sardinia, Sicily and others (Casini in Meschini and
Frugis 1993) (Figure 5.26). Dispersal and movement of owls occurs in autumn and
winter, and some autumn movements can be considered as migration (M. Mastrorilli
personal communication). Younger individuals are more mobile and could move for
tens of kilometers (Spina and Volponi 2008).
The species breeds mainly in lowlands and lower hills; more than 80% of presence
locations are found below 300 m a.s.l. and more than 50% below 100 m a.s.l. The Little
Owl becomes less frequent with the increase of elevation. It rarely breeds above 1000 m
a.s.l., with the higher records between 1300 and 1450 m a.s.l in both the Alps and
Apennines (G. Calvi personal communication). It is a well-distributed bird of prey in
lowland areas and hillsides, with rare breeding records in the lower mountains. This

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 Chapter 5: Distribution, Population Estimates and Trends

Breeding evidence
Confirmed (434)
Probable (370)
Possible (881)

Figure 5.26 Little Owl distribution in Italy 2010–2016 (www.ornitho.it/archive).

explains the gaps in the distribution along the Alpine arc. Nevertheless, some breeding
pairs have been recorded at higher elevations in recent years: 1000 m in Liguria
(Andreotti 1989), at 1100 m in Piedmont (Mostini in Mingozzi et al. 1988), and the
highest in western Europe has been recorded in the Orobic Alps, Lombardia at 1230 m
(Mastrorilli 2001). Other important factors driving the species distributions are climate
and vegetation: the Little Owl prefers a warm climate and avoids wide and
continuous woodlands.
In Italy the Little Owl is widely spread in the Po Plain and in other lowlands; it is also
quite common in low hills, both on Tyrrhenian and Adriatic Apennine ridges. It also
breeds on the main islands, Sardinia and Sicily, but avoiding higher reliefs. For the same
reason, the species is absent from the Alpine arc and from the higher Apennine reliefs,
with the exception of the main valleys (e.g., Valtellina, Aosta valley). The Italian
distribution of the Little Owl in 2010–2016 (www.ornitho.it/archive) follows that of
the early 1980s (Casini in Meschini and Frugis 1993) with the main exception of the
northern Apennines and other low reliefs where forest expansion is the most likely
driver of local extinction.
Between 1988 and 2004 the Little Owl was recognized as the most common bird of
prey in Italian towns due to the high number of urban parks and wooded boulevards
(with owls showing a preference for species such as Poplar Populus sp., Plane Platanus
sp. and Horse-chestnut Aesculus hippocastanum), and also the urbanization of country
buildings and the favorable characteristics of historical city centers (old buildings with

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 5.5 Overview per Country

plenty of hollows). Roof spaces in recently built industrial buildings are often used as
breeding sites (Mastrorilli 1999). During several urban surveys, the presence of Little
Owls has been observed in 52 major Italian towns.
The Little Owl breeds mainly in various types of rural areas, preferring sites with
traditional farming where it can rely on a wider prey spectrum and on greater availabil-
ity of nesting sites, represented by farmhouses, ruins and tree holes (e.g., pollarded
willows, poplars, mulberries). Small villages, especially if close to farmlands, are also
often occupied. Over the past few years the Little Owl has bred more frequently in
industrial facilities. In southern Italy and on the main islands lacking buildings and
trees, the Little Owl nests in piles of stones. The main limiting factors for the species in
Italy are loss of biodiversity in farmlands with intensive farming, cutting down of old
trees with holes, demolition or renovation of rural buildings, and roadkills (G. Calvi
personal communication).
Due to the different weather conditions along the Peninsula, there are several areas
that can be colonized by the species. On the Po Plain the species distribution is strongly
affected by severe winters (i.e., snow and relatively low temperatures) causing signifi-
cant population decreases. In central and southern Italy the environment and weather
conditions are typically Mediterranean and thus more suitable to the owl’s ecology and
for this reason populations are more stable there. The most favorable conditions for
Little Owls (suitable weather, food availability, suitable habitat) are encountered in
southern Italy (M. Mastrorilli personal communication).

5.5.12.2 POPULATION ESTIMATES

A strong decrease was observed during 1960–1980, followed by a rapid recovery
(Brichetti 1997), yielding stable populations recently estimated at 10 000–50 000 pairs
(Manez in Tucker and Heath 1994) and 10 000–30 000 pairs in 1996 (Brichetti 1997).
BirdLife International (2004, 2015) estimated the total population at 30 000–
50 000 pairs around 2000 and 40 000–70 000 pairs in 2006. The Italian breeding popu-
lation was estimated at 40 000–70 000 territories by Brichetti and Fracasso (2006) on the
basis of researchers’ experience. The European Environment Agency (2021) reported
40 000–70 000 breeding pairs in 2007–2018. There is a lack of wide-scale monitoring
programs and, as a consequence, a lack of monitoring data for reliable and more precise
population estimates (www.ornitho.it/archive).

5.5.12.3 DENSITIES
The following densities have been reported in various areas of Italy: in mixed farmland
(Ticino Park, Pavia, northern Italy) 1.1 territories/km² (Cesaris 1988); rice fields (Pavia)
0.4 territories/km² (Galeotti and Pirovano unpublished data); urban areas (Pavia) 1.25
territories/km² (Galeotti and Morimando 1991). Intensive farmland on the Po Plain,
Lombardy, northern Italy) produced 1 territory/km² (Galeotti and Sacchi 1996) and
the Low Alps (Lombardy) >0.5 territories/km² (Galeotti and Sacchi 1996). Table S5.5
shows density data for the owl in some Italian towns. Table S5.6 reflects densities of
Little Owls in an array of Italian countryside locations. Densities are highly variable,

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 Chapter 5: Distribution, Population Estimates and Trends

depending on habitat types and can reach 8 territories/km² (Brichetti and Fracasso
2020), usually ranging from 0.5 to 1.5 pairs/km².

5.5.12.4 TRENDS
The only available trend for this species refers to the period 2000–2011 and shows a
moderate decline for the species (mean annual variation = –3.6%; Campedelli et al.
2012). It is worth noting that this trend derives from the national common breeding
bird monitoring program, which is not aimed at monitoring owls. Some recent local
atlases seem to confirm the negative trend, especially in northern Italy: in South Tyrol
the species, rare during the period 1987–1991, has disappeared and has not been
detected in 2010–2015 (Unterholzner et al. 2018). In Venice Province a decrease in
occupied units between 1996–1998 and 2008–2012 has been recorded, during both
breeding (–16%) and wintering (–42%; Bon et al. 2014). This pattern is far from being
universal: in Umbria region the species has widened its distribution increasing from
49% occupied units in the period 1988–1993 to 85.3% between 2012 and 2017 (Velatta
et al. 2019). In the Lazio Region and the Treviso Province the species seems stable
(Mezzavilla and Bettiol 2007, Brunelli et al. 2011). The species is considered to be Least
Concern in the national Red List (Peronace et al. 2012) and its conservation status is
judged as favorable (Gustin et al. 2016).

5.5.13 Switzerland
5.5.13.1 DISTRIBUTION
The species is present in very low numbers in four small regions : (1) in the Canton de
Genève (Geneva), (2) in Ajoie, north of the Jura canton, (3) in Ticino and (4) in the
Seeland region (Fribourg canton). The three main ones are in border areas with France
(1), France and Germany (2) and Italy (3). A conservation initiative in the Basel region is
trying to restore a link with the neighboring German and French populations through
improvement and restoration of habitat, without a positive result so far for the Swiss
side, but the surrounding populations are increasing (C. Meisser personal
communication).
The evolution of the distribution of the species is illustrated in four maps with the
respective distributions of 1950–1959, 1972–1976, 1993–1996 and 2013–2016
(Figure 5.27).

5.5.13.2 POPULATION ESTIMATES

The species has undergone a marked decrease, due especially to habitat changes. The
Swiss population was estimated at 185 pairs in 1980 (Juillard 1980), 30–40 pairs in
1986–1991 (Manez in Tucker and Heath 1994), 70 pairs in 1992–1996 (Meisser and
Juillard 1998) and 58 in 2002 (Meisser et al. 2016). But the trend has been slowly
recovering since then: 90–120 in 2011–2014 (Meisser et al. 2016), 115–150 in
2013–2016 (Knaus et al. 2018) and 153 in 2021 (compilation made by BirdLife
Switzerland).

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 5.5 Overview per Country

Figure 5.27 Little Owl distribution in Switzerland in 1950–1959, 1972–1976, 1993–1996 and
2013–2016 after Knaus et al. 2018. Reproduced with permission from Vogelwarte, Sempach,
Switzerland. www.vogelwarte.ch.

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Around Geneva, 77 territories were counted in 2021, compared to 32 in the worst

year, 20 years ago (Meisser 2022). The increase is largely due to strong growth in one
area, covering half of the suitable habitats of the canton de Genève. In Ajoie, the trend
is similar: from 13 territories in 2003 to 47 in 2021 (Collectif Chevêche Ajoie,
N. Apolloni personal communication). Tessin has 23 pairs in 2021, which is six times
more than in 2005 (Ficedula, R. Lardelli personal communication). The fragile return of
the species to the Seeland region began in 2005. The number of territories has recently
increased, reaching six by 2021 (Swiss Ornithological Institute Sempach personal
communication).
The recent increases in the three main (and still very small) populations are moder-
ate and have not significantly changed the distribution of the species in Switzerland
compared to 30 years ago, with the exception of the incipient re-colonization in the
Seeland region. This is probably because potential for new populations in Switzerland
remains scarce, given the low level of current suitable habitats elsewhere in the
country.

5.5.13.3 DENSITIES
Around Geneva, the mean density is 1.2 calling males/km² (about 60 km² of suitable
habitats). The highest densities are 2.5 territories/km² (for example 24 territories over
9.5 km² in 2021; Meisser 2022).

5.5.14 Austria
Since the drastic decline in the population of the Little Owl in Austria, the species has
evolved from a very common species up to the 1970s to Critically Threatened and Red-
Listed (Dvorak et al. 2017). Recently there have been intensive efforts to preserve this
species. Targeted protective measures were preceded by systematic population and
reproduction controls, as well as analyses of the habitat, which also accompany all
measures for the purpose of evaluation.

5.5.14.1 DISTRIBUTION
In Austria there are currently three more or less interacting Little Owl subpopulations
(Figure 5.28). The westernmost subpopulation mainly breeds in the Machland, around
Ried/Riedmark, in the Eferdinger Becken as well as the lower Mühlviertel in the federal
state of Upper Austria, with a population size of 35–65 breeding pairs between 2013 and
2020 (F. Kloibhofe personal communication). The main areas of distribution in
Lower Austria are the districts of Krems, St. Pölten, Tulln, Wiener Becken, Wagram,
eastern Mostviertel and northwest Weinviertel, ranging between 20 and 76 breeding
pairs (59 to 119 occupied territories) in the period 2013–2020 (F. Grinschgl,
W. Hovorka and R. Ille personal communication). The third subpopulation breeds
in the easternmost federal state of Austria, in the Burgenland. While Little Owls were
found both west and east of Lake Neusiedl up to 2000, the settlement is currently
limited to the east side, with a few exceptions. From 2013 to 2020 between 15 and

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 5.5 Overview per Country

Figure 5.28 Little Owl distribution in Austria (Österreichischer Bericht gemäß Artikel 12 der
Vogelschutzrichtlinie, 2009/147/EG Berichtszeitraum 2013 bis 2018. Ergebnisbericht Wien, Oktober
2019. Bearbeiter: Michael Dvorak, BirdLife Österreich) (after European Conservation Agency 2021).

18 breeding pairs were reported (15 to 23 occupied territories; F. Grinschgl and

R. Ille personal communication).

5.5.14.2 POPULATION ESTIMATES

The species is decreasing (Glutz Von Blotzheim and Bauer 1980) and 40–60 pairs
were counted in the 1990s (Manez in Tucker and Heath 1994) and 60 pairs in 1997
(Ille and Grinschgl 2001). Based on systematic searches, the population size between
1996 and 2001 was estimated at about 50 pairs and 20 single individuals (nearly
exclusively males) (R. Ille personal communication). BirdLife International (2004)
estimated the total population at 70–100 pairs. BirdLife International (2004, 2015)
estimated the total population at 70–100 pairs around 2000 and 130–170 pairs
in 2001–2012. The European Environment Agency (2021) reported 140–190 breeding
pairs in 2007–2018. The minimum population size is estimated at 70–159 breeding
pairs during the period 2013–2020, and the number of breeding pairs increased during
this period (R. Ille and F. Grinschgl, personal communication). Higher numbers are
suspected because male responsiveness to playback can be very low and pairs may be
very hidden during the breeding season, especially in sparsely populated areas or if
predators like Tawny Owls, Long-Eared Owls or Eagle Owls settle nearby (F. Grinschgl
and R. Ille personal communication). Solitary owls in general are males (Ille and
Grinschgl 2014, 2017).
In recent years, local initiatives in each of the four federal provinces led to a reversal
of the trend. Today the compiled estimates of breeding pairs have increased to just
below 300 breeding pairs (Table 5.6) (R. Zink personal communication) making the
short-term overall population trend positive.

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Table 5.6 Overview of population estimates for the Austrian Little Owl population in Austria between 2019 and 2021, monitoring efforts in
Lower Austria have been highest in 2020

2019 2020 2021

Known Checked Known Checked Known Checked

breeding breeding Successful breeding breeding Successful breeding breeding Successful
pairs pairs broods pairs pairs broods pairs pairs broods

Burgenland (Wurm 40 24 24 69 40 32 93 63 59
and Paar 2021)
Lower Austria (Pegler, approx. 103 >64 approx.
Stefan and 100 100
Geringer unpubl.
data/Grinschgl
2020/Steiner et al.
2020)
Upper Austria 67 56 54 83 71 67 93 69 67
(Lugmair and
Kloibhofer 2021)
Vorarlberg (Engel- 7
Klien, Eulenhilfe
Vorarlberg
e. V. pers. comm.)

Total 191 78 255 163 293 126

(compilation)

Compilation by R. Zink personal communication.

 5.5 Overview per Country

5.5.14.3 TRENDS
Despite short-term fluctuations, the numbers indicate an upward trend for the Little
Owl in Austria, despite the fact that the factors responsible locally for the rapid decline
have turned out to be more negative in the meantime. These factors are still intensifi-
cation of agriculture, decline in the keeping of farm animals, expansion of settlement
areas and traffic routes, as well as loss of small-scale structured habitats that lead to a
lack of suitable food-rich hunting grounds and safe breeding sites. The decline in insect
populations (especially of profitable large insects) has led to an even greater dependence
on the Common Vole, associated with greater fluctuations in reproductive success
(R. Ille and F. Grinschgl personal communication).
One reason for the population recovery in Austria is the increasing acceptance
of (Stone Marten) safe nesting boxes placed in trees or on buildings, which on the
one hand compensates for the lack of suitable natural breeding sites and on the
other hand leads to lower breeding losses. The acceptance of nesting boxes is also
increasing as the young owls that have grown up in nesting boxes accept or even
prefer it for their own broods (Ille R. personal communication). In northwest
Weinviertel, in the year 2009, nestboxes were only used for 22% of broods, 91.7% in
2014 and 75% in 2020. In 2020, in the Wagram area, all broods were raised in
nestboxes. The use of nestboxes in Burgenland developed similarly from 20% in 2008
to 100% in 2017. Of course, this also has the disadvantage that the population can only
be preserved and consistently stabilized with constant maintenance of the nesting
boxes. Furthermore, protective measures by offering nestboxes may only work up to a
certain population density. However, in the area around Krems, nearly all pairs success-
fully bred in loess caves, showing that natural breeding sites can also be safe.
Another factor that could have a positive influence on population development is
global warming and the decreasing amount of precipitation, which corresponds to the
climatic conditions in Little Owl primary habitats associated with lower winter losses
(see Chapter 10, Population Regulation).

5.5.14.4 DENSITIES
Densities vary greatly, depending on habitat type, availability of food and nest sites. In
Austria, Little Owl habitats are orchards (e.g., Mostviertel, Machland, Ried/Riedmark),
vineyards with cellar lanes and old press-houses (e.g., northwest Weinviertel), outskirts
of villages with old barns and other old farm buildings (Wagram district in Lower
Austria, Burgenland), isolated courtyards (Burgenland, Lower Austria) and loess walls
(e.g., district of Krems, Wagram). For Upper Austria Kloibhofer and Lugmair (2018)
reported population densities between 0.4 breeding pairs/km2 (Eferdinger Becken) and
2.6 breeding pairs/km2 (Machland, Ried/Riedmark) for 2018. In the density center of
Burgenland, Grinschgl and Ille (personal communication) found a population density
of about 1.3 breeding pairs/km2 in 2017 and 1 breeding pair/km2 in 2020. In northwest
Weinviertel in Lower Austria, population density was 0.8 breeding pairs/km2 in
2014 and 2.5 breeding pairs/km2 in its density center (Pulkautal) in 2017. In the
Krems area, with 8 breeding pairs/km2, and Wagram, with 10 breeding pairs/km2,
density was locally very high in 2017.

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5.5.15 Czech Republic

5.5.15.1 DISTRIBUTION
The Little Owl was the most abundant and widespread owl in Bohemia in the 1940s
(Stastny et al. 1996). However, in recent decades its distribution range has clearly been
reduced. Between 1973–1977 and 1985–1989, the number of occupied squares
decreased by 43% (Stastny et al. 1996; each survey square was 132 km² in size). The
Little Owl’s highest abundance was recorded in areas up to 500 m elevation. The main
populations live south of Plzen town, in the Labe valley near Decin town and in Ohre
valley near Lovosice town. In southern Moravia, which was quite densely inhabited by
owls in the past, just a few pairs remain. The Little Owl breeds mostly in farm buildings
(55.7 %), followed by residential buildings (39.1 %), industrial buildings (2.1%) and
churches or castles (1.5 %) (Chrenková et al. 2017). Breeding in tree hollows is
extremely rare. In previous monitoring programs the percentage of unoccupied study
areas increased from 0% in 1993–1995 to 59% in 1998–1999 and 66% in 2005–2006
(Schröpfer 1996, 2000, Šálek and Schröpfer 2008). The percentage of unoccupied study
areas in our monitoring across the country (63%) is comparable with the last monitor-
ing program in 2005–2006, despite the fact that the total area of Little Owl monitoring
increased by about 57%. A contraction in the Little Owl’s range is also shown by the
abandoned quadrants (Figure 5.29) compared to Stastny et al. (1996), showing a decline
of about 91% of occupied quadrants.

5.5.15.2 POPULATION ESTIMATES

Owl numbers have been declining since the 1950s. Between survey efforts conducted in
1973–1977 and 1985–1989, a lack of confirmed breeding records occurred in 43% of the

Figure 5.29 Occupied and unoccupied mapping squares recorded during nationwide monitoring
program for the Little Owl in the Czech Republic (2015–2016) and Slovakia (2009–2014).

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 5.5 Overview per Country

surveyed squares (Stastny et al. 1996). In 1985–1989 the population size was estimated
at 700–1100 pairs (Stastny et al. 1996). An abundance and distribution survey was
carried out in 1993–1995 in the Czech Republic. On the basis of the results from
16 study areas, the population size was estimated at 1000–2100 pairs (Schröpfer
1996). The same area was again surveyed in 1998–1999. A noticeable decline to
500–1000 pairs was recorded (Schröpfer 2000). The main reasons for the decline since
1993–1995 were most probably the severe winters of 1995–1996 and 1996–1997, with
long-lasting snow cover. BirdLife International (2004) estimated the total population at
200–400 pairs. During the period 2009–2016, Chrenková et al. (2017) monitored
1968 potential breeding localities across the Czech Republic. In total, the presence of
94 males at 84 localities was recorded in 40 mapping squares, which represented 20.8 %
of the mapping squares that were controlled for Little Owls ðn ¼ 192Þ, but only 6.4 % of
overall number of mapping squares at the country level ðn ¼ 628Þ. The total breeding
population size was estimated at 130 breeding pairs in 2015–2016 (Chrenkova et al.
2017). BirdLife International (2004, 2015) estimated the total population at 200–400
pairs around 2000 and 250–500 pairs in 2001–2012. The European Environment
Agency (2021) reported 100–150 breeding pairs in 2007–2018.

5.5.15.3 DENSITIES
In total, 16 of 27 surveyed study areas were not occupied; in the remaining 13 areas the
average density was 0.012 pairs/km². The highest densities were recorded in western
Bohemia south of Plzen town, where the density was 0.145 pairs/km2 (Schröpfer 2000).
The average population density in 2015–2016 was 0.09 calling males/10 km². The
population density in individual study areas also differed markedly (Figure 5.30) from

13 2
14 3
23 25 18 19
17 22
4

24 5
7
6
15 16 Czech Republic 12
21
20
8 26 28 30 48 55
10
11 27 49
9 54
29 34 Slovakia 52
46
33 35
50 51 53
32 44 47
45
36
Study areas 31
43 40 41
Population density – calling males/10km2 42
37
0 country border 39
0.01–0.1 rivers 38
0.1–0.3
N
0.3–0.5
0 25 50 100 150 Km
> 0.5

Figure 5.30 Distribution of individual study areas and area-specific population densities of the Little
Owl recorded during nationwide monitoring program in the Czech Republic (2015–2016) and
Slovakia (2009–2014).

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 Chapter 5: Distribution, Population Estimates and Trends

0.02 (study area = 22) up to 0.33 (study area = 23) calling males/10 km² (Chrenkova
et al. 2017).

5.5.15.4 TRENDS
The national 2015–2016 population density of 0.09 calling males/10 km² (2015–2016)
is comparable with the density recorded in 2005–2006 (0.1 calling males/10 km²; Šálek
and Schröpfer 2008) but had declined by about 73% compared to monitoring in
1993–1995 (0.33 calling males/10 km²; Schröpfer 1996).

5.5.16 Slovakia
5.5.16.1 DISTRIBUTION
The Little Owl is a common breeding species in lowlands and basins of southern
Slovakia (Danko et al. 2002). The species is distributed along rivers in north Slovakia
too, in several areas up to 600 m elevation. In lowlands it usually occurs between
100 and 300 m. Its typical habitats in Slovakia include small field patches surrounding
barns and other agricultural buildings, and villages. These habitats are usually situated
at lower elevations – in lowlands and hilly areas. The species avoids forests and big
cities. The Little Owl can sometimes be found at the margins of industrial zones
surrounded by fields. The highest densities in Slovakia were recorded in the lowlands
of eastern Slovakia, the lowest in the basins and hills of north Slovakia. The Little Owl
breeds mostly in farm buildings (55.7%), followed by residential buildings (39.1%),
industrial buildings (2.1%) and churches or castles (1.5 %) (Chrenkova et al. 2017).
Breeding in tree hollows is extremely rare. The Chrenkova et al. (2017) study also
indicates that large-scale extinctions and contractions of the distribution have occurred
over recent decades, although this pattern was more obvious in the Czech Republic. In
particular, the occurrence of the Little Owl was not confirmed in 63% of the study areas
in the Czech Republic and 44% in Slovakia.

5.5.16.2 POPULATION ESTIMATES

Between 1980 and 1999 the total number of breeding pairs in Slovakia was estimated at
800–1000 pairs; the number of wintering Little Owls between 1980 and 1999 was
estimated at 2000–3000 individuals. The population slightly decreased and the geo-
graphic distribution decreased by 20% (Danko et al. 2002). BirdLife International (2004,
2015) estimated the total population at 800–1000 pairs around 2000 and 400–700 pairs
in 2012. The European Environment Agency (2021) reported 300–600 breeding pairs in
2007–2018. During the period 2009–2016, Chrenkova et al. (2017) monitored
877 potential breeding localities in Slovakia through playback. In total, 231 males at
218 localities in Slovakia were recorded. Occurrence was confirmed in 61 mapping
squares, which account for 37.7% of controlled mapping squares (n = 162) and 14.2%
of all available mapping squares (n = 429) in the country (Figure 5.29). The average
population density of the Little Owl in all study areas across Slovakia was 0.36 calling
males/10 km², which is significantly higher than in the Czech Republic (0.09 calling

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 5.5 Overview per Country

males/10 km²). The total breeding population size of the Little Owl was estimated at
550 pairs in Slovakia.

5.5.16.3 TRENDS
A slight long-term decline of the total population is evident (Danko et al. 2002).
In Slovakia, the lack of long-term data on population dynamics prevents detailed
assessment of population density changes. However, the population density in 2009–
2014 (0.36 calling males/10 km²) is markedly lower than densities in 1995–1999
(0.75–0.1 calling males/10 km²; Pačenovský 2002) or local-scale studies performed
during the 1980s and 1990s (e.g., 1.1–3.5 calling males/10 km²; Danko et al. 1995).

5.5.16.4 DENSITIES
The highest densities in Slovakia were recorded in the lowlands of eastern Slovakia
(12–15 pairs/121 km² or 0.09–0.12 pairs/km²), the lowest in basins and hills in north
Slovakia (1–3 pairs/121 km²) (Danko et al. 2002). The population density in individual
study areas in 2009–2014 (Chrenkova et al. 2017) differed markedly (Figure 5.20).
Occurrence was not confirmed in 44% of the research areas. The highest recorded
density was 1.19 (study area = 43), the lowest occupied study area (study area = 34)
had a density of 0.09 calling males/10 km².

5.5.17 Croatia
As of 1991, the population of Croatia was estimated at 6000–8000 pairs (although the
data quality for this estimate was not provided; Manez in Tucker and Heath 1994).
BirdLife International (2004, 2015) estimated the 2002 population at 500–1000 pairs
and 4000–6000 pairs in 2010. The European Environment Agency (2021) reported
4000–6000 breeding pairs in 2007–2018.

5.5.18 Slovenia
5.5.18.1 DISTRIBUTION
In 1769, J. A. Scopoli described the Little Owl to science from a specimen collected in
Slovenia. In the past, the species was very common and widely distributed in Slovenia
(Ponebšek 1917, Reiser 1925). Later a marked decline was noticed and the species
vanished from several parts of the country, with the bulk of the population distributed
in the southwestern (Mediterranean) and northeastern (sub-Pannonian) parts of the
country (Berce 2019). In the southern part of Slovenia there is a hybrid zone between
the subspecies A. n. noctua and A. n. indigena (Keve et al. 1960). The Little Owl is mainly
a lowland species in Slovenia, below 630 m (Tome 1996a, b). However it has also been
found at higher elevations (e.g., from 690–900 m) in the alpine region (Vogrin 2001),
but these reports were viewed as reflecting nonbreeding individuals (Božič and Vrezec
2000, Vogrin 2001).
The Little Owl (Čuk) is a rare nesting bird now (Berce 2019), most common in the
Primorska region, namely in the Vipava valley in the fields of Goriška, in the northern

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 Chapter 5: Distribution, Population Estimates and Trends

part of the Karst and in the Koper hills, which is similar to what was reported by Geister
(1995) (Figure 5.31). Individual breeding pairs can also be found in the Tolmin region.
Additional records are from Goričko, and in some places in the fields of Drava and Ptuj
and in the Celje basin. It is rare in the Ljubljana Marshes, where up to five pairs
allegedly nest (Berce et al. 2014) and from where, according to Tomé et al. (2005), it
disappeared in the late 1980s, and in Bela krajina, where it appears singly. It is
interesting that there are records from Čatež ob Savi near Brežice in May 2016 and
from Veržej on the Pomurska Plain in June 2017, as the species had not yet been
recorded in Krško-Brežiška (G. Bernard personal communication). In the southern
and southeastern parts of the country the species has been reported during the
breeding season in the vicinity of Mirna in the Dolenjska region and the vicinity of
Veliki Lašče, which is quite distant from other occurrences of the species. Compared
to Geister (1995), the species is less widespread, with almost complete extinction in
the northeastern part of the country, where it lived in Slovenske gorice, Dravsko polje
and Pomurska ravina, Ribniško-Kočevski podolje, Pivka, Dolina Reke and Brkini.
The owl also disappeared from Gorenjska. It is a resident of a predominantly lowland
and hilly landscape, as the majority of the nesting population (>90%) is found at
altitudes up to 400 m.
The species lives mainly in agricultural landscapes with tall orchards, shrubs with
individual trees, and its occurrence is strongly associated with human habitation, where
it nests in hollows, buildings, abandoned buildings and less often in churches (Slana
2000, Štumberger 2000, Surina 2000, Berce and Kmecl 2008, Berce et al. 2014). There is
also one known example of nesting in the nest of a white stork, in Župečja vas on the
Drava Plain (Bračko and Štumberger 2017). It is a typical open landscape species that
avoids dense forest areas and has gradually adapted to living near human habitations
such as in Karst, where owls were recorded in areas of the cultural landscape and near
human dwellings (Berce and Kmecl 2008).

Figure 5.31 The Little Owl in Slovenia 2002–2017 (European Environment Agency, 2021).

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 5.5 Overview per Country

5.5.18.2 POPULATION ESTIMATES

In the second half of the twentieth century the Little Owl breeding population was
estimated at 500–800 pairs (Geister 1995). A strong negative trend was noticed in the
later part of the twentieth century, especially in northeastern Slovenia (Vogrin 1997),
and the beginning of the twenty-first century (Berce 2019). The species is now probably
extirpated from the central part of Slovenia. According to recent surveys and popula-
tion estimates the population is estimated at 100–150 breeding pairs, with the
bulk of the population concentrated in the Mediterranean region of southwestern
Slovenia (e.g., Karst area; Berce 2019). In the Red Data Book of Slovenia, the species is
listed as an Endangered species with intensification of agriculture and traffic collisions
listed as the main reasons for the decline and continual threat to its population (Vrezec
and Vrh Vrezec 2011, Berce 2019). BirdLife International (2004, 2015) estimated the
total population at 150–200 pairs in 2000 and 100–150 pairs in 2002–2012. Berce
(2019) and the European Environment Agency (2021) reported 100–150 pairs in
2002–2017.

5.5.18.3 DENSITIES
The density in Dravsko polje, south of Maribor town was 0.048 pairs/km² (total area 210
km²) (Vogrin 1997). In central Slovenia, in Ljubljansko barje in an area of about 160
km², Trontelj (1994) estimated the population of Little Owls at 5–10 pairs or 0.03–0.06
pairs/km². Local densities have been found to be low in central and northeastern
Slovenia, with 0.03–0.1 pairs/km² (Trontelj 1994, Vogrin 1997), but in southwestern
Slovenia in the Karst region in the Mediterranean part of Slovenia are much higher with
0.3–4.0 pairs/km² (Berce 2019).

5.5.19 Serbia and Montenegro

5.5.19.1 DISTRIBUTION
The species occurs in the Pannonian peri-Pannonian parts of Serbia (south of the rivers
Sava and Danube) and in mountain regions with basins. In Montenegro (Crna Gora) it
is found in maritime Montenegro and mountain regions with basins (Stevanović and
Vasić 1995).

5.5.19.2 POPULATION ESTIMATES

Estimates for Vojvodina (northern part of Serbia – Pannonian Plain) are 1500–2000
pairs. Estimates for the central part of Serbia, excluding Vojvodina and Kosovo are
2500–5000 pairs. The Kosovo region (Kosovo and Metohija) is estimated at
4000–6000 pairs, Serbia (including Vojvodina, Central Serbia and Kosovo) at
8500–13 000 pairs and Montenegro at 1500–2000 pairs (S. Puzovic and D.V. Simic
personal communication). An estimate of 5000–10 000 pairs was made for 1962
(Manez in Tucker and Heath 1994). BirdLife International (2004) estimated the total
population for Serbia and Montenegro at 10 000–15 000 pairs. BirdLife International
(2015) estimated the total population of Montenegro at 1000–2000 pairs in 2002–2012
and of Serbia at 10 300–15 000 in 2008–2012.

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 Chapter 5: Distribution, Population Estimates and Trends

5.5.20 Albania
BirdLife International (2015) estimated the total population of Albania at 4000–8000
pairs in 2002–2012.

5.5.21 Greece
5.5.21.1 DISTRIBUTION
The species is a fairly common and widespread resident owl in Greece. Little Owls have
been familiar birds to people living in Greece for at least 2500 years. They appear in
ancient Greek art and in classical times they were associated with the goddess Athena.
They still breed locally within Athens and other urban centers, e.g., Lamia, Chalkis,
Eubä, Parnas, Zygos and Peleponnese Mountains, though they are more typically found
in rural areas. They occur widely over the whole mainland and on most islands, i.e., the
Ionian and Aegean Isles, the Cyclads and Naxos (Schönn et al. 1991), in a wide variety
of open habitats, including farmland, phrygana and the surroundings of villages and
shepherd’s huts (Eakle 1994). Little Owls have been recorded as high as 1650 m on
Mount Oiti (Handrinos and Akriotis 1997) and occur at relatively high elevations in
other areas, provided suitable open habitat is available. It appears that Little Owls have
declined since around 1970 (Cramp 1985). This is most evident in agricultural areas,
where the increased use of insecticides has reduced their food supply, and in coastal
areas, where urbanization has resulted in increased disturbance and loss of habitat, such
as derelict buildings.

5.5.21.2 POPULATION ESTIMATES

Although they have been a protected species for many decades, they are disliked and
persecuted in many rural areas due to a superstitious belief that their call is a bad omen.
However, despite this level of undocumented background impact, the owls remain
quite numerous and widely distributed. Their total population ranged between
5000 and 15 000 pairs in the 1990s (Handrinos and Akriotis 1997). BirdLife
International (2004) estimated the total population at 5000–15 000 pairs. BirdLife
International (2015) estimated the total population at 5000–15 000 pairs in
2008–2012. The European Environment Agency (2021) reported 5000–15 000 pairs in
2007–2018.

5.5.22 Cyprus
Considered a common but decreasing species, with 2000–4000 pairs as of 1988 (Manez
in Tucker and Heath 1994). BirdLife International (2004) estimated the total population
at 5000–15 000 pairs. BirdLife International (2015) estimated the total population at
4000–10 000 pairs in 2006–2012. The European Environment Agency (2021) reported
4000–10 000 pairs in 2007–2018.

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 5.5 Overview per Country

5.5.23 Hungary
5.5.23.1 DISTRIBUTION
The spatial distribution of the Hungarian Little Owl population is well known but data
are lacking on temporal changes/tendencies in population size. The species is present
throughout Hungary, especially east of the river Danube (Schönn et al. 1991, Gorman
1995) (Figure 5.32). The core population inhabits the Hungarian Plain, although breeding
pairs can be found in hilly parts of the country, in smaller densities (Gorman 1995).
Typical nesting sites are the lofts of farmhouses and barns (Schmidt 2000). The core
population of Little Owls is located in the Great Plain. The bird is found almost every-
where in the plains and hills, but it avoids the higher, closed, forested areas of the mid-
mountains. It is typically connected to the lowland landscape, where it prefers grazed
grasslands and the immediate vicinity of livestock farms. It is less common in the south-
western part of the country. Based on probability modeling of occurrence (Hámori 2019)
for the nesting period, the species is the most common between the Danube and the Tisza
and in the Trans-Tisza. Within these geographical areas, the most significant breeding
population is located in Bács-Kiskun, Csongrád, Békés, Hajdú-Bihar and Szabolcs-Szatmár-
Bereg counties. It is also common in the Kisalföld, in the outskirts of western Hungary, in
the eastern areas of the Transdanubian Mountains and the Somogy Hills, in the Mezőföld
and Cserehát regions. The probability of occurrence is increased by the proportion of
different agricultural areas and adjoining sandy steppe, saline and closed grasslands, as
well as the number of buildings in the given area (Hámori 2019).

5.5.23.2 POPULATION ESTIMATES

The population was estimated at 2000–2500 pairs as of 1994, and is probably stable
again after a decline (Gorman 1995). BirdLife International (2004) estimated the total

Figure 5.32 Little Owl distribution in Hungary (after Nagy in Szép et al. 2021).

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 Chapter 5: Distribution, Population Estimates and Trends

population at 1500–2500 pairs. BirdLife International (2015) reported 1500–2500 pairs

during 2000–2012, the European Environment Agency (2021) reported 3000–4000
pairs during 2007–2018. The estimated size in 2021 was approximately 1500–4000
breeding pairs, and the population tendency was considered to be negative (MME
Nomenclator Bizottság 2008, Hámori and Csortos 2016, Hámori 2019).

5.5.23.3 DENSITIES
Density modeling shows the highest densities in the Transdanubian Mountains, the
northern periphery of the Northern Central Mountains, the Upper Kiskunság and other
parts of the Danube–Tisza region, as well as in the Trans-Tisza region. In the central and
southern parts of the latter, as well as in the northern part of the Bereg Plain, the
density data show close to 2 individuals/km². Densities increase with the proportion of
agricultural land characterized by arable land and other permanent crops, as well as
with the proportion of closed grasslands, but decreases somewhat with higher forest
cover in the given area and sandy soil cover in the habitat.

5.5.24 Bulgaria
5.5.24.1 DISTRIBUTION
Ignatov and Popgeorgiev (2021) presented historical and newly collected distribution
data in order to update the distribution map with the help of members of the Bulgarian
Society for the Protection of Birds (BSPB) – consisting of unpublished field observations
covering the entire Bulgarian territory. All previously published data were reviewed and
used, to give an updated view on the presence of the species. The distribution is shown
in Figure 5.33 in a 10
10 km grid. Of a total of 1240 squares, 1126 were found to be
occupied, 398 had been published and re-confirmed, 8 were new records and 720 were
published but not re-confirmed. The elevation distribution reveals that most Bulgarian
Little Owls are breeding up to about 300–400 m a.s.l. No more than 9% prefer higher
altitudes from 400 to 800 m. Just 5% are above 900 m and several localities up to about
2300 m, mainly in mountain huts. Ignatov and Popgeorgiev (2021) confirmed that the
species prefers to breed and winter in human settlements (90%).
The species is widespread everywhere in Bulgaria occurring up to about 900 m a.s.l.
(Patev 1950, Simeonov 1983), about 98% in human settlements (Ignatov 1993), mostly
in villages and cities, such as in 1983, in Sofia (capital of Bulgaria), with 140 reported
breeding pairs (Iankov 1983). In the following decades, no changes have been reported,
including the last previously published data from the Atlas of Breeding Birds in Bulgaria
(Iankov 2007). The population of Little Owls is found to be occupying villages, towns
and the city of Sofia (Iankov 1983, Simeonov 1983).
There are many breeding pairs within the villages and outside as well in the regions
of the middle part of the Struma valley, and the valleys of the rivers Russenski Lom,
Provadiiska, Rossitca, Suha Reka, Ogosta, Kamchia, Arda and Iskar. For regions like
lowland mountains – Eastern Rhodopi Mountain, Strandja Mountain and Southern
Dobrudja – most breeding pairs are found in villages. The species is widespread from
0 to 2300 m a.s.l. Most territories are around 400 m a.s.l.; 95% of the population is

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 5.5 Overview per Country

(B)
23˚0'0''E 24˚0'0''E 25˚0'0''E 26˚0'0''E 27˚0'0''E 28˚0'0''E
N
44˚0'0''E

44˚0'0''E
43˚0'0''E

43˚0'0''E
42˚0'0''E

42˚0'0''E

Presence confirmed and published till 2007

Presence reconfirmed in this study
New in this study
Elevation, m
41˚0'0''E

00

41˚0'0''E
00

50

00

92
-5

-1

-1

-2

-2
0

00

00
50

0
10

15

20

23°0'0''E 24°0'0''E 25°0'0''E 26°0'0''E 27°0'0''E 28°0'0''E

Figure 5.33 (A) Distribution of the Little Owl in Bulgaria (after Hristov and Stoychev in Iankov (2007)).
(B) Distribution of Little Owl in Bulgaria, showing published data, confirmed data and newly collected
data in the standard 10
10 km UTM grid covering Bulgaria (1240 grid cells) (after Ignatov and
Popgeorgiev 2021).

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 Chapter 5: Distribution, Population Estimates and Trends

(C)

1
0,8
0,5
0,2
0

Figure 5.33 (C) Probability of occurrence of Little Owl in Bulgaria using Maxent (after Ignatov in
press).

found up to 800–900 m a.s.l. (Simeonov 1983, Ignatov 1993). One confirmed record of
breeding at about 2300 m has been published (Simeonov 1983) and there are four new
records (Rhodopi Mountain – 2, Vitosha Mountain – 1 and Central Balkan range – 1) in
the mountains, including one at an altitude of 2400 m on Pirin Mountain. More than
50% of the population is breeding at 0 to 400 m in villages on the Black Sea coast, low
mountains villages and those around river valleys. There are just a few pairs breeding in
high mountains, in mountain huts – on Rila Mountain (Partizanska poliana), Pirin
Mountain (Banderitca), Rhodopi Mountain (Perelik and Persenk) and Stara Planina
Mountain (Sinanitca). In the autumn–winter period, the highest place that the species
has been recorded is 1100 m a.s.l. Birds inhabiting settlements do not usually displace
for winter. Most of the owls leaving their breeding areas are young birds or those
breeding outside of the settlements.
Ignatov (in press) modeled the potential distribution and ecological requirements of
Little Owls within part of its range using MaxEnt. The Environmental Niche Model had
a high degree of predictability (AUC >0.902). Ignatov estimated that suitable habitats
comprised 90.08% of Bulgaria. The climate changes are affecting the distribution, i.e.,
the vertical distribution of the species expanded by 100 m. With a high number of
breeding pairs relative to elsewhere in Europe, Bulgaria has an opportunity and obliga-
tion to help preserve the species.

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 5.5 Overview per Country

5.5.24.2 POPULATION ESTIMATES

Nankinov (2002) characterized the Little Owl population in Bulgaria as exceeding
10 000 pairs. In 1982 about 140 pairs were encountered in Sofia (Iankov 1983).
According to Kostadinova (1997) the breeding population was estimated at
4000–10 000 pairs. BirdLife International (2004) estimated the total population at
5000–8000 pairs. BirdLife International (2015) reported 10 000–14 000 pairs during
2005–2012 and the European Environment Agency (2021) reported 5000–8000 pairs
during 2013–2018.

5.5.25 Romania
The Little Owl is the most common owl species in the country. It can be found
throughout the country with the exception of the forested mountainous areas. As an
adaptable species, the owl can be found regularly in villages and towns. In Bucharest
the species breeds in the Opera building. The species occurs at up to 800 m (Schönn
et al. 1991).

5.5.25.1 POPULATION ESTIMATES

The population is estimated at 20 000–40 000 pairs and was considered stable as of 1988
(Manez in Tucker and Heath 1994). BirdLife International (2004) estimated the total
population at 40 000–60 000 pairs. BirdLife International (2015) estimated the total
population at 15 000–40 000 pairs in 2001–2013. The European Environment Agency
(2021) reported 15 000–40 000 pairs in 2014–2018.

5.5.26 Moldova
5.5.26.1 DISTRIBUTION
The Little Owl is one of the most widely distributed owl species in Moldova. It is also
considered the most numerous of the nocturnal raptors (Ganya and Zubkov 1975). In
parks of Askania-Nova, Little Owls willingly occupy the empty nests of Magpies, Pica
pica, in insular forests in waterless valleys with groves of the white acacia, far from
human dwellings. On the plot at the site “Florike,” on an area of about 50 ha, about
100 Magpie nests were counted and Little Owls were nesting in 42 of them, as if in a
colony (Uspensky 1977).

5.5.26.2 POPULATION ESTIMATES

The population was estimated to be 5000–7000 pairs in 1988 (Manez in Tucker and
Heath 1994). BirdLife International (2004, 2015) estimated the total population at
3200–4200 pairs.

5.5.26.3 TREND
Both the short-term and long-term trends are stable, with casual fluctuations that do
not influence the overall trend.

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 Chapter 5: Distribution, Population Estimates and Trends

5.5.27 Turkey
5.5.27.1 DISTRIBUTION
The Little Owl is a common and widespread species throughout the country with high
densities in natural steppes and villages throughout central and eastern Turkey
(Ornithological Society of Turkey 1969, Eakle 1994). It is less common to rare in
forested areas, e.g., along the Black Sea coast. Due to its common status the species
has not been mentioned in recent Turkish Bird Reports (Ornithological Society of the
Middle East and Central Asia, OSME). However, Kasparek (1992) reviewed the species as
a widespread, common breeding resident in nearly the whole of Turkey, only local in
the Black Sea Region and other coastal regions. Further, it was mentioned that the
species is rare in alpine areas, though it had been recorded up to 4400 m. In Aegean and
Mediterranean coastal areas the species has been recorded scarcely and is mainly
confined to villages with surrounding agriculture. However, it is again more commonly
found at higher elevations where rocky habitat, macchia and extensive agriculture are
prevalent. These studies also revealed that the Little Owl is certainly much more
common than estimated (E. Vaassen personal communication). Local populations of
the Little Owl as well as the other insect-eater, Eurasian Scops Owl Otus scops, have
nearly disappeared from some habitats (villages and ruins) in the Göksu and Çukurova
Deltas in southern Turkey, probably due to the massive application of insecticides
(E. Vaassen personal communication).

5.5.27.2 POPULATION ESTIMATES

Kasparek and Bilgin (1996) estimated in their Checklist of Turkish Vertebrates that the
total Little Owl population was 5000–50 000 pairs. More recent field studies have
revealed that the owl is certainly much more common than estimated and therefore a
population of 30 000 pairs for the whole of Turkey (782 700 km²) is likely to be the
absolute minimum (E. Vaassen personal communication). Estimates for Turkey may
prove to be very low, given the density of 500 pairs in 80 km² for Emirdag. The national
population may reach 200 000–250 000 pairs (E. Vaassen personal communication).
BirdLife International (2004, 2015) estimated the total population at 300 000–-
600 000 pairs in 2000–2013.

5.5.27.3 DENSITIES
Recent field studies have revealed that the species is locally very common, with dens-
ities of 8–10 pairs/km² in rocky highland areas with marginal cultivation. Similar
densities have been located in the surrounding districts of large cities like Ankara and
Konya, where building sites and patches of wild-grown steppe are prevalent (Vaassen
2000).

5.5.28 Poland
5.5.28.1 DISTRIBUTION
Until the end of the 1960s, the Little Owl was one of the most common owls in Poland
(Taczanowski 1882, Sokołowski 1953). In the 1960s, one or two pairs were nesting in

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almost every village in the area of Legnica and Leszno in southwestern Poland
(Tomiałojć 1972, Bednarz et al. 2000). The first data concerning the decrease in popula-
tion numbers of the Little Owl appeared in the 1970s and 1980s when it was considered
to occur generally in small numbers, and only locally in medium numbers (Tomiałojć
1990). The disappearance of this species started in western Poland. In the Gorzów
Wielkopolski Region, this bird was relatively frequently observed in the 1970s, whereas
at present it is practically extirpated (Jermaczek et al. 1995, A. Jermaczek personal
communication). A similar situation is observed in the Poznań Region of western-
central Poland and the Silesian Region of southwestern Poland, where the Little Owl
currently occurs in very small numbers (Dyrcz et al. 1991, Bednarz et al. 2000). The
tendency towards the gradual disappearance of the species from western Poland was
noted in the West Pomeranian Region of northwestern Poland, where at present there
are probably only two nesting sites remaining (R. Kościów personal communication). In
the nineteenth century the Little Owl occurred in about 130 larger and smaller towns in
Poland, and more recently, from 1990–2000, it inhabits over 60 towns, mainly in
southern and southeastern Poland (Grzywaczewski and Kitowski 2000). In Poland,
the Little Owl mainly occurs in cities and upland areas, i.e., the uplands of Lublin,
Cracow, Cracow-Czestochowa and the Sub-Carpathian Highlands. The cities and rural
areas located in these regions of south and southeastern Poland can be considered the
last strongholds for this species (G. Grzywaczewski personal communication).

5.5.28.2 POPULATION ESTIMATES

The species is scarce in most areas, with marked declines after severe winters (Tomiałojć
1976). Based on the available information, the number of Little Owls in Poland was
estimated at 1000–1500 breeding pairs in 2003 (Grzywaczewski 2006). Little Owl
estimates in different regions of the country are given in Table S5.7 and Table S5.8.
Detailed data in the 1990s indicated 10–20 pairs in the Białystok Region of northeastern
Poland (M. Polakowski personal communication), 150–200 pairs in the Lublin Region
of southeastern Poland (G. Grzywaczewski unpublished data), 10–100 pairs in the
Łódź Region of central Poland (R. Włodarczyk personal communication); 50–200
pairs in the Cracow Region of southeastern and southern Poland (K. Kus personal
communication) and 120 pairs in the Radom and Kielce Regions of south-central
Poland (J. Tabor personal communication). BirdLife International (2004) estimated
the total population at 1000–2000 pairs. BirdLife International (2015) reported
500–1000 pairs during 2008–2012 and the European Environment Agency (2021)
reported 500–1000 pairs during 2007–2018. Population estimates in 1985–2004
were 1000–2000 pairs (Chylarecki and Sikora 2007), in 1999–2000, 1000–1500 pairs
(Grzywaczewski 2006), in 2008–2012, 500–1000 pairs (Chodkiewicz et al. 2015),
in 2013–2018, 500–1000 pairs (Chodkiewicz et al. 2019) and in 2020, 1400 pairs
(Wilk et al. 2020).

5.5.28.3 DENSITIES
In the 1980s, the average density of the Little Owl population in the agricultural
landscape was 0.17 territories/km². A relatively rapid change took place in the 1990s,

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 Chapter 5: Distribution, Population Estimates and Trends

when population density dropped to 0.07 territories/km². In 2004 in urban areas,

especially in southern and southeastern Poland, an increase was observed in the owl
population density to 0.26 territories/km². In Chełm, southeastern Poland, the density
of Little Owls is among the highest in Poland, reaching on average 0.4 territories/km².
However, after the snowy and frosty winter of 2001–2002, the density dropped to 0.22
territories/km² (G. Grzywaczewski personal communication). In the Kutnowska and
Łowicko-Błońska Lowlands (central Poland, a total area of 4352 km2) in 2016–2018 the
density varied from 0.6 to 4.0 territories/10 km2. Little Owls inhabit vast areas of
intensively used farmland (Jabłoński 2019) (Table S5.9).

5.5.29 Russian Federation

There have been substantial changes in the political boundaries of the former USSR,
complicating analysis of the older literature on the Little Owl. Below, we briefly describe
the owl distribution in the former USSR, and then the owl situation in the respective
countries. In the European territory of the former USSR, the Little Owl is distributed
mainly in anthropogenic landscapes, in southern Kazakhstan and Middle Asia, mainly
in the deserts, dry steppes and in mountainous regions. From the western border the
species spreads eastward to the Zabaikalie (Trans-Baikalia) (Sretensk). To the north it
reaches Lithuania and Latvia (in Estonia the species is a vagrant bird), Pskov, Moscow,
Ryazan Region, Kazan, Buguruslan, Orenburg, from where the border stretches
via central Kazakhstan to Zaisan Lake. Further eastwards the species occurs only
in isolated populations in the southeastern Altai (Chuiskaya steppe), in Tuvinskaya
(Tyva Autonomous Soviet Socialist Republic) and in Trans-Baikalia (Selenginskaya
Dauria, Aginskaya, Borzinskaya and Nerchinskaya steppes). In Tien-Shan and
the Pamir, the Little Owl breeds at high elevations (up to 4200 m). The southern
border of the Little Owl’s range is situated outside the borders of the former USSR
(Ivanov 1976).
In Russia, the Little Owl is resident, or rarely migratory. The total number in
European Russia is estimated to be 10 000–99 999 pairs (Mishchenko 2004).
BirdLife International (2004) estimated the total population at 10 000–100 000 pairs.
BirdLife International (2015) reported 50 000–70 000 pairs during 2008–2012. While
there is some information on owls moving northwards along human-transformed
areas, fluctuations in owl numbers are difficult to determine, and some researchers
report that the population is currently decreasing in Russia. The population in the
Republic of Bashkortostan (Karyakin 1998) is estimated at 10 breeding pairs, with
winter numbers at 10–50 individuals. At least 10–15 pairs breed in the whole of
the south of the Kaliningrad Region (Grishanov 1994). The Saratov Region of
100 000 km² likely supports about 200–300 pairs (A. Antonchikov personal commu-
nication). At present the species is comparatively stable in the Republic of Dagestan,
and in some places can even increase due to an increase in human-transformed
landscapes in the lowland regions of the Republic and restoration of formerly aban-
doned cattle-farm complexes. In the Lower Cis-Volga River area Zav’yalov et al.

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 5.5 Overview per Country

(2000) illustrated the intra-year dynamics of abundance for two different environ-
ments of Saratov city. For 1993–1998, they found Little Owl densities in old apart-
ment buildings and the forst-park zone to be respectively 0.2 and 0.05 individuals/
km² in winter, 0.1 and 0.2 in prespring, 0.3 and 0.3 in the prenesting period, 0.6 and
0.4 during nesting, 0.5 and 0.4 postnesting and 0.3 and 0.2 for the migration period
(Zav’yalov et al. 2000).

5.5.30 Kaliningrad Region

As a whole, for eastern Prussia, the Little Owl is considered a rare breeding species,
whose numbers decrease markedly as a result of severe winters (Grishanov 1994).
Population densities increase slightly from the northwestern to the southeastern parts
of the region (Tischler 1941). At present, the status in the Kaliningrad Region looks
similar to that occuring before 1940, where the Little Owl was extremely rare to the
south of the Neman River, east of the Gulf of Kursiu (Courish Gulf ) and on Samland
Peninsula, and was slightly more common in the central part of the region. In the
south, the species occurs most frequently along the border with Poland.

5.5.31 Republic of Dagestan

5.5.31.1 DISTRIBUTION
The Little Owl is resident in the Republic of Dagestan, spread widely from the shore of
the Caspian Sea and lowlands, reaching to 2500 m in the mountain-alpine regions. Due
to the arid climate, lowland (northern) desert regions and insignificant forest coverage
(7–8% of total area) of the region, the distribution of the owl is noncontinuous. The
species inhabits mountain steppes in small numbers, being scattered among other types
of landscapes, where it prefers the slopes of southern exposures, with rocks, big and
small fragment screes, and outcrops of stones that provide vertical topography. The
species often settles near humans that raise cattle and poultry, because the latter
supports the concentration of insects and small rodents. In the open landscapes of
northern (lowland) Dagestan, including the regions of the central and southern
Caspian Sea, it is especially attracted by small human settlements and cattle farms.
In the lowlands it inhabits the floodland forests along rivers such as the Terek and
Sulak, and forest belts of the tape-grove type. Part of the population inhabits big cities
and suburbs, nesting under the roofs of old buildings. While the major part of the
population is resident, birds that inhabit the mountain-alpine plots migrate in the
autumn–winter period to the foothill and flat regions (E.V. Vilkov personal
communication).

5.5.31.2 DENSITIES
Densities of Little Owl have been reported at 16–22 individuals/km² in the open
landscapes of northern (lowland) Dagestan, including areas of the central and southern
Caspian Sea region (Vilkov 2003).

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5.5.32 Republic of Bashkortostan

At the northern border of its range, the Little Owl is a very rare breeding species in
Bashkiria. Only one breeding site for this species is known in the rocky massif of Belaya
River near Kuznetsovskiy khutor. However, it is likely that the number of Little Owls is
under-estimated because settlements in the steppe and forest-steppe zones of the
Republic have not been investigated (Karyakin 1998).

5.5.33 Republic of Tatarstan

In Tatarstan, the Little Owl is recorded mainly in the western and southern districts, i.e.,
Apostovskiy, Buinskiy, Tetyushskiy, Zelenodolskiy, and also in the Leninogorskiy,
Aznakievskiy and Sarmanovskiy districts (Rakhimov 1995). Recent research points at
some shift of the border of the Little Owl breeding range in Tatarstan in a northerly
direction (Rakhimov and Pavov 1999). This northerly shift seems connected to the
rapid rate of forest logging in the area. Following humans, the species spreads into
anthropogenic landscapes in the northern half of the Republic.

5.5.34 Republic of Ossetia

Komarov (1998) reported winter bird population counts of up to 4.1 individuals/km² in
the settlements of the Ossetia Plain.

5.5.35 Tyva (Formerly Tuva)

The Little Owl is a rare, probable breeding species in Tyva. In 1983, two specimens were
sighted in the upper parts of Barlyk River at Kurgak-Sai. In August of 1984, the species
was observed at the Khapshi site and on a macro-slope of the Tsagan-Shibetu Mountain
ridge at Oruktyg and Kedyrorug. It is a rare wintering species in Tyva, observed at the
Semigorka site (Popov 1991).

5.5.36 Latvia
5.5.36.1 DISTRIBUTION
The Little Owl is very rare in Latvia (Lipsberg 1985), locally distributed in different
regions of the Republic, most often in the western part. The northern border of the
species’ breeding range crosses the territory of the country. According to most authors,
in the nineteenth century the species nested more often in Kurland than in Livland
(Meyer 1815, Russow 1880). A specific nesting place was found in 1896 and several
subsequent years in the Rembate settlement (not far from the Lielvarde settlement)
(Sawitzky 1899). In subsequent publications, authors confirm the more frequent occur-
rence of the species in Kurland than in the rest of Latvia. Nonbreeding season records in
the country are from areas surrounding the towns of Riga, Elgava, Dobele and also the
settlements Inchukalns, Rembate, Puze Misa, Olainr, Ropaji and Burtnieki (Graubitz

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 5.5 Overview per Country

1983). In the first half of the twentieth century, the species was described as a very rare
breeding species (Loudon 1906, Grosse and Transehe 1929, Transehe and Sinats
1936, Taurinsh and Vilks 1949) and found in Kurzem. Breeding was confirmed at
only two sites in Rembate, not far from Lielvarde (around 1896 and several subsequent
years) (Sawitzky 1899) and near Elgava town (in 1949) (Lipsberg 1985). All other
references deal with birds found outside the breeding season and without clear
breeding evidence.

5.5.36.2 POPULATION ESTIMATES

The Little Owl is considered rare, with an estimate of 10–30 pairs in 1988 (Manez in
Tucker and Heath 1994). BirdLife International (2004) estimated the total population at
10–30 pairs. BirdLife International (2015) and the European Environment Agency
(2021) reported extinction as of 2007.

5.5.37 Lithuania
5.5.37.1 DISTRIBUTION
The Little Owl is a rare resident breeder in Lithuania (Zhalakevichius et al. 1995) and is
considered to have a decreasing breeding population. It is assumed that only isolated
pairs breed.

5.5.37.2 POPULATION ESTIMATES

Manez (in Tucker and Heath 1994) gave an estimate of 10–50 pairs for the period
1985–1988 and considered the population stable. BirdLife International (2004) esti-
mated the total population at 5–10 pairs. BirdLife International (2015) reported 1–10
pairs during 2008–2012 and the European Environment Agency (2021) reported 5–10
pairs during 2013–2018.

5.5.38 Estonia
Although the breeding range of the Little Owl reaches western Latvia it has been
recorded only once in Estonia. Around 1880–1885, one specimen was shot at
Kolovere, Laanemaa district (Stall 1904, Lilleleht and Leibak 1992).

5.5.39 Belarus
The Little Owl is a rare resident breeding species throughout all of Belarus (Nikiforov
et al. 1997) and is estimated at 400–1000 pairs, with some decline in 1970–1980.
BirdLife International (2004, 2015) estimated the total population at 400–1000 pairs.

5.5.40 Ukraine
5.5.40.1 DISTRIBUTION
The species is a resident bird in Crimea. Some short-distance winter migrations have
been observed. On the southern coast of Crimea the species probably occurs to the east

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 Chapter 5: Distribution, Population Estimates and Trends

of Alushta town. Numbers in steppes and foothills are moderately high and distributed
relatively evenly. Along the southeastern shore (south coast) it occurs sporadically. The
owls breed in walls of the fortress on Arbat(skaya) Spit (Kostin 1983).

5.5.40.2 POPULATION ESTIMATES

Manez (in Tucker and Heath 1994) estimated 11 000–12 000 pairs in 1988, and con-
sidered the population to be increasing. BirdLife International (2004, 2015) estimated
the total population at 15 000–22 000 pairs in 2000.

5.5.41 Armenia
In Armenia, the Little Owl occurs in semi-desert, steppe and mountain-meadow zones.
Dementiev and Gladkov (1951) consider the species to be a typical bird for Caucasia.
Athene noctua indigena is characteristic of different, but predominantly open habitats,
which are situated in the piedemont climatic zone of the river basins of both the
Arax and Kura (K. Jenderedjian personal communication). Lyaister and Sosnin (1942)
mentioned the species as a resident bird, occuring mainly in places of lower elevation
and in the foothills (although up to 2000 m sporadically). It breeds in ravines and
gorges, and also in stony ruins of disused settlements and houses. The species has
also been observed in gardens. The species was recorded in the Oktemberyan district,
in the surroundings of Novyi Shakhvarut, in typical semi-desert, at an elevation of
about 900 m (Sosnikhina 1950). It was discovered in the cave of the lava mountain
Karkhan (Sosnikhina 1950). The species is distributed in three State Nature Reserves
in Armenia (Khosrov, Erebuny and Shikahogh). It was also recorded in rock outcrops
of the Goravan sands sanctuary, and in the vicinity of Yerevan (Sokolov and
Syroechkovsky 1990). BirdLife International (2004) estimated the total population at
800–1500 pairs. BirdLife International (2015) estimated the total population at
1000–1500 pairs in 2002–2012.

5.5.42 Georgia
5.5.42.1 DISTRIBUTION
The Little Owl is a year-round resident in the lowlands, plains, plateaus, foothills and
low- and middle-mountain belt of Georgia. It is uncommon in central parts of the
country, and common in eastern and southeastern areas. It is more widespread in the
Iori plateau, Karthli Plain, Alazani Plain, Gardabani Lowland, in lower parts of the Iori,
Alazani, Khrami River valleys, Shiraki and Eldari semi-deserts, Udabno and Kvernaki
ridges and in some other points in the Kura River basin. The western boundary of its
breeding range remains uncertain. It prefers dry open and semi-open habitats with
ravines, precipices and rock outcrops. The species often utilizes older buildings and
ruins for breeding. In semi-desert, nests have been found in holes and in cracks in
rock precipices. Owls sometimes nest in settlements, including the environs of Tbilisi
(A. Abuladze personal communication).

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 5.5 Overview per Country

5.5.42.2 POPULATION ESTIMATES

The present Georgian population is estimated at several thousand breeding pairs
(4000–4500 pairs at the end of the 1970s, about 3500 pairs in the mid-1980s and
3500–4000 pairs in the middle to late 1990s). During the last 25 years no alarming
trends have been detected. Numbers fluctuate annually. A slow decline was registered in
some areas in the first half of the 1980s, but the population and distribution have been
stable throughout the twentieth century (A. Abuladze personal communication).

5.5.43 Kazakhstan
The Little Owl is spread unevenly in Kazakhstan, and occurs sporadically at
certain sites. The highest breeding densities are observed in the desert zone of south
Kazakhstan, in the regions of south Pribalkhashie, in the foothills of Karatau, in the
Syr Daria River valley, along the Aral Sea shore and on the Mangyshlak Peninsula.
The northern border for Little Owl nesting sites in Kazakhstan is roughly the 49th
parallel, and only in the Ural River basin does it extend up to the 51st parallel.
It breeds in Ustyurt, where it is common in breaks in this plateau, but seldom sighted
in its central parts. On the Mangyshlak Peninsula it is common, but further north-
wards, in the northeastern part of the Near-Caspian Lowland, data regarding
its presence (or absence) are missing. It is found in the northern Cis-Caspian Sea
area, in the lower parts of the Ural and Volga Rivers and further north, and in the
Volga–Ural steppes near Urda. Along the Ural River valley it has spread north to the
Ilek River mouth. The Little Owl is extremely rare north of Ustyurt, in the desert
steppes of the Emba River basin, and in the region of Mugodzhary. The species
is well distributed and breeds in high numbers around the Aral Sea. It has been
found considerably northwards, i.e., near Chelkar Station and in the Irgiz settlement
(Gavrin 1962).
Little Owls are common, and at certain sites even numerous, in Kyzyl-Kumy, Syr-
Daria River valley and in the foothills of Karatau. In the plains of southern Pribalkash’ie,
it is spread more evenly and at certain sites it is a common bird. It is also common in the
deserts of northern Pribalkhash’ie (Balkhash Lake area), where it normally breeds north
of the 48th parallel. Northwards, in the region of the Kazakh small-knoll area, the Little
Owl is very rare and is spread extremely sparsely. In the Kyzylrai Mountains it has been
found in the stony knolls of Kotur-Kyzyltau (4920’ N), which is the northernmost
nest record of the species in eastern Kazakhstan. In the Tien-shan Mountain region,
the Little Owl breeds on the slopes of Ugam ridge, and is also found to the south of
Chimkent on the Kazgurt ridge.

5.5.44 Uzbekistan
The Little Owl is widely spread in Uzbekistan. It occurs at oases, in mountains, semi-
deserts and deserts (Sagitov 1990), in settlements, cemeteries, tugais, sands, on rock
outcrops and “kyrs” of Ustyurt (Kostin 1983). The species is considered to be a resident

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 Chapter 5: Distribution, Population Estimates and Trends

bird in Karakalpakia (Ametov 1981). On the northern slopes of the ridge in the lower
belt of mountains, the species occurs in low densities (Meklenburtsev 1936). According
to Salikhbaev et al. (1970), the species is common in the foothill plains or “adyrs.” It is
also common in the entire Zarafshan valley (Bogdanov 1956). Population densities
depend mainly on the habitat; in the Kashkadarya valley the owls are more numerous
in the foothills, and in clayish and sandy deserts (Meklenburtsev 1958). In the Nuratau
Ridge, it is common on “adyrs” and in the foothill plain. It is found in lower numbers in
the lower belt of mountains (Salikhbaev et al. 1970).

5.5.45 Turkmenistan
The Little Owl is spread throughout the country and occurs all year round. It breeds
in rock holes, in rodent burrows, on clayish precipices, in tree hollows, in walls and
under roofs of buildings, and in cornices. The distribution and population density of
the species are determined by the available food, which mainly consists of small
mammals (Bel’skaya 1992). In the western part of Turkmenistan, the Little Owl is a
resident species of the southeastern Cis-Caspian Sea area. It has been recorded
during mid-January in Krasnovodsk town, below the Nebit-Dag Mountain, and
in the Janga settlement (Khokhlov 1995). In Badkhyz, the Little Owl is a common
species, distributed throughout the region. The highest numbers are recorded on
clayish precipices of the Kyzyldzhar ravine. It is rarest in the desert steppe (Simakin
2000). This species is found throughout the area north of the river bed of Zapadnyi
(western) Uzboi. The natural and northern border of the region are the precipices
(cliffs) of the Ustyurt Plateau (Kaplankyr chink). The western boundary is the coast of
Kara-Bogaz-Gol Gulf and from there, south along the Caspian Sea to Iran (Zarkhidze
and Loskutova 1999).

5.5.45.1 DENSITIES
In northwestern Turkmenistan, Athene noctua bactriana occurs everywhere. In
1989–1991 the population density in different habitats ranged from 0.4–5.0 individ-
uals/km² (Zarkhidze and Loskutova 1999). In the Repetek Nature Reserve, fluctuating
densities were found in Black Saxaul thickets (0.7–1.8 individuals/km²), and in White
Saxaul thickets (0.3–1.2 individuals/km²) (Ataev 1977).

The global distribution of the Little Owl indicates a presence in the following countries,
but no data on the population was available for this project.

5.5.46 Iran
Ilanloo et al. (2020) built the first country-level habitat suitability map for the Little Owl
and identified the most influential environmental predictor of its distribution in Iran.
Using 177 distribution points of species presence data (Figure 5.34) and eight uncorrel-
ated environmental variables, a species suitability habitat model was estimated using
the maximum entropy approach (MaxEnt). Results showed that suitable habitats for
this species are located in the northwest, parts of the northeast and in southwest Iran.

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 5.5 Overview per Country

Figure 5.34 Little Owl observations in Iran 2012–2021. After GBIF (Global Biodiversity Information
Facility) Occurrence Download https://doi.org/10.15468/dl.mnkcux.

Distance to forests, distance to human settlements and distance to agricultural lands are
the most important determinants of the Little Owl distribution in Iran. Human migra-
tions from rural areas to the cities and availability of many vacant human settlements
surrounded by traditional and semi-modern agricultural landscapes produce suitable
habitats for Little Owl in Iran.
Results of estimating area of suitable habitats of the species within the agricultural
lands showed that 151 718 km² of species-suitable habitat are within agricultural areas
in Iran. Most of the suitable agricultural habitats are in northwest and northeast of the
country (Figure 5.35).

5.5.46.1 DENSITIES AND POPULATION ESTIMATES

When using an average density of one observation per km² we would get a population
estimate of 150 000 occupied 1
1 km grid cells for all suitable agricultural areas in Iran.

5.5.47 Tajikistan
While the species is widely spread, it is mainly associated with desert landscapes. In the
northern part of the Republic it inhabits suitable habitats of Mogol-Tau, in “says” of the
Kuraminskiy Mountain ridge, in the Syr-Dar’ya River valley (in the vicinity of Leninabad,
Kairak-Kum and further to the border with Uzbekistan), and on the northern slope of the
Turkestan ridge (surroundings of Shakhristan, in the valley of Ok-Tanga at 2400 m). It also
occurs on the southern slope of the Turkestan Mountain ridge. The species was found in the
Iskander-Kul’ Lake area (Ivanov 1940), and in Gissar-Karategin (Popov 1959), where it is
most commonly found in the foothills. It is found throughout the Gissar Valley, occupying
habitats suitable for nesting (Ivanov 1940, Popov 1959). In southern Tajikistan the Little

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 Chapter 5: Distribution, Population Estimates and Trends

Figure 5.35 Little Owl-suitable habitats within agricultural lands in Iran (after Ilanloo et al. 2020).

Owl nests in the mountains of Rangin-Tau, Aruk-Tau, Ak-Tau, Buri-Tau, Kara-Tau and
Khazratishokh, and from the banks of Pyandzh River to the foothills of the Darvaz
Mountain ridge. The species has been sighted frequently in the valleys of the Vakhsh and
Kafirnigan Rivers, and on the lower portions of the Yakh-Su and Kizyl-Su streams (Ivanov
1940). In Badakhshan the species was found in the upper parts of Shakhdara (Meklenburtsev
1946), in the upper parts of Vakhan-Darya (Bazai-Gumbez) (Ivanov 1940), in the Drukhmatz
River valley, and in the surroundings of the Pamir Botanical Garden (Popov personal
communication). In all probability, the Little Owl is widely distributed in Badakhshan, but
due to small numbers it remains unrecorded by ornithologists in other areas. The species
occurs in the southern part of Pamir too, i.e., in the Beik River valley, Ak-Su tributary (Ivanov
1940), in the surroundings of Zor-Kul Lake and the Bash-Gumbez site (Abdusalyamov 1971),
and near the Soviet farm “Pamir” on Bulun-Kul (Meklenburtsev 1936).

5.5.48 India (Ladakh)

5.5.48.1 DISTRIBUTION
Ladakh is an area through which the line of control (as per the 1972 SIMLA Agreement)
between Pakistan and India passes. Although a major part of Ladakh lies in Indian-held
Kashmir, a sizeable part is present on the Pakistani side of Kashmir. Further in the north

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above Saltoro Range lies the Siachen Glacier where the two armies are fighting each
other at the highest battlefield in the world. The region contains all the Little Owls in
India. It lies above 4000 m and includes some extremely desolate areas (N. Paklina and
C. van Orden personal communication).
Limparungpatthanakij et al. (2017) observed a Little Owl at Se La Pass, Arunachal
Pradesh, India, situated near the eastern border of Bhutan (27.51N, 92.11E) at 4200 m
a.s.l. The habitat was open with rocky terrain and scattered trees. So far the only docu-
mented record in Arunachal Pradesh was a specimen collected in the Mishmi Hills (Baker
1926), about 400 km to the east, the same paper wherein Baker described Athene noctua
ludlowi. Further details of the Mishmi specimen are unavailable and it is unclear where
exactly in the Mishmi Hills it was recorded, and the season when it was collected. However,
the Little Owl is treated as resident in northeastern Sikkim (Ganguli-Lachungpa et al.
2011), with photographic evidence, and the species was recently added to the Bhutan list,
based on a record from Nub Tshonapatra in far western Bhutan (Wangdi 2015).
The Little Owl may be resident at Se La Pass, as it is known to be resident on the
Tibetan Plateau and the nearby mountainous regions of the Himalayas (Grimmett et al.
2011). The subspecies ludlowi is stated to be present throughout the year, at all elevations,
between Gyantse and Phari (Baker 1926). Another sighting at the same site, on May 1,
2017, by Jainy Maria, Yann Muzika and Rufikul Islam seemingly supports its likely
resident status. Nevertheless, further investigation, and more reports of sightings from
the area are needed to confirm its seasonal status (Limparungpatthanakij et al. 2017).

5.5.48.2 POPULATION ESTIMATES

The region of Ladakh held some 120 pairs in 2003 (N. Paklina and C. van Orden
personal communication).
The global distribution of the Little Owl indicates a presence in India, Pakistan,
Nepal and Bhutan, but no data on the population were available for this project.
However a map of the Little Owl’s distribution in these countries, from Grimmett
et al. (1999) is shown in Figure 5.36.

5.5.49 Bhutan
The Little Owl has only recently been recorded in Bhutan. The Little Owl is resident in
Baluchistan and the trans-Himalayas, living on cliffs and ruins in semi-desert (Wangdi 2015).

5.5.50 Mongolia
The Little Owl occurs in natural habitats and anthropogenic environments and is
considered rare everywhere in Mongolia. Breeding was observed in the Chentej
Mountains near Boroo, near Chudshirt and in the Gobi Desert. Other records are from
Somon, Buucagaan, Jarantaj (border with China), Conocharajch-gol, Ulaan-uul,
Chvod, Ajmak (southern Gobi) (Schönn et al. 1991). In Mongolia, the Little Owl is
considered a rare south-Eurasian species, occuring as a breeding species in the desert
mountains and stony deserts of northwestern Mongolia and southeastern Altai

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Afghanistan
Iran

Kuwait Pakistan

India
Nepal Bhutan
Bahrain
di Arabia United Arab Emirates
Bangladesh

Oman
Myanmar

Yemen

outi

Figure 5.36 Map of Little Owl distribution in India, Pakistan, Nepal and Bhutan (after Grimmett et al.
1999).

(Rogacheva 1988). The Little Owl is probably a vagrant species. The only specimen
observed within the borders of the Krasnoyarsk Territory was in 1908 in the Us settle-
ment. Overall, the owl breeds in stony deserts, semi-deserts and dry steppes with
separate rocks or stony ridges; in the mountains, it occurs up to 2800 m (Sushkin 1938).

5.5.51 Tibet
5.5.51.1 DISTRIBUTION
The Little Owl is common in a large area of northern China (Fu-Min Lei personal
communication), and the Tibetan Plateau features some isolated breeding populations
above 4000 m in extremely desolate areas (N. Paklina and and C. van Orden personal
communication).

5.5.51.2 POPULATION ESTIMATES

In 2003, the population of the Tibetan Plateau was estimated at 50 000 pairs after
extrapolation from the density in Ladakh (N. Paklina and C. van Orden personal
communication).

5.5.52 China
The northern border of the Little Owl distribution range crosses northeastern China,
the southern goes through Tschajul, Yaan, Daxian, Nanyang to the south of Xinhailian
and the Yellow Sea. Qingdqo was registered as the eastern-most point of occurrence
(Schönn et al. 1991).
Little Owls are mainly distributed in foothills near the forest, prairie and hillside
fields. The most suitable habitats in Qishan (34.4 N, 107.6 E) are found to be in open
country with scattered trees, earth banks and crevices. The population density differs
according to the season; the average density is 0.04 pairs/km² in winter (January and
February) and 0.21 pairs/km² in summer (June). The average distance between nests is

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 5.5 Overview per Country

Figure 5.37 Distribution of Little Owl subspecies in China (after Lei et al. 1997).

189 m, and the shortest distance between two nests is less than 100 m. However, they
are now rarely found in the Weinan, Meixian and Huxian counties of the Shaanxi
Province, and in the Laishui and Yixian counties of the Hebei Province (Figure 5.37).

5.5.53 Korea
The most recent record of the Little Owl dates from 1960, when a dead Little Owl was
found by one of the employees of the Army Post Exchange in Seoul, Korea. It was said to
have been found in the folds of a tarpaulin on a supply truck that had arrived that
morning from the Port of Inchon on the Yellow Sea, approximately 40 km west of
Seoul. According to Austin (1948), only three records of the species exist for Korea, all
based on specimens taken by Won in North Korea (Pyongan Namdo) in November
1931 and February 1932.

5.5.54 Syria
The Little Owl can be observed in most parts of Syria. In and around Damascus the
species is relatively rare; it may be absent from the intensively farmed land of Ghouta.
In the fertile regions around Homs, Hama, Idlop and Aleppo, the species is common
and occupies houses, barns and stables. Historical ruins (e.g., Palmira) are a favorite

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 Chapter 5: Distribution, Population Estimates and Trends

breeding place. In the Euphrates River valley and in the desert, the Little Owl breeds in
stone piles (Schönn et al. 1991).

5.5.55 Jordan
5.5.55.1 DISTRIBUTION
A fairly common resident in the western parts of Jordan, the Little Owl is scarce to rare
in the east. It is found in the highlands, rift margins and parts of the Jordan Valley, and
in true desert areas. In the highlands, it is fairly common in open habitats with rocks
and low vegetation, but never in woods or deep gorges.
In the rift margins the species is a common resident from the northern areas,
southward to the slopes that face the Dead Sea, where it becomes scarce. It is also rather
uncommon in the rift margins/slopes facing Wadi Araba, the Aqaba Mountains and
Wadi Rim. In the eastern desert, the Little Owl is common in the areas just east of the
highlands, which slope gently towards the eastern desert plateau. This area is character-
ized by rolling open country with rock boulders and low steppe vegetation. In the
eastern desert it is only locally present, usually in the few limestone hills in the flat
hammada/flintstone desert and around farms. It is more common and widespread in
the northeastern desert, which is covered by basalt stones and rocks (Andrews 1995,
Khoury personal communication).

5.5.55.2 POPULATION ESTIMATES

The population size is estimated at several thousand pairs (Andrews 1995, Khoury
personal communication).

5.5.56 Israel
5.5.56.1 DISTRIBUTION
The Little Owl is resident in most areas and is found as territorial pairs during the
breeding season. Outside the breeding season it normally occurs as singles, but some
pairs remain together throughout the year. Immature birds wander and disperse short
distances. The most continuous and dense population occurs along the central moun-
tain range, including Golan, Galilee, Carmel, Shomron and Judea. In Hermon, it is a
rare breeder up to 500 m (Figure 5.38). There is a quite dense population in the Jordan
River Valley, especially on cliffs along the Jordan River itself. It is more widely dispersed
in the northern valleys and along the Coastal and Judean Plains (in the Dan region,
mainly in the eastern hills). It is common throughout much of the Judean Desert,
mainly northwestern parts, near the Judean Hills. It is almost totally absent in the low-
lying arid areas of the Judean Desert, in the Dead Sea Depression and along the Arava.
In Negev it occurs continuously, mainly in mountainous regions north of Nahal Paran,
in the Makhteshim, Nahal Zin area and in northeastern Negev between Arad and
Yeroham. The densest concentration in the desert and semi-desert regions occurs in
northwestern Negev, Holot Halutza, Nizzana, Shivta and eastwards to Mashabbe Sade.
In the Eilat Mountains and the town of Eilat, the species appears occasionally as a

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 5.5 Overview per Country

Figure 5.38 Little Owl distribution in Israel (after Shirihai 1996).

wanderer or migrant, though a few breeding pairs are scattered in “wadis” (an annually
dry riverbed or valley) north of the Eilat Mountains and northwards in the area of Biq’at
Sayyarim and Shizzafon. In all parts it breeds mainly outside human settlements or on
fringes of villages and towns. A few breed in local enclaves within cities such as

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 Chapter 5: Distribution, Population Estimates and Trends

Jerusalem, where there were 30–40 pairs within 68 km² (0.44–0.59 pairs/km²) in the
1980s and a similar number in Haifa (Shirihai 1996).

5.5.56.2 POPULATION ESTIMATES

The total population in Israel was estimated in the 1980s at several thousand pairs
(Shirihai 1996).

5.5.57 Arabian Peninsula

5.5.57.1 DISTRIBUTION
The Little Owl is a widespread, locally common resident in the deserts of Arabia
(Figure 5.39). There is no information to suggest seasonal or other movements and
it is generally thought of as sedentary, occuring in all states except Bahrain. It has
occurred on Das Island in the middle of the Arabian Gulf, evidence that some dispersal/
movement is taking place. There are a number of records from the periphery of the
Empty Quarter, but it appears to be absent from the core area. This is likely due to the

Figure 5.39 Little Owl distribution in the Arabian Peninsula (after M. Jennings personal
communication).

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 5.5 Overview per Country

lack of suitable roosting/nesting sites rather than the species itself not being tolerant of
hyper-arid environments. The owl is scarce in the highland areas of the southwest,
except for the drier eastern fringes. It is resident on Farasan Island, but generally scarce
at other coastal sites and is not known from Masirah or Socotra Islands. It is a common
bird in the United Arab Emirates, where it features a much higher breeding density than
is likely in other states except possibly northern Oman (M. Jennings personal
communication).

5.5.57.2 POPULATION ESTIMATES

The total Arabian population may be of the order of 5000–6000 pairs. There is no
evidence to suggest populations are changing in any way, but the huge expansion in
agriculture in central and northern Arabia in recent decades is likely to have allowed
many more to breed than did previously. In the United Arab Emirates the population
has been estimated at 300–1000 pairs, giving a much higher breeding density there
than is likely in other states, except possibly northern Oman. This population includes
an estimated 50 pairs at Jebel Hafit on the United Arab Emirates/Oman border.
Population numbers in pairs are: Kuwait 50, Oman 1000, Qatar 50, Saudi Arabia
3500, United Arab Emirates 600, Yemen 500, Bahrain – not available (M. Jennings
personal communication).

5.5.58 North Africa

The species is widespread to the north of the Sahara from Morocco to Egypt, with an
extension to the south along the Red Sea coast to eastern Ethiopia and Somalia. The
upper elevational limits in its distribution are 700 m in the Beni Aros Mountains,
1100–1200 m in the High Atlas, 1500 m at Touggana, 1250 m at Iframe, 2200 m to
the west of Bon Iblanc and above 2000 m near Khenifra (Heim de Balsac and Mayaud
1962).
The exact southern distribution range remains unclear in several countries in north-
ern Africa. The species was reported to Bojador Cape (Western Sahara), Ain Najla,
Smara, Tifariti, Tarfaia, Seguiet et Hamra and Guelta Zammour. It probably reaches
south of Rio de Oro and further to Mauretania. It is believed to occur south of the 30th
parallel in Morocco. In Algeria the owl is found even more southerly to Zirara, l’Qued
Mya. In Libya, the southern boundary of its range remains unclear, around Tripoli and
Cyrenaica. In the southern area of northern Africa, the distribution is made up of
isolated population clusters, e.g., around Timbuktu (Mali), the Ahaggar Mountains,
Aïr (Azbine), Serin Tibesti, Ennedi and Jebel Marra (Sudan) (Schönn et al. 1991).

5.5.59 Egypt
In Egypt, the northwestern border of the Little Owl’s distribution remains unclear. In
eastern Egypt, the distribution area covers the delta of the river Nile up to the Sinai, to
the Red Sea coast through Sudan and Ethiopia to Somalia. Following the river Nile, the
species breeds further south than the Assuan Dam (Schönn et al. 1991).

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 Chapter 5: Distribution, Population Estimates and Trends

5.5.60 Libya
While the Little Owl is common in the northern part of the country, the southern limit
of its distribution remains unclear, but is probably around the border with the Oubari
Desert. In the east, the species is found from Ajadabia to the Egyptian border. The
species is believed to breed around Fezzan. Owls observed in Goddua, Traghen, Hun
and Sebha confirm the sporadic occurrence and migratory behavior in this region
(Schönn et al. 1991). Resident breeders are common in open areas throughout
Tripolitania extending, eastwards to the Sirte area (old olive groves and arid, rocky
desert) (Isenmann et al. 2016). Further south, breeding is claimed only as probable in
Fezzan (occurrences recorded at Al Qurdah, Goddua, Ghat, Hun, Qatrun, Sebha). In
Cyrenaica Little Owls were recorded on May 26, 2008 near Lake Ain Azziana, Benghazi
and on May 27, 2008 at Old Al Marj and Cyrene. Two birds found dead on a road
on January 5, 2011 near Kabao referred to saharae. At Djanet/Algerian Sahara, close to
Ghat, the diet included 70% insects and 17% rodents (Sekour et al. 2011).

5.5.61 Tunisia
The Little Owl is a sedentary breeding species occuring from the Mediterranean coast to
the Sahara Desert. It was considered the most common owl species in Tunisia, occuring
in open or scarcely forested areas along fields with walls or rocks (Isenmann et al. 2005).
The species is widespread across the country in both the dry, unfertile desert mountains
of the south and the green fertile regions of the north. In central and northern Tunisia,
the species breeds in olive plantations (Schönn et al. 1991), widely distributed from the
coast to the Saharan regions. In 2004, 18 pairs were recorded in a 2 km² area, surround-
ing a northern village with fields, piles of stones, and occupied and abandoned houses
(B. A. Hajer personal communication). A brood of four 1-week-old pulli was observed in
Khanget Oum Ali on May 26, 1993 (H. Dlensi personal communication). The diet is
mostly insects, scorpions, lizzards and gerbils, in the Thyna salt-pans also bats and birds
and in Ichkeul mainly insects (Isenmann et al. 2005). It has been observed as far south
as Borj Bourguiba and Dehiba.

5.5.62 Algeria
The species is fairly widespread from the Mediterranean coast to deep into the desert,
reaching the Ahaggar where it is rare (Isenmann and Moali 2000). There are records of
this owl up to 1400–1700 m on the Holm Oak-covered southern slopes of the Aurès,
and up to 1600 m on scree and broom slopes in the Djurdjura Mountains. It is less
strongly linked to trees than the Eurasian Scops Owl Otus scops, and takes advantage of
more open areas with walls, wells, rocks and screes (Isenmann and Moali 2000).

5.5.63 Morocco
The Little Owl is a resident breeder in Morocco (A. n. glaux in nondesert areas and
A. n. saharae in the desert). It is common and widespread in Rif, the “North Atlantic,”

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 5.5 Overview per Country

Central Plateau and “Mid Atlantic” (Thévenot et al. 2003). It is also locally common in
eastern Morocco from the Plains to the Saharan Atlas. It can be found frequently up to
1500 m in the Middle and High Atlas and is rarer above. The owl occurs south of the
High Atlas, is common in Souss, Anti-Atlas and “East Sahara” (A. n. glaux, but saharae
recorded twice in Tafilalt and once in Souss at Massa).

5.5.64 Western Sahara

In Western Sahara, it is locally common along the Atlantic coast in Lower Draa, Tarfaya
and south to Boujdour in Saquiat Al-Hamra, but rare inland at Aïn Nakhla, Smara,
Tifariti and Aïn Tahla. Rare further south in Oued Ad-Deheb and only reported from
near Guelta Zemmour, Sbayera, Negyir and Awserd (22 40’ N) in the far south
(Thévenot et al. 2003).
The species occurs in good densities in the entire coastal region of Bas Draa up to the
Oued Draa valley, e.g., more than 20 birds between Plage Blanche and Foum Assaka
(May 15, 1994) and five birds between Ras Oumlil and Aoreora (January 27, 2011). It is
less common in mainland Bas Draa with observations in Jbel Bani and Assa (Bergier
et al. 2017), and common in coastal Saquiat Al Hamra, with observations for the
Tan-Tan plateau, the gueltas of Oued Chebeika, the hinterland of Akhfenir, Rhouiba
plateau, Oued Ez Zehar, Khaoui Naam, Ténouchad, region of Khnifiss, Cap Juby, north-
east from Tarfaya and Amgriou, south of Sebkha Oum Dba and Oued Khwiy As-Saraq,
the Dawra, from the valley of Saquiat Al Hamra to Dchira with pairs along the Laâyoune
Lake and the hinterland of the port in high densities. In the interior Saquiat Al Hamra,
owls have been observed in Hassi Amit south of Ouarkziz, east of Zag, south of M’Sied,
near Abatteh near the Aydar massive, between Laâyoune and Smara, in the valley of
Saquiat Al Hamra in Aïn Najla and east of Sidi Ahmed Laroussi, more south between
Laâyoune and Bou Kra, halfway between Bou Kra and Guelta Zemmour and in Sbayera.
Little Owls are much rarer in northern coastal Oued Ad Deheb, with five birds between
Boujdour and Dakhla, one at Lamjayir and one at the mouth of Oued Kraa. They are
even more rare to the south, with observations around El Argoub, Gleib Jédiane and
Imlili, and very local in interior Oued Ad Deheb in Graret Ouchfegt, Negjyr massive,
Tiris, Oued Jenna and Aousserd. The southernmost observation is east of the Boulariah
dam (2226’45”N 1431’20”W) on October 21, 2014 (Bergier et al. 2017).

5.5.65 Mauretania
The Little Owl is a resident breeder in sandy and rocky locations in Saharan Mauretania
from Tiris-Zemmour south as far as 17N (Wadab, Alké Awana). It has not been recorded
in Senegal.

5.5.66 New Zealand

The Little Owl was introduced to New Zealand by the Otago Acclimatisation Society
(Otago Fish and Game Council) in 1906 when 28 birds were imported from Germany

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 Chapter 5: Distribution, Population Estimates and Trends

(subspecies Athene noctua vidalii), hence it is often referred to as the German Owl in New
Zealand rather than the Little Owl. They were released in an effort to control the
numbers of small introduced birds, which were considered a pest by orchardists.
Another 39 owls were brought to Otago and released in 1907, 80 more in 1908 and
another 72 in 1910 (14 were released and the rest sold to farmers). All owls were
thought to originate from Germany. Some may have been released in North
Canterbury. A pair was released in Rotorua on the North Island in 1908. Some were
said to have been transferred unsuccessfully to the North Island, possibly after 1908.
The species is now widespread in the farmlands of Nelson, Marlborough, Canterbury,
Otago and Southland. None appear to exist on the North Island. On the South Island
they are to be found east of the main divide, although there have been reports of them
in Westland and Fiordland. They are plentiful in eastern Otago, and Otago and
Southland contain the greatest number of Little Owls anywhere in New Zealand
(C. Sawyer personal communication).

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 Chapter 6: Habitat

Chapter Summary
Habitat is one of the main drivers of species presence, and it needs to be able to support entire
populations of Little Owls year round. For Little Owls, habitats will typically include open
hunting ground rich in small prey, hunting perches, day roosts and nest holes, with a benign
climate and land management regimes that give reasonable long-term continuity. These
habitat aspects can be met within a wide diversity of natural and anthropogenic landscapes.
Recent declines in numbers and distributions across much of Europe show that tolerable
limits are being exceeded. These results are consistent with the view that agricultural change
has influenced birds through changes in food quality or quantity (Benton et al. 2002).
In this chapter we review the parameters that are of importance for the species, its prey
species and its predators. The favored habitat for the Little Owl varies from the natural
landscapes of steppes and arid deserts to anthropogenic areas. The common features are
open areas with low grass, perches and cavities in the ground, rocks and trees or build-
ings. The species avoids forests, fallow land and large parcels of arable land. A mosaic
effect seems to be important for Little Owls, due to the use of habitat edges, in particular
for the richness of prey found there. The relations between the landscape factors will
determine local owl densities and demographics. All quantitative studies available were
done on anthropogenic habitats. For natural habitats, only qualitative descriptions were
available. We first discuss natural habitats in general terms, then we give an overview of
different types of occupied anthropogenic habitats, followed by the actual preference of
the species toward certain habitat parameters. The latter studies entail both occupied and
unoccupied habitats, while habitat typology studies consider only occupied habitats.
Understanding the parameters that have an influence on habitat suitability for Little
Owls is not always easy, since some studies yield contrasting results. The reasons for the
different outcomes of studies are mostly related to the different local circumstances per
study, with the size of the area studied and with the spatial scale at which the research
was done. We illustrate contrasting results at different landscape scales. While it is
recognized that models are just an approximation of the truth, modeling Little Owl
presence with different techniques can help to interpret and understand the behavior
of the species at different spatial scales. Multiscale habitat assessments enhance our
understanding of habitat selection, especially when combined with long-term demo-
graphic data. Further, models allow prediction of habitat suitability and help to illu-
minate conservation priorities. Interpretation and communication of the results is

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 Chapter 6: Habitat

enhanced by transparent and repeatable methodologies. The results of such examin-

ations allow us to view Little Owl habitat use from an array of perspectives, and offer
insights into key landscape components and associated management options.
In Plates 9, 11, 14–19 (plates ludlowi, orientalis, plumipes, bacteriana, impasta, spilogas-
tra, somaliensis, saharae), we offer illustrations of natural habitats taken in various
locations across the Little Owls’ range. We have selected these to reflect important
and typical habitat conditions that have been demonstrated to be of significance to
Little Owl populations. Readers will easily notice striking differences in ecological
aspects of the landscapes used by Little Owls.
In Plates 3–8, 10, 13 (plates noctua, vidalii, indigena, sarda, lilith, glaux) we offer
illustrations of anthropogenic habitats and habitat elements taken in various locations
within the global distribution of the Little Owl. We have selected these to reflect habitat
conditions that have been demonstrated to be of significance to Little Owl populations.

6.1 Natural Habitats

Natural habitats in moderate and warm European regions, northern Africa and Asia are
comprised of ravines, gorges, gullies, walls of river terraces, precipitous cliffs and dry
unwooded mountains, as well as dry hilly steppes, semi-deserts, and sandy or clay deserts.
The few vertical structures, containing cavities that are available in these habitats are
exploited by the species for nesting. In steppe-like areas the species compensates for the
lack of these upstanding structures by occupying cavities in the ground dug by mammals.
In Morocco, north of the High Atlas, the species inhabits open land, wadi edges,
matorral, palm groves, coastal cliffs, inland cliffs and rocky hillsides. At elevations of
2000–2600 m at Jbel Yagour and the western High Atlas, the species occurs on upland
meadows with scattered boulders and low herbage. In predesert and desert areas it is
found on open, spiny bush-covered slopes (Crassicauletum with Euphorbia), palm and
Argan groves, open acacia woodlands, cliffs and embankments (Thévenot et al. 2003).
In Somalia the species breeds in holes in steep dry-river banks. Termite heaps are
used as perches and to breed in (Schönn et al. 1991).
In the Arabian Peninsula, Little Owls are found in rocky or broken country or in
open well-wooded areas, spending the day in rock crevices and tree holes. They do not
frequent precipitous cliffs and although they have been found up to 2600 m in Yemen
and 2200 m in the northern Oman highlands they seem more numerous in nonmoun-
tainous habitats (M. Jennings personal communication).
In Afghanistan, the species is mostly found in the middle-high mountains and along
the valleys in the steppe regions where willows, poplars, tamarisks and loam walls offer
breeding sites (Schönn et al. 1991).
In Uzbekistan, isolated pairs occupy old wells in the deserts and nest in the holes of
rocks, in clayish and sandy deserts, river valleys, on loess precipices and in the hollows
of trees (Sagitov 1990). In the desert they nest in the burrows of Great Gerbils Meriones
opinus and Thin-toed Ground Souslik Spermophilopsis leptodactylus or in the nests of
Rock Nuthatch Sitta neumayer (Zarudnyi 1896). The species probably digs burrows

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 6.2 Anthropogenic Habitats

independently, as they improve existing burrows at the end of the tunnel, creating an
internal room or chamber (Pukinskiy 1977).
On the Kazakh Plain, the Little Owl inhabits various habitats, occuring equally often
in sandy desert, clayish desert and in semi-desert (Osmolovskaya 1953). The main
requirement is the availability of shelter, such as a burrow or deep hole, in which the
birds can roost and breed. Most often nests can be found in the vertical walls of wells, in
which they use burrows dug by Rollers Coracias garrulus. In sands they settle in aban-
doned burrows of Steppe Tortoises, foxes and badgers (Spangenberg and Feigin 1936).
In the desert zone the Little Owl willingly settles in river valleys with precipitous banks
and tugai groves. The link with the cultural landscape and human settlements is
different in the separate Kazakh subspecies. With rare exceptions, the desert subspecies
A. n. bactriana clearly avoids nesting in settlements, occupied houses and in industrial
constructions. The south European A. n. indigena and the Asian mountain A. n. orientalis
subspecies more frequently occupy cultural landscapes and human dwellings (Gavrin
1962). In the northern steppes, the species is absent because the tall vegetation inhibits
its ability to hunt on the ground (Schönn et al. 1991).
In Crimea, the owl is a typical rocky-steppe species and inhabits rocky outcrops in
steppes and foothills, shell open mines and coastal precipices (Kostin 1983). The species
inhabits the desert mountains and stony deserts of northwestern Mongolia and south-
eastern Altai (Rogacheva 1988). In Tadjikistan the species occupies bank precipices and
also settles in gullies of foothill areas. Loess precipices are the most favorable nesting
habitat. In the mountains, the Little Owl breeds in niches of rocks on pebble slopes
(Abdusalyamov 1971). In the Volga-Kama territory, the species is widely spread in open-
steppe or forest-steppe habitats (Kulaeva 1977). On the Kyzylkum sands, the species
inhabits practically all wintering sites on the gray-ground plain and in the depths of
sands. The species breeds in small numbers in “ostantsakh” or isolated remains of
mountains, occupying empty cavities in the ground (Gubin 1998).
In the mountains of central and east Asia the species is found on the treeless and
shrubless steppes, where the Little Owl is a clear ground dweller. It has adapted to the
environment by mainly roosting and breeding in burrows of Himalayan Marmots
Marmota himalayana, which seem to be a limiting factor for the owl’s distribution
(Schönn et al. 1991).
In Asian desert areas, the Saxaul sand deserts are frequently occupied by Little Owls. In
the Mongolian northern Gobi Desert, large undulating sand planes with Saxaul trees
(Haloxylon ammodendron) intermingle with steep sand cliffs where holes created by erosion
offer nesting cavities. These holes are complemented by burrows dug by mammals like
pikas Ochotana, Siesel Spermophilus citellus, gerbils Rhombomys and voles Microtus in north-
ern Gobi (Schönn et al. 1991). Smaller mammals serve as prey for Little Owls.

6.2 Anthropogenic Habitats

The essential criteria for Little Owl habitat can be found in many different combin-
ations of landscape parameters. The types of habitat described here reflect only those
habitats actually occupied by owls.

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In central Europe, pastures and meadows flanked by pollard trees provide ample nest
sites, hunting perches and year-round short herbage with plenty of invertebrate prey.
These components reflect optimal habitat and are characteristic of conservative small
farming economies (Exo 1983, Schönn et al. 1991).
Around the Mediterranean Sea maquis vegetation areas are widespread and widely
used by the owls. This vegetation is limited to 3 m in height and features different
species of oak (e.g., Quercus ilex, Q. coccifera, Q. suber) that offer holes that are frequently
used by the owls. Olive plantations are favored in northern Africa (Schönn et al. 1991).
On the outskirts of Ankara (Turkey) the Little Owl inhabits, at very high densities,
apartment buildings that are under construction (E. Vaassen personal communication).
The species occurs in settlements, rocky gorges, on forest edges, inhabiting solitary
buildings and isolated trees with hollows amidst fields in Bulgaria. Modern urban
landscapes provide good conditions for the species (Iankov 1983).
The Little Owl occupies a great variety of habitat types from sea level up to 1000 m in
Portugal. Frequented habitat types include a great variety of agricultural fields with
hedges of different types and in certain regions of extensive crop fields with some trees
or rocky outcrops. It is found in different kinds of agro-forest systems, mainly the
extensive systems of cork and Holm oaks and of chestnut woods. In these habitat types
it is normally more common in areas with old trees featuring cavities, especially in cork
and chestnut. It also frequents parks and gardens in cities, small towns and villages. In
Lisbon it is not a rare species, frequenting the great urban parks and some gardens
(L. Reino personal communication).
In Spain, the species shows a preference for open habitats, being found in olive
groves, cereal fields, orchards, vineyards, vegetable gardens, wastelands, scattered
wooded areas and populated areas. It avoids excessively wet zones, dense forests and
high mountain habitats (Muntaner et al. 1983, Olea 1997).
In Britain, the species is found in agricultural countryside well endowed with hedge-
row trees and farm buildings, old orchards in parkland, drained fenland with lines of
pollard willows Salix, and marginal landscapes such as industrial waste ground, sand
dunes, moorland edges, old quarries, sea cliffs and inshore islands, treeless rising
ground and settlements (Sharrock 1976). It is found only infrequently and imperman-
ently within major cities (Sharrock 1976).
Old orchards are particularly valuable to Little Owls, and their use is widely recog-
nized (Juillard 1984, Fuchs 1986, Génot 1990c). Even under intensive management,
occupancy of orchards by owls may continue, provided a nucleus of old trees remains.
In areas of West Flanders densely populated by Little Owls, modern low-stem orchards
with integrated fruit growing (reduced pesticide use) offer good habitat when nestboxes
are provided.
In Iran the most suitable habitats are found in agricultural areas (Ilanloo et al. 2020).
Half of Iranian lands are covered with high mountain ranges and large sections of its
central areas are covered by arid plains where the owl does not occur. Suitability of
agricultural lands was modeled and showed that distance to forests (51.2%), distance to
human settlements (16.2%) and distance to agricultural lands (15.5%) were consecu-
tively the most important variables in habitat selection in Iran. Avoidance of forest and

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 6.2 Anthropogenic Habitats

forest edges was recognized as a response to the presence of the Tawny Owl (Zabala et al.
2006, Michel et al. 2016). Human settlements include villages featuring traditional
agricultural lands and traditional and semi-mordern cultivation systems. Ilanloo et al.
(2020) found that the species is more commonly observed in human settlements
compared to natural areas for breeding. Agricultural lands are mostly used for foraging
and rural areas for breeding. Considering the enhancement of immigration from rural
to urban areas, vacant human dwellings surrounded by agricultural lands provide
suitable habitats.
The Little Owl mainly occupies agricultural landscapes with tree lines and farm
buildings in New Zealand. When shrubs and bushes are present, the species also
occupies mountains of middle to high altitudes. Farmland and plantations with Pinus
radiata, poplars and willows offer hunting grounds with commanding perches and
breeding cavities. In cities, the owls can easily forage on the ground in parks due to
the short vegetation. Around Alexandra, Central Otago, apricot plantations are charac-
teristic habitats. The plantations are mostly surrounded by pine or poplar trees. Near
cities, old trees and sand cliffs are occupied when available. Goldmining activities
during previous centuries created new cliffs and holes in the landscape (Schönn et al.
1991).

6.2.1 Farmland Heterogeneity

The amount of high-quality habitat patches in farmlands, their distribution and the
resource accessibility therein play a key role in regulating habitat effects on reproduct-
ive success. Heterogeneous habitats offer nonsubstitutable resources (e.g., nest sites and
food) and substitutable resources (e.g., different types of food) in close proximity,
thereby facilitating landscape complementation and supplementation. Michel et al.
(2017) studied how spatial resource separation in homogeneous agricultural landscapes
affects reproductive success see Chapter 10). Decreasing farmland heterogeneity con-
strains population productivity by increasing separation of food resources from nest or
roost sites, resulting in low landscape complementation, and reducing alternative food
resources, limiting landscape supplementation. Michel et al. (2017) suggest that struc-
tural heterogeneity affects landscape complementation, whereas the heterogeneity and
management of farmland affect landscape supplementation.

6.2.2 Mosaic-Like Landscapes

Landscape heterogeneity is the only determining factor for the species in some areas of
France, highlighting the importance of optimal mosaic structure (Ferrus et al. 2002),
rather than a minimal occurrence of some landscape elements. Little Owls prefer
landscapes in Flanders with intermediate openness while they avoid extremely open
(e.g., lakes, dunes, large parcels) and extremely closed (e.g. forests, built-up) landscapes.
Mosaic-like landscape structure was found to be more important than the actual
land-cover types. The perimeter of the grassland parcels or the number of parcels rather
than their area have a positive influence on habitat suitability (Van Nieuwenhuyse and

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Bekaert 2001b). Landscape heterogeneity (convergence of points for three or four areas
having different agricultural land uses) and structuring of grassland were related to
population density in Austria (Ille and Grinschgl 2001). The number of such parcels per
25 ha had a positive impact on habitat suitability in Deux-Sèvres, western France
(Bretagnolle et al. 2001). Spatio-temporal rotation in the management of landscape
patterns can offer optimal habitats year-round, as the species can very quickly react to
changing environments by shifting its hunting ranges, even beyond its territories
(Finck 1990).
The species displays a considerable plasticity in its habitat requirements. Plasticity
here refers to the capability of the Little Owl populations to remain viable, given the
relatively wide range of open and semi-open habitats that it occupies across its 9000
km-wide range.
Some research stresses heterogeneity in the habitats that the owl occupies (Van
Nieuwenhuyse and Bekaert 2001b). A multitude of habitat typologies for the Little
Owl in Europe (Table 6.1) clearly confirms the plasticity of the species in most of the
European part of its range. Research results also confirm that the small size of the scale is
more important than the actual land use. All typologies feature the essential criteria for
suitable Little Owl habitat which is met within a wide diversity of natural and anthro-
pogenic landscapes and ecosystems (Génot and Van Nieuwenhuyse 2002).
This heterogeneity illustrates that certain landscape parameters are interchangable.
Splitting the habitats into discrete groups or habitat types allows us identify and
quantify the number of different habitat typologies and furthermore it allows us to
select representative sampling points for surveillance of the Little Owl in its environ-
ment. In terms of sustainability of inventory and monitoring actions, most of the
resources should go to assessing the main habitat types. Monitoring these types allows
the tracking of conditions in an objective, representative and optimal way since both
optimal and suboptimal habitats are included and distinguished.

6.3 Habitat Preferences

Essential criteria for suitable Little Owl habitat are year-round prey availability, prey
accessibility, vertical landscape structures with cavities and limited predation pressure.
An optimal mosaic of short vegetation and commanding perches for prey observation,
tall vegetation for prey reproduction and availability of secure cavities for roosting and
nesting is preferred (Génot and Van Nieuwenhuyse 2002).
Habitat preference can be studied through a multitude of statistical methods.
Classification methods use information on occupied and unoccupied habitats in their
tests. This way the methods look for parameters that best explain the difference
between both types of habitats. Regression methods predict the number of pairs or
territories as a continuous outcome rather than a probability of occupation or not.
Telemetry data use only occupied habitats, but analyses the differences between the
used parts of the habitat and those that are not used, to distinguish positive and
negative impacts of the habitat features.

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Table 6.1 Overview of clearly distinguished types of occupied Little Owl habitats in Western Europe

Author Region Types of habitat

Van Nieuwenhuyse and Flanders (North Belgium) Semi-open grassland

Leysen (2001) Grassland around farms
Cattle-breeding landscape without farm buildings
Horticultural landscape
Rural areas with cattle breeding
Cereals and orchard far away from farms
Cattle-breeding landscape in the vicinity of farm buildings
Van Nieuwenhuyse and West-Flanders (Belgium) Humid pasture areas with many pollard willows
Nollet (1991) Large isolated meadows with very few trees
Fields at slightly higher altitude with many farm buildings
Juillard et al. (1992) Causse Méjean (Lozère, France) Pasturelands grazed by sheep with typical mounds of stones
“clapas”
Hay meadows and cereal fields with “clapas”
Abandoned farms with ruined buildings
Active farms
Disused quarries
Linear habitats of coppiced trees (Fraxinus excelsior)
Mastrorilli (2001) Bergamo (northern Italy) Woods and zones with a particularly closed tree cover
Urban area including villages and industrial areas
Residential area with parks
Gardens at the perimeters of the urban areas
Arable lands with or without irrigation and cultivated or
uncultivated grasslands
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246

Table 6.1 (cont.)

Author Region Types of habitat

Centili (1996) Tolfa Mountain area (Italy) Open meadows

Partially wooded
Covered meadows and cultivated (grass)land
Barthelemy and Bertrand Garlaban hills (15 km from Marseilles city, Suburban area with houses and gardens
(1997) southern France) Undisturbed hill area with low shrubs and rocky patches
Fajardo et al. (1998) Southern Spain Marshland
Grassland
Sunflower fields
Olive tree plantations
Orchards
Urban areas
Eucalyptus plantations
Loske (1986) Germany Orchard meadow
Grassland with pollard trees
Borders around villages
Grassland with solitary trees
Isolated farms
Bridges
Churchyards
“Lössgrube”
Ferrus et al. (2002) Northern Vosges (northeast France) Urban-type
Mixed landscape
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Scarpe-Escaut Plain (north France) Pastures (20–40% of the area) and arable land in about the
same proportions
Arable land, pastures and woodlands are intermingled
Arable land and pastures and built-up areas representing the
built-up and related areas
Causse Méjean Arable land and pastures in equal proportions (40–60% of
the area)
Large proportions of pastures (60–80%) and a clearly lower
proportion of arable land (up to 20%)
Ille (1996) Austria 46% arable land and 17% grassland in Marchfeld
34% arable land, 35% grassland and 11% vineyards northern
Burgenland
Blache (2004) Plaine de Valence; Drôme, France Mixture of fields, farms and isolated houses, hedges, isolated
trees and roads
Mixture of fields and roads
Small-scale mixture of fields, farms and isolated houses, hedges,
isolated trees, roads, tree nurseries and urban zones
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6.3.1 Classification Methods

Recent studies have clarified the preference of the Little Owl for specific habitat param-
eters such as areas of land use (e.g., grassland, fields, forest, built-up areas, orchards,
surfaces with short/tall vegetation), lengths of linear elements (e.g., roads, tree-lines,
fences, field and meadow edges) and absolute number of specific habitat elements (e.g.,
individual trees, number of cavities, number of perches, number of abandoned build-
ings) (e.g., Van Nieuwenhuyse and Bekaert 2001b, Génot and Van Nieuwenhuyse
2002). An array of statistical techniques are used to detect actual habitat preferences
from the range of conditions that are available in landscapes. All these techniques try
to distinguish occupied from unoccupied landscapes and hence they need to be com-
pared. For simplicity, cartographic representations of landscapes are cut into square grid
cells. The landscape within each grid cell is then quantified and used for analysis (Ferrus
et al. 2002, Blache 2004). The parameter that will be explained is Little Owl presence
using observed associations between the landscape and the owls’ occurrence.

6.3.2 Regression Methods

Onkelinx and Everaert (2018) modeled the Little Owl distribution and population
numbers in a 3500 ha area in 200  200 m grid cells in Gooik, Belgium, to aid the
planning process for land re-allocation and optimization. The aim of the model was to
explain and quantify the observed landscape factors that have an impact on the Little
Owl. Next, the model was used to simulate possible implications of changes in the
landscape for the species. In this way different scenarios could be assessed upfront.
Playback (Van Nieuwenhuyse et al. 2001e) was used to assess the local owl popula-
tion. The landscape was quantified as area of forest, high-stem and low-stem orchard,
arable fields, fallow land, grassland, hedges, low vegetation, tall vegetation, total parcel
perimeters and urban. Strongly correlated variables were removed. The model high-
lighted an increase in observed owls with increasing area of fallow land, tall vegetation,
grasslands, hedges, high-stem orchard or arable field edges. Negative impacts were low-
stem orchards, forest and urban areas.
Different scenarios of possible land re-allocation plans were scored using the model
and the possible impact on the Little Owl was estimated. An ecological status quo
scenario (nature values should be equal or better) as a compromise between nature
and landscape performed well and even led to small population increases. This plan
would secure and improve the most valuable nature areas in the long term and develop
a connected ecological network in the whole area. Pollard willow rows, hedges and
high-stem orchards would be secured and further restored or extended. The high-stem
orchard extension of the plan, aiming at a full restoration of traditional fruit orchards
for the nearby Brussels market did not perform well due to over-estimations of the
impact of the current orchards in the model. Territoriality which was not taken care of
in the model started to play a role in this scenario. The purely agricultural optimizing
scenario, where agricultural parcels were taken together and arable lands sometimes
exchanged for grasslands and vice versa also performed well. This scenario amplified

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 6.3 Habitat Preferences

traditional pastoral land use and induced a partial move of the owl population from
suboptimal arable-dominated locations towards high-quality grassland and high-stem
orchard areas.

6.3.3 Telemetry
Apolloni et al. (2018) studied habitat selection and range use of adult owls in relation
to landscape configuration, habitat structure and resource distribution via telemetry
in orchard-rich Baden-Württemberg, Southern Germany, in a population of about
220 breeding pairs on 250 km². Of all telemetry fixes, 47.8% were located in orchards,
39.2% in cropland, 9.2% in field margins, 10.2% in grassland and 3.5% in other habitat
types. The proportion of major habitat elements over the whole study area comprised,
on average, 33.4% cropland, 19.1% forest, 17.1% human settlements, 9.8% orchards,
9.7% open grassland, 7.1% vineyards, 2.8% woods/shrubs and 1% other habitats.
Comparision of proportions of available habitat elements versus those used shows
significant differences. Orchards were by far the most preferred habitat element in the
home ranges, with significantly higher values. The same held for habitat patch use,
with field margins the second preferred habitat, followed by grassland and cropland.
Roads and wooded areas were strongly under-proportionally used.

6.3.4 Cross-Border Comparison

Tschumi et al. (2020) compared land cover, land-use intensity and resource availability
between plots of highest habitat suitability for Little Owls among two neighboring, but
politically separated areas in southwestern Germany versus northern Switzerland. The
land cover and land-use richness did not differ between German and Swiss plots, yet
there were marked differences in terms of land-use intensity and the availability of
resources. Land-use intensity was significantly higher and resource availability lower in
Swiss compared to Germand plots. While accounting well for remotely sensed data such
as land cover, habitat suitability models may fail to predict land-use intensity and
resources across borders. The relationship between geodata used as proxies and eco-
logically relevant resources may differ according to history, policies and socio-cultural
context, constraining the viability of habitat selection models across political borders.
Tschumi et al. (2020) stress the need for fine-scale resource assessments comple-
menting landscape-scale suitability models. Conservation measures need to consider
the availability of crucial resources and their socio-economic moderators to be effective.
The following habitat parameters were recorded for each sampling plot: permanent
meadow area, land-use richness (number of different land-cover types), low-intensity
meadows (occurrence of permanent meadows of low management intensity, after
Jenny et al. 2011), grazing (occurrence of grazed areas), different cutting regimes (occur-
rence of different regimes), small structural elements (sum of occurrences of dry stone
walls, stone piles, stacks of wood, brush piles, hedges, summer houses, unmaintained
buidings and equipment shelter buildings), tree cavities (total number of tree cavities
>6 cm diameter and >20 cm depth), roosting sites (occurrence of potential roosting sites

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 Chapter 6: Habitat

in buildings, pallet stacks, excluding nestboxes, tree crowns and tree cavities), small
rodent index (total predicted small rodent traces per ha after Apolloni et al. 2018), number
of trees (all trees standing freely, in groups or orchards, excluding trees in hedges),
sampling date and cavity occurrence in individual trees (cavities >6 cm diameter and
>20 cm depth after Bock et al. 2013).
The discrepancy between habitat suitability model predictions and effective resource
availability at a fine scale may explain the absence of Little Owls from areas with high
suitability scores in Switzerland. The Swiss plots had a reduced occurrence of small
structural elements and a decreased structural diversity compared to the German plots.
Grazing intensity in the Swiss plots was high, offering low plant diversity and reduced
accessibility due to dense and monotonous swards. The likelihood of cavities decreased
after reaching a maximum at 53.9 cm (German plots) and 63.9 cm (Swiss plots)
diameter at breast height (dbh), suggesting selective removal of large damaged trees.
Lower availability of roosting sites and a lower small rodent index in Swiss plots
compared to German further explains the difference in ultimate habitat quality.
Animal numbers per area are lower in southwestern Germany than Switzerland (45
compared to 65 cows, sheep and goats per km²) resulting in higher intensification
pressure in Switzerland. Differences in property rights result in orchards or allotments
being owned more often by nonfarmers, resulting in lower constraints for cost-effective
management in southwestern Germany. Major socio-cultural drivers contribute to the
differences between Germany and Switzerland, symbolized by Swiss farmers’ unwilling-
ness to invest in conservation, often fed by their concerns about being considered
unproductive and reduced tolerance towards nonproductive land uses.

6.4 Habitat Suitability Factors

Habitats will typically include open hunting ground, rich in small prey, hunting
perches, day roosts and nest holes, and with a benign climate and land management
regimes that give reasonable long-term continuity.

6.4.1 Prey Accessibility Through Short Vegetation

A Little Owl habitat needs to have sufficient food for both territory owners and their
offspring. As Little Owls eat live prey (or sometimes recently dead prey), these prey
animals also have to be able to live in a sustainable way, maintaining viable populations
over time. Besides having prey at their disposal, Little Owls also need to be able to access
and catch the prey. As nearly all of the Little Owls’ prey are ground-dwelling species, the
owl needs to catch its prey in short vegetation. To catch prey in an energy-efficient
manner, the owl uses the “perch-and-pounce” hunting strategy. It also uses the terres-
trial “on foot” hunting method. While hunting for rodents and insects, the birds tend
to be perched on a relatively low perch (1–3 m) to obtain a commanding view over their
immediate surroundings. Once they discover prey, they pounce on it from the perch.
Being posted above the ground allows the owls to discover prey visually. Kestrels
(Village 1990) and Barn Owls (Shawyer 1998) are able to hunt more in flight rather

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than being perched and have less problems catching prey in tall grass. Due to its limited
ability to fly above tall grassland, its nocturnal lifestyle, and its more limited hearing
capabilities, the Little Owl is more dependent on its sight for hunting.
In Western Europe, earthworms make up an important part of the owls’ diet,
allowing it to hunt successfully on bare ground or in short grasslands. Ideal hunting
grounds for the species are to be found at the borders of parcels. These borders function
as ecotones with a much higher prey diversity than in the center of parcels. Ideal
habitats are small-scale pastures with year-round grazing and very small parcel sizes
(e.g., <1 ha), which have a lot of parcel border for a given area. These small-scale
heterogeneous conditions combine prey presence with prey accessibility. Larger grass-
lands with tall vegetation favors voles and hence is more preferred by kestrels, harriers,
Barn Owls and Short-eared Owls.
Prey accessibility for the species is maximized by short vegetation with commanding
perches that allow either “perch and pounce” or “walking” hunting techniques. The
owl hunts mainly on the ground for earthworms or insects and prefers short vegetation
or bare ground for this. Short vegetation can be found in grazed grasslands of all kinds,
preferably with an average plot size <0.6 ha (villages with the highest population
densities had an average plot size of 0.55 ha; Dalbeck et al. 1999a,b). Preference for
pastures over other types of grasslands, especially hay fields, shows the importance of
year-round prey accessibility (Dalbeck et al. 1999a,b). During the summer months,
utilization of arable land by owls in Germany is disproportionate to its availability. In
the summer, territories are reduced in size, with the owls concentrating on allotments
that support optimal accessibility of food (Finck 1990, Orf 2001). As the habitat quality
increases, active home ranges tend to decrease in size, with the owls using grassland
with short vegetation within a radius of 400 m around the nest (Orf 2001). Depending
on the relative areas of fields and grasslands, both are sustainable habitats as long there
is heterogeneity in time and space. Sometimes even gardens provide suitable foraging
habitat (Orf 2001). During some periods of the year, road edges provide a large propor-
tion of short or absent vegetation, making hunting owls very vulnerable to vehicle
collisions (Hernandez 1988, Clec’h 2001). In Germany, newly created vineyards
(Weinberg area) have been extremely attractive to Little Owls in the first years after
creation. The vineyards were characterized by bare ground and poles shorter than 2 m
placed at regular intervals (Eick 2003). Vegetation cover in foraging grounds of southern
Spain generally does not exceed 2 cm in height; use of areas with cover higher than this
was not observed. The most favored hunting grounds consisted of a heterogeneous
mosaic of bare ground and short grass patches (Fajardo et al. 1998).

6.4.1.1 VEGETATION HEIGHT

Recent research has shown that owls generally select for areas that are associated with
high prey densities and allow for successful foraging, i.e., comparatively short-
vegetation pastures.
Telemetry analysis uses observed versus expected use of landscape elements in
relation to their availability in the landscape (Finck 1990, Génot and Wilhelm 1993,
Orf 2001, Grzywaczewski 2009). If there were no real preference of the species for a

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certain vegetation height, then all vegetation heights would be used as proportionally
available. A special liking of the species for specific heights of perches can be studied by
calculating how many perches are available per height class, compared to how many of
these are used by the owls. Grzywaczewski (2009) measured the vegetation height of an
occupied Little Owl territory situated near Lublin. Counting the available grid cells per
vegetation height yielded the expected use by the owl if no preferential vegetation
height existed. Counting the observed use of the grid cells by the owls yielded the
observed use.
Šálek and Lövy (2012) studied the spatial ecology and habitat selection in anthropo-
genic landscapes in Central Europe. The most important foraging habitat during the
entire breeding season was grassland (especially pastures), reaching 90% for all loca-
tions. Vegetation height and cover were the main factors determining habitat selection;
Little Owls significantly preferred sparse and short-sward vegetation patches that
enabled hunting of ground-dwelling prey. Utilization and availability of different
habitat types in home ranges changed markedly during the breeding season. In April,
Little Owls predominantly hunted in gardens, whereas from May to July they preferred
short-sward pastures or lawns. Total utilization of grassland habitats (pastures/lawns,
meadows and gardens) was highest during the nesting and fledging periods (June) and
reached up to 90% of all locations, while later in the fledging period (August) these
areas were represented less and use decreased to 51%.
Mean vegetation height and vegetation cover were both found to influence the
selection of foraging habitats of Little Owls. Both vegetation characteristics showed a
significant relationship with hunting preferences. Average vegetation height of foraging
sites during the breeding season was 4.7 cm (SD = 5.9, n = 667) and total vegetation cover
was 51.0% (SD = 34.6, n = 667), in contrast with 31.6 cm and 82.8% respectively in
control locations. This pattern held consistently during the whole breeding season. Little
Owls predominantly hunted from elevated perches (87%), such as trees, poles and
buildings that were on average 4.5 m above ground. In August they shifted their foraging
activity to cereal fields after stubble ploughing and hunted on the ground (13%).
Mayer et al. (2021) conducted a case study on the Little Owl to investigate how high-
frequency GPS data can be used to investigate spatio-temporal movement patterns and
habitat selection to inform conservation practices. They equipped six Danish food-
supplemented little owls and six nonsupplemented owls in the Czech Republic with
high-resolution GPS loggers that recorded one position per minute. A new approach to
avoid highly spatio-temporally auto-correlated data is to simulate a random movement
trajectory with the same auto-correlation structure as the observed animal movement
trajectory, using correlated random walks (Quaglietta and Porto 2019). Mayer et al.
(2021) compared nightly Little Owl space use, spatio-temporal movement patterns and
habitat selection in surviving territories in Denmark with a local population in the
Czech Republic that was also declining, but not food-supplemented. Both were located
in intensively cultivated farmland, but differed in the composition of crop types and
regarding supplementary feeding. Mean field sizes were comparable in the Czech
Republic and Denmark (4.8  12.6 ha versus 4.9  5.9 ha; unpaired t-test: t = –0.02,
df = 162, p = 0.98). Similarly there was no statistical difference regarding habitat

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diversity between Czech and Danish owl territories (Simpson’s Index of Diversity: 0.68 
0.03 versus 0.62  0.11, Mann–Whitney test: W = 8, p = 0.63). Available land-cover types
based on buffers around the nest location changed with distance from the nest, with the
Czech area dominated by maize, cereals and built-up areas, and in Denmark by cereal,
pastures and rape. Habitat use also changed with distance from the nest, with built-up
areas being mostly used in close nest proximity, whereas cereal, pastures and road verges
were used with increasing distance from the nest. In general, built-up areas were by far
the most used land-cover type, making up 79% of all GPS positions in the Czech
Republic, followed by fallow (5%), maize (4%), rape (3%), road verges (2%), pasture
(2%), cereal (2%) and other land cover (3%). In Denmark, built-up areas were also the
most used land-cover type (53%), followed by pasture (12%), cereal (12%), forest (11%),
maize (6%), road verges (3%) and other land cover (3%).
Comparing used positions to available positions obtained from correlated random
walks, all owls (of both sexes and study areas) strongly selected for built-up areas,
avoided cereal and showed no selection or avoidance for pastures and road verges. For
the other land-cover types, habitat selection partly differed between females and males,
and between the two study areas. In both areas, rape was avoided by males, but selected
by females. Maize and forest were selected by Danish males and avoided by Czech
males, and other arable crops were neither selected nor avoided by males of both
populations. Fallow, maize, and other arable crops were either avoided by females (in
both populations) or not available in proximity to the nest. Forest was neither selected
for nor avoided by Danish females and not available to Czech females. Danish males
selected for proximity to roads, whereas Danish females avoided proximity to roads,
and all owls, apart from Czech females, selected for proximity to field edges. Finally,
male owls in Denmark increasingly selected for maize with increasing distance from the
nest at the expense of the other land-cover categories, whereas there was no clear
pattern for males in the Czech Republic.
Habitat use changed with distance from the nest, with built-up areas being mostly
used in close nest proximity, whereas cereal, pastures and road verges were used with
increasing distance from the nest. The furthest distance category in Denmark was
500–2000 m from the nest compared to 500–1000 m for Czech owls, because Danish
owls moved further from the nest.
In general, little owls in both populations used foraging areas with either compara-
tively short vegetation (e.g., lawns, gardens and orchards in built-up areas) that enabled
hunting of ground-dwelling prey (Šálek and Lövy 2012; Grzywaczewski 2009) or areas
with high structural diversity (close to field edges or fallow areas in the case of Czech
males) that potentially support higher prey abundance (Apolloni et al. 2018). The
proportion of ruderal vegetation around farmsteads was higher in the Czech Republic
than Denmark, potentially providing better rodent habitat (Šálek et al. 2018). Road
surfaces also provide opportunities for owls to localize and capture large insects like
carabid beetles in dry weather, and earthworms and amphibians in wet weather, as well
as invertebrate and small mammal road-kill carrion. Favorable foraging conditions close
to roads may, however, increase collision risk with vehicles, which is an important
additive source of Little Owl mortality (Šálek et al. 2019).

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6.4.2 Prey Accessibility Through Commanding Perches

Tomé et al. (2011) compared the diurnal hunting behavior of Little Owls in two structurally
different habitats: a treeless pseudo-steppe and a Holm Oak Quercus ilex rotundifolia wood-
land in Portugal. While the overall prey availability was similar, perch and vegetation
characteristics differed between habitats. Using focal sampling in 29 territories, the type
and height of utilized perches, time spent on perches, and attack and giving-up distances
involved in insect hunting were determined. Moreover, they analyzed the inter-
relationships between behavior and habitat-specific vegetation characteristics. Little Owls
adapted their hunting strategies to habitat features, achieving similar prey detection rates
and the same hunting success in the two habitats. In the pseudo-steppe, where only low
stone piles were available as perches, owls selected the highest spots (average 0.77 m). In this
area, a positive correlation was found between perch height and distance to prey attacked,
indicating that perching higher resulted in increased prey detectability. In the woodland,
owls selected perches that were lower than maximally available (averages 2.67 m versus 4.99
m, respectively). Attack distance did not differ between habitats, despite the use of higher
perches in the woodland. These results indicate that the more developed ground vegetation
in the woodland possibly hampered detection of prey, counterbalancing the possible
advantages for prey detection due to the use of higher perches. In such situations, Little
Owls seemed to select an intermediate perching height to maximize prey detectability.
Apolloni et al. (2018) studied perch use in relation to vegetation height using
artificial perches on 1.5 m poles with horizontal branches of 25 cm placed in parcels
of at least 50  50 m. Perches were placed at least 50 m away from natural perches and
>10 m away from field borders to avoid edge effects. Thirty percent of fixes were
recorded within high grass vegetation (>20 cm), independent of habitat type and
70.0% within low grass vegetation (<20 cm); 29.4% of the fixes were recorded in
grassland areas and 70.6% in cropland. Little Owls preferentially visited plots with
low vegetation, irrespective of habitat type. The visits to perches decreased with increas-
ing vegetation height. Surprisingly, the number of visits was higher in cropland than in
grassland and decreased with distance to the nest. During the nestling period and late
summer the perches were more frequently visited than during the fledgling period. The
explained variance was low, indicating that other factors that might have an effect on
the frequency of visits to perches were not included in the model.

6.4.3 Prey Availability

Insect prey density and diversity is related to the population density of owls in Austria,
and the density of small mammals showed a positive association with average repro-
ductive success (Ille and Grinschgl 2001). Grimm (1986) found more and larger prey in
a meadow edge beside a brook than in grasslands. The impact of cattle trampling
yielded more Carabid ground beetles. Tall vegetation is crucial for small mammals, with
edges tending to have greater diversities and densities (Dalbeck et al. 1999a,b).
Earthworm densities are always higher in grassland than fields because plowing leads
to higher oxidation rates and the associated reduction of organic material, and hence

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 6.4 Habitat Suitability Factors

less food for earthworms due to reduced humus. Higher use of pesticides, predation and
mortality are typically associated with plowed fields. Densities in grassland depend on
the fertility of the soil (a balanced nutrient influx yields higher densities than unfertil-
ized grasslands), and higher densities are found with organic fertilizer than inorganic
due to direct intake of the first by earthworms (Edwards and Bohlen 1996).
Actions to conserve vole numbers has no effect on either productivity or population
regulation of Little Owls in the lower Rhine area, Germany (Exo 1987), perhaps because
the species predominantly feeds its young on earthworms (Lumbricidae) and beetles
(Coleoptera) (Juillard 1984) or due to the fact that voles found there are in lower
densities due to the regular flooding of the Rhine (M. Exo personal communication).
Research by Génot (2005) in the northern Vosges of France did not reveal any relation-
ship between vole numbers and the Little Owl. Zwölfer et al. (1981) showed that hedges
and other vegetation gradients have a significant positive impact on prey species in
terms of both density and diversity.
Permanent tall vegetation is needed as the breeding grounds for owl prey (and thus
indirectly for prey availability) but reduces the accessibility to the owl. Hence a com-
bination of short and tall vegetation (with accessibility from commanding perches)
yields the highest diversity of prey numbers and species (Van Nieuwenhuyse et al.
1999). Edges have a special importance for the owl. In this context edges reflect hedges,
walls and a high proportion of edge structures relative to the area (Dalbeck et al. 1999a,b),
meadow edges, orchard edges (Van Nieuwenhuyse and Bekaert 2002), field edges (Van
Nieuwenhuyse and Bekaert 2001), tree lines and field edges (Van Nieuwenhuyse et al.
2001c), roads (Hernandez 1988), pasture fences (Loske 1986), hedged open fields
(Zuberogoitia and Campos 1997), length of hedges in the Netherlands (Visser 1977)
and vertical structuring of grasslands (Ille and Grinschgl 2001). This is probably due to
a mix of different parameters – a high diversity of prey breeds in the tall vegetation at
high densities, prey becomes visible and reachable in the shorter vegetation nearby, and
can be caught easily from fence poles along the edges of two adjacent parcels.
Apolloni et al. (2018) studied prey availability in relation to vegetation height. The
abundance index of voles in typical vole habitat was significantly and strongly related
to season and vegetation height. During the year, the abundance index of voles showed
a peak in March and Mid-October. There was a significant positive correlation between
vegetation height and vole abundance. The abundance index in orchards was highest
(5.93  7.38) and showed the strongest increase in relation to vegetation height. In
comparison, grassland (4.48  7.6) and field edges (1.72  8.70) has smaller abundance
indexes for voles.

6.4.4 Snow
Most Little Owls have difficulties surviving more than three weeks of snow cover
(Helbig 1981, Schönn 1986). In the former Soviet Union, Gusev (1952) showed a sharp
correlation between snow cover depth and hunting success. With a snow depth of
0.5–3 cm, the hunting success of the Little Owl for rodents was higher than in snowless
periods. With shallow snow cover, rodents cannot dig through it and have to move to

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the surface, where they become clearly detectable on the white background and there-
fore are easily caught. Friable snow 7–9 cm deep sharply decreases the hunting success
of Little Owls because small rodents dig tunnels in the snow and seldom emerge on the
surface. Snow cover from 12–70 cm further reduces hunting success. Despite deeper
snow, the Little Owl may hunt more successfully, e.g., when melting water freezes
again, inhibiting rodents from making tunnels in the snow and forcing them to
the surface.

6.4.5 Weather and Season

The degree to which animals use habitats in relation to their relative availability is a
widely used index to assess the apparent importance of alternative habitats. However,
habitat selection is a dynamic process that is likely to be influenced by a variety of
temporally variable factors such as seasons or weather. As a possible temporally variable
external driver on habitat selection by foraging owls, weather is known to influence
spatial behavior and foraging decisions and affect diet composition and prey-specific
predation rates of Little Owls (Van Harxen 2021). By failing to quantify the influence of
temporal drivers on habitat selection, one may therefore risk missing information about
the ecological and behavioral basis for observed species–habitat relationships. Despite
the dynamic nature of habitat selection, temporal variation arising from factors such as
weather are rarely quantified in species–habitat relationships. Sunde et al. (2014) ana-
lyzed habitat use and selection (use/availability) of foraging in 27 radio-tagged Little
Owls to see how this varied as a function of weather, season and availability.
On an annual basis, foraging little owls spent 54% of their time in cultivated fields
(95% CI: 47–62), as compared to a mean availability of 76% in the home ranges. These
mean figures covered a considerable seasonal variation with a peak from September to
April and a low in June. Gardens/buildings was the second most frequently used land-
cover type with an annual mean of 25% (95% CI: 19–32) as compared to 2.9% mean
coverage. Garden/building use peaked in May–July, and dipped in September–April.
Pastures represented 7% (95% CI: 4–12) of all use (minor seasonal variation) as opposed
to 8% mean availability. Use as well as availability of gardens/buildings decreased and
of cultivated fields increased with increasing distance from the nest. For the 20 owls with
access to “moist” surfaces in their home ranges, mean use was 4% (95% CI: 2–11) as
opposed to 6% available. Little Owls selected general land-cover types (p <0.0001), as well
as ground moisture categories (p = 0.008) differently according to month, but did not
select areas in relation to tree vegetation in general (p = 0.71) nor with month (p = 0.64).
Temporal predictors, including weather, explained significant amounts of the vari-
ation in use of all four focal land-cover categories – but the amount of variation, as well
as the specific, influencing variables differed. Overall, weather and seasons explained
the variation in the use most for cultivated fields and gardens/buildings and least for
pastures and “moist” ground. Life history variables (sex, breeding phase) explained
negligible variation after seasonal effects had been accounted for.
After having accounted for monthly variation and habitat composition at home-
range level, weather influenced the use of cultivated fields and gardens/buildings

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 6.4 Habitat Suitability Factors

through linear effects of wind and quadratic effects of temperature. Cultivated fields
were used more (p = 0.0005) and gardens/buildings less (p = 0.0003) with increasing
wind speeds. In relation to temperature, use of cultivated fields peaked (adjusted for
month: p = 0.026), and use of gardens/buildings dipped (p = 0.0047) at intermediate
temperatures of 3–9 C. In a seasonal context, use of cultivated fields correlated posi-
tively with temperature in winter and negatively with temperature in summer, whereas
the opposite was the case for gardens/buildings. Use of pastures and “moist areas”
increased with increasing temperature (p = 0.0097 and p = 0.0003, respectively).
Sunde et al. (2014) found that the use of the two most frequently used land-cover
types, gardens/buildings and cultivated fields, varied more than threefold, as a simple
function of season and weather through linear effects of wind and quadratic effects of
temperature. Even when controlling for the temporal context, both land-cover types
were used more evenly than predicted from variation in availability (functional
response in habitat selection). The use of two other land-cover categories (pastures
and moist areas) increased linearly with temperature and was proportional to their
availability. The study shows that habitat selection by generalist foragers may be highly
dependent on temporal variables such as weather, probably because such foragers
switch between weather-dependent feeding opportunities offered by different land-
cover types. An opportunistic foraging strategy in a landscape with erratically appearing
feeding opportunities in different land-cover types may possibly also explain decreasing
selection of the two most frequently used land-cover types with increasing availability.
For opportunistic foragers like Little Owls experiencing temporally variable foraging
quality of different habitats, functional responses in selection for individual habitat
types is perhaps exactly what we should expect as a rule rather than as an exception. As
a consequence, the most favored habitat composition for generalist foragers may be a
mixture of a range of alternative habitat types that varies in profitability rather than
maximum availability of a single ‘‘best’’ one.

6.4.6 Perches
The availability of hunting perches has been reported as varying in importance. In
central Europe, the species seems to be mainly a ground forager (Exo 1991, Schönn et al.
1991) probably because of the relative importance of earthworms in their diet (Exo
1991). Mud-covered entrances to nestboxes in West-Flanders during the nestling period
suggest the importance of walking as a hunting technique. Some German studies show
no correlation between the presence of fence poles and the species (Dalbeck et al.
1999a,b), while other studies show a clear positive relationship (Loske 1986). This
difference might indicate that other landscape structures, e.g., trees or shrubs, are
interchangeable for fence poles to serve as perches. In Vosges du Nord (northeastern
France) males spend less time on perches at the end of spring and beginning of summer
(Génot and Wilhelm 1993). Up to 82% of perch use is on trees, 14% on poles and 4% on
hedges year-round. Fruit trees offer shelter in addition to perches. One radio-tracked
female used fence poles significantly more during the nestling than the courtship
(prebreeding) period. Modern low-stem orchards with integrated fruit growing (reduced

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 Chapter 6: Habitat

pesticide use) in Flanders (Génot and Van Nieuwenhuyse 2002) and Germany (Orf
2001) and vineyards in Austria (Ille and Grinschgl 2001) and Germany (Eick 2003) offer
good hunting grounds due to the short vegetation and the vine stocks or poles as
commanding perches. These areas also serve as breeding habitats as long as nest cavities
are available.
In Spain, the Little Owl forages mostly from perches (Fajardo et al. 1998), probably
because of the limited importance of earthworms in the diet. Anti-predator strategies
might lead owls to avoid perching on the ground, where they are more vulnerable to
predators. Perching sites above 3 m are seldomly used in southern Spain. No change in
height of perches was observed during the summer months, when road mortality is at
its maximum.
R. Tomé (personal communication) studied the diurnal foraging behavior of Little
Owls in two Mediterranean habitats (a pseudo-steppe and a Holm-oak woodland) with
different perch characteristics in Portugal. During 1999, 19 observation periods involv-
ing Little Owls hunting insects were recorded for each area. He investigated which type
of signals Little Owls used for detecting prey, with the birds using both visual and
acoustic cues while hunting. The use of visual hunting was indicated by the selection of
the highest possible perches in the pseudo-steppe, by the positive relationship between
perch height or vegetation coverage and giving-up times in the woodland area, and by
the positive association between perch height and attack distance in both sites. The use
of acoustic hunting was indicated by similar giving-up times in both areas. In spite of
major inter-areal differences in perch height, owls apparently were able to optimize
their foraging behavior in both habitats, maintaining similar detection times and
hunting success. Hunting success was probably affected by prey-specific characteristics.
In the district of Modena (Italy), a study of 41 song perches showed that 46.3% were
rural buildings, and the number of frequently used perches was positively correlated
with the extension of orchards (Selmi et al. 2005).

6.4.7 Cavities
Cavities are crucial for the species, as Little Owls use cavities or crevices to breed in and
to roost in during the day. Relatively small holes (e.g., 6–7 cm diameter) to the nest
cavities are preferred. Since the owls do not build their own nests, they are dependent
on natural or artificial cavities. Competition for cavities, with other bird and mammal
species, forces the owls to take cavities with small entrance holes. The size of the
entrance to the cavity is a balance between ease of passage by the owls and limiting
access to predators.
Positive correlations have been found between Little Owl densities and the age of
trees (rather than the number of trees) in Düren (Northrhine-Westphalia) (Dalbeck
et al. 1999a,b), with the number of pollard trees in Northrhine-Westphalia (Loske
1986), and with potential breeding cavities and grasslands (Exo 1983). The Little Owl
adapts to nestboxes easily, but this had no influence on population density in
Northrhine-Westphalia (Dalbeck et al. 1999a,b). The absence of impact might have
been related to the availability of natural cavities. No correlation was found between

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 6.4 Habitat Suitability Factors

the number of nest sites and density in the Netherlands (Fuchs 1986). In Germany,
juvenile Little Owls utilize the crevices in piles of fire wood regularly for daytime
roosting (Eick 2003).
The addition of nestboxes can have a profound effect on Little Owl populations
(Stange 1999, Bultot et al. 2001, Génot 2005). The provision of nestboxes can initially
appear to increase the local population, as birds without any nest sites readily occupy
the boxes (Stange 1999, Bultot et al. 2001). Over a period of several years, it appears that
the resident owls switch from using natural cavities to nestboxes. Further, the social
dynamics of the species (i.e., the apparent desire of the species to form population
clusters) occurs; if the availability of natural nest sites is low, owls from outside the local
study area will abandon their territories and shift towards the nestbox-rich area. These
types of movements were observed by owls in an area of circa 200 km² in northern
France (Génot 2006) over a 20-year period.

6.4.8 Roosts
Roost-site selection can be influenced by predation risk (Tillmann 2009), parasite
occurrence (Rohner et al. 2000), disturbance probability (Peters and Otis 2007) and
thermal properties (Kendeigh 1961). Predator avoidance during roosting should result
in preferences for sites inaccessible to most predators, offering escape routes or enabling
detection of approaching predators. Sheltered or enclosed roost sites may be invisible
and inaccessible to large predators (Mainwaring 2011). Open roost sites may allow more
effective predator detection and escape response, but may be more costly if predation
risk is increased and the energetic costs of escape behavior are high. The benefits of
different sites are likely to be species specific and dependent on the local predator
community. The availability of adequate roost sites may be an important aspect of
habitat quality, and understanding roost-site selection as a component of habitat use
throughout the year and across life-history stages provides a crucial link between
behavioral ecology and conservation biology.
Bock et al. (2013) studied roost-site selection throughout the year, providing infor-
mation about year-round habitat requirements at different life-history stages of Little
Owls by repeated daytime location of 24 adult and 75 juvenile radio-tagged individuals
from July to November. Little Owls preferred sheltered roost sites such as tree cavities
with multiple entrances. They increasingly used sheltered sites from summer to winter
and preferentially used sheltered roost sites with low ambient temperatures. Juveniles
used significantly less sheltered sites during dispersal than before and afterwards, and
used less sheltered sites than adults within their home range. The survival probability of
birds roosting frequently at exposed sites was reduced. Roost-site selection is probably
driven by the two mechanisms of predator avoidance and thermoregulation, and the
costs of natal dispersal may include increased predation threat and higher energy
expenditure for thermoregulation. We suggest that adequate roost sites, such as multi-
entrance tree cavities, are an important habitat requirement for Little Owls and that
habitat quality can be affected by manipulating their availability.

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A total of 1223 Little Owl locations were recorded by Bock et al. (2013) between mid-
July and the end of November 2010, including 318 roost sites of adults and 905 roost
sites of juveniles. Little Owls used open (42%) as well as sheltered sites (58%) for
roosting. Roost sites were in or on trees in 84% of cases and apple trees were the most
frequently used tree species (apple: 45%, pear: 18%, cherry: 2%, other tree species:
15%). Nearly one-third of roosting sites (29%) were in fruit tree cavities. Nestboxes
(17%), stacks of wood (10%) and small buildings (2.5%) were also used. During the
period of postfledging dependence, Little Owls roosted most often within fruit tree
crowns, in tree cavities with multiple entrances, in nestboxes and in wood stacks. When
roosting in sheltered sites, juveniles used tree cavities more frequently than adults,
whereas adults used wood stacks more frequently than juveniles. Little Owls used tree
crowns less then expected from their availability, buildings and tree cavities with one
entrance, according to their availability, and nestboxes, tree cavities with multiple
entrances and wood stacks more frequently than expected from their availability.
Temperature was associated with use of sheltered roost sites and preference indices.
Sheltered sites were used more frequently at lower temperatures. In particular, preference
for tree cavities with multiple entrances and for wood stacks increased at low tempera-
tures and preference for fruit tree crowns decreased significantly. In frosty weather
conditions, more than 80% of adults and more than 50% of juveniles roosted in sheltered
sites. Precipitation had no effect on roost-site selection. Irrespective of temperature, roost-
site selection and preference indices differed between seasons. Adult Owls increasingly
used sheltered sites and tree cavities as the season progressed from summer to winter.
Of the 98 individuals tracked after mid-July, 25 individuals were recorded as dead
during the 25 weeks of the study. Weekly apparent survival rate was similar for birds
that used sheltered sites infrequently (natal dispersal of juveniles in autumn; Φ = 0.986;
95% CrI = 0.975–0.994) and those that used them regularly (adults and juveniles in
summer, juveniles in winter; Φ = 0.986;95% CrI = 0.978–0.992), but was substantially
higher for birds that frequently used sheltered sites (adults in autumn and winter;
Φ = 0.997; 95% CrI = 0.989–1). Cryptic roosting behavior was associated with increased
survival rates.

6.4.9 Predator Avoidance

The structural diversity in the landscape needs to offer enough security for Little Owls
to cope with predators. Landscapes with a multitude of cavities limit the chance of
predation. The breeding success in piles of stones near human settlements in Portugal
might be higher due to some possible predator avoidance (Tome et al. 2004).
Apparently suitable but unoccupied habitat exists in Biscay (northern Spain) within
the distribution area of the Tawny Owl. Attacks on Little Owls by Tawny Owls have
been observed on three occasions (Zuberogoitia and Campos 1997). Mikkola (1983)
reported Little Owls in the Tawny Owl diet, and Schönn (1986) reported the absence of
overlapping territories between the two species. In Spain, perches are nearly always
used in response to possible ground predators (Fajardo et al. 1998). This argument is
also used to explain variations in daytime activity in southern Spain (Negro et al. 1990).

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 6.4 Habitat Suitability Factors

Nest predation is considered to be a key factor influencing the evolution of avian

breeding life-history traits (Roff 1992). Sexual displays may increase predation risk
through an increase in the conspicuousness of the breeders (e.g., Møller et al. 2006).
Predators may also easily locate breeders because during reproduction, the latter limit
their movements around a fixed-location nest. Several studies have provided evidence
supporting plasticity in nest-site selection and reproductive investment after a predation
event and/or a change in the predation risk during reproduction. Little Owls were found
to change nesting cavities for their replacement brood after nest predation (author’s own
observation). Parejo and Aviles (2010) investigated whether an experimentally increased
predation risk prior to reproduction affected breeding territory selection and subsequent
reproductive strategies in three Mediterranean cavity-nesting birds, i.e., the Little Owl,
the European Roller Coracias garrulus and the Scops Owl Otus scops.
Parejo and Aviles (2010) experimentally increased predation risk by presenting
realistic models of the Spanish Montpellier snake Malpolon monspessulanus at nest sites.
Snakes constitute the most important group of nest predators for open-nesting birds,
and evidence suggests that snake predation is also an important determinant of nest
failure for cavity-nesting birds (Hensley and Smith 1986, Neal et al. 1993). In 2009 they
paired nestboxes in the area by proximity (n = 66 nestboxes in 33 pairs). Each nest of a
paired nestbox (nest pair) was assigned to one of the following two treatments: a risky
treatment (i.e., Montpellier snake presentation) and a safe (control) treatment (i.e., flat
stone presentation). Stones were used as neutral, harmless and unusual objects above
the nestboxes to control for the nonpredatory object effect, as well as for the unfamiliar-
ity of objects positioned on nestboxes.
Territories used in 2008 were more likely to be re-occupied in 2009 when they
belonged to the safe rather than to the risky treatment group (with the safe treatment
14/15 occupied territories in 2008 were re-occupied in 2009 versus 8/14 with the risky
treatment; generalized linear model: Chi2 = 5.59, p = 0.018). In 2009, birds initiated
reproduction in 29 of the 33 pairs of nestboxes. In these 29 nest pairs, first breeders in
each pair of boxes showed a significant preference for safe nestboxes over risky ones
(generalized linear mixed model (GLMM); F1,28 = 5.21, p = 0.03, n = 29 nest pairs). This
pattern did not vary among the different species (generalized linear model (GLM);
Chi2 = 2.43, p = 0.30, n = 29 nest pairs) nor among periods through the breeding period
(GLM; Chi2 = 1.50, p = 0.47, n = 29 nest pairs). Considering all reproduction efforts
throughout the breeding season (i.e., final occupation), birds also occupied more safe
than risky territories (GLMM; F1,28 = 13.53, p = 0.001, n = 29 nest pairs. This pattern also
did not vary among the different species. Laying date, clutch size and the probability of
successfully raising at least one chick (success versus failure) did not differ between safe
and risky territories occupied by Little Owls. However, the average egg volume of little
owls was 8% larger in risky (mean  standard error egg volume = 14.2  0.21 cm, n = 14
eggs) than in safe territories (egg volume = 13.0  0.13 cm, n = 41 eggs; linear mixed
model (LMM); laying date effect: F1,40 = 2.09, p = 0.16; treatment effect: F1,40 = 7.44,
p = 0.009; random effect of the nest: Z = 1.90, p = 0.03).
Hence, Parejo and Aviles (2010) found that birds in their cavity-nesting bird com-
munity effectively assessed predation risk before selecting nest sites. Occupied

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territories in 2008 were more likely to be re-occupied in 2009 when they were in the safe
rather than in the risky treatment group, suggesting that predation risk may induce a
decrease in nest-site fidelity by birds, as has been already reported for other species. The
experiment seemed to have a long-term effect because final occupation of territories
depended on the manipulated predation risk at the beginning of the season, indicating
that nest-site quality is probably assessed early in the breeding season and that birds can
remember the value conferred on each site.
Little Owls breeding in risky nest sites laid bigger eggs than those nesting in safe
territories, possibly as a consequence of a nonrandom distribution of individuals
among territories differing in predation risk or alternatively because individuals might
perceive the threat imposed by snakes as enormous in risky territories and decide to
make a final investment in the current reproduction, which would be reflected in large
eggs. However, as Parejo and Aviles only had this type of result for one of the three
species i.e. Little Owl, this result may be due to chance.
Parejo and Aviles (2010) concluded that Little Owls and their bird community
display a rapid preventive response to a short-term manipulation of the predation risk
in breeding sites, highlighting the major role of nondirect effects of predation in
modulating the community structure of prey. The fear factor should always be con-
sidered and added to the direct predation effects when analyzing the consequences of
predation on prey population and community dynamics.

6.4.10 Edge Effects and Landscape Patterns

A large number of studies have examined the patterns and processes of landscape
fragmentation (e.g., Turner and Gardner 1997, Turner et al. 2003) and the impact of
habitat edges on bird communities (e.g., Hudson 1991, Sparks et al. 1996, Fuller et al.
1997, Baker et al. 2002, Berg 2002, Schroth 2004).
Hedges are an important landscape element that can have a favorable impact on species
diversity (Hinsley and Bellamy 2000). Increasing the structural complexity of a hedgerow
and its associated habitat elements may reduce the incidence of predation on birds.
Hedgerows also provide physical shelter and are an important source of winter food supplies,
especially berries and other fruits.
The reproductive success in the Loggerhead Shrike (Lanius ludovicianus) was lower
from nests along fence lines than from nests found free standing and distributed at
random (Yosef 1994) due to greater rates of predation. Where nests of breeding species
are concentrated in a linear fashion, it should be most profitable for a predator to search
along such corridors. Hole-nesting species were found to be more at risk at the edge of
forests than within the forests due to a higher predation risk (Sandstrom 1991).
Similarly, increased nest predation was found in neotropical migrant birds at a forest–
stream ecotone (Gates and Giffen 1991).
Since Little Owls are both predator and prey, they need to seek a profitable trade-off.
When predators are limited, breeding in coppiced tree lines may be extremely positive
since both prey abundance and detectability are favorable. To minimize the risk of nest

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 6.4 Habitat Suitability Factors

predation on the owl itself, the habitat should offer plenty of linear elements rather
than just a few. Hence, mosaic-like landscapes which feature a maximal amount of
edges will reduce the risk of predation on the owl.
Some landscape elements have different impacts on the species in different concen-
trations. Farm buildings and their surroundings had a positive impact on Little Owls in
France (Bretagnolle et al. 2001). In the absence of natural cavities in Flanders, the
species is attracted to farm buildings for breeding and during winter. In small numbers,
the presence of farm buildings had a positive impact on Little Owl occurrence, but
when this land cover surpassed 0.21 ha per 25 ha, the impact became negative (Van
Nieuwenhuyse et al. 2001c). A differential impact of roads on the presence of species
was found, according to traffic loads, in Haut-Léon, western France, where Clec’h
(2001b) found that a 2-km-wide area bordering dual carriageway roads and a 500 m
area around a major road was not occupied, while roads with limited traffic were
not avoided.

6.4.11 Negative Habitat Relationships

The density of Little Owls is suppressed by factors that decrease the accessibility and
availability of prey or the number of nesting cavities, or increase the influence of
predator or human actions. The increase in the amount of arable land has had a
negative impact in Germany; further, the presence of trees was positively correlated
with meadows and negatively correlated with arable land (Loske 1986, Van
Nieuwenhuyse and Bekaert 2001). The owl avoids large rape and mustard seed fields
in Deux-Sèvres, western France (Bretagnolle et al. 2001). It also avoids larger forests,
thus the spread of Little Owls in Europe only occurred after the principal deforestation
period of the ninth and tenth centuries (Schönn 1986). The cessation of
pastoral farming practices (e.g., husbandry of sheep and cattle) resulted in a
reduction in the amount of short vegetation (Juillard et al. 1992). The owl avoids
larger cities in the UK (Sharrock 1976), larger built-up areas in Flanders (Van
Nieuwenhuyse and Bekaert 2002), traffic (Fajardo et al. 1998, Clec’h 2001b), forest
(Schönn et al. 1991, Van Nieuwenhuyse and Bekaert 2001, 2002), and villages with
few but large grassland areas (Dalbeck et al. 1999a,b). The only exceptions are when
these areas are surrounded by “traditional” farming practices, i.e., environments where
different land uses cover small areas, providing heterogeneous habitats within small
areas (Ferrus et al. 2002). In the southern part of its distribution, the Little Owl is also
found in large cities (Rome, Italy; Budapest, Hungary; Bucarest, Romania; Sofia,
Bulgaria; Ankara, Turkey). Built-up areas and maize (Zea mays) culture in Flanders
(Belgium), up to an optimal coverage of 0.21 ha per 25 ha and 0.65 ha per 25 ha
respectively, have had a positive impact on the presence of owls. The probabilities of
finding Little Owls in areas larger than this decrease again (Van Nieuwenhuyse et al.
2001c).
Little Owls negatively associate with forest, and tend to avoid forest edges up to
150 m (Michel et al. 2016).

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6.5 Assessments at Multiple Spatial Scales

The multiscale approach to studying habitat selection is based on the conceptual
framework suggested by Johnson (1980), whose basic assumption was that animals
are capable of making decisions regarding resources at consecutively smaller scales.
Consequently, general habitat selection can be regarded as a hierarchical process, e.g.,
a fitting patch for breeding at a small scale and appropriate areas for foraging at a larger
scale (Martinez and Zuberogoitia 2004b). The multiscale approach may be especially
useful in identifying key factors involved in habitat preferences of owls because they
have large home ranges, usually comprised of different patches for breeding and
foraging (Mikkola 1983, Taylor 1994). Importantly, some of the apparently conflicting
results of habitat preference identified in different research areas are potentially due to
researchers working at different spatial scales. Analysis of habitat preferences at differ-
ent spatial scales within the same research area can reveal this. The idea of different
scales was introduced by Schönn et al. (1991) who identified three aspects of species
ecology that a habitat must provide for successful survival, i.e., reproduction, foraging
and roosting. The three areas do not necessarily have to be situated in concentric circles.

6.5.1 Multiscale Habitat Assessments

The multiscale approach is especially useful in identifying key factors of habitat prefer-
ences for owls, as they have large home ranges, usually comprised of different patches
for breeding, foraging, roosting and dispersing (e.g., Mikkola 1983, Taylor 1994,
Kavanagh 2002).
Multiscale habitat assessments have become a common analysis tool in owl conser-
vation and management. Most often, these assessments reflect the enumeration of
habitat conditions within circles of different sizes that are centered over a point in
the landscape (e.g., the nest site) and determined from vegetation maps, aerial photo-
graphs or satellite imagery. The smallest circle size is often 0.1 ha and it is common
practice for researchers to go into the field and physically measure the types and size of
vegetation or other structures at this scale. Subsequent larger circles reflect the average
home-range size of a pair of the owl species under review, and a circle that reflects a
larger landscape size (e.g., to assess the characteristics of watersheds, soils, climate).
Particularly valuable are assessments where comparisons are made between habitat
conditions around owl sites and those around random points in the landscape.
Sufficient sample sizes are critical in determining the statistical significance of the
observed differences.
Not all nesting territories are of equal benefit to the local owl population (see
Newton 2002, 1998 for a review of limiting factors in owls and birds, respectively).
Researchers have observed that for some species of owls, relatively few nesting territor-
ies provide most of the reproduction to a given population. The results of multiscale
habitat assessments benefit significantly by linking long-term reproductive data from
specific nest locations. This type of information is particularly relevant in the develop-
ment of specific habitat management recommendations for the species of interest.

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 6.5 Assessments at Multiple Spatial Scales

Table 6.2 Different scales of study in habitat selection of Little Owls

Van Martinez and

Nieuwenhuyse Zuberogoitia
et al. 2008 2004b Blache 2004 Šálek et al. 2016

Nest-site 282 m 25 ha 200 m 10.5 ha 250 m 19.6 ha 168 m 8.9 ha

scale
Home- 564 m 100 ha 309 m 30 ha 500 m 78.54 ha 284 m 25.3 ha
range
scale
Landscape 2000 m 1256 ha 5600 m 100 km² 1000 m 314.15 ha 5600 m 100 km²
scale

A growing number of multiscale habitat assessments have been undertaken on Little

Owls (i.e., Blache 2004, Martinez and Zuberogoitia 2004b, Zabala et al. 2006, Cornulier
and Bretagnolle 2006, Zabala et al. 2006, Van Nieuwenhuyse et al. 2008; Šálek et al.
2016, Apolloni et al. 2018). The different spatial scales applied are given in Table 6.2.
Martinez and Zuberogoitia (2004b) assessed habitat conditions within three plot
sizes, centered on 78 Little Owl nests and 55 nonoccupied but apparently suitable
territories. A territory was considered as suitable for nesting when it contained mature
carob trees, derelict or inhabitated country houses or ephemeral rivers, according to the
general preferences of the Little Owl in the study area. Habitat selection was studied
using GLMs using a logit link to identify the difference between occupied and unoccu-
pied habitats. At the nest-site scale, they found significantly more arid plantations with
high availability of linear structures in the proximity of villages in occupied habitats
than unoccupied ones. At the home-range scale, Little Owls were found more often in
habitats with high percentages of arid plantations, linear structures, the presence of
neighboring Little Owls and the vicinity of cities. At the landscape scale, the model
predicts occupancy of habitats in areas with arid plantations. At all spatial scales, arid
plantations is the most discriminating parameter between occupied and unoccupied
habitats, stressing the importance of traditional land uses.
Blache (2004) used three different methods to study the habitat preferences of Little
Owls in Plaine de Valence, France. First he delimited seven types of landscapes using a
multiple correspondance analysis of 817 grid cells of 250  250 m with a hierarchical
clustering of the first two axes. Jacob’s Index (Jacobs 1974) was used to determine if
some landscape types, at the smallest scale, were disproportionately used by the owls.
The habitat type that contained the highest number of houses and isolated farms but
also a significant proportion of fields, hedges and isolated trees was preferred by
Little Owls.
Second, the importance of 15 individual landscape elements was studied using
univariate logistic regression of occupied and unoccupied grid cells. At two spatial
scales (250  250 m and 750  750 m gird cells), isolated farms and mulberry trees
were positively associated with Little Owl presence. Intensively managed orchards and

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 Chapter 6: Habitat

forests showed a negative association with Little Owls. At the smallest scale, roads,
grasslands, the number of isolated houses and hedges showed moderate association
with Little Owl presence.
Blache (2004) finally studied the importance of the parcel size around occupied (n =
52) and unoccupied farms (n = 23). T-tests were used to compare the mean parcel area
between occupied and unoccupied farms at the three spatial scales given in Table 6.2. In
contrast to most other studies, the habitat diversity around farms was not a discrimin-
ating factor between occupied and unoccupied farms. The species prefers farms with
only a few large parcels. Farms that had an occupation of several years by owls were
found to have the smallest diversity of habitat. The species does not show a preference
for a fragmented habitat in Valence. In the Netherlands, van’t Hoff (2001) found that
seemingly good habitats that were isolated were abandoned sooner than clustered
habitats of lower quality. Further research on the role of social interactions in this area
might explain this interesting behavior of the species.
Zabala et al. (2006) modeled the distribution of a Little Owl population in Spain
using data on land use and landscape composition in occupied and unoccupied areas.
Their statistical analyses (logistic regression and multiple linear regression) showed that
variables such as topography, altitude, road density and urban areas had a negative
effect at the lower scale, whilst conifer plantations had a negative effect at every spatial
scale. The density of predator species, in particular the Tawny Owl, linked to the
plantations, had an effect only over habitat selection, but not over occupancy. The
negative effect of Tawny Owls on Little Owls is only highlighted in the habitat
selection analysis.
Cornulier and Bretagnolle (2006) used a recently developed method generalizing
Ripley’s K function for nonhomogeneous point patterns to test the aggregation of
the nests of Little Owls exhibiting heterogeneous distributions in response to landscape
structure. Initially the Little Owl was found to form clusters at some scale; taking spatial
heterogeneity into account revealed that territorial Little Owls showed no clustering
of territories when habitat availability was considered. When assuming homogeneity,
Little Owls maintained some regular spacing at small scales (ca. 150 m), as expected
from a territorial species, whereas territories were clustered at medium scales, between
500 and 1000 m. Consequently, opposite patterns can be found in a single dataset
at different scales. This result highlights that it is recommended to account for non-
stationarity when testing for aggregation. The authors demonstrated that accounting
for large- or small-scale heterogeneity affects the perception of spacing behaviors
differently.
Martinez and Zuberogoitia (2004b) found that at the nest-site scale, Long-eared Owls
preferred wooded areas with few paved roads while Little Owls preferred arid planta-
tions. Furthermore, the probability of finding an occupied territory increased with the
proximity of another occupied territory in the surroundings. The home-range scale
models mirrored the feeding requirements of the owls. Long-eared Owls occupied areas
with high percentages of forest, arid plantations, edges between these two land uses,
short distances between nests, the presence of conspecifics and little human disturb-
ance. Little Owls occupied arid plantations with high availability of linear structures

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 6.5 Assessments at Multiple Spatial Scales

and proximity to villages. At the landscape scale, Long-eared Owls avoided extensive
forests, and Little Owls preferred arid plantations.
Van Nieuwenhuyse et al. (2008) examined the relationship between the presence of
Little Owls and the landscape in three different sized circular plots (25 ha, 100 ha,
1256 ha) in East-Flanders (northern Belgium) with discriminant analysis. They used
the number of parcels, the length of the parcel edges and the parcel area of rowcrops,
cereals, maize, built-up areas, farm buildings, forests. The lengths of waterways, tree
lines and roads were also included in the analysis.
The predicted probability of Little Owl presence dropped with the area covered by
farm buildings at the nest-site scale and increased with these areas at the home-range
and the landscape scales. The perimeter of the farm buildings associated positively at
the nest-site scale, but negatively at larger scales. At the three spatial levels, a positive
association of Little Owls with maize edges was found, while it was negative with the
area of maize or number of maize parcels. This indicates that maize parcels do not have
a negative influence on the species as long as the parcel size is very limited, confirming
the preference by the owl for mosaic-like landscapes. For a certain period during the
growing of maize, the parcels remain mainly bare land, which is favorable for Little
Owls. Large maize parcels were negatively associated with Little Owls. Roads and
waterways featured a negative association with the species at all levels, in contrast to
Barn Owls that seem to be attracted by waterways. Little Owls were negatively associ-
ated with forests at nest-site level, but positively at larger scales. Railways were posi-
tively associated with the owls at the nest-site and home-range scales but negatively at
the landscape scale. The species can tolerate smaller amounts of railway. Larger
amounts of railways are most associated with urban centers and are hence avoided.
To compare the habitat selection of Little Owls with Barn Owls, Van Nieuwenhuyse et al.
(2008) quantified for each landscape element the difference between the coefficients of
both discriminant functions (both good and bad habitats). As they are a common landscape
feature, railways are associated with both species, as is shrubbery and tree-growing parcels at
the nest-site scale. Barn Owl presence was mainly associated with the amount of cereal grain
fields. Tree lines discriminate for Little Owls but not for Barn Owls. At the 100 ha scale,
waterways become very distinctive for Barn Owls and not at all for Little Owls. On the other
hand the area of fallow land is distinctive for Little Owls. High-stem orchard are positively
discriminative for both species. Finally, at the landscape level, Barn Owls showed a positive
association with urban areas and railways, while for Little Owls these landscape elements
have a negative association with the species presence. The occurrence of elm forests is
positively associated with Little Owls and negatively with Barn Owls. Earlier research at
the nest-site scale revealed a positive impact of conspecifics on the probability of finding
Little Owls (Van Nieuwenhuyse and Bekaert 2002).
Šálek et al. (2016) studied habitat selection of Little Owls in well-studied research
areas in the Czech Republic, Slovakia and Hungary at three spatial scales. The popula-
tion densities in the study regions differ markedly with the lowest density in the Czech
Republic (0.29 calling males/10 km²) and Slovakia (0.92 calling males/km²) compared
to 5.01 calling males/10 km² in Hungary. Per country 50 occupied and 50 unoccupied
habitats were randomly selected using repeated playback visits.

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 Chapter 6: Habitat

Habitat selection was studied using presence–absence data to compare drivers for
occupation. Nine habitat variables that might be important predictors of occurrence
were selected: proportion of grasslands, arable land, residential buildings, gardens and
orchards, artificial structures, farm buildings, woodland, reeds and marshes, and water
area. Principal componant analysis (PCA) was applied to study correlating independent
variables. Generalized additive models (GAMs) and GLMs were calibrated to assess the
probability of Little Owl presence per region. GAMs used forward-stepwise selection.
GLMMs were used to allow comparison of GAMs with their parametric counterparts.
The percentage cover of grasslands, arable land, gardens and orchards, and farm
buildings were generally the most important habitat attributes at smaller spatial scales.
At large spatial scales the owl was positively associated with open habitats in terms of
arable fields.
After controlling for regional effects, the percentage cover of gardens and orchards at
the home-range scale was shown to be another generally important habitat attribute.
The association with gardens and orchards varied inter-regionally with respect to the
spatial scale. While this factor correlated positively at the nest-site and home-range
scales, it did not with other habitat attributes and appeared to reflect a unique land-
scape feature. As owls prefer to breed in artificial constructions such as farm buildings in
Central Europe, the association with gardens and orchards may rather reflect a prefer-
ence for high-quality foraging habitats composed of a mosaic of trees and short vegeta-
tion. Woody habitats had a negative impact on the probability of occupation due to a
possibly higher predation pressure. As expected, a stronger association of Little Owls
with arable land in the Czech Republic and Slovakia than in Hungary, indicated a lower
availability of suitable open habitats in the former two regions.
Farm buildings were found to be important habitat factors at smaller spatial scales
across all study regions. Farm buildings are known to be the main nesting locations in
Central Europe. They offer a high diversity of nesting sites and nearby feeding grounds.
Active dairy farms are especially attractive due to grain spillage, silage holes, grain
storehouses, manure heaps, and a variety of short-sward pastures and hayfields, which
provide a high diversity of food during the entire year.
Apolloni et al. (2018) studied habitat selection and range use of adult owls in relation
to landscape configuration, habitat structure and resource distribution via telemetry.
Large- and fine-scale distribution patterns of voles were assessed by transect counts of
signs of vole presence. An experiment using artificial perches was carried out to deter-
mine the fine-scale adjustment of the owls’ range use in relation to prey abundance and
vegetation height. Habitat selection and resource exploitation were structured at three
spatial levels. At the landscape scale, orchards were highly preferred over other areas,
correlating with the patchy large-scale occurrence of voles, which were absent in crop-
land but abundant in orchards and grassland, and perch use, in combination with prey
abundance and vegetation height. Within home ranges, the spatial distribution was
highly inhomogeneous and structures with high prey abundance were used over-
proportionally. At the scale of foraging sites, the owl preferred patches with low
vegetation over those with high prey abundance, establishing that prey availability
is crucial.

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 6.6 Multilevel Resource Selection

All levels of habitat selection and range use are related to farming practices and
affected by current cultivation. Conservation measures should focus on the conser-
vation and restoration of orchards at the landscape level and habitat management
measures should focus on grasslands as the main food providers by creating a mosaic
of patches with short and tall grass. Together with other habitat structures providing
food resources such as field edges, wildflower areas and structures facilitating access to
prey, the quality of habitat patches in terms of food availability could be greatly
improved.

6.6 Multilevel Resource Selection

Habitat models are fundamental tools for designing evidence-based conservation
measures, particularly for locating sites with high potential for promoting species
re-colonization and occupancy. However, it remains challenging to respond to both
the need for large-scale general rules, and for fine-scale information concurrently.
In the classic hierarchical nested design (Johnson 1980) as presented above, infer-
ences are specific to each level and sometimes conflicting. Factors operating at the
broader scales may constrain habitat availability at the finer scales, and conversely,
large-scale patterns can also result from individual behavioral processes at finer scales
(Mayor et al. 2009).
Multilevel habitat models provide all-in-one surfaces that explicitly account for
conditional dependencies among single-level selection probabilities. These conditional
relationships among the hierarchical levels of selection have recently been explicitly
included in multilevel models (McGarigal et al. 2016) that transcend single-level
models by integrating the different levels of selection. Such integrations are relatively
easy to achieve, as the hierarchical conditional probabilities collapse to a simple equa-
tion that is the product of the relative probabilities (DeCesare et al. 2012). Such scale-
optimized, multilevel models provide a more integral insight into animal–habitat
associations than single-level models, but they have rarely been applied in a conser-
vation context (DeCesare et al. 2012). An all-in-one habitat suitability model provides a
better basis to inform conservation strategies over large spatial extents as well as local
conservation measures, thereby responding simultaneously to the needs of both policy-
makers, who call for large-scale general rules, and to practitioners in the field who call
for fine-scale information (DeCesare et al. 2012).
Fattebert et al. (2018) developed multilevel habitat suitability models for the Little
Owl over a large area, comprising Switzerland and the neighboring Baden-Württemberg
state in southwestern Germany, aiming to identify the most promising conservation
areas. Candidate explanatory covariates were based on previous studies: elevation,
slope, Normalized Difference Vegetation Index, percentage croplands, percentage
forests, percentage orchards, percentage meadows, percentage hedge, percentage
built-up and distance to the forest edge. They evaluated habitat suitability for the
species over a large area by using publicly available, high-resolution governmental
land-use data and Little Owl occurrence data obtained from citizen-science species

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 Chapter 6: Habitat

observations in combination with radio-tracking data. Their study encompassed large

geographical areas of two European countries, differing in their agricultural policies.
The all-in-one model resolves two main issues in the conservation of an endangered
species: identification of large-scale suitability patterns to develop conservation strat-
egies and identification of fine-scale clusters of high-quality habitat patches where
conservation measures can be applied. Level integration enhanced the estimates of
suitability by considering home-range selection rules at both the landscape and the
home-range selection levels, and habitat preferences within home ranges. The large-
scale evaluation of Little Owl habitat suitability showed that suitability values were
generally lower in Switzerland than in Baden-Württemberg, and the distribution of
good habitat areas differed between the two countries. Single-level habitat selection
models showed that the occurrence of Little Owl populations was based on topographic
factors and the large-scale avoidance of forests, while within Little Owl landscapes,
preferences for home-range placement and range use were based on the availability of
important vegetation and land-use types.
The results of the three individual scales of habitat selection analyses confirm that
good Little Owl habitat is located in widely open agricultural landscapes that often exist
in the vicinity of settlements or even suburban areas. Such open landscapes can offer a
high diversity in the form of stands of trees (orchards, tree rows, tree groups) with
associated meadowland. Within these areas, home ranges occurred at some distance
from forests, and are preferentially located in meadowland with scattered trees. While
slopes within the home range might not be of direct importance for Little Owls, they
may correlate with fine-scale characteristics of preferred open areas, such as food
availability or tree cavities. On the other hand, at the landscape level, steeper slopes
have a negative effect on owl occurrence, as they may not offer these preferred fine-
scale resources. The final multilevel model validated well over the broad landscape,
demonstrating good predictive ability and transferability. It is accurate in identifying
areas and patches of good habitat at a larger scale, taking into account fine-scale
information too.
In conclusion, Fattebert el al. (2018) showed that the simultaneous integration of
different spatial scales of habitat selection behavior into a multilevel habitat suitability
map creates a promising tool for conservation planning of endangered species over
large geographical areas. Level integration adjusts areas of good habitat in the landscape
by considering home-range selection processes, and corrected suitability values, and
thus the distribution of high-quality patches, by considering habitat use within home
ranges. The single habitat suitability layer allows for adequate identification of both
large-scale suitability patterns to develop conservation strategies, and fine-scale clusters
of high quality where conservation measures can be applied.

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 Chapter 7: Diet

Chapter Summary
The Little Owl has a generalist diet and takes a high diversity of small prey. It catches
and eats a variety of small-sized prey across its entire distribution area. The diet of the
owl varies with the season and the geographical area. From north to south and from
winter to summer, an increase in the numbers of insects in the diet has been observed.
Regardless of the numbers of insects, small mammals remain the key prey category by
biomass and energetic yield, and they contribute significantly to the ecology and welfare
of the species. As with some insect-eating owl species, it is very difficult to get a true
picture of its diet by only studying its pellets or prey remains in nestboxes, because Little
Owls catch lots of prey of which the remains are hard to find and identify in pellets
(e.g., insects and especially earthworms, caterpillars and larvae). We present their relative
proportion. In this chapter we look at this prey diversity through time and space, and
focus on the hunting method of the owl, as well as caching behavior in larders. We use
the term larder to refer to food stored in the nest site unless otherwize indicated. We offer
a thorough review of the owl’s diet and individual prey species, with attention given
to the importance of voles and other large prey. Pellet contents are described in detail
and comprehensive results of camera-observed breeding seasons are presented.
Much of the new insights stem from the research of Van Harxen and Stroeken
(personal communication) who examined 17 breeding seasons by camera observation
in 2002–2020, totaling 34 916 prey items.

7.1 Hunting
The typical nocturnal activity pattern of the Little Owl has two peaks of about 1–2
hours each, occurring just after sunset and before sunrise, separated by a period of less
activity around midnight (Exo 1989). Van Harxen and Stroeken (personal communi-
cation) found a different pattern between males and females during the nestling phase.
Both sexes showed a clear evening peak (between 22:00 and 23:00), but the morning
peak was much less pronounced in males compared to females (Figure 7.1).

Supplementary resources are available online at www.cambridge.org/littleowl. These include

appendices as well as tables which previously appeared in the first edition (prefixed with ‘S’).

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 Chapter 7: Diet

3500

3000
Male Female
2500

2000

1500

1000

500

0
20–21
21–22
22–23
23–24
00–01
01–02
02–03
03–04
04–05
05–06
06–07
07–08
08–09
09–10
10–11
11–12
12–13
13–14
14–15
15–16
16–17
17–18
18–19
19–20
Figure 7.1 Prey supply during the nestling phase per hour for 17 camera-observed breeding seasons,
2002–2020; 34 916 prey items (Van Harxen and Stroeken personal communication).

The Little Owl also hunts by day, especially during the nestling phase (Hibbert-Ware
1938, Van Zoest and Fuchs 1988, Ryabitsev 2001) and during the warm season in the
Mediterranean (Zerunian et al. 1982, Lo Verde and Massa 1988, Negro et al. 1990) and in
eastern Palearctic areas (Osmolovskaya 1953). Van Harxen and Stroeken (personal commu-
nication) found that both sexes deliver a substantial part of the prey to their nestlings by
day, 18.2% of prey items. Males (21.1%) contributed more in daytime than females (15.9%).
The Little Owl’s crepuscular and nocturnal feeding habits (Hibbert-Ware 1938,
Haverschmidt 1946, Juillard 1984, Van Zoest and Fuchs 1988, Schönn et al. 1991,
Fajardo et al. 1998, Ryabitsev 2001) are linked to prey activity (Zerunian et al. 1982,
Fattorini et al. 2000). Van Harxen and Stroeken (2010, 2011b) found insects, frogs and
birds disproportionally being caught by day, and rat-tailed maggots (Eristalis tenax),
moths and salamanders almost exclusively in the evening and night. They saw also a
remarkable difference between Wood Mice (Apodemus sylvaticus) and Common Voles
(Microtus arvalis), the latter being caught by day twice as much, reflecting the activity
pattern of both species (Table 7.1).
The method of hunting is related to the type of vegetation in the foraging area
(Schönn et al. 1991). In low vegetation, Little Owls hunt mainly beetles, earthworms
and micromammals on the ground from a perch, dropping onto prey below or nearby
(Hibbert-Ware 1938, Van Zoest and Fuchs 1988, Schönn et al. 1991, Fajardo et al. 1998).
In Belgium, Libois (1977) reported that the Little Owl caught 65% of its prey (n = 1782)
on the ground.
When the vegetation is high, it catches flying insects from a perch, using a twisting
and turning flight pattern (Haverschmidt 1946, Beven 1979). In the Netherlands, the
type of perch is related to the type of prey chosen. The time spent on a perch is variable
and depends on the perch height (Van Zoest and Fuchs 1988). In open Holm-oak
woodland and treeless pseudo-steppes in the south of Portugal, Little Owls had similar
hunting success in woodland vegetation, despite it being higher and more dense than
the pseudo-steppe. Little Owls corrected for this by choosing higher perch sites in
woodland (average 2.76 m versus 0.77 m in the pseudo-steppes). The fact that they
did not choose higher available sites (maximum available 5 m) indicates an optimal

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Table 7.1 Percentage of different prey items caught during three periods (night, day, evening) (n = 40 754)

Small Larvae and Rat-tailed Total

Time Insects Frogs prey caterpillars Cockchafer maggots Earthworms Salamanders Shrews Moths Birds Mice Voles count

00:00–06:00 36.0 35.9 33.7 59.4 15.5 72.2 53.0 52.9 51.2 47.3 16.3 51.7 38.2 17 140
06:00–20:00 28.6 36.8 22.3 13.0 21.2 2.3 18.0 2.9 12.2 2.7 65.2 15.2 33.0 7296
20:00–24:00 35.4 27.3 44.0 27.6 63.3 25.6 29.1 44.1 36.6 50.0 18.5 33.1 28.8 16 318
Total count 1503 315 5679 14 787 10 340 528 5020 68 41 930 92 650 801 40 754

Van Harxen and Stroeken personal communication.

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foraging height, above which prey detectability decreases due to vegetation character-
istics, despite the higher visual field (Tomé et al. 2011). Little Owls choose lower perches
when the vegetation is higher to increase the efficiency of acoustic detection in the
pseudo-steppes.
The owl can catch beetles on the ground by approaching in a short low flight, followed
by rapid runs or a few hops (Beven 1979). A Little Owl was reported to have run across a lawn
in very rapid bursts and at the end of each burst stopped with its head cocked as if listening
and occasionally made a grab at something with its beak, then hopped like a thrush and ran
rapidly again (Cameron-Brown 1975). The Little Owl hunts earthworms by hopping over
the ground, stopping suddenly, bending forward and seizing a worm with its bill, and
pulling it, often flapping its wings to maintain its balance, sometimes falling onto its back
as the worm comes loose (Haverschmidt 1946). Earthworms are predominantly caught
when it is dark (76% between 22:00 and 06:00), but on rainy days also during daytime. In
dry periods they can suddenly appear in the nestling supply when the lawn is sprayed.
Rat-tailed maggots (larva of Eristalis tenax) are caught when they crawl over the
ground searching for a proper place to pupate. As they do this in great numbers in
suitable weather conditions, Little Owls sometimes return to the same spot over and
over again. They can pick them up between pavement stones where they try to hide
(Pauline van Marle personal communication).
Cockchafers (Melolontha melolontha) are caught in various ways, by picking up those that
fall to the ground when they are pairing in oaks alongside the road, by snatching them from
branches and leaves and, as cockchafers fly in a very sluggish way, by chasing them in flight.
After sunset, cockchafers are, like many insects, attracted by artificial light. Beetles that hit the
lamp fall on the ground and are easy prey. Cockchafers are caught throughout the day, but
the majority of them (57%) are captured between 21:00 and 23:00. Before they are fed to
nestlings, the elytra of cockchafers are removed by the female. When they are consumed in
great numbers the bottom of a nestbox is strewn with discarded elytra. Besides vertebrates,
cockchafers are almost the only type of prey that is stored in or at the entrance of the nest
(Van Harxen and Stroeken personal communication) in massive numbers, i.e., 50 beetles.
The method of transporting captured prey differs per prey item. In Switzerland,
Juillard (1984) determined that for 4918 prey brought to the nests, almost all inverte-
brates were transported in the bill, 98% of birds in the bill (the rest in the foot) and 65%
of small mammals were transported in the bill and 35% in the foot. There was almost
no significant difference between males and females in the method of transport, but the
male brought more prey by the head than the female who prefered carrying them by
the neck (Juillard 1984).
During summer nights, the owls like hunting under lamps, well-lit houses, yards,
streets and roads, where plenty of insects are concentrated, especially moths (Nankinov
2002) or grasshoppers on asphalt roads (Adam 1973). Besides fields, the Little Owl
hunts on dunghills, in gardens and on lawns by dusk and during the first hours of
the night in Denmark (Laursen 1981). The species is able to catch bats (Reinard 1977).
Van Harxen and Stroeken (personal communication) found four bats in a larder in a
nestbox during their research between 1998 and 2021, of which two were Common
Pipistrelle (Pipistrellus pipistrellus).

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 7.2 Capturing Prey

The owl raids the nests and nest cavities of birds such as Starlings (Sturnus vulgaris;
Tinbergen and Tinbergen 1932, Elliott 1940, Anderson 1949, Burton 1983), House
Sparrows (Passer domesticus; Desmots 1988) and Tree Sparrows (Passer montanus;
Steffen 1958). One owl regularly stole worms from a Blackbird (Turdus merula; Tricot
1968). Birds can be taken from the water (Barber 1925) or ground (Ellis 1946). The Little
Owl also hovers like a Kestrel (Gregory 1944, Tayler 1944, Gyllin 1968, Malmstigen
1970, Wahlsted 1971, Martin and Rollinat 1982).
The Little Owl can combine different types of hunting according to available prey.
Thus, by daylight in southern France, a nervous bird hunted by perching, hovered
sometimes like a harrier, caught beetles on the ground with its bill or claws, changed
perches, hovered and walked on the ground picking up some beetles (Génot and
Lecomte 2002).
In the Cis-Balkhash Lake area (Russia) or in Kazakhstan, the Little Owl is a frequent
visitor of Gerbil (Gerbillinae) burrows (Shtegman 1960, Gavrin 1962). At night, they catch
Gerbils as they sleep in their nests and eat them immediately. They use certain burrows as
larders and can settle in the colonies of underground Gerbils, an interesting parallel with
the behavior of the Burrowing Owl (Athene cunicularia). This special “underground”
behavior of the Little Owl visiting and living in burrows is also demonstrated by the
wear patterns on their feathers from skins in museum collections in Central Asia and
Middle Asia (Van Nieuwenhuyse et al. 2008). The upper parts of the feathers are missing,
especially on the head, back and shoulders, and the feather’s blades are nude stems
jutting like needles. This underground behavior might help to explain the mystery
regarding the bad condition of feathers from the eastern subspecies (Shtegman 1960).

7.2 Capturing Prey

The ontogenetic development of catching and handling prey in the Little Owl was studied
by Ille (1983) in an aviary. The capture of prey by an adult owl entails different steps, i.e.,
locating the prey, approaching, attacking the prey and eating or storing the prey, and also
nonobligatory acts such as ruffling the feathers or mantling the prey with its wings. These
different steps of catching prey appear to depend on the condition of the owl (age, state of
hunger) or/and on environmental factors (size and resistance of prey, competition for
food). The ruffling of the feathers and mantling of prey are due to threat behavior towards
the prey and competitors for food. The beating of wings on a large prey that is still moving
serves only for maintaining balance. Every kind of movement of the prey, real or artificial,
triggers an act of catching the prey, explaining the aggressive behavior of the Little Owl.
The owl reacts to the form of prey according to a process of conditioning. In captivity, the
species prefers to seize a mouse by the body and strike it with its beak (Csermely et al. 2002).
The complete act of catching prey appears after a maturation process involving
several stages. The fledglings go through this process at an average age of 62–76 days.
The choice of prey and the foraging method is not instinctual but rather a learned
process (Sageder 1990). Although experiments in captivity showed that Little Owls
prefer catching live prey over dead, the species can eat dead prey such as Hedgehogs

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 Chapter 7: Diet

120

male / numbers male / biomass

100
Percentage prey supply by sex

female / numbers female / biomass

80

60

40

20

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35
Days after first hatching

Figure 7.2 Percentage prey supply in the nestling phase by the male and female (for 15 camera-
observed breeding seasons by observation of prey items with gender identified, 2002–2020; 33 579
prey items (Van Harxen and Stroeken personal communication).

(Erinaceus europaeus) that have been victims of vehicle traffic (Van de Velde and
Mannaert 1980). While carrion is very seldom used as food (Hibbert-Ware 1938), the
Little Owl can be necrophageous in some cases, feeding on dead adult chickens (Blache
2001) or 1-day-old chicks (Berghmans 2001a) or dead mice intentionally deposited near
the nestbox by the landowner. They also take different bait, purposely put in the
vicinity of the nestbox; not only voles and mice but also 1-day-old chicks and even
meat meant for human consumption like mince (when put in the nestbox). Some
worried landowners provide extra food for the juveniles by putting cat food in the
nestbox, which is always eaten.
Van Harxen and Stroeken (personal communication) showed that despite the male
being indispensable during the first 10 days of the nestling phase with respect to prey
supply, from day 1 the female increased her share in the supply (Figure 7.2). Around day
10 she started delivering more prey than the male (based on numbers) up to more than
70% in the second half of the nestling phase. With respect to biomass, the turning
point lay a few days later (day 14).
In total, the males delivered 46.5% of the prey items and 53.5% of the biomass. The
difference was due to the fact that males catch 55.5% of the heavier, vertebrate prey. Females
caught the greater part of the small invertebrate prey and earthworms (60.8% each).
Cockchafers were caught in more equal proportions, with females slightly more (52.1%).

7.3 Prey Caching

Larders contain the remains of prey taken before feeding it to the nestlings. The term
“larder” is only partly correct, as the primary purpose is a deposit location for surplus

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 7.3 Prey Caching

Figure 7.3 Wood Mice in a Little Owl larder.

food, rather than for formal food storage (Hibbert-Ware 1938a). The male brings food
from the larder to the female at the nest site as required (Owen 1919). The food reserves
hidden in holes and niches are often eaten in one day (Gavrin 1962). Certain Little
Owls take their food directly to the nest as they probably have no other suitable hole for
a larder (Hibbert-Ware 1938).
During the breeding season, most prey is brought directly to the breeding female or to
the nestlings. Vertebrate prey in particular is then placed in a pile in a corner of the nest
and stored for later. In years where micromammals are plentiful, such a supply can consist
of many dozens of prey items of various species. The largest larder Van Harxen and
Stroeken (2011b) found contained 13 Wood Mice (Apodemus sylvaticus), 21 Bank Voles
(Myodes glareolus), 11 Common Voles (Microtus arvalis) and 1 unidentified mouse, i.e.,
46 specimens in all stages of decomposition. The highest number of Wood Mice found
during a nestbox check was 30 (Figure 7.3), 37 Common Voles were once found in one pile,
25 for Bank Voles and 14 for House Mice (Mus musculus). The highest number of large
Yellow-Necked Mice (Apodemus flavicollis) during one check was 3, Water Voles (Arvicola
amphibius) 5, Brown Rats (Rattus norvegicus) 6, and Common Shrews (Sorex araneus) and
Greater White-Toothed Shrews (Crocidura russula) 3 each (Van Harxen and Stroeken
personal communication). The largest larder, with 57 specimens, was found in Brabant
in the Netherlands (Anita van Dooren/Christien Hermsen personal communication).
Larders are used in winter too, e.g., in Switzerland containing 22 voles (Blanc 1958)
and in a nestbox with 95 rodents (Juillard 1984). In Russia, a Little Owl built reserves of
30 or more animals (Koblik 2001). In Britain, a cache of 167 headless Storm Petrels
(Hydrobates pelagicus) was recorded by Lockley (1938). Short-term storage of food in
larders is a behavior shown by a large number of owl species, especially Pygmy Owls
(Glaucidium sp.) (Del Hoyo et al. 1999).

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7.4 Prey Species

In the western Palearctic, the diversity of prey species can be summarized as follows: 54
mammals, 87 birds, 15 reptiles, 14 amphibians, 2 fishes, 350 insects and 27 other
invertebrate species, totaling 544 different prey species. In Supplementary Appendix
A we offer a complete list of prey species, as found in the literature.
Before legal protection was extended to birds of prey and owls in European countries,
analysis of the diet was done only to classify the Little Owl as a “useful” species (Mercier
1921, Farsky 1928, Sosnikhina 1950, Blagosklonov 1968, Barabash-Nikiforov and
Semago 1963). In Britain, after its introduction, a big debate took place about the extent
of Little Owl predation on game and poultry, as individual owls occasionally feed on
chicks (Hibbert-Ware 1938b).
Through the diet of the Little Owl, we can study the impact of changes in agricul-
tural techniques. Some prey species are linked to the diversity of habitat and are
sensitive to biocides. Before the 1950s, many authors stressed the importance of
cockchafers in the May, June and July diet of the owl, when this extremely abundant
beetle was fed to the young (Farsky 1928, Madon 1933, Hibbert-Ware 1938). The young
in the cavities could be found standing on a thick layer of cockchafer remains
(d’Hamonville 1895). Nowadays, cockchafers have become very rare and therefore do
not appear frequently in the diet of the Little Owl. This situation is similar for crickets,
that, with the owls, have disappeared from very intensively used landscapes in
Switzerland (Juillard 1984).
In the Netherlands, on the sandy soils in the eastern and southern parts of the
country, during the last decade, cockchafers have made a comeback, especially in
regions where there are many oaks. In some years, thousands of cockchafers can be
found in just one farmyard. Camera observations revealed that hundreds of them are
fed to the nestlings, as many as 1389 in 2019 and 1459 in 2020, during the incubation
and nestling period (Van Harxen 2021).
The higher the diversity of the habitat, the wider the trophic niche of the owl. In
very intensively managed agricultural landscapes, habitat diversity is low and very
often the Little Owl is more “specialized,” as it is limited to one kind of prey, such as
earthworms (Juillard 1984, Blache 2001), which must be taken in large quantities.
However, the problem is that earthworms contain less energy value, with 3.01 kilocal-
ories per gram and is the poorest in fat, with 1.49% (versus 19.75% for Gryllotalpa
gryllotalpa) and is less than 40% protein (Juillard 1984). By comparison, a Wood Mouse
is 70% protein and is richer in mineral salts. While earthworms can be abundant,
especially after rain, their low weight and their weak energetic yield is not as beneficial
as a rodent for feeding nestlings. Thus, the presence of earthworms in the diet may well
be more an expression of poor prey diversity in very intensively managed agricultural
landscapes rather than the adaptative capacity of the Little Owl in foraging. Moreover,
earthworms contain water, and when adults feed the nestlings only that kind of prey,
the nest becomes very wet and dirty (Luder and Stange 2001). Feeding earthworms also
causes strong gaseous ammonia, which has a negative impact on the climatic

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 7.6 Seasonal Variation

conditions in the nest (Van Harxen 2021). In Flanders, there are indications that the
abundance of the species is related to parameters that have a positive impact on
earthworms (Van Nieuwenhuyse and Leysen 2004).

7.5 Large Prey

The Little Owl is capable of catching prey that weighs as much as the owl itself
(Ryabitsev 2001). It catches large prey because of its compactness and strength, and
in particular its leg muscles seem unusually powerful. Its thighs are thick, and the
bundles of flexors in the toes yield extra power. The flexor of the metatarsus is strongly
developed, and serves to pull and hold the prey (Shtegman 1960).
Although the Little Owl is small, it is an energetic and very skillful hunter. Its flight is
low to the ground and swift, and the owl uses elements of topographic relief to attack
suddenly. These are some of the reasons why its prey can be relatively large. The owl,
however, prefers to hunt small mammals and Zerunian et al. (1982) consider that it is
limited by the size and the activity of its prey. When the prey is large, it catches nearly
exclusively young individuals (91.6% young Siesel (Citellus citellus), 8.4% adult; n = 71).
When prey is small, the owl successfully catches both young and adult specimens
(23.4% young Midday Gerbil (Meriones meridianus), 76.6 adults; n = 128)
(Osmolovskaya 1953). Finally, the species has a wide trophic niche ranging from ants
(0.01 g), up to rats or young rabbits and hares. Van Harxen and Stroeken (personal
communication) found two headless juvenile hares (Lepus europaeus) in a nestbox of
which the biggest weighed 89.5g. The daily food requirement for a Little Owl is 59–75 g
at 0 C and 23–30 g at 30 C, and averages 46.7 g for the whole year (Manez 1983b).

7.6 Seasonal Variation

In many cases, the diet shows seasonal variations due to the climate, but there is no
universal trend and patterns vary according to the studies. In most studies, however,
invertebrates are important throughout the year. Since camera-aided research has shown
to give a clearer view of soft prey these results should be interpreted with caution.
In Britain, the seasonal percentage by number of main prey shows the following
trend: rodents represent between 30% and 50% of the diet in winter but between 25%
and 30% during the other months, birds account for 8–10% in winter but under 5% in
the other months, and insects comprise around 30% every season with a maximum of
34% in spring (Collinge 1922). Hibbert-Ware (1938) confirmed the importance of
insects in the diet at all times of the year, with the owls catching the dominant species
according to their seasonal abundance, i.e., weevils (Curculionoidea) in spring,
cockchafers (Melolonthidae) in summer, earwigs (Dermaptera) and dor beetles
(Geotrupidae) in autumn, and even rove beetles (Staphylinidae) in winter.
In northeastern France, insects are numerically represented more in spring (79.8%),
summer (75.1%) and autumn (75.9%) than in winter (28.5%), while rodents follow the

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 Chapter 7: Diet

opposite trend: 15.9% in summer and 70.4% in winter (Génot and Bersuder 1995).
However, in southern France, invertebrates (number of prey and biomass) are numer-
ous year-round, but mammals remain the primary food source by biomass (Génot
1992c).
In Germany, beetles remain important numerically in the diet (73%) all year (Kuhn
1992). In Austria, the number of insects in the diet reaches its highest percentage (67%)
in summer, but remains low in biomass (2%), and even though the proportion of
mammals is reduced in summer, their role in the diet, in terms of biomass, remains
high all year (Sageder 1990).
In Switzerland, the seasonal variation of prey composition is not significant, except
for an increase of Cantharid larvae during winter (Schmid 2003).
In southern Italy, Coleoptera are present year-round but are more important in
biomass in autumn (27.9%). Ground and rove beetles are mainly represented in spring
and winter, respectively. Scarabidae and longhorn beetles (Cerambycidae) are found in
summer, while histerids (Histeridae) are only recorded in autumn. Mammals and birds
represent the main diet by biomass in spring; mammals, birds and reptiles in summer,
and birds, mammals and Coleoptera in winter. Given the numbers of prey in the winter
diet, earwigs are rather strongly represented (22.3%) (Moschetti and Mancini 1993).
In central Italy, the main seasonal difference in the diet is reflected by changes in the
amount of birds, scarabid beetles (Scarabaeidae) and earwigs (Dermaptera). In summer,
birds are more important (31.9% of the biomass) as opposed to 10–17% for other
seasons. Earwigs are only found in spring and are scarce in autumn–winter, and
practically absent in summer. A similar situation exists in the seasonality of scarabid
beetles. Finally, vertebrates represent 8.66% of prey numbers and 80% of biomass in
summer, and 2.74% of prey numbers and 60.2% of biomass in autumn–winter
(Zerunian et al. 1982).
In Spain, even if rodents represent the main part of the diet in spring, arthropods are
important all year long (Manez 1983a).
In the Crimea, the relative proportions of rodents and beetles are 40% and 39% in
April, 38% and 53% in May, 54% and 34% in June, and 11% and 48% in September,
respectively (Kostin 1983).
In Turkmenistan, rodents are most significant in winter (91% by number of prey
items) and then decline in spring (65%) and summer (56%), while birds, reptiles and
insects, mainly beetles, increase (Sukhinin et al. 1972). In the same country, the diet
can change during winter and in different years, with invertebrates representing 26.8%
by number in 1959 and 41.1% in 1960 (Bel’skaya 1992).
In Uzbekistan, rodents dominate the diet almost all year, and especially in winter.
During the other seasons, birds, reptiles and insects comprise the remainder of the food
spectrum (Allanazarova 1988).
Finally, the seasonal change in the diet depends on the density and availability of
certain prey. The pattern is clear that rodents are the base by biomass everywhere,
but depending on the region, insects, mainly beetles, grasshoppers, and earwigs,
increase in importance during the summer nestling period and in autumn by number
and biomass.

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 7.7 Geographical Variation

7.7 Geographical Variation

Variations in the diet of the Little Owl are linked to overarching localized modifications
in habitat conditions. For example, in southern Portugal, R. Tomé (personal communi-
cation) found more invertebrates (mainly beetles, in particular Scarabid) in steppe areas
(71.4% by biomass), and more rodents in Holm-oak woodland areas (58.8% by bio-
mass). In central Italy, the owls’ diet was studied at five sites, with insects comprising
91.1% of the prey items by number; Staphylinidae and Tenebrionidae were mostly taken
in urban sites while Scarabaeidae were caught at a rural site (Fattorini et al. 2000).
Manganaro et al. (2001) compared the percentage of prey numbers in the diet of owls
at two sites (farmland and urban) in Rome. They found more earwigs, grasshoppers and
beetles, and fewer reptiles, birds and small mammals at the farmland site compared to
the urban site (Manganaro et al. 2001).
In Spain, the owls’ diet was analyzed at 11 sites located in three main climatic zones:
wet climate in the north, “cold” Mediterranean in central Spain, and Mediterranean.
The diet was based on invertebrates and locally on reptiles in the Mediterranean area,
on small mammals in the wet area and on a mixture between both in the cold
Mediterranean area, where small mammals always reflected the larger part of the
biomass, but not as much as in the wet area (Manez 1983a).
In Kazakhstan, a comparison of the Little Owl’s diet was made between desert and
semi-desert environments. More Carabidae, Cerambycidae, Tenebrionidae and
Buprestidae, and fewer Scarabaeidae were found in the desert. This is related to the
occurrence of these beetles in steppe and sand habitats. For example, Little Owls caught
a lot of Buprestidae, in particular Julodis variolaris, in the steppe, with bushes of Tamarix
sp. because the larvae of these beetles develop in the roots of these shrubs (Formozov
and Osmolovskaya 1953).
In the Danube Delta in Romania, Pocora et al. (2012) found a high proportion of the
Harvest Mouse (Micromys minutus), in 103 pellets collected from April to June 2009,
representing 42% of vertebrate prey and 3% of all prey items. The major part of the
items found were insects (92.7%) of which Coleoptera species formed the most import-
ant order (71.9%), ground beetles (Carabidae) (22.3%) the most important family, and
Mole Crickets (Gryllotalpa sp.) (15.2%) the most abundant species.
On two locations in Israel (Charter et al. 2006), mammals were found in 93.2% and
73.4% of pellets, respectively, and accounted for 21.0% and 12.8% of prey by number.
Typical species for the region, such as the Lesser White-toothed Shrew (Crocidura
suaveolens), Etruscan Shrew (Suncus etruscus), Kuhl’s Pipistrelle (Pipistrellus kuhlii),
Social Vole (Microtus socialis) and Tristram’s Jird (Meriones tristrami) could be identified.
Chenchouni (2014) found, in 48 pellets collected in northeast Algeria, remains of
471 prey items from 38 different species. Although insects accounted for 90% of the
prey items, they contributed only 8% to the biomass. However, insects are a rich source
of energy especially in arid regions and a very important component of micronutrients
(Ghenam and Si Bachir 2011, Sekour et al. 2011).
Zarei et al. (2021) found in Iran that 63% of 22 pellets that were examined contained
remains of rodents, 11% reptiles and only 7% contained remains of insects. This could

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 Chapter 7: Diet

partially be due to the fact that only 36% of the pellets were collected in spring
and summer.
Zhao et al. (2011) examined 644 pellets collected from November to April (non-
breeding season) in the desert in northwestern China, and found that mammals
accounted for 55.7% by number and 95% by biomass. Insects (beetles) accounted for
42% by number and 2.3% by biomass, consisting of species that were active both in
daylight and at night.
In an arid habitat in northern Qatar, Abu Baker and Yamaguchi (2017) examined 87
pellets collected in April–May in 2014 and 2015, containing 193 prey items. They found
that beetles and rodents were the most common items, respectively 45% and 37% by
numbers. Of all pellets, 70% contained remains of rodents and 58.6% of beetles. Of
71 rodents 43 could be identified, including Sundevall’s Jird (Meriones crassus) as the
most common (20) and the Pygmy Gerbil (Gerbillus henleyi), a new species for Qatar.
Other species found were the Lesser Jerboa (Jaculus Jaculus), Sundevall’s Gerbil (Gerbillus
crassus) and the Balochistan or Dwarf Gerbil (Gerbillus nanus). Eight Baluch Rock Geckos
(Bunopus tuberculatus) and five Egyptian Spiny-Tailed Lizards (Uromastyx aegyptia)
showed that Little Owls do not only hunt by night, but even in the heat of the day
in Qatar (up to 50 C).
The geographic variation of the diet shows that the species is a generalist from
the trophic point of view. The Little Owl can occupy three to four main trophic
niches: small mammals in central Europe, insects in the Mediterranean basin, both
kinds of prey in the middle regions, and small mammals, insects and reptiles in
arid regions.
The Little Owl is a nonselective predator, catching prey groups according to their
relative abundance and biomass, depending on the region, season, local habitat condi-
tions and occasion. Studies show that the Little Owl is more of a generalist than the
Barn Owl (Tyto alba), globally recognized as a small-mammal specialist (Gotta and
Pigozzi 1997, Manganaro et al. 2001). Delibes et al. (1983) consider the Little Owl to
be more of a generalist than the Long-eared Owl (Asio otus) in Spain, while Romanowski
(1988) showed that in a suburban area of Poland, the Little Owl and the Long-eared
Owl share the same trophic niche breadth with a high degree of specialization. In this
study, Romanowski suggested that the actual contribution of insects to the Little Owl’s
diet was generally over-estimated. In such cases, competition could happen between
both owls, but the generalist nature of the Little Owls’ trophic niche generally avoided
an overlap with other predators. There may be exceptions in particular situations, such
as in Uzbekistan where competition exists between the Little Owl and snakes (Blunt-
nosed Viper Vipera lebetina and Saw-scaled Viper Echis carinatus) (Ishunin 1965). Van
Harxen and Stroeken (2021) found that Little Owls are capable of exploiting certain
types of prey when suddenly abundant, In particular, cockchafers can be delivered to
the nest in great numbers in a short period of time. For instance, on May 6,
2015 between 22:00 and 22:48 hours, 43 cockchafers were taken to nestlings, almost
one per minute, in co-operative action by the male (n = 16 ) and the female (n = 27).
The same applies to earthworms after rainfall, or even when the lawn is sprayed
overnight in times of drought.

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 7.8 Different Methods

7.8 Different Methods

Most studies on the diet of Little Owls rely on counting prey remains found in pellets.
The main limitation of this is that prey that is digested in its entirety is not taken into
account. Van Harxen and Stroeken (personal communication) showed that soft prey
such as larvae, caterpillars and earthworms (which leave no detectable remains in
pellets) make up more than half of the prey supply in the breeding and nestling phase.
Research on pellets alone thus yields an incomplete picture of the actual consumption
in that period. To improve comparability we present studies by the type of data collec-
tion used; we zoom in on pellets, nestbox litter and nestbox prey delivery from
camera observations.

7.8.1 Pellets
Pellets are pale gray in color when they contain rodent remains (Thomas 1939), and
black (Hibbert-Ware 1938) or brick-red when their content is only insects (Festetics
1959) (Figure 7.4). The shiny parts of beetles are clearly visible on the outside edge of
the pellets (Thomas 1939).
They are usually rounded at the ends, though occasionally one tip is like a thread
(Hibbert-Ware 1938). They look like the pellets of the Kestrel (Falco tinnunculus), but
those of the Kestrel are narrower, usually harder and more compact, never contain sand
from earthworms (Vendrig and Groen 2001), and consist mainly of fur from mammals.
Captive Little Owls evacuate one or two small pellets daily, about 11.5 hours after
each meal, with the pellet produced when digestion is still incomplete (Hanson 1973).
In nature, the pellets are evacuated after the night’s feeding and before daytime
roosting (Hibbert-Ware 1938). The time between eating prey and producing pellets

Figure 7.4 Little Owl pellet.

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 Chapter 7: Diet

depends on the season and the type of prey (Schönn et al. 1991). Thus, a pellet can be
produced four hours after catching grasshoppers and 12 hours for vertebrates (Angyal
and Konopka 1975), or, in another case, 5.5 hours after the owl ate a mouse and 7.5
hours after eating a bird (Oles 1961). An incubating female observed by cameras
evacuated one pellet each day. Sometimes she dropped the pellet between the eggs,
taking it away later (Geert Sterringa personal communication).
Normally, Little Owls spread pellets randomly in the field, except in winter when
it is possible to collect up to 40 pellets in the same place because the species often
uses the same roost (Génot and Bersuder 1995), and sometimes up to hundreds in
barns where cavities are occupied for decades (Van Harxen and Stroeken personal
communication).
A pellet can measure up to 6.5  1.6 cm, but most of them are (much) smaller. In
Austria, Sageder (1990) found an average pellet volume of 5 cm³ (0.9–20.1 cm3; n = 433)
in the Seewinkel region, with grassland and more prey diversity, an average pellet
volume of 5.91 cm3 (1.8–14.5 cm3; n = 165). Hámori et al. (2019) found an average
volume of 5.62 cm3 for 218 undamaged pellets collected from 20 nestboxes at three
different sites in Central Hungary.
The number of prey items per pellet have been reported as 1.2 in Bulgaria (Simeonov
1968) and 10.5 (min 1 to max 60) in Italy (Moschetti and Mancini 1993). In France,
Génot and Bersuder (1995) found in 234 winter pellets: no prey (n = 12), one prey (n =
102), two prey (n = 109) and three prey items (n = 3). In Moldavia, pellets contained an
average of 1.93 prey (0.87 insects and 1.06 vertebrates; n = 634), with the following
seasonal composition: 1.59 prey (0.39 insects and 1.19 vertebrates) in autumn–winter
and 2.17 prey (1.19 insects and 0.98 vertebrates) in spring–summer (Ganya and Zubkov
1975).
The content of pellets also gives insight into the foraging area of the Little Owl. In
this way, analysis of beetles caught by the owl showed that the species hunted along
forest edges and in very open woodlands (Kuhn 1995). Sometimes information about
climate conditions can be retrieved, as was shown in Central Hungary where, in 2015,
dry weather reduced the water-covered areas of the Upper Kiskunság Lakes and made it
necessary for Little Owls to shift from Common Spadefoots (Pelobates fuscus) – a
dominant species in nearby sites and in 2016 (Hámori et al. 2019) – to Common
Voles (Microtus arvalis).
Pellets consist mainly of fur and bones of rodents – vertebrae and ribs, skull frag-
ments, metacarpus and phalanges, femur, tibia, jaw bones with molars, humerus and
radius, bird bones – skull fragments, beaks, scapula, clavicles, tarsometatarsus, phal-
anges and claws, vertebrae and ribs (Hibbert-Ware 1938a), bones of frogs or snakes,
fragments of snail shells (Lancum 1925, Berghmans 2001d) and chitin remains of
insects. A German-based project reported insect remains in 352 pellets (Kuhn 1992)
in which 9893 insect body parts were recorded; 4027 (41%) were heads, 2329 (23%)
were left elytra, 2250 (23%) were right elytra and 1287 (13%) were thorax. The average
size of beetle parts recovered from the pellets was 1.2 cm, with a range of 0.2–3.5 cm
(Kuhn 1992). Earthworms can be identified in pellets thanks to their bristles, but a
detailed quantification is not possible. Consuming earthworms can also be uncovered

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 7.8 Different Methods

Figure 7.5 Geographical distribution of 87 pellet studies across Europa, Asia and northern Africa.

by the presence of sand, as most earthworms contain a certain amount (Onrust and
Piersma 2017). In two German study areas, 57% and 80% of 75 pellets contained
earthworms (Koop 1996).
Pellets can also contain grass as vegetal fibers or seeds of weeds (Collinge 1922,
Hibbert-Ware 1938, Quadrelli 1985), soil (Hounsome et al. 2004), stones (Hulten
1955, Sagitov 1990, Bel’skaya 1992), cherry stones (Misonne 1948) or eggshell
(Sagitov 1990). Eggshells may stem from hatched but also unhatched eggs.
Sometimes females eat (parts of ) of unhatched eggs or feed them to their juveniles, as
observed by cameras (Van Harxen and Stroeken personal communication). Grass is not
eaten as food, but facilitates the formation of pellets with the chitinous parts of insects
(Madon 1933, Libois 1977, Quadrelli 1985), but may be taken coincidently when
hunting in grasslands. The same explanation may be valid for the ingestion of stones,
as given by Thiollay (1968).

7.8.2 Prey Remains in Pellets

In recent decades, many studies have been conducted on the diet of Little Owls, using
prey remains in pellets throughout Europe and, in recent years, China, Qatar, Israel,
Egypt, Syria and Iran) (Table S7.1; Figure 7.5).
After insects (on average, 69.6% of the number of prey found per study), mammals
form the most important group with 21.5%. Birds, amphibians, reptiles and unspeci-
fied invertebrates together make up less than 10%. Per study, the composition can
change from 100% mammalian in two study areas in China to almost 100% insects
in multiple other studies (Table 7.2). Overall, we see an increase in the proportion of
reptiles and insects and a decrease in the proportion of mammals with decreasing
latitude.

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 Chapter 7: Diet

Table 7.2 Average and frequency of different prey types in 87 pellet studies across
Europa, Asia and northern Africa

Average Frequency Mimimum Maximum

Mammals 21.2 100.0 0.2 100.0

Birds 3.8 94.3 0.0 40.0
Amphibians 0.6 27.6 0.0 18.3
Reptiles 2.5 65.5 0.0 28.4
Insects 69.6 97.7 0.0 99.4
Other invertebrates 2.0 57.5 0.0 26.3

The composition of pellets depends not only on the location, year and season, but
also on the life cycle of prey itself. Hibbert-Ware (1938a) examined 2460 pellets and
found large numbers of insects: earwigs (Forficulidae), ground beetles (Carabidae), rove
beetles (Staphylinidae), weevils (Curcilonidae), click beetles (Elateridae), scarab beetles
(Scarabaeidae) and earth boring beetles (Geotrupes). However, Hibbert-Ware (1938a)
did not include cockchafers (Melolonthidae), although they are amongst the most
abundant insects eaten by the Little Owl, appearing in small numbers in May, being
extremely abundant in June and July and disappearing in August, because their season
is short, making it less important throughout the year.
The specific type of prey is very dependent on the region and habitat conditions. In
Western and Central Europe it is mainly Common Voles (Microtus arvalis) and Wood
Mice (Apodemus sylvaticus) when it comes to mammals, in China, Shao and Liu (2008)
found Midday Gerbil (Meriones meridianus), Tamarisk Gerbil (Meriones tamariscinus),
Striped Hamster (Cricetulus barabensis), Gray Hamster (Cricetulus migratorius), Desert
Hamster (Phodopus roborovskii), Great Gerbil (Rhombomys opimus), Northern Three-toed
Jerboa (Dipus sagitta), Mongolian Five-toed Jerboa (Allactaga sibirica), Hotson’s Jerboa
(Allactaga bullata), Long-eared Jerboa (Euchoreutes naso) and Kozlov’s Pygmy Jerboa
(Salpingotus kozlovi). In Qatar, Abu Baker and Yamaguchi (2017) found Egyptian Gerbil
(Jaculus jaculus), Sundevall’s Jird (Meriones crassus), Wagner’s Gerbil (Gerbillus dasyurus),
Pygmy Gerbil (Gerbillus henleyi) and Dwarf Gerbil (Gerbillus nanus). The discovery of the
remains of a Pygmy Gerbil was the first sighting of the species in Qatar. The same applies
to the other prey groups, where the reptiles also provide local flavor with Arabian Desert
Gecko (Bunopus tuberculatus) and Egyptian Mastigure (Urosmastyx aegyptia) in Qatar (Abu
Baker and Yamaguchi 2017), Przewalski’s Toadhead Agama (Phrynocephalus przewalskii) in
China (Shao and Liu 2008) and Black-headed Ground Snake (Rhynchocalamus melanoce-
phalus), Mediterranean House Gecko (Hemidactylus turcicus) and Ocellated Skink
(Chalcides ocellatus) in Israel (Charter et al. 2006).

7.8.3 Prey Remains in Nestbox Litter

In the period 1998–2021, Van Harxen and Stroeken (personal communication) counted
every vertebrate prey they found in the nest during the incubation and nestling period

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 7.8 Different Methods

resulting in a total of 4959 prey items. Mice and voles made up more than 85% of the
prey, followed at a considerable distance by birds, shrews and amphibians (Figures 7.6
and 7.7). Other prey types did not play a significant role (Figures 7.8 and 7.9). Within
the mice, it was mainly the Wood Mouse (Apodemus sylvaticus) (84.9% of the mice and
35.2% of the total number of prey) and within the voles it was mainly the Common
Vole (Microtus arvalis) (76.3% of the voles and 33.8% of all prey) that occurred. Wood
Mice and Common Voles accounted for more than two-thirds of all prey remains.
Both the absolute number of prey remains per year (adjusted for the number of
nestboxes visited and the time of visit) and the relative share of each prey group varied
significantly from year to year. Rodents make the difference at the population level.
Van Harxen and Stroeken (2009, 2011b) found in their analysis of 24 years (1998–2021)
of research on larders during nestbox controls a clear relationship between the cor-
rected number of rodents per year (micromammals corrected for nests visited and age of
nestlings (<11 days)) and reproduction parameters such as clutch size and the stand-
ardized condition of nestlings (the condition index).
In particular, it is the mice and voles that determined the differences between the
years and the height of the corrected number of micromammals.
Prey remains show variation between the years, the visiting time, the age of the
young, the number of visits in the nestling period, the weather conditions and the
mowing regime, making it difficult to compare territories at a detailed level. Hence,
across territories, and even in the same territory, large differences can be observed in the
number of mice and voles found. Taking into account the number of years that a nest
was visited in the nestling period with young up to 10 days old there is a large variation
in frequency (in how many years prey was found) and the average number of prey per
year (Table 7.3), e.g., in the number of nests that were monitored for nine years, the
extremes were a nest in which prey was found in all nine years (n = 95 items) with an
average number of 10.6 prey items (minimum = 1, maximum = 35) and a nest in which
prey was found in only four of the nine years (n = 14 items) with an average number of
1.6 prey per year (minimum = 0 and maximum = 10).
The territories also differ greatly from each other when it comes to the diversity in
prey species. The highest diversity was found in a nestbox where 16 different species
were found in the nestbox over the years: eight mammals, seven birds and one
amphibian (Van Harxen and Stroeken personal communication).
Sometimes changes in the local micromammal fauna can be illustrated by means of
prey remains in nestboxes, for example, the area expansion of the Yellow-necked
Mouse (Apodemus flavicollis) in the southeast Achterhoek. Until the 1990s, the species
only occurred in a few forests in Zuid-Limburg in the Netherlands. In the following
years, a steady expansion followed in various places along the eastern border with
Germany (Broekhuizen et al. 2006). In 2005, the species was first found in
Winterswijk (Regelink and Bosch 2007) 1.5 km from the German border. In 2013, the
first Yellow-necked Mouse was found in the nestbox of a Little Owl, 7 km away from the
2005 location and 6 km from the German border. Since then, Yellow-necked Mice have
been found at 16 locations, the furthest in 2021 at 14 km from the German
eastern border.

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 Chapter 7: Diet

Figure 7.6 Little Owl larder with intact prey.

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 7.8 Different Methods

Figure 7.7 Little Owl larder with decapitated prey.

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 Chapter 7: Diet

0.08 % Lagomorphs
0.20 % Bats 0.60 % Moles
2.54 % Amphibians
3.63 % Shrews
7.10 % Birds

44.30 % Voles

41.54 % Mice

Figure 7.8 The relative shares of each prey group in the prey remains found (n = 4959) (Van Harxen
and Stroeken personal communication).

16
Number of prey remains corrected nests visited and time of visit

14

12

10

0
1998

1999

2000

2001

2002

2003

2004

2005

2006

2007

2008

2009

2010

2011

2012

2013

2014

2015

2016

2017

2018

2019

2020

2021

Year
Mice Voles Shrews Bats Moles Lagomorphs Birds Amphibians

Figure 7.9 The number of prey remains per prey animal group 1998–2021 corrected for the number
of nests visited and the time of visit (Van Harxen and Stroeken personal communication).

7.8.4 Nest Camera-Observed Breeding Seasons

Thanks to recent technical developments and fast internet it is nowadays much easier
to get a good view of the prey supply to the incubating female and the nestlings than
ever before.
This section presents some data and analysis of a long-term study in the Netherlands
(Winterswijk) during 17 camera-observed breeding seasons from 2002 until 2021 across

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Table 7.3 Variability in number of micromammal prey remains based on number of years visited in the first 10 days of the nestling period

Maximum frequency Minimum frequency

Years Years
with with
Years prey prey
visited n Location found Total Mean Min Max SD Location found Total Mean Min Max SD

2 29 Roerdink 2 24 12.0 6 18 8.5 De Vossevelder 0 0 0.0 0 0 0

3 22 Hammerman 3 49 16.3 5 32 14.0 Helmerdink 0 0 0.0 0 0 0
4 20 Kapee 4 9 2.3 1 4 1.5 Beenekussteeg 1 1 0.3 1 1 0.5
5 19 Steengroeveweg 5 58 11.6 2 32 12.3 Brandt 2 2 0.4 1 1 0.5
6 7 Walien 6 56 9.3 4 17 4.6 Kloosterdijk 2 16 2.7 5 11 4.6
7 8 Vondermansdijk 7 62 8.9 1 25 8.6 Haflingerhoeve 2 10 1.4 2 8 3.0
8 7 Piepersweg 8 74 9.3 2 30 9.1 Tankink 3 9 1.1 3 3 1.6
9 6 Wessels 9 95 10.6 1 38 12.2 De Grutter 4 14 1.6 1 10 3.2
10 4 Bergstraat 8 96 9.6 1 35 11.9 te Brake 6 76 7.6 2 20 9.3
11 5 Heelweg 10 86 7.8 1 23 10.3 Lindeboomweg 7 49 4.5 1 35 10.3
Vragender
12 6 Schoppenweg 11 104 8.7 1 24 7.3 Corleseweg 7 68 5.7 3 21 7.7
14 3 Olivier 13 104 7.4 1 41 10.3 Klein Gebbink 11 45 3.2 1 10 3.3
15 3 Schurink 12 40 2.7 1 8 2.6 Stroette 9 38 2.5 1 14 3.8
17 1 Overkamp 15 103 6.1 1 36 9.6

Van Harxen and Stroeken personal communication

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the egg laying/incubation and nestling period (Van Harxen and Stroeken personal
communication), with, respectively, 5123 and 35 528 incoming owl movements with
prey brought in.
The differences between the camera-observed breeding seasons are large, both in
terms of the number of prey and biomass with the most extreme ranging from
1149 prey items in Dg17 to 4579 in Hv09, and the biomass from 2618 g in Hv12 to
9617 g in Hx18 (Table 7.4).
To a large extent, these differences are caused by the number of young in the nest
(adjusted for mortality) and the number of days that the nest has been observed.
Nestlings multiplied by nest observation days yields the number of prey days. The
number of prey items per prey day varies from 11.6 in Hv07 to 34.8 in Hv08, and
across all observations an average of 25 per prey day. The average biomass per day per
young varies from 38.7 g in Dg17 to 81.8 g in Hv12. The average per prey day is 60 g.

7.8.4.1 EGG LAYING AND INCUBATION PERIOD

Prey Frequencies
During incubation, the female is almost entirely dependent on what is provided by the
male. With a few exceptions, all 4727 prey from 13 camera-observed breeding seasons
were delivered by the male (97.7%). Even if the female came in with prey, she probably
received it from the male, out of sight of the cameras.
Invertebrate prey such as larvae, caterpillars, cockchafers and earthworms made up
the majority of items; almost 93% of the total supply. Within these, larvae/caterpillars
and cockchafers accounted respectively for 43% and 40% of invertebrate prey. Between
the camera-observed breeding seasons, large differences in prey numbers are observed
from 135 items in camera-observed breeding season Dg17 to 639 in Hx16, i.e., fivefold
(Table 7.5).
Prey frequencies show a significant linear relation with the clutch start (r = 0.6354,
n = 13, p <0.05), earlier in the year and fewer prey items. To a large extent, this is
because cockchafers, in terms of numbers, are an important prey for the investigated
locations Hv and Hx, usually appearing in the second half of April. Early breeders that
start at the end of March/beginning of April (hatching at the end of April/beginning of
May) then miss the increasing numbers up to the hatching of the first young
(Figures 7.10 and 7.11).
The difference is most extreme when comparing camera-observed breeding season
Hx15 (clutch start April 25) with Hx21 (clutch start March 25), 315 and 4 cockchafers,
respectively, but the pattern is also highly significant over all years (r = 0.7092, n = 12,
p <0.01).
A similar effect occurs in larvae and caterpillars (Figure 7.12). By laying very early
(late March/early April), females miss the high prey numbers later in the season. This
seems to be a downside of early breeding. Caterpillars advance in warm springs with the
earlier sprouting of trees and shrubs. Whether this also applies to larvae is not known,
but in any case there is no significant pattern in the number of caterpillars or larvae. For
incubating females, caterpillars and larvae make up a smaller part of the prey supply
than for nestlings (Van Harxen and Stroeken personal communication).

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Table 7.4 Prey supply for 17 camera-observed breeding seasons (2002–2021)

Prey items and biomass

Laying and incubation

Reproduction phase Nestling phase

First Hatching Clutch Young Obs. Obs.

Year Location egg date size Hatching Unhatched died Fledged days Freq. Biomass days Freq. Biomass

2002 Hw02 Apr 11 May 15 6 5 1 0 5 34 92 655 15 1325 2904

2003 Hw03 Apr 15 May 15 4 4 0 0 4 29 59 462 21 1244 3656
2004a Hw04 May 06 Jun 05 4 3 1 0 3 30 94 368 23 1402 3893
2007 HV07 Apr 09 May 10 4 3 1 0 3 15 151 316 42 1432 5480
2008 Hv08 Apr 13 May 14 4 4 0 3 1 31 418 809 38 2991 6087
2009 Hv09 Apr 12 May 13 4 4 0 0 4 31 334 631 39 4579 9319
2010 Hv10 Apr 21 May 21 4 2 2 0 2 30 400 1221 40 2318 4558
2011 Hv11 Apr 18 May 20 4 2 2 0 2 32 552 1007 39 1516 5892
2012b Hv12 Apr 20 May 24 3 1 2 1 0 34 230 707 31 546 2618
2015 Hx15 Apr 25 May 26 5 4 1 0 3 31 621 1146 36 3600 8142
2016 Hx16 Apr 20 May 21 4 3 1 0 3 30 639 1130 34 2568 5430
2017c Hx17 Apr 14 May 16 4 3 1 3 0 32 260 1301
2018 Hx18 Apr 11 May 11 4 4 0 0 4 30 297 464 43 4488 9617
2019 H19 Mar 31 Apr 30 3 2 1 0 2 29 214 513 39 2681 4260
2020 Hx20 Mar 30 Apr 30 3 3 0 0 3 31 343 440 44 3689 7643
2021d Hx21 Mar 25 3 0 3 0 0 73 773 2010
2017 Dg17 Apr 01 May 02 3 3 0 0 3 31 135 1369 32 1149 3712

Average May 04 Jun 07 3.9 2.9 0.9 0.4 2.5 30 5123 13 393 34 35 528 83 206

Total 66 50 16 7 42
a
replacement cluctch, b juvenile dead on day 31, c female killed by a stone marten, d
293

eggs did not hatch.

Biomass in grams.
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Table 7.5 Prey frequency during incubation and in the breeding period for 13 camera-observed breeding seasons

Hv08 Hv09 Hv10 Hv11 Hv12 Dg17 Hx15 Hx16 Hx17 Hx18 Hx19 Hx20 Hx21 Total

First egg Apr 13 Apr 12 Apr 21 Apr 18 Apr 20 Apr 01 Apr 25 Apr 20 Apr 14 Apr 11 Mar 31 Mar 30 Mar 25

Smal prey item 48 17 23 15 4 21 61 46 21 11 23 24 7 321

Larva/caterpillar 235 90 42 258 97 40 127 313 60 151 128 211 182 1934
May beetle 108 208 230 233 100 352 219 73 121 40 94 4 1782
Moth 1 1 7 1 6 1 2 19
Earthworm 4 79 7 4 5 63 43 64 6 6 10 73 364
Salamander 6 1 7
Frog 4 3 13 20
Bird 2 1 1 3 1 8
Micromammal 3 11 2 2 3 2 1 24
spp.
Shrew spp. 1 1 2 1 3 1 1 10
Hamster 1 1
True mouse spp. 15 8 10 6 10 45 2 8 6 9 1 2 122
Vole spp. 6 3 11 5 12 11 4 14 30 2 4 1 12 115

Total 418 334 400 552 230 135 621 639 260 297 214 343 284 4727

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Number of cockchafers per day

0
1
2
3
4

0.5
1.5
2.5
3.5
4.5

25 March
27 March
29 March
31 March
2 April
4 April
6 April
8 April
10 April
12 April

first egg
14 April
16 April
18 April
20 April
22 April
24 April
26 April
28 April
30 April
2 May
4 May
6 May

8 May

(May 13) are indicated (Van Harxen and Stroeken personal communication).
10 May

12 May

Date
14 May
16 May
first young

18 May
20 May
22 May
24 May
26 May
28 May
30 May
1 June
3 June
5 June
7 June
9 June
11 June

13 June
15 June
17 June
19 June
21 June
23 June
25 June
27 June
29 June
Figure 7.10 Supply of cockchafers per day for 13 camera-observed breeding seasons (2008–2021). The mean clutch start (April 11) and average hatching date
 Chapter 7: Diet

Figure 7.11 Cockchafers in Little Owl larder.

The differences might be related to weather conditions too. The years 2018–2020
were characterized by pronounced dry springs. Earthworms appeared little above
ground in those years, with the result that the females were only given small numbers,
i.e., on average in those three years only 7, compared to almost 61 on average in the
four other years in the camera-observed breeding seasons in location Hx.

Prey Biomass
Diversity is even more important in total biomass, ranging from 1369 g in camera
observed breeding season Dg17 to 464 g in Hx18 (Table 7.6). There is no relationship
between the start of laying and total biomass. Differences between the years of camera-
observed breeding seasons at the Hx location (2015–2021) are mainly caused by the
number of earthworms (reference weight 5 g) and micromammals (reference weight
19 g). The higher number in 2017 is partly caused by the fact that the meadows adjacent
to the territory were mown at the end of the breeding period. In two days, 13 voles were
brought in. This accounts for almost half of the biomass in that year. But also during
the rest of the breeding period, a few more voles were brought in than before the
mowing peak. Other years lack such a peak, although 2016 shows a small peak on
May 1 and 2 (five voles). Striking is the small number of voles in the years 2018–2020

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297

Number of larvae/caterpillars per day

0
1
2
3
4
5
6

25 March
27 March
29 March
31 March
2 April
4 April
6 April
8 April
10 April
12 April

first egg
14 April
16 April
18 April
20 April
22 April
24 April
26 April
28 April
30 April
2 May
4 May
6 May
8 May
10 May
12 May

Date
14 May
16 May
first young hatched

hatching date (May 13) are indicated (Van Harxen and Stroeken personal communication).
18 May
20 May
22 May
24 May
26 May
28 May
30 May
1 June
3 June
5 June
7 June
9 June
11 June
13 June
15 June
17 June
19 June
21 June
23 June
25 June
27 June
Figure 7.12 Supply of larvae and caterpillars per day in 13 camera-observed breeding seasons (2008–2021). The mean clutch start (April 11) and average

29 June
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Table 7.6 Prey supply in biomass during incubation and in the breeding period at 13 camera-observed breeding seasons

Hv08 Hv09 Hv10 Hv11 Hv12 Dg17 Hx15 Hx16 Hx17 Hx18 Hx19 Hx20 Hx21 Total

First egg Apr 13 Apr 12 Apr 21 Apr 18 Apr 20 Apr 01 Apr 25 Apr 20 Apr 14 Apr 11 Mar 31 Mar 30 Mar 25

Smal prey item 36 16 17 13 2 20 52 41 19 10 19 18 7 268

Larva/ 235 90 42 258 97 40 127 313 60 151 128 211 182 1934
caterpillar
May beetle 108 208 230 233 100 352 219 73 121 40 94 4 1782
Moth 1 1 7 1 6 1 2 19
Earthworm 20 395 35 20 25 315 215 320 30 30 50 365 1820
Salamander 30 5 35
Frog 50 38 163 250
Bird 100 50 50 150 50 400
Micromammal 57 209 38 38 57 38 19 456
spp.
Shrew spp. 10 10 20 10 30 10 10 100
Hamster 25 25
True mouse 285 152 190 114 190 855 38 152 114 171 19 38 2318
spp.
Vole spec. 114 57 209 95 228 209 76 266 570 38 76 19 228 2185

Total 809 631 1221 1007 707 1369 1146 1130 1301 464 513 440 856 11 592

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 7.8 Different Methods

(two, four and one, respectively) probably caused by mowing in those years outside of
the breeding period (Van Harxen and Stroeken personal communication).

7.8.4.2 NESTLING PERIOD

Prey Frequencies
Nestlings are mainly fed with invertebrate prey such as larvae, caterpillars, earthworms
and cockchafers. Vertebrate prey accounts for only 4.4% of the supply, mainly mice and
voles (73.4%) (Table 7.7). Shrews make up only 2.2% of vertebrate prey (less than 0.01%
of the total supply). Birds and salamanders are also almost negligible with respectively
4.6% and 4.4% of vertebrate prey and 0.2% of the total supply. Frogs are fed more often,
i.e., 15.4% of vertebrate prey; at 0.7%, they account for a modest share of the
total supply.
At almost 39%, larvae and caterpillars make up the most important fraction of
invertebrate prey. Cockchafers make up just over 23% of the supply and earthworms
almost 14%. Together with a large group of unidentified small prey (almost 18%) they
form by far the largest part of the invertebrate prey in the picture (84%). This makes the
Little Owl, at least in the nestling phase, more of an insect and worm rather than a
micromammal hunter.
Almost no earthworms were fed in the first week (Figure 7.13). The few earthworms
that were delivered in this period were eaten by the female herself. At day 8, an
earthworm was fed to the nestlings for the first time (Van Harxen and Stroeken personal
communication).
Earthworms not only contain a relatively large amount of water, but also sand and
feces (Onrust and Piersma 2017), components that are not digested, but have to be
excreted in their entirety. Especially for small nestlings, this costs lots of energy. The
excreted water, in combination with the feces, also creates conditions for ammonia
development, creating an unfavorable living environment in the nest. In the first week
there are clearly less earthworms supplied, probably only when there are no
alternatives.
In Switzerland, Juillard (1984) examined prey supply to nestlings at 10 locations
using camera traps, with earthworms accounting for an average of 65.3% of the supply
(numbers) with peaks of 88.9% and even 95%. Many earthworms were also supplied in
the first week after hatching at almost all locations. As a result, the young were often
wet, dirty and in poor condition.

Biomass
In terms of biomass, a different pattern is observed, i.e., vertebrate prey represents from
4.4% by frequency to 35.8%, and invertebrates drop from 95.6% to 64.2%. Among
vertebrate prey, voles and mice account for 68.6% of the biomass of the vertebrate prey
and 24.6% of the biomass of all prey.
Within the invertebrate prey, it is mainly earthworms that provide the biomass, 29%
of the total biomass and just over 45% of invertebrate prey. Larvae and caterpillars make
up 16% of the total biomass and almost 25% of invertebrate prey. Cockchafers still
account for almost 10% of the biomass of all prey and just over 15% of invertebrates.

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Table 7.7 Prey frequency during the nestling period during 12 camera-observed breeding seasons

Hv07 Hv08 Hv09 Hv10 Hv11 Hv12 Dg17 Hx15 Hx16 Hx18 Hx19 Hx20 Total

Small prey item 237 188 766 462 192 16 278 1188 909 1153 249 706 6344
Moths 86 155 58 156 36 8 32 36 190 15 105 877
Larva / caterpillar 441 2199 2597 981 411 187 402 614 489 1402 931 1005 11659
May beetle 414 181 224 641 60 39 250 819 798 897 1389 1359 7071
Earthworm 124 218 712 74 560 184 89 885 228 747 17 382 4220
Frog 45 18 45 21 43 24 8 3 5 1 213
Salamander 6 16 4 17 2 1 5 7 2 60
Bird 1 9 6 10 6 1 1 4 2 16 4 4 64
Shrew 2 3 2 4 2 3 10 3 2 31
Micromammal spp. 30 5 6 4 1 8 2 1 3 2 4 66
Mouse spp. 43 44 25 24 17 15 81 7 21 24 22 5 328
Vole spp. 95 36 22 35 46 40 27 26 81 45 47 114 614
European water vole 1 2 2 3 1 1 10

Total 1432 2991 4579 2318 1516 546 1149 3600 2568 4488 2681 3689 31 557

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 7.8 Different Methods

300

250
Number of earthworms per day

200

150

100

50

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44
Days aer first hatching

Figure 7.13 Supply of earthworms per day in the nestling period in 12 camera-observed breeding
seasons (Van Harxen and Stroeken personal communication).

In the 12 camera-observed breeding seasons, larvae, caterpillars, cockchafers, earth-

worms and other small prey frequencies made up the largest proportion of prey sup-
plied to the nestlings, but based on the biomass it was mainly earthworms and mice
that contributed (together accounting for 56%), followed at a distance by larvae,
caterpillars and cockchafers (together almost 26%) (Van Harxen and Stroeken personal
communication) (Table 7.8).

Micromammals
Supply of mice and voles plays an important role when correcting for the number of
young and the number of observation days. The number of rodents is related to the
total number of the prey items supplied (r = 0.0651, p <0.05, n = 12) and the proportion
of biomass of the rodents (r = 0.7587, p <0.01, n = 12), hence, the more rodents that are
supplied, the less invertebrate prey is needed (Figure 7.14).
The complementarity of rodents and invertebrates in the diet is most extreme
between camera-observed breeding seasons Hx15 and Hv07. The contribution of
rodents to the biomass ranges from 9.4% in Hx15 to 58.6% in Hv07. Inversely propor-
tional to this is the number of other prey, such as larvae, caterpillars and other
small prey.

7.8.5 Other Camera-Observed Breeding Seasons

Throughout Europe
In recent decades, the prey supply in the incubation and nestling period has been
studied in various places in Europe with the help of photo or video images. This has

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Table 7.8 Prey supply in biomass in grams during the nestling period during 12 camera-observed breeding seasons

Dg17 Hv07 Hv08 Hv09 Hv10 Hv11 Hv12 Hx15 Hx16 Hx18 Hx19 Hx20 Total Pecentage

Small prey item 272.5 180 165.5 713 418.5 151 10.5 1161.5 879.5 1107 227 684.5 5970.5 8.2
Moths 8 86 155 58 156 36 32 36 190 15 105 877 1.2
Larva/caterpillar 402 441 2199 2597 981 411 187 614 489 1402 932 1005 11 660 16.0
May beetle 250 414 181 224 641 60 39 819 798 897 1389 1359 7071 9.7
Earthworm 445 620 1090 3560 370 2800 920 4425 1140 3735 85 1910 21 100 29.0
Frog 562.5 225 562.5 262.5 537.5 300 100 37.5 62.5 12.5 2662.5 3.7
Salamander 5 30 80 20 85 10 25 35 10 300 0.4
Bird 50 50 450 300 500 300 50 200 100 800 200 200 3200 4.4
Shrew 30 20 30 20 40 20 100 30 20 310 0.4
Mouse/vole spp. 152 570 95 114 76 19 38 19 57 38 76 1254 1.7
Mous spp. 1539 817 836 475 456 323 285 133 399 456 418 95 6232 8.6
Vole spp. 513 1805 684 418 665 874 760 494 1539 855 893 2166 11 666 16.0
European water 45 45 90 90 135 45 450 0.6
vole

3711.5 5479.5 6086.5 9318.5 4558 5891.5 2617.5 8141.5 5429.5 9616.5 4259.5 7643 72 753 100.0

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 7.8 Different Methods

resulted in a total of 30 published references that provide a detailed picture of what

incubating females and nestlings are fed (Table 7.9).
There were differences between registrations regarding the start and duration, but
most started at the beginning of incubation and continued until the nestlings were at
least 20 days old. Registrations took place in theory from 0:00 to 24:00, but due to
technical defects, occasionally a few hours were missing, in exceptional cases a few days.
Earthworms, larvae and caterpillars together make up almost 50% of the diet. Along
with beetles, spiders, grasshoppers, centipedes, snails, moths and unnamed insects,
invertebrate prey make up 94% of the prey count, and vertebrate prey, with mammals
as the main group, only 6%. Between locations and years we see large differences with
remarkably large numbers of earthworms in the Czech and especially the Swiss loca-
tions (up to 95%). In the Dutch locations, the proportion is a lot lower, at 15.6%, with
camera-observed breeding season GP00 as an outlier at 45.3%. The portion of inverte-
brate prey is filled with beetles (21.6%) and especially larvae and caterpillars (36.8%). In
the non-Dutch locations, the share is only 5.5% and 10.3%, respectively. Other

Table 7.9 Overview of camera studies

Period Locations 1 2 3 4 5 6 Total number

Czech Republic 2019 2 2 2

Belgium 2005 1 1 1
Switzerland 1979–1980 7 7 2 8
Netherlands 1983–2020 7 4 1 1 1 19
Total number 14 2 0 1 1 1 30
of years
Proporon of rodents in biomass, corrected for number of young

70

60
y = –0.0096x + 56.125
R² = 0.573
50
and observaon days

40

30

20

10

0
0 500 1000 1500 2000 2500 3000 3500 4000 4500 5000
Number of prey items

Figure 7.14 Linear relationship between prey frequency and the proportion of rodents in biomass,
corrected for the number of young and observation days (Van Harxen and Stroeken personal
communication).

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 Chapter 7: Diet

Table 7.10 Prey composition (%) in 30 camera registrations

Average SD Min Max Frequency

Earthworms 31.30 26.97 0.79 94.98 30

Larvae and caterpillars 27.08 20.17 0.21 72.07 30
Beetles 15.71 13.79 0.00 49.36 27
Unidentified insects 13.08 9.78 1.02 38.40 30
Moths 6.36 10.73 0.00 34.95 27
Mammals 4.73 3.54 0.21 14.64 30
Amphibians 0.74 1.35 0.00 5.53 19
Birds 0.43 0.63 0.00 3.39 29
Spiders 0.29 1.00 0.00 5.39 11
Grashoppers 0.13 0.63 0.00 3.52 3
Leech 0.08 0.40 0.00 2.24 3
Moles 0.04 0.19 0.00 1.07 1
Reptiles 0.02 0.09 0.00 0.41 2
Millipeds 0.02 0.05 0.00 0.19 3
Snails 0.01 0.04 0.00 0.20 1

vertebrate prey such as birds (0.43%), moles (0.02%), amphibians (0.74%) and reptiles
(0.02%) together account for less than 1.5% of the supply (Table 7.10).

7.8.6 Soft Prey

In order to get a good insight into the diet of Little Owls, pellet research might not be
sufficient, because important prey species cannot be demonstrated or insufficiently
quantified. Soft prey such as larvae and caterpillars leave no or barely recognizable
remains in pellets and for earthworms, although the chaetae made from chitin can be
found – Hounsome et al. (2004) found frequencies of 94% to 100% earthworms in
pellets. These are, however, hard to quantify (Romanowski et al. 2013). With eight
chaetae on each segment and between 100 and 150 segments per worm, the consump-
tion of just one worm yields many hundreds of chaetae. The assumption by Altringham
et al. (1994) that the presence of chaetae in a pellet is equivalent to the consumption of
one or more earthworms of at least 5 g (based on chaetae <0.5 mm) may be considered
rather arbitrary, but nevertheless deserves further investigation. It remains relevant to
gain more insight into the extent to which soft prey can be found in pellets and,
especially, quantified, because during the incubation and nestling period, larvae, cater-
pillars and earthworms in studies where the prey supply has been observed with
cameras (Van Harxen and Stroeken personal communication) make up a substantial
part (22%–95.2%) of the prey supply (Figure 7.15). However, it is conceivable that in
warm and dry areas in North Africa and the Middle East their share is less and that
analyzing pellets provides a more accurate picture of reality.

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 7.8 Different Methods

100

90
Percentage of undetectable, soft prey items in pellets

80
(larvae, caterpillars and earthworms)

70

60

50

40

30

20

10

Locations
Undetectable Detectable

Figure 7.15 Proportion of undetectable, soft prey items in pellets (larvae, caterpillars and
earthworms) in the prey supply in the incubation and nestling period in 30 studies where prey
supply was recorded with cameras (n prey = 41 140).

7.8.7 Voles and Mice

Even if the diet includes insects at all times of the year, voles, in particular the Common
Vole (Microtus arvalis), are very important to the Little Owl in central Europe. This has
been shown in eastern France, where voles represented 70% of the biomass from
8181 prey items recovered in pellets (Génot and Bersuder 1995), Poland (74% by
biomass; n = 288 prey items; Romanowski 1988; and 47.2% by biomass; n = 1953 prey
items; Grzywaczewski et al. 2006), Germany (with 76% of the number of mammal prey;
n = 1805 prey; Lack 1946), Switzerland (51% by biomass; n = 2557 prey items; Schmid
2003), Moldavia (with 51% of the number of vertebrates; Ganya and Zubkov 1975),
Romania (with 42% of the number of mammals; Popescu and Negrea 1987) and
Caucasia (with 85% of the content of 175 pellets; Afanasova and Khokhlov 1995).
Even in southern France, the Common Vole represents 6.5% of the numbers and
50% of the biomass (n = 2345 prey). Further, even if it is a new species in the region,
the Little Owl quickly incorporates small mammals into its diet (Blache 2001).
The difference in energetic yield of voles in relation to that of insects or earthworms
is well illustrated by a nest in Austria where a brood of three, 1-month-old nestlings
were fed only with insects up to 50 times per hour (Ille 1992). In Austria, Ille (1996)
observed the lowest reproductive rate in 1993 after a Common Vole population decline.
In Britain, Leigh (2001a) reported small mammal populations to be a key factor
affecting the breeding performance of Little Owls. In Germany, Illner (1991) showed
a correlation between the reproductive rate of owls and the percentage biomass of the
Common Vole (Microtus arvalis) and the Mole Cricket (Gryllotalpa gryllotalpa) in the

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 Chapter 7: Diet

owls’ diet. Koop (1996) found voles to be an important prey when the owls were laying
eggs and rearing young; in particular, their availability was important not only in the
meadows but also along habitat edges (hedges, waysides, field edges). According to
Romanowski (1988), the percentage of the Common Vole in the owls’ diet does not
depend on the yearly fluctuation of this vole. As with the Long-eared Owl (Asio otus),
when the Common Vole population is low, the Little Owl may be able to focus its
hunting effort on small, isolated colonies of this vole (Goszczynski 1981). In an effort to
avoid short-term fluctuation of tropic resources, this situation is supported by the
“super territory” foraging behavior shown by the Little Owl (Finck 1990).
The importance of micromammals is also evident from the long-term study of the
prey supply to nestlings by Van Harxen and Stroeken (personal communication) in
their study area in Winterswijk (the Netherlands).
Voles and mice, in particular Common Voles (Microtus arvalis) and Wood Mice
(Apodemus sylvaticus), are an important part of the diet there. Especially in the breeding
season, when there are young, micromammals are important as a staple food, not so
much in terms of numbers, but as biomass. Rodents, unlike soft prey such as larvae,
caterpillars and earthworms, can be kept in the prey supply and fed to the young by the
female throughout the day.
Camera-observed breeding seasons in which the prey supply is recorded 24 hours a
day during incubation and the nestling period provide a good picture of the importance
of rodents and also provide surprising insights.
As an example, on May 1, 2019 at observation Hx19, the young hatchlings were
1 day old. In the early morning and the second part of the evening, five prey items were
brought in (three caterpillars and two voles). Nestlings were fed 16 times throughout
the day, of which 11 (for 47 minutes) were from the larder. Two Common Voles and a
Wood Mouse had been brought in the night before (Table 7.11).
If there are enough rodents to feed the young, the female can use other prey that is
brought in during the day for her own consumption. Even the male can then afford
to eat.
On May 2, 2019, when the young were 2 days old, 23 prey items are brought in,
including three voles and one Wood Mouse. All were put in stock. Of the remaining
19 prey (1 larva, 5 caterpillars, 11 cockchafers, 1 earthworm and 1 small, unclear prey),
16 were eaten by the female, 2 consumed by the male and only 1 (caterpillar) was fed to the
young. The young were fed 15 times throughout the day, of which 12 were from the rodent
supply. Only once did the female herself take a bite of one of the mice (Table 7.12).
For adult owls, rodent prey is beneficial too, reducing the effort of making flight
movements to provide sufficient biomass for small prey. Figure 7.16 shows prey supply
per day during two camera-observed breeding seasons, Hv09, where only 55 micromam-
mals were supplied for the four young, 0.35 per nestling day, over 39 observation days,
and Dg17, with 116 rodents over 32 observation days for three young or an average of
1.2 per day per young, i.e., more than three times that for Hv09. The parents in Hv09
had to compensate for the low supply of micromammals with plenty of (small) prey,
resulting in no less than 4579 prey flights. The adult birds in Dg17 needed only 1149, or
a quarter of the number in Hv09 (Van Harxen and Stroeken personal communication).

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 7.8 Different Methods

Table 7.11 May 1, 2019: prey supply, and feeding frequency and duration from the
prey supply (mice), young 1 day old

Time Feeding Duration (mins) Action

03:43 Stock 4 Female feeds, remains stored in larder

05:17 Stock 5 Female feeds, remains stored in larder
05:46 Vole 6 Female feeds, remains stored in larder
07:53 Stock 5 Female feeds, remains stored in larder
08:25 Vole 2 Female feeds, remains stored in larder
08:37 Stock 5 Female feeds, remains stored in larder
09:52 Stock 1 Female feeds, remains stored in larder
12:18 Stock 6 Female feeds, remains stored in larder
12:43 Stock 2 Female feeds, remains stored in larder
14:16 Stock 3 Female feeds, remains stored in larder
16:40 Stock 6 Female feeds, remains stored in larder
17:42 Stock 3 Female feeds, remains stored in larder
20:07 Stock 7 Female feeds, remains stored in larder
22:00 Caterpillar 1 Female feeds
22:24 Caterpillar 1 Female feeds, eats the remains
23:08 Caterpillar 1 Female feeds, eats the remains

Table 7.12 May 2, 2019: prey supply, and feeding frequency and duration from the
prey supply (mice), young 2 days old

Time Feeding Duration (mins) Action

01:12 Stock 8 Female feeds nestlings

01:40 Earthworm Male hunts, female eats prey
01:49 Larva Male hunts, female eats prey
01:54 Caterpillar Male hunts, female eats prey
02:00 Caterpillar Male hunts, female eats prey
03:46 Stock 6 Female feeds nestlings
04:57 Caterpillar Male hunts, female eats prey
05:40 Vole 4 Male hunts, female feeds nestlings
05:58 Vole Male hunts, prey put in stock
06:42 Unclear Male hunts, female eats prey
07:15 Cockchafer Male hunts, female eats prey
07:18 Caterpillar Male hunts, female eats prey
08:15 Stock 7 Female feeds nestlings
08:55 Cockchafer Male hunts, female eats prey
08:56 Stock 2 Female feeds nestlings
09:56 Cockchafer Male hunts, female eats prey
10:27 Stock 10 Female feeds nestlings

continues
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 Chapter 7: Diet

Table 7.12 (cont.)

Time Feeding Duration (mins) Action

10:37 Cockchafer Male hunts, female eats prey

10:58 Cockchafer Male hunts, female eats prey
12:00 Stock 8 Female feeds nestlings
13:11 Stock 3 Female feeds nestlings
13:33 Stock 4 Female feeds nestlings
14:55 Cockchafer Male hunts, female eats prey
15:39 Cockchafer Male hunts and eats prey
15:44 Cockchafer Male hunts and eats prey
16:06 Stock 6 Female feeds nestlings
17:10 Cockchafer Male hunts, female eats prey
17:17 Cockchafer Male hunts, female eats prey
17:28 Cockchafer Male hunts, female eats prey
18:47 Stock 6 Female feeds nestlings
19:17 Caterpillar 1 Female feeds nestlings
19:24 Stock 5 Female feeds nestlings
20:31 Mouse 4 Female feeds nestlings
22:31 Stock 3 Female feeds nestlings
23:05 Vole Male hunts and eats prey

250

200
Number of prey items

150

100

50

0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39
Days as of the first egg
Hv09 Dg17

Figure 7.16 Number of prey per day in Hv09 and Dg17 (Van Harxen and Stroeken personal
communication).

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 7.8 Different Methods

Of the 32 observation days in Dg17, parents only had to fly up and down more than
50 times per day on 11 days (34%, on average 71 times). The parents in Hv09 had to
deliver prey more than 50 times per day in 32 out of 39 days (82%, on average 140
times). In fact on five days, they brought in more than 200 prey items, with the highest
number being 236 on day 13. On those days they entered the nestbox clearly tired at
the end of the evening; sometimes they barely managed to land on the approach board
without tumbling off again.
The much larger number of mice in Dg17 was mainly due to the fact that a lot of
Wood Mice were supplied (81 of the 108 micromammals brought in). In the territory (a
large garden) many branches and hay stacks were present that yielded an extremely
high density of Wood Mice. The nestlings in Hv09 had to be fed by the natural
population with 25 Wood Mice, 22 Common Voles, 2 voles and 6 nonspecified rodents
(Van Harxen and Stroeken personal communication).
In particular, the presence and catchability of Common Voles is very dependent on
the weather conditions and the mowing scheme. In Hx20, plenty of voles showed up
when the meadows around the nestbox were mown (Figure 7.17).
After the first mowing on May 5, 73 Common Voles were brought (7.3 per day) in a
timeframe of 10 days. In the 36 days before (from the first egg on March 30) there were
only 7 (0.2 per day). For the three juveniles, the voles came at the right time; in the
10 days between age 6 and 15, they each got an average of 2.4 voles per day. During the
31 days of the laying and breeding period, the female had to be satisfied with a total of
one vole and one Wood Mouse. Due to the mild winter and the warm spring, the grass
growth started earlier than in other years and the grass was already so high at the

18

16

14

12
Number of voles

10

8 mowing

4
first young
first egg hatched
2

0
30 March
1 April
3 April
5 April
7 April
9 April
11 April
13 April
15 April
17 April
19 April
21 April
23 April
25 April
27 April
29 April
1 May
3 May
5 May
7 May
9 May
11 May
13 May
15 May
17 May
19 May
21 May
23 May
25 May
27 May
29 May
31 May
2 June
4 June
6 June
8 June
10 June
12 June

Days as of the first egg

Figure 7.17 Supply of voles in camera-observed breeding season Hx20 from the first egg (Van Harxen
and Stroeken personal communication).

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 Chapter 7: Diet

14
Number of micromammals corrected for nests visited and

12

10
age of nestlings (<11 days)

0
1998

1999

2000

2001

2002

2003

2004

2005

2006

2007

2008

2009

2010

2011

2012

2013

2014

2015

2016

2017

2018

2019

2020

2021
Year

Figure 7.18 Micromammals corrected for nests visited and age of nestlings (<11 days) 1998–2021
(Van Harxen and Stroeken personal communication).

beginning of April that the voles had sufficient cover and were poorly available until it
was mown.
Rodents also make the difference at the population level. Van Harxen and Stroeken
(e.g., 2009, 2011b), in their analysis of 24 years (1998–2021) of research on larders
during nestbox controls, found a clear relationship between micromammals, corrected
for nests visited and age of nestlings (<11 days), and reproduction parameters such as
clutch size and the standardized condition of nestlings (the condition index).
The average number of micromammals (Wood Mice, Common Voles, House Mice
and Bank Voles being the four most important rodent species in the research area)
found in larders is corrected for the number of nests visited and the age of the nestlings
(<11 days) (Stroeken and Van Harxen 2021) (Figure 7.18).
The corrected number of micromammals varied from 1.00 in the extremely low
rodent year, 2004, to 12.44 in the absolute peak year, 2014, with remarkable bad years
(2009–2011). The last few years were above average, mainly because of Common Voles
and Wood Mice having alternating peaks. In 2014 it was mainly the Wood Mouse that
peaked and in 2019 the Common Vole. The two other species played an inferior role.
A positive significant quadratic relationship (y = –0.0071x2 + 0.1497x + 3.5442, R2 =
0.4688, p <0.01, n = 24) between clutch size and micromammal abundancy
(Figure 7.19) was observed.
A quadratic relationship was observed between the number of young per started nest
(Figure 7.20) and the rodent abundance (y = –0.0102x2 + 0.2194x + 1.5679, R2 = 0.5437,
p <0.01, n = 24), indicating more rodents meant more young.
A third parameter that was influenced by the number of rodents was the condition of
the nestlings, expressed as the percentage deviation of the measured weight on a certain
age day compared to the average (expected) weight on that day. If both are equal, the
condition index is 1.00. If the measured weight is higher than expected, the index

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 7.8 Different Methods

4.6

Average clutch size per started nest 4.4

4.2
y = –0.0071x2 + 0.1497x + 3.5442
4 R² = 0.4688

3.8

3.6

3.4
0 2 4 6 8 10 12 14
Number of micromammals corrected for nests
visited and age of nestlings (<11 days)

Figure 7.19 Quadratic relationship between micromammals corrected for nests visited and age of
nestlings (<11 days) (x-axis) and the average clutch size per year (y-axis) 1998–2021. (Van Harxen and
Stroeken personal communication).

2.9
Average young per started nest

2.7

2.5
y = –0.0102x2 + 0.2194x + 1.5679
2.3 R² = 0.5437

2.1

1.9

1.7

1.5
0 2 4 6 8 10 12 14
Number of micromammals corrected for nests visited and
age of nestlings (<11 days)
Figure 7.20 Quadratic relationship between micromammals corrected for nests visited and age of
nestlings (<11days) (x-axis) and the number of young per started nest (y-axis) 1998–2021 (Van Harxen
and Stroeken personal communication).

exceeds 1.00. If a nestling is lighter, the index is below 1.00. An index of 1.05 thus
means that the young is 5% heavier than would be expected on the basis of age, an
index of 0.95 that it is 5% lighter than expected. The positive quadratic relationship
between micromammals corrected for nests visited and age of nestlings (<11 days) and
the condition index showed a significant quadratic relationship (Figure 7.21) (y =
–0.001x2 + 0.0245x + 0.9373, R2 = 0.5877, p <0.01, n = 24): the more rodents, the
better condition the young have. Almost all the years in which the condition index
exceeded 1.0 were years in which the micromammals corrected for nests visited and age
of nestlings (<11 days) was 3 or more.

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 Chapter 7: Diet

1.1

Average condition nestlings 1.08

1.06

1.04 y = –0.001x2 + 0.0245x + 0.9373

R² = 0.5877
1.02

0.98

0.96

0.94
0 2 4 6 8 10 12 14
Number of micromammals corrected for nests visited
and age of nestlings (<11 days)
Figure 7.21 Quadratic relationship between micromammals corrected for nests visited and age of
nestlings (<11 days) (x-axis) and condition-index 1998–2021 (Van Harxen and Stroeken personal
communication).

The fact that all three reproduction parameters show a positive significant relation-
ship with the micromammals corrected for nests visited and age of nestlings (<11 days)
shows that rodents play an important role for Little Owls. The owls manage to raise
their young, even in years with few rodents, but yielding fewer offspring and in poorer
condition (Stroeken and Van Harxen 2021).

7.8.7.1 RELATIONSHIP BETWEEN COMMON VOLES AND WOOD MICE

In the prey remains research of Van Harxen and Stroeken (2021), Wood Mice (n = 1746)
and Common Voles (n = 1677) turned out to be by far the most important species
found in the larders. In larders in which at least three mice were identified, 84.4% of
them had Wood Mice and 78.5% Common Voles; both Wood Mice and Common
Voles were found in 75% of the territories. The latter is not only important because
there are two common species that contribute highly to the biomass in the prey supply
to the nestlings, but also because their peak years alternate.
Figure 7.22 illustrates the relative proportions from 1998 to 2021 of the numbers of
micromammals corrected for nests visited and age of nestlings (<11 days). In 2014, it
was mainly voles that made up the numbers, but in 2019 it was the Wood Mice.
Territories in which only one of the two species were available would certainly have
done worse in those years. Even in good micromammal years, like 2014 and 2019, we
see the benefits of this larder effect.
Alternating peaks happen not only between the years, but also at the seasonal level.
Summing the catches of voles and mice over 17 camera-observed breeding seasons at
four different locations and relating them against time, Common Voles show two
peaks, one around May 10 and the other around June 3 (Figure 7.23) coinciding with
the mowing of the grasslands in the vicinity of the nests.

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 7.8 Different Methods

7
Number of voles corrected for nests visited and age of

5
nestlings (<11 days)

0
1998

1999

2000

2001

2002

2003

2004

2005

2006

2007

2008

2009

2010

2011

2012

2013

2014

2015

2016

2017

2018

2019

2020

2021
Year
Common Vole Wood Mouse

Figure 7.22 Wood Mouse and Common Vole numbers corrected for nests visited and age of nestlings
(<11 days) per year for 1998–2021 (Van Harxen and Stroeken personal communication).

4.5

3.5
Percentage of voles as prey items

3 Average Hatching
Date
2.5

1.5

0.5

0
29 March
31 March
3 April
6 April
9 April
11 April
13 April
15 April
17 April
19 April
21 April
23 April
25 April
27 April
29 April
1 May
3 May
5 May
7 May
9 May
11 May
13 May
15 May
17 May
19 May

2 June
4 June
6 June
8 June
10 June
12 June
14 June
16 June
18 June
20 June
22 June
24 June
26 June
28 June
30 June
21 May
23 May
25 May
27 May
29 May
31 May
25 March
27 March

Date

Figure 7.23 Number of voles per day as a percentage of the total (n = 801), summed over 17 camera-
observed breeding seasons, with the average hatching date indicated in red (Van Harxen and Stroeken
personal communication).

Wood Mice show a gradual build-up to May 24, followed by a slow decline with no
peaks (Figure 7.24).
Both patterns alternate, i.e., the Wood Mouse reaches the highest numbers between
the two vole peaks. The result is that micromammals are constantly available, of one
species or the other.

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 Chapter 7: Diet

Table 7.13 Weight of intact micromammals in larders 1998–2021

Average Min Max.

Species Frequency weight weight weight

Common Shrew Sorex araneus 17 8.39 1.2 19.6

Greater White-toothed Shrew 33 11.31 6.8 18.5
Crocidura russula
Bank Vole Myodes glareolus 60 19.67 7.9 33.7
Euopean Water Vole Arvicola 17 43.21 17 83.6
amphibius/scherman
Common Vole Microtus arvalis 398 19 3 40.7
Wood Mouse Apodemus sylvaticus 296 18.72 3.3 40.7
Yellow-necked Woodmouse 9 41.61 28 56
Apodemus flavicollis
House Mouse Mus musculus 81 15.74 4.9 34.1

Total 911 18.86 1.2 83.6

3.5
Percentage of wood mice as prey items

2.5

2
Average Hatching Date

1.5

0.5

0
4 June
6 June
8 June
19 April
21 April
23 April
25 April

2 June

10 June
12 June
14 June
16 June
18 June
20 June
22 June
24 June
26 June
28 June
30 June
29 March
31 March
3 April
6 April
9 April
11 April
13 April
15 April
17 April

27 April
29 April
1 May
3 May
5 May
7 May
9 May
11 May
13 May
15 May
17 May
19 May
21 May
23 May
25 May
27 May
29 May
31 May
25 March
27 March

Date

Figure 7.24 Number of Wood Mice per day as a percentage of the total (n = 650), summed over 17
camera-observed breeding seasons, with the average hatching date indicated (Van Harxen and
Stroeken personal communication).

7.8.7.2 WEIGHT AND SEX OF MICE

In Winterswijk (the Netherlands), the average weight of 911 intact micromammals
found in larders was 18.86 g. The lightest (a young Common Shrew) was 1.2 g, the
heaviest (an adult European Water Vole), 83.6 g (Table 7.13).

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 7.8 Different Methods

Percentage of 296 intact Wood Mice and 398 intact 30

25
Wood Mouse

20 Common Vole
Common Voles

15

10

0
0–5 5–10 10–15 15–20 20–25 25–30 30–35 35–40 40–45
Weight classes of intact voles

Figure 7.25 Percentage distribution in weight classes of 296 intact Wood Mice and 398 intact
Common Voles (Van Harxen and Stroeken personal communication).

Wood Mice and Common Voles in particular are supplied in all size classes and
therefore the supply of both species seems to be a reflection of what is currently in the
field and not driven by a preference of the owl for larger specimens (Van Harxen and
Stroeken 2019) (Figure 7.25).
Male mice are caught substantially more than females. In a one-year study in
Winterswijk (Van Harxen and Stroeken 2021), 69.5% of Common Voles (n = 154)
and 57.9% of Wood Mice (n = 190) were male. The larger proportion of males probably
reflects the chances of catching them: males stay above ground more often when there
are young than the female, who feeds the young underground.

7.8.8 Birds
Birrer (2021) studied birds as the prey of owls, despite the fact that food of most owl
species is composed primarily of mammals. Birds seem to be of minor importance and
are often considered an alternative prey. Because food availability is a basic factor for
ecology and conservation, it is important to have quantitative data, not only on the
main prey but on alternative prey species as well. Birrer visualized the bird proportion
in 122 Little Owl diet papers with >50 vertebrates or >500 invertebrate prey items and
excluding lists with >10% vertebrates that were not determined at least to class level,
through Lorenz-like curves (Damgaard and Weiner 2000) indicating the inequality of
birds as prey in literature studies. Prey lists were sorted by increasing proportion of birds
in vertebrate prey and then visualized as cumulative distribution curves showing the
proportion of prey in relation to the proportion of birds (Figure 7.26).

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 Chapter 7: Diet

Figure 7.26 Graph showing the proportion of birds in the prey lists of the Little Owl (n = 122) in
relation to the percentage of prey lists and some important key metrics: Point A: percentage of prey lists
with no birds (exact data point: 8.2%); Point B: maximum percentage of birds in a prey list (exact data
point: 43.2%) (Birrer 2021). (a) Percentage based upon prey frequency. Small box: median (50% on x-
axis; percentage birds y-axis exact point 4.3%); large box: 90% quantile (90% on x-axis; percentage
birds y-axis exact point 18.5%). (b) Percentage based upon prey biomass. Small box: median (50% on
x-axis; percentage birds y-axis exact point 18.5%); large box: 90% quantile (90% on x-axis; percentage
birds y-axis exact point 17.8%).

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 7.8 Different Methods

Camera observations for 17 camera-observed breeding seasons in the breeding

period, out of a total of 41 140 prey, only 92 birds were brought in, an average of
5.4 per observation and 0.22% of all prey. The frequency ranged from 1 to 16 and
as a percentage of the total supply frequency per location it ranged from 0.03%
to 1.55%. Out of the total of 2085 vertebrate prey, birds made up 4.4%, with
large outliers both high (38.1%) and low (0.59%) (Table 7.14; Van Harxen and
Stroeken personal communication). Birds account for a very small part of the total food
supply.
Figure 7.27 illustrates the complementarity between birds and other vertebrates (y =
0.0003x2 – 0.1378x + 15.939, R2 = 0.4838, p <0.01, n = 17).
The same pattern is observed from the analysis of the prey remains in nestbox litters.
At only 199 locations were (remains of ) birds found during the nest visits (13.8%). With
352 of the total number of the 4959 vertebrate prey remains found during those checks,
birds accounted for 7.1% (Table 7.15; Van Harxen and Stroeken personal
communication).
Predominantly common species were caught, especially Starlings (24%), Blackbirds
(22%) and House Sparrows (15%), followed at a considerable distance by Great Tits (9%)
and Song Thrushes (5%). Other species were only found occasionally.
Birds as prey fluctuate across the years when correcting the frequencies for the
number of locations per year and the locations that were visited in the first 10 days
after hatching.
The number of birds was in most cases limited to one or two (altogether 83%,
Figure 7.28), but sometimes there were more in larders. On May 23, 2015, in a nest
with four 7-day-old nestlings, six newly fledged Starlings of the same age were found.
Three intact Starlings weighed an average of 60 g each, totaling a biomass of around
360 g.
In the 78 nests where more than one bird was found, 42 cases involved one species
(53.9%), 32 times it was two species (41%) and 4 times three species (5.1%). One pair of
owls in Barlo, the Netherlands on May 23, 2009 had a Chiffchaff, a Yellowhammer and
a Song Thrush in their larder.
The highest number of birds of the same species was found in a nest with eight birds
stored: one Blackbird and seven House Sparrows.
Table 7.15 gives an overview of the diversity of bird species found in larders
(1998–2021) by Van Harxen and Stroeken (personal communication).
Birds do not seem to be an important prey group for Little Owls, at least in the
breeding season, but in some years they are a significant addition to the menu.

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 Chapter 7: Diet

Table 7.14 Bird as prey seen during 17 camera-observed breeding seasons

(2002–2021) as frequency and as a percentage of the total number of prey supplied and
the number of vertebrate prey

Frequency of Total number Number of

birds of prey Percentage vertebrates Percentage

Hw04 1 3409 0.03 190 0.53

Hv07 2 4913 0.04 224 0.89
Hx16 2 3207 0.06 141 1.42
Dg17 1 1496 0.07 143 0.70
Hx20 4 4032 0.10 128 3.13
Hx21 1 773 0.13 169 0.59
Hx19 4 2895 0.14 109 3.67
Hv12 2 1303 0.15 90 2.22
Hx15 7 4221 0.17 164 4.27
Hv09 6 2068 0.29 110 5.45
Hw03 5 1583 0.32 74 6.76
Hv08 9 2718 0.33 126 7.14
Hx18 16 4785 0.33 42 38.10
Hx17 1 260 0.38 107 0.93
Hv11 7 1417 0.49 96 7.29
Hw02 12 1284 0.93 136 8.82
Hv10 12 776 1.55 36 33.33
Total 92 41140 0.22 2085 4.41

18

16

14 y = 0.0003x2 – 0.1378x + 15.939

R² = 0.4838
Birds prey frequency

12

10

0
0 50 100 150 200 250
Vertebrate prey frequency

Figure 7.27 Negative quadratic relationship between number of birds (y-axis) and other vertebrates
(x-axis) in prey supply in 17 camera-observed breeding seasons (Van Harxen and Stroeken personal
communication).

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Table 7.15 Overview of birds in larders in nestboxes 1998–2021

1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 Total

Bird species 1 6 1 1 1 3 7 3 2 3 2 2 1 2 15 4 54
Common Blackbird 3 2 4 1 5 2 3 2 8 1 4 1 2 5 2 5 3 11 5 3 1 1 3 77
Turdus merula
Song Thrush 1 2 2 2 2 1 2 1 1 1 1 2 18
Turdus philomelos
Common Starling 2 1 2 4 2 6 1 4 1 1 2 4 2 2 1 12 1 5 3 20 10 86
Sturnus vulgaris
Yellowhammer 1 1 1 1 1 1 6
Emberiza citrinella
Great Tit 1 1 2 1 3 1 5 1 1 3 2 1 1 5 2 30
Parus major
Sparrows 2 1 6 2 1 3 1 1 4 2 12 1 36
Passer spp.
Eurasian Tree 5 5
Sparrow
Passer montanys
House Sparrow 12 1 1 1 15
Passer domesticus
White Wagtail 1 1
Moctacilla alba
Common Chifchaf 1 1
Phylloscopus collybita
Common Chaffinch 1 1 2
Fringilla coelebs
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320

Table 7.15 (cont.)

1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 Total

Eurasian Jay 1 1 2
Garulles glandarius
Carrion Crow 1 1
Corvus corone
Great Spotted 1 3 4
Woodpecker
Dendrocopos major
Common Redstart 1 1 2 4
Phoenicuris
phoenicurus
Black Redstart 1 1
Phoenicuris ochruros
Stock Dove 1 1
Columba oenas
Common Wood 1 1
Pigeon
Columba palumbus
Barn Swallow 1 1 4 6
Hirundo rustica
Common Parakeet 1 1
Melopsittacus
undulatus
Total birds 5 6 19 13 10 20 4 12 13 7 7 9 7 22 13 14 9 33 11 9 13 13 60 23 352
Total prey 51 132 85 157 68 107 38 115 109 193 149 64 57 64 157 201 623 248 250 231 269 623 432 536 4959
Percentage birds 9.8 4.5 22.4 8.3 14.7 18.7 10.5 10.4 11.9 3.6 4.7 14.1 12.3 34.4 8.3 7.0 1.4 13.3 4.4 3.9 4.8 2.1 13.9 4.3 7.1

Van Harxen and Stroeken personal communication

 7.8 Different Methods

Table 7.16 Proportion of prey groups mentioned in prey lists (n = 122) with >50
vertebrates or >500 invertebrate prey items and excluding lists with >10% vertebrates
that are not determined at least to class level

Prey lists Median of 90% of Max

without this vertebrate prey vertebrate prey vertebrate
prey group by prey group by prey group prey by prey
Group (%) (%) (%) group (%)

Birds (frequency) 8.2 4.3 18.5 43.2

Birds (bodymass) 8.2 5.2 17.8 51.4
Reptiles 65.6 0 12.2 50
(frequency)
Reptiles 65.6 0 3 13.2
(bodymass)
Amphibians 61.5 0 5.9 27.8
(frequency)
Amphibians 61.5 0 2.6 19.2
(bodymass)

After Birrer (2021)

100
Bird prey percentage corrected for number of locaons

90

80
and visits within 10 days aer hatching

70

60

50

40

30

20

10

Year

Figure 7.28 Bird prey frequencies corrected for number of locations and number of checks within
10 days after hatching (Van Harxen and Stroeken personal communication).

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 Chapter 7: Diet

7.8.9 Other Prey

Birrer (2021) studied amphibians and reptiles as prey of owls and their respective
contribution to prey frequency and biomass. Figures 7.29 and 7.30 illustrate the
inequality of distribution of amphibians and reptiles across 122 prey lists. Exact points
are given in Table 7.16.

Figure 7.29 Graph showing the proportion of amphibians in prey lists of Little Owl (n = 122) in
relation to the proportion of prey lists: (a) frequency (b) biomass. (After Birrer 2021).

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 7.8 Different Methods

Figure 7.30 Graph showing the proportion of reptiles in prey lists of Little Owl (n=122) in relation to
the proportion of prey lists: (a) frequency, (b) biomass. (After Birrer 2021).

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 Chapter 8: Breeding Season

Chapter Summary
This chapter will cover the entire Little Owl breeding cycle. The breeding season is
obviously a critically important period, during which reproduction can be influenced
by many different factors, such as weather, food, habitat, density, geographical location
and parental experience. The season begins in January or February (Glue and Scott 1980,
Exo 1987) with the affirmation of territorial boundaries and onset of courtship. The Little
Owl does not have high productivity due to very few replacement clutches, moderate
fledging success and relatively high egg failure. According to the mortality rate of adults
and juveniles, each pair should produce between 1.7 and 2.34 fledged young per year to
compensate for mortality (Exo 1992) and actually most of the long-term breeding studies
across Europe show results ranging between both values. Analysis of consistently organ-
ized long-term demographic data took place to enhance our understanding of Little Owl
population dynamics. Further, this demographic data was linked to specific habitat
conditions at the nest-site, home-range and landscape scales.
In our preparation of this chapter, we became aware that clarification of terminology
related to nesting success and reproduction was needed. Clarification of these terms is
important to provide an accurate and consistent foundation for the data that will be
used to assess the reproductive performance of the owls, as well in long-term monitor-
ing of status and trends.

8.1 Pair Formation and Pair Bond

Still little is known about pair formation. Globally, two situations can occur. Either both
partners settle during the dispersion period in a new territory that was unoccupied in the
previous breeding season or one of them takes the place of a lost partner. The latter
situation can occur at any time of the year, even during the breeding season if there are
already eggs or young. There are countless examples where, after the disappearance of
one of the two birds, a new partner is reported within a week or even less. Van Harxen
and Stroeken (2021) described a situation in 2017 in which a female taken by a Stone
Marten (Martes foina) was replaced by a new partner, born in 2016, less than 200 m from
her new place and that had not bred before, within four days. Even a male can be
replaced quickly (Van Harxen and Stroeken 2005), e.g., by a non-breeding floater.

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 8.1 Pair Formation and Pair Bond

Table 8.1 Turnover in occupation of adult males and females in territory 343 in
Vragender (Netherlands)

Territory 343 2013 2014 2015 2016 2017 2018 2019 2020 2021

Male identification #466 #017 #182 #182 #170 #199 #052 #052 #640
Female identification #467 #018 #398 #398 #398 #173 #173 #759 #060

Van Harxen and Stroeken personal communication.

A new partner does not always lead to a match, as observed with cameras in 2013.
After the male was last seen on March 29, a new partner appeared on April 14, followed
by a second one on April 19. Although the female, who had bred there since 2012,
made attempts to bond with both, it did not lead to pair formation. In 2014 the nestbox
remained unoccupied (Van Harxen 2021), in 2015 there was a brood by two
new partners.
New settlements are even less known. Van Harxen (2021) described a situation in
early October in which a male of 5 months occupied a nestbox that was installed in
early August. In the months that followed, he regularly hooted. On February 1, a second
owl was observed for the first time at 8 months of age. The same day a mating is
observed and on April 9, the first egg was laid. During the first nestbox check in early
May, it appeared that both birds were not only from the same year, but also from the
same neighborhood, i.e,. the female was born in a nestbox 365 m away and the male
3.5 km further north.
Once formed, pairs usually stay together for the rest of their lives (Van Harxen
personal observation). Van Harxen and Stroeken (2020b) observed ringed birds that
established pair bonds of 4 or 5, and up to 8 years. However, a substantial proportion of
the partners were bonded for a shorter period because one of them disappeared. Despite
the fact that mortality can rarely be demonstrated and because brood dispersion is low
in a vital population, the lost partner is assumed to have died. Seven of the 12 pairs, in
which the date of birth of both partners was known, were born in the same year. For the
other five, the age difference was a maximum of 5 years, with males always older. With
such differences, the chance of losing a partner soon is much bigger. Therefore, in some
territories, the turnover was considerable. From 2013 to 2021, seven different males
occupied a territory in Vragender (Netherlands), five of them once, the other two twice
each. The turnover of the females in this territory was also very high, with six individ-
uals over the same period. Four times it was a completely new pair that bred (Table 8.1).
After the young have fledged the pair bond loosens, but both partners remain in the
territory and regularly stay close by. During camera observations, the female re-appeared
in the nestbox fairly soon after the breeding season and stayed there for most of the
day. The male was initially not tolerated in the nestbox, but as of mid-November
allowed to enter and stay. The male usually left the nestbox in the middle of the

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 Chapter 8: Breeding Season

Figure 8.1 Mutual affections during establishment of a pair bond.

Figure 8.2 A Little Owl calling.

afternoon to come back after dusk and called his partner from the branch on which the
nestbox was placed. Shortly after, the female came out of the nestbox. Sometimes both
flew away immediately, alternatively the feathers were first brushed or mutual affec-
tions exchanged (Figure 8.1). Towards the start of egg-laying, mating followed regularly.
Vocal activity increases and paired owls indulge in frequent calling (Figure 8.2).
Haverschmidt (1939) reported a duet in broad daylight, where both birds were sitting
on an exposed branch of a tree constantly calling and answering each other.
Technically this is not “dueting” but rather call–response vocalizations. This behavior
increases and intensifies until it leads to the more advanced stages of courtship with
nest visits and copulation. Nest visiting and showing consists of flying as a pair to

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 8.2 Life Expectancy

30

25
Long-term percentage

20

15

10

0
1 2 3 4 5 6 7 8 9 10 11 12
Age in years

Figure 8.3 Long-term percentage age distribution in years of 209 females with known birth year
2015–2021 (Van Harxen and Stroeken personal communication).

various nest sites which are preselected by the male within his territory. Nest choice
begins with a male calling loudly trying to lure the female into the nest cavity. In each
cavity, the male designates the potential nesting place by scratching his paws in loose
material lying on the bottom. If the female enters the nest cavity and takes over the
scratching, the action is successful but is often preceded by several failed attempts. This
behavior is also observed in places where cavities are limited, e.g., where only one
nestbox is present as a suitable nesting place. Usually, such a visit ends with the male
flying away first and the female shortly after.

8.2 Life Expectancy

The overall average female age distribution as a percentage, as monitored by Van
Harxen and Stroeken (personal communication) in 2015–2021, considering only the
females with known age, i.e., ringed as nestlings, is shown in Figure 8.3.
Roughly half of females were 1 or 2 years old in their first or second breeding season,
one-quarter were 3 or 4 years old, and one-quarter more than 4 years old, indicating
that new recruits contributed substantially to the breeding population and that habitat
conditions were favorable, allowing females to survive for many years.
In 2015, the average clutch size was smaller than average (3.68 versus 4.02 long-term
average), due to a flea plague, more eggs than usual did not hatch and more nestlings
died, resulting in fewer fledglings (1.67 versus 2.28 long-term average). This effect was
still visible in the low number of 2-year-old females in the following year. Between
subsequent years a similar pattern was observed (Figure 8.4).
Males were caught less often and therefore it was not possible to present accurate
data about age distribution compared to females. However, representation based on

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 Chapter 8: Breeding Season

120

100
Percentage per age

80

60

40

20

0
2015 2016 2017 2018 2019 2020 2021
21 24 31 23 33 37 40
Year
Number of Females
1 2 3 4 5 6 7 8 9 10 11 12

Figure 8.4 Annual age distribution of 209 females in years in 2015–2021 (Van Harxen and Stroeken
personal communication).

40

35

30
Percentage per age

25

20

15

10

0
1 2 3 4 5 6 7 8 9
Age in years

Figure 8.5 Long-term percentage age distribution in years of 181 males with age as the timespan
between ringing date and the last re-capture date in 1988–2021 (Van Harxen and Stroeken personal
communication).

the difference between the ringing and last re-capture date showed a similar distribu-
tion to that of females (Figure 8.5). The lower percentage of males older than 4 years
might be a methodological artifact rather than an actual lower age of males compared
to females.

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 8.3 Copulations

8.3 Copulations
Haverschmidt (1939) undertook a very detailed study and described the copulation
behavior as follows: “The session begins with the male owl alighting on to his favourite
perch and then continues with a questioning call ‘hoo,hoo?’ which he repeats continu-
ously increasing intensity and volume, building up to a ‘hoo-ee, hoo-ee?’ The male
shifted position a few times on the branch during the calling session. Suddenly a second
owl appeared and alighted near to the male, the female in contrast with the male was
silent. Without any further uttering the male jumped up onto the female’s back, the
female leaned forward so her body became horizontal and lifted up her tail a little, to
which the male sunk down onto her, soon after copulation occurred, with frenzied
wing flapping. The female then produces a shrill shrieking sound. Sometimes the male
continued to call while sitting on the females back, and often flew round in a semi-
circle to alight into a cavity at the rear of the tree, as if to show the cavity to the female.
He then flew back to his favourite perch again.”
Van Harxen (2021) describes the typical mating behavior as follows: “It’s ten to
seven in the evening and just dark when the male lands on the branch. As usual, the
male comes to pick up his partner for the nightly hunting. He calls and waits patiently
for her to join him. In the distance, another Little Owl gently calls, a reason to answer
loudly. Meanwhile, his partner has also settled on the branch, shakes her feathers a bit
and both seem to be getting ready to leave. For a moment they are a bit indecisively
dragging but then suddenly – without a clear visible announcement – he jumps on top
of her. She presses herself flat on the branch, sticks her tail up as far as she can
(Figure 8.6) – the cloaca is located directly below the base of the tail – and after some
somewhat clumsy-looking aerial acrobatics, it is done. The male uses his wings to
maintain balance. The entire ritual takes barely ten seconds and the actual copulation –
the pressing of the cloaca’s – between two and three seconds.”
The time just before the copulation, when the male is on the back of the female, can
last 30 seconds (average 27 s, n = 10), while the copulation time is shorter, with an
average of 3.1 s (n = 18) (Etienne 2003). The mating includes the “male on the female’s
back” and the copulation. It lasts on average 32.9 s (n = 56) and there are 0–4 copula-
tions per evening, 66% of copulations occuring during the evening (n = 64) (Ancelet
2004). The maximum mating time can reach five minutes (Torregiani 1981). The
mating season lasts from the beginning of February to the beginning of May. Some
copulations have been seen as late as November 28 (Etienne 2003) or December 6 (Exo
1987), but this could have a social function, to decrease the aggressiveness between the
two partners (Exo 1987). Ancelet (2004) reported 53 matings at the same place, on a
particular branch of an apple tree.
In 2016–2019, camera observations of Beleef de Lente (Van Harxen and Stroeken
personal communication) allowed meticulous monitoring of the moment of courtship
and mating. Most copulations took place on the branch in front of the nestbox and
were clearly visible (Figure 8.7). Occasionally, judging by sounds elsewhere, they copu-
lated out of sight of the camera. Hence, the number of registered copulations should be
considered a minimum. A total of 223 copulations were registered, an average of 56 per

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 Chapter 8: Breeding Season

Figure 8.6 Little Owls mating.

year (Table 8.2). Registration began when the cameras were switched on, ranging
between February 15 and March 7. Copulations that took place before that time were
missed. All observed copulations took place outside, no mating was observed in the
nestbox, despite the fact that it was spacious enough. In two out of three days with a
copulation, they mated only once, but on some days up to four times. In all cases the
male stayed the same, the female of 2018–2019 was different from that of 2016–2017,
with no effect on the number of copulations.
Most copulations took place in the evening hours after dusk (Figure 8.8), frequently
when the male came to pick up its partner from the nestbox at dusk to go hunting
together. Before flying away the pair often copulated. Occasionally they mated during
the day, sometimes as a second mating of that day, but usually as a first mating.
In all years, the number of copulations increased towards egg laying and continued
during the egg-laying phase, until a few days after the clutch was completed
(Figure 8.9). In 2021 when copulations were no longer regularly observed, a copulation
took place 19 days after the laying of the last egg.

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 8.4 Nest Sites

Table 8.2 Frequency of daily copulations (n = 223) through camera observations in

2016–2019

2016 2017 2018 2019 Total %

Once 36 31 37 39 143 64.1

Twice 18 10 14 13 55 24.7
Three times 6 5 4 6 21 9.4
Four times 1 1 1 1 4 1.8

Total 61 47 56 59 223

Van Harxen, Stroeken and Sterringa personal communication.

Figure 8.7 Illustration of nestbox equipped with webcam outside and inside (Beleef de Lente).

8.4 Nest Sites

Little Owls will use a variety of nest sites (Figure 8.10), however the specific require-
ments are a chamber with a large enough capacity to hold the eggs and subsequent
young as they grow; the chamber has to be able to protect from wind and rain, should
preferably also be dark and the entrance must be accessible to the owl but prevent
access by predators as much as possible. A pollard willow can typically offer all these
features (Figure 8.10a).

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 Chapter 8: Breeding Season

50

45

40

35
Number of copulaons

30

25

20

15

10

0
0-1

1-2

2-3

3-4

4-5

5-6

6-7

7-8

8-9

9-10

10-11

11-12

12-13

13-14

14-15

15-16

16-17

17-18

18-19

19-20

20-21

21-22

22-23

23-24
Time in hours

First Second Third Fourth

Figure 8.8 Hourly distribution of copulations (n = 223) through camera observations in2016–2019
(Van Harxen, Stroeken and Sterringa personal communication).

(a)

4.5
Egg-laying dates
4

3.5

3
Number of Copulations

2.5

1.5

0.5

0
10-3
11-3
12-3
13-3
14-3
15-3
16-3
17-3
18-3
19-3
20-3
21-3
22-3
23-3
24-3
25-3
26-3
27-3

10-4
11-4
12-4
28-3
29-3

13-4
30-3
31-3

14-4
15-4
16-4
17-4
18-4
19-4
20-4
21-4
22-4
23-4
24-4
25-4
26-4
27-4
28-4
29-4
30-4
1-3
2-3
3-3
4-3
5-3
6-3
7-3
8-3
9-3

4-4
5-4
6-4
7-4
8-4
9-4
1-4
2-4
3-4

Date

(b) (c) (d)

Figure 8.9 Daily copulations (n = 223) through camera observations in 2016–2019 (Van Harxen, Stroeken
and Sterringa personal communication). Egg-laying dates in black. (a) 2016 (b) 2017 (c) 2018 (d) 2019.

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 8.4 Nest Sites

Figure 8.10 (a) Pollard willow offering all the neccessary features for a suitable nesting cavity. (b)
Examples of nest-site cavities in trees and walls, (c) examples of nest site cavities in houses and piles
of stones.

Little Owls will use any suitable, available site within their territory, close to their
foraging habitat. When available, Little Owls prefer to nest in tree cavities, and splits in
trunks or branches that give the owls excellent opportunities for nesting. In Western
and Central Europe, buildings such as stables, haylofts, and other used and disused farm
buildings are also often exploited (Schönn et al. 1991).
Among many alternative nest sites are crevices in ruins and down wells, adobe
buildings in the Sahara Desert, holes in quarries, walls, sand-pits, stick nests
(Uspensky 1977, Sagitov 1990) and disused rabbit burrows (Bannerman 1955). In the
former USSR, breeding was recorded in windmills, mud tombs, granaries, haylofts,
haystacks, below overhanging rocks, burrows of large gerbils, and nests of Rock
Nuthatches Sitta neumayer; some were tunnelled by the birds themselves (Dementiev
and Gladkov 1951). “Clapas,” walls, ruins, rocks and “chazelles” offered cavities at
Causse Méjean, France (Juillard et al. 1992). In Haut-Léon, western France, only build-
ings are used as nesting sites, in 55% of cases under the roof (Clec’h 2001). Specific
nicknames relate to nesting places, such as “Chouette des tuiles” (tile owl) in southern
France (Barthelemy and Bertrand 1997), “Baumkauz” (tree owl) (Weimann 1965) and
“Stockeule” (willow owl) (Schönn et al. 1991) in central Europe. Buildings and mounds
of stones are used in Tolfa, Italy (Centili 1996) and fruit trees (especially apple), pollard

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 Chapter 8: Breeding Season

(b)

Figure 8.10 (cont.)

trees, man-made constructions and nestboxes in Betuwe (Netherlands) (see Fuchs 1986
for proportional distribution of the use).
Glue and Scott (1980) studied a sample of 482 nest sites and found that 24% were in
oak, 23% in ash, 18% in fruit trees and 15% in willows. Of 172 nests in Germany, 49%
were in fruit trees, 31% in pollard trees, 27.5% in buildings and 18% in assorted places,
e.g., quarries, and nestboxes (Schönn 1986). Of 530 nests in France, 18% were in fruit
trees, 11% were in pollard trees, 12% in other trees, 32% in buildings and 26% in other

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 8.4 Nest Sites

(c)

Figure 8.10 (cont.)

places like nestboxes and rock faces (Génot 1992c). Of 100 nests in buildings in
Brittany (western France), 46% were in agricultural buildings, 6% in manors, 9% in
inhabited houses, 33% in isolated or abandoned houses, 5% in hangars and 1% in
pigeon houses (Clec’h 2001). Of 144 nests in Austria, 17 were in trees, 62 in barns, 53 in
wine cellars, 6 in farms, 4 in stacks of straw bales and 2 in churches (Ille and Grinschgl
2001). In the Mediterranean region, agricultural practices provide nesting opportunities
by clearing all large rocks and stones from the land. Farmers pile them up at the side of
the land they have cleared. These stone piles provide alternate nest sites in areas that
lack suitable nesting trees (Juillard et al. 1992). Centili (2001) recorded 39 nests in Italy:
25 were in stone piles, 13 in buildings and 1 in an iron pole. In Britain, at high
elevation, tree-lined hedgerows give way to stone walls that are utilized by Little Owls
as nest sites. In a study area in Cheshire, England, a pair of owls had nested for several
years in a nestbox situated 5 m up an oak tree; storms damaged the box just prior to the
breeding season one year and the owls vacated the nestbox and nested in a rabbit
burrow close to the tree. The following year the nestbox had been replaced and the owls
returned to the box. The burrow was inspected and it seemed to be in good condition,
showing that the owls preferred the elevated site in this case (Leigh 2001a). The diver-
sity of buildings as nesting places has been shown in Belarus, where Little Owls use
barns, windmills, water towers, silo towers, building blocks, churches, cemetery graves,
old beehives and even more unusual places such as haystacks (Nikiforov et al. 1989). In
the Netherlands, Van Harxen and Stroeken (personal communication) found 162 cav-
ities other than nestboxes: 130 (80%) in all sorts of buildings (most of them in barns of

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 Chapter 8: Breeding Season

different sizes and for different uses, but also in farmhouses and hay stacks), 22 (14%) in
fruit trees, mainly high-stem apple trees and 10 (6%) in pollard trees such as willows,
poplars and black alder. In riverine areas, the majority of the Little Owls bred in trees,
e.g., pollard willows and poplars in floodplains and fruit trees in high-stem orchards on
the levees (Fuchs and Van de Laar 2010). Since the large-scale uprooting of the old
orchards and environmental development in the floodplains, their numbers have
decreased and the remaining pairs have shifted to nestboxes.
In natural habitats, e.g., the deserts and semi-deserts of the former Soviet Union, the
species uses nesting sites such as holes in rocks and cliffs in river valleys, in loess
precipices or, in sandy deserts, hollows of trees, burrows of Great Gerbils (Rhombomys
opimus) or Thin-toed Sousliks (Sagitov 1990), abandoned burrows of foxes, badgers and
steppe tortoises in hilly sands, burrows of Rollers (Coracias garrulus) in the banks of
rivers or slopes of ravines, and burrows of the Rock Nuthatch in clefts of rocks (Gavrin
1962). In Moldavia, the Little Owl occupies empty nests of the Magpie (Pica pica), far
from human settlements. Over about 50 ha 42 of 100 Magpie nests were occupied by
Little Owls for breeding, in a kind of “nesting colony” (Uspensky 1977). In the
Caucasian area, Little Owls have also been found to breed in old nests of Carrion
Crows (Corvus cornix) (Il’yukh 2002). In the past, Little Owls in the Netherlands bred
in rabbit holes in coastal dunes and the big sand drifts of the Veluwe (Bijlsma et al.
2001). Nowadays it is very rare due to the closing-up of the vegetation in the dunes
following the collapse of the rabbit population.
Nestboxes are widely used as a conservation tool in areas where nest-site availability is a
limiting factor, and in these cases nestboxes will be readily used. In areas where the natural
nest sites are not a limiting factor, it can take a number of years for the owls to adopt a
nestbox. The nest height will vary according to availability; in a study in France, Génot
(1990a) found the range of heights of 25 nests to be 1–4.3 m high in fruit trees. The average
height was 3.9 m in trees, buildings and nestboxes in Switzerland (Juillard 1984) (n = 59)
and 2.4 m in trees in Germany (Exo 1981) (n = 28). In buildings, the owls mainly use
cavities in or under the roof space, often under loose or missing roof tiles. Agricultural
buildings offer further nesting opportunities; barns with ledges are readily used, as are hay
barns. Nesting places in buildings can be very big (several m3), but also very small (10  10
 8 cm). In the Netherlands many pairs breed under the corrugated sheets of pigsties. The
owls enter the nest by crawling under the curve of a sheet. As the opening is very narrow
they have to slide with their belly over the wooden beam the sheets rest upon. By doing so
they leave a clear track on the beam which makes it easy to determine where the nest will
be. The nest is mostly found where the eggs cannot roll away at the base of the roof. Little
Owls can be very steadfast in their choice of a nesting place, using the same place over and
over again. Van Harxen and Stroeken (2011a) reported several nesting places being used
for decades, e.g., under the roof of a small shed for cattle that was used for at least 27 years
(1988–2014), probably much longer, as the shed was built in the 1950s. The respective
owls used different places to nest: front side left, front side middle, front side right, back
side left and back side middle. Sometimes the same place was used for a few years
sequentially, but mostly nest sites switched between years, possibly to limit the impact
of parasites (see Chapter 10 on Population Regulation).

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 8.6 Eggs

8.5 The Nest

Nest dimensions recorded in British cavities were, on average, 20 cm wide (10–50 cm;
n = 74). Ten entrance passages had an average length of 80 cm (50–130 cm) (Glue and
Scott 1980). Génot (1990a) undertook a similar exercise in France and found that the
average cavity depth of 25 nest sites was 77 cm (32–200 cm). The positions of entrance
holes and cavities inside trees were analyzed by Exo (1981) and Génot (1990a). When
the species uses burrows, the nest has been reported to have a diameter of 20–30 cm,
located 1–3 m from the entrance (Gavrin 1962) or a diameter of 20–25 cm, 1–1.5 m
from the entry (Sagitov 1990). One nest chamber was recorded at 25  14 cm
(Abdusalyamov 1971). When the dimensions of the nest do not provide enough space
for the young to move around, they leave the nest early and frequently suffer increased
mortality (Ancelet 2003).
Different types of holes have been distinguished, including horizontal and vertical
(Exo 1981, Génot 1990a) . In Switzerland, only 1 in 110 fruit trees had a cavity that was
suitable for the Little Owl (n = 561, including 250 apple trees) (Juillard 1984); in France,
there was one suitable cavity per 60 fruit trees (n = 8839, including 2386 apple trees)
(Génot 1990a). Natural cavities, such as in apple trees, used for a limited number of
years, gave rise to more frequent changing of nest sites (Fuchs 1986). On sunny days,
overheating can occur, as the temperature inside a nestbox can be 7–9 C warmer than
outside, and similarly for the nests under roofs, putting the development of the
embryos at risk (Beersma and Beersma 2000). It is advised to put nestboxes in trees in
the shadow of leaves. Little Owls may not yet be able to distinguish, or may not have a
choice, between safe nest locations and those that are likely to overheat (Van Den Burg
et al. 2003).
In terms of construction, no nest building is carried out, although the female will
form a scrape in the litter in the chamber (Ullrich 1973). A bed of pellets forms the base
for the eggs in previously used nest sites. Normally, there is no litter in the nest, only a
layer of food remains or pellets. In some nests, stems of dry grass have been discovered
(Sagitov 1990) and pieces of pine wood, which have a positive effect on the nestlings
by providing drier conditions in the nest during wet weather (Robiller 1987).

8.6 Eggs
The eggs are white, with a silky smooth finish. The shape is a slightly elliptical sphere
(Figure 8.11). The newly laid eggs average approximately 15.6 g, and range from
12.5–19 g (n = 95) (Illner in Glutz Von Blotzheim and Bauer 1980) and measure
33–40 mm by 27–31 mm. For Athene noctua noctua, the mean dimensions: 34.4 
28.8 mm (n = 140) (Schönwetter in Schönn et al. 1991), for Athene noctua vidalii: 36 
30 mm (n = 100) (Witherby et al. 1938), for Athene noctua indigena: 34  28 mm (n = 38)
(Makatsch 1976) and for Athene noctua bactriana: 30.9–34.5  26.1–28.9 mm (Sagitov
1990).
Il’yukh (2002) examined Little Owl eggs in the Caucasian area (Table 8.3) spanning
1999–2001 and found that measurements in the size and shape of the eggs did not

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 Chapter 8: Breeding Season

Table 8.3 Clutch size and characteristics of eggs in the Little Owl in the Pre-
Caucasian area

Indices n Range Mean  SD

Clutch size 31 0–8 4.870.22

Length, mm 89 32.2–36.1 34.320.09
Width, mm 89 26.9–31.1 29.100.09
Volume, cm3 89 12.5–17.8 14.860.12
Width–length ratio, % 89 78.2–88.5 84.810.20

After Il’yukh 2002.

Figure 8.11 Egg of the Little Owl.

significantly differ. There was a very low variation in the dimensions, which is typical
for birds that breed in the limited, stable spaces of closed nest sites.
However, there was a significant inter-year variability in some egg measurements
(Table 8.4). In 2001, a year with a rainy and cool spring, the Little Owl eggs were
significantly larger and more round than in the dry, hot season of 1999 and during
the spring of 2000. The difference in Little Owl egg lengths was significant between
1999 and 2000 (t = 2.89, p< 0.01) and 2000 and 2001(t = 3.91, p <0.001). The egg width
in 2001 was significantly higher than in 1999 (t = 2.19, p <0.05) and 2000 (t = 4.35,
p<0.001), and in 1999 higher than in 2000 (t = 2.67, p <0.01). By volume, eggs in
2000 were considerably smaller than those from 1999 (t = 3.55, p<0.001) and 2001
(t = 5.78, p <0.001). In 2001 the eggs were more round than in 2000 or 1999 (t = 2.13,
p <0.05). It is interesting to note that the length and width of eggs varied to a lesser
extent in cool and rainy seasons, than in hot and dry ones.

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 8.6 Eggs

Table 8.4 Inter-year variability of size and form of Little Owl eggs in the Pre-
Caucasian area

Years n Range Mean  SD

Length, mm
1999 38 32.2–36.1 34.360.16
2000 12 32.5–35.0 33.550.23
2001 39 32.8–36.0 34.530.10
Width, mm
1999 38 27.5–31.1 29.030.14
2000 12 26.9–29.5 28.310.23
2001 39 28.1–31.1 29.420.11
Volume, cm³
1999 38 12.5–17.8 14.800.21
2000 12 12.7–15.3 13.720.22
2001 39 13.5–17.5 15.260.15
Width–length ratio, %
1999 38 81.2–87.7 84.510.25
2000 12 78.2–87.9 84.441.02
2001 39 83.0–88.5 85.220.22

After Il’yukh 2002.

Van Harxen and Stroeken (personal communication) measured 604 eggs over a
period of 8 years: 272 incubated eggs (1998–2000) and 332 unhatched eggs
(2017–2021). The average length was 34.2 mm, the average width 28.8 mm and the
average volume 14.44 mm3. They found no significant differences between years nor
between incubated and unhatched eggs (Table 8.5). The most elongated egg measured
34.4  22.6 mm (ratio l:b = 0.67) and the most rounded egg 32.2  32.1 (ratio l:b =
0.97). The average ratio was 0.84. The largest egg had a volume of 18.14 mm3 and the
smallest 8.7 mm3. Unlike Il’yukh (2002), they found no significant difference in volume
per clutch size (Table 8.6).
The weight of an egg is determined not only by its fresh weight, but also by its
measuring moment. From the moment an egg is incubated, it steadily loses weight due
to metabolism. Three eggs that were about to hatch (picked up, squeaking young)
weighed 11.3, 11.4 and 12.2 g. The same eggs weighed 13.5, 13.8 and 14.7 g six days
after the clutch was started. In 22 days, these eggs lost an average of 2.4 g of weight. The
average daily decrease of 100 eggs was 0.085 g. Over the whole of the breeding period
(26 days), this amounts to 2.2 g. However, the variation per egg is large, ranging from
0.02 g to 0.15 g per day.
Il’yukh (2002) reported seasonal changes in the size and form of eggs (Table 8.7). In
early clutches eggs were significantly smaller than those from later clutches in length
(t = 4.36, p <0.001), width (t = 2.76, p <0.01) and volume (t = 3.42, p <0.01).

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Table 8.5 Annual variation between length, width and volume of 272 incubated (1998–2000) and 332 unhatched (2017–2021) eggs

Length Width Volume

Year n Mean Min Max Mean Min Max Mean Min Max

Incubated 1998 67 34.4 28.4 40.4 28.6 22.6 30.4 14.40 8.68 18.14
1999 120 34.2 31.3 38.5 28.8 27.0 31.1 14.51 11.70 17.72
2000 85 34.2 30.5 36.6 28.5 26.1 30.8 14.19 11.89 16.62

Unhtached 2017 54 34.6 29.6 37.0 28.9 26.2 30.4 14.77 10.33 17.16
2018 53 34.0 31.2 35.9 28.7 26.9 30.5 14.25 11.78 16.67
2019 64 34.6 31.6 37.8 28.9 26.3 30.7 14.77 11.13 18.03
2020 75 33.9 30.6 37.7 28.8 26.8 30.8 14.35 11.64 17.73
2021 86 34.0 29.3 39.1 28.8 25.5 32.2 14.35 9.95 17.77

n/mean 604 34.2 28.4 40.4 28.8 22.6 31.18 14.44 8.68 18.14

Van Harxen and Stroeken personal communication.

 8.6 Eggs

Table 8.6 Clutch size and characteristics of eggs of the Little Owl in Zuidoost-
Achterhoek, the Netherlands

Clutch size

2 3 4 5 6

No. of eggs 14 93 148 80 60

Average volume 14.56 14.56 14.39 14.34 14.59
Average length 34.8 34.2 34.2 34.3 34.6
Average width 28.7 28.9 28.7 28.6 28.8

Van Harxen and Stroeken personal communication.

Table 8.7 Seasonal variability of size and form of Little Owl eggs in the Pre-
Caucasian area

Eggs laid n Range Mean  SD

Length, mm
Early 55 32.2–35.6 34.040.12
Late 34 33.7–36.1 34.780.12
Width, mm
Early 55 26.9–30.1 28.910.10
Late 34 27.5–31.1 29.430.16
Volume, cm³
Early 55 12.5–16.3 14.530.14
Late 34 13.0–17.8 15.390.21
Width–length ratio, %
Early 55 78.2–88.5 84.940.27
Late 34 81.2–87.8 84.600.27

After Il’yukh 2002.

In addition, links between the quantitative indices of Little Owl eggs and clutch size
(Table 8.8) are obvious. The biggest eggs were observed in clutches consisting of six
eggs, the smallest in clutches of seven or eight eggs. The lengths of eggs in eight-egg
clutches were significantly less than those in four- (t = 3.54, p <0.01), five- (t = 3.17,
p <0.01) and six-egg (t = 3.75, p <0.001) clutches. Likewise, the widths of eggs from
eight-egg clutches were significantly less than in four- (t = 5.76, p <0.001), five-
(t = 3.56, p <0.01) and six-egg (t = 6.6.7, p <0.001) clutches. Seven-egg clutches
contained narrower eggs than four-egg (t = 2.54, p <0.05) and six-egg clutches
(t = 2.78, p <0.05). By volume, eggs of eight-egg clutches were significantly lower than

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 Chapter 8: Breeding Season

Table 8.8 Characteristics of Little Owl eggs in different clutch sizes in the Pre-
Caucasian area

Number of eggs in clutch n Range Mean  SD

Length, mm
4 28 32.8–35.4 34.400.12
5 25 32.5–36.1 34.390.19
6 18 33.0–36.0 34.610.20
7 7 33.0–35.0 33.930.32
8 8 32.2–34.4 33.290.29
Width, mm
4 28 28.1–31.1 29.340.14
5 25 27.5–31.1 29.090.20
6 18 28.6–30.3 29.420.12
7 7 26.9–29.5 28.310.38
8 8 27.6–28.7 28.240.13
Volume, cm³
4 28 13.5–17.5 15.120.18
5 25 12.8–17.8 14.880.28
6 18 13.8–16.5 15.290.21
7 7 12.7–15.3 13.880.33
8 8 12.5–14.4 13.540.23
Width–length ratio, %
4 28 83.0–88.5 85.310.30
5 25 81.2–87.9 84.580.34
6 18 83.1–87.7 85.040.27
7 7 78.2–87.6 83.531.64
8 8 83.0–86.9 84.850.50

After Il’yukh 2002.

four- (t = 5.41, p <0.001), five- (t = 3.70, p <0.001) and six-egg (t = 5.62, p <0.001)
clutches; likewise seven-egg clutches consisted of eggs significantly smaller by volume
than four- (t = 3.30, p <0.01), five- (t = 2.31, p <0.05) and six-egg (t = 3.60, p <0.01)
clutches. Finally, in rainy springs, Little Owl eggs were significantly bigger, the eggs
hatch later and the clutch size was smaller.

8.7 Laying of Eggs

In western Europe, egg-laying generally begins in the last week of April (Knötzsch 1978,
Ullrich 1980, Génot 1992c, Lederer and Kämpfer-Lauenstein 1996) or the second half
of April (Bultot et al. 2001). In the arid regions of the owl’s distribution, laying starts
in the second week of March (Uzbekistan; Sagitov 1990) or the third week of March

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 8.7 Laying of Eggs

(Turkmenistan; Ataev 1977). In the Saratov region (Russia), laying occurs from late May
to early June (Zav’yalov et al. 2000). Taking into account very early clutches and
replacement clutches, egg-laying is extremely extended through the distribution area
and can last for three months, as in Kazakhstan (from beginning of April to the end of
June; Gavrin 1962) or even 5 months, as in Turkmenistan (from the first week of March
to the middle of August; Bel’skaya 1992).
The earliest recorded laying dates are March 16, 1950 in Britain (Glue and Scott
1980), March 19, 1959 in Tajikistan (Abdusalyamov 1971), March 22, 1974 in
Turkmenistan (Ataev 1977), April 4, 1938 in France (Labitte 1951), April 6 (year
unknown) in Germany (Schönn 1986) and France (Blache 2004), April 9, 1977 in
Switzerland (Juillard 1984) and March 4, 2020 in the Netherlands (camera observation,
personal communication H. Wanders). Unusually early records for eggs were December
1949 in Hungary (Solymosy 1951) and January in France and Turkey (Pardieu 1927,
E. Vaassen personal communication). Framis and Alonso (2009) found, on January 29,
2007, an almost 4-week-old juvenile in a nestbox in the gardens of the passenger
terminal at Reus Airport (near Tarragona) in Catalonia, Spain. This implies an estimated
laying date in the first half of December.
In the Netherlands, the number of clutches that started in March accounted for
4.48% of a total of 15 341 clutches with known laying dates between 1976 and 2021
and showed a gradual increase (Table 8.9). An exceptional year was 2014, with almost
24% due to high numbers of voles and mice (Van Harxen et al. 2023).
The laying date is influenced by the food, with early laying dates corresponding to
peak vole years (Illner 1979, Ullrich 1980, Finck 1989, Sill and Ullrich 2005). Climatic
conditions also have a relationship with the laying date. If the duration of snow cover
in winter is long, the laying date can be delayed (Illner 1979, Ullrich 1980, Exo 1992).
Lederer and Kämpfer-Lauenstein (1996) reported that the laying date was earlier with
higher amounts of rainfall in spring (March and April), i.e., during wet springs, the
availability of earthworms is greater and the female can reach a better physical state.
However, Bultot et al. (2001) found the contrary; for each 100 mm of extra rainfall
laying was delayed an average of six days.

Table 8.9 Proportion of years with early breeders per decade in 1976–2021 in the
Netherlands (n = 15 341)

Percentage of Years without Percentage without

Decade March mean March clutch March clutch

1970 0.00 4 100

1980 0.00 10 100
1990 2.80 4 40
2000 2.58 1 10
2010 5.30 0 0
2020 4.22 0 0

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 Chapter 8: Breeding Season

29-4
y = –0.1962x + 44674
R² = 0.3412
24-4

19-4
Egg Laying Initiation Date

14-4

9-4

4-4

30-3

25-3
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017
2018
2019
2020
2021
Year

Figure 8.12 Linear advancement of egg-laying initiation date in the Netherlands 1973–2020 (n =
18 146). Source: Meetnet Nestkaarten van Sovon Vogelonderzoek Nederland (Nienhuis et al. 2015).

External factors may significantly affect the breeding dynamics. In other owl species,
when negative factors are prevalent, e.g., when prey cycles are extremely low and the
weather factors are not good, the amount of resources is too low and the owls’ poor
physical condition will not allow them to lay eggs that year. Additional research is
needed on the status and physical condition of nonbreeding Little Owls.
In the Netherlands, a systematic advance of egg-laying initiation was observed
(annual average egg-laying initiation date = 23/4 in 1970 – 0.1962 days per year, p =
0.00002, R2 = 0.34) from the mid-1970s (Van Harxen et al. 2023) to the early 2020s,
possibly as a consequence of climate change (Figure 8.12).
Van Harxen et al. (2023) reported a substantial difference in the mean laying
initiation date of 20 days between two consecutive years (April 27 in 2013 and April
7 in 2014; mean 1998–2021: April 17), due to the mean temperature in early spring,
2013 being cold (March, 2.5 C versus 6.2 C long-term average, first 10 days of April,
4.2 C versus 7.9 C long-term average). Voles and other prey species (e.g., cockchafers)
appeared later than usual. 2014 was the opposite (March, 8.4 C and April, 12.1 C). As a
consequence, grass started to grow extremely early and thus voles were available early
and abundant, causing excellent feeding conditions. Due to a very good mast year in
2013, Wood Mice also were extremely abundant (see Chapter 6, Diet).

8.8 Laying Interval

Schönn et al. (1991) reported an interval of one to three days between the laying of two
eggs, predominantly two days, but sometimes larger intervals. Ullrich (1973, 1975b)

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 8.9 Clutch Size

and Exo (1983) also mentioned an interval of one day for different breeding pairs.
Camera research allowed the precise moment of laying to be determined (in hours and
minutes). The average laying interval of 19 pairs varied between 47:58 and 60:16 hours,
with an overall average across all nests of 54:33 hours. At the egg level, the shortest
interval was 44:56 hours and the longest 68:08 hours (Table 8.10).
The average egg-laying interval for three-egg clutches was 56:17 hours and for four-
egg clutches 54:04 hours. The time difference between the first and last egg was on
average 5.11 days (5 days and 2 hours) for three-egg clutches and 7.76 days (7 days and
18.5 hours) for four-egg clutches. The only five-egg clutch took almost eight days to
complete, which was relatively quick due to significantly shorter intervals between the
individual eggs (47:47 hours).
In two clutches, significantly longer intervals between the eggs were noted: 121:39
hours for a two-egg clutch and 84:18 hours between the second and third egg of a three-
egg clutch. The impression was that in both cases there would have been an egg in
between, which for some reason it was not laid (or was laid outside the nesting cavity).
Both females showed egg-laying behavior. Both clutches were very early (first eggs
March 23 and March 25, respectively) and for both broods the eggs did not hatch.
Due to these special circumstances, both clutches were not included in the calculation
of the averages.

8.9 Clutch Size

While the main studies about breeding have been focused on western European coun-
tries, some data on clutch sizes have come from countries of the former Soviet Union.
Clutch sizes vary, and most commonly contain between one and seven eggs, but
clutches of eight and nine eggs have been recorded in eastern countries (Gavrin 1962,
Abdusalyamov 1971, Ataev 1977, Sagitov 1990, Nemeth in Schönn et al. 1991, Hudec
et al. in Schönn et al. 1991, Rakhimov and Pavov 1999, Il’yukh 2002) and even 10 in
the Netherlands (Willems et al. 2004) or 12 in the desert of Turkmenistan (Sopyiev
1982) (Table 8.11). High numbers of eggs have been attributed to two clutches in the
same nest in some cases in the Netherlands where a nest with eight eggs belonged to
two joint breeding females. Six young could be divided into two groups of three with an
age difference of four days, where the average weight of the young in the oldest group
on the first control day was 1.8 times as high as that in the youngest group (Stroeken
and Van Harxen 2018). In the exceptionally good mouse year, 2019, a nest with 11 eggs
was found in Brabant, also with two breeding females (Van Dooren and Hermsen 2019).
Also in a nest with 13 eggs in the Betuwe, two females were involved, possibly even a
third (Fuchs unpublished).
The average clutch size in Europe varied from 2.65 (Leigh 2001a) to 5.24 (Kohl in
Schönn et al. 1991) (Table 8.12). Remarkably high clutch sizes were found in Central
Asia: 6.2 (n = 12) in Kazakhstan (Gavrin 1962), 6.67 (n = 15) in Turkmenistan (Ataev
1977) and 6.0 (n = 6) in Uzbekistan (Bakaev in Sagitov 1990). It remains unclear
whether these were exceptions due to small sample sizes, exceptional prey-rich years
or due to excellent habitat quality (i.e., high numbers of mice and voles).

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346

Table 8.10 Laying intervals for 19 camera-observed breeding seasons

Interval between eggs in hours and minutes Interval between first and last eggs in days

Nest Clutch size Mean Second Third Fourth Fifth First to last Three eggs Four eggs Five eggs

Hw05-I 4 55:28 56:11 49:30 60:45 166:26 6.93

Hw05-II 3 53:18 51:13 55:23 106:36 4.44
Hw08 3 56:16 54:54 57:39 112:33 4.69
Rd07 4 55:30 55:30 52:30 58:30 166:30 6.94
Rd08 4 51:56 53:29 50:14 52:07 155:50 6.49
Hv07 4 52:12 55:12 46:48 54:36 156:36 6.53
Hv08 4 58:39 52:35 55:15 68:08 175:58 7.33
Hv09 4 53:32 56:07 48:51 55:39 160:37 6.69
Hv10 4 47:58 46:51 44:56 52:08 143:55 6.00
Hv11 4 53:31 57:22 48:37 54:36 160:35 6.69
Hv12 3 60:16 59:55 60:38 120:33 5.02
Hx15 5 47:47 49:16 44:57 45:06 51:49 191:08 7.96
Hx16 4 55:15 52:30 50:05 63:10 165:45 6.91
Hx17 4 58:13 57:03 54:42 62:54 174:39 7.28
Hx 18 4 52:34 51:00 49:27 57:17 157:44 6.57
Dg17 3 54:55 55:02 54:49 109:51 4.58
Dg 18-II 3 52:42 51:09 54:16 105:25 4.39
Hx19 3 58:04 56:31 59:38 116:09 4.84
Hx20 3 58:31 61:02 56:01 117:03 4.88

Total 70 19 19 19 12 1 19 8 11 1
Mean 3.68 54:33 54:21 52:19 57:04 6.64 5.11 6.76 7.96

Van Harxen, Stroeken and Sterringa personal communication.

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Table 8.11 Clutch sizes of the Little Owl recorded across Europe

No. of
Country 1 2 3 4 5 6 7 8 9 10 Mean clutches Author

Britain 1.00 10.00 35.00 39.00 13.00 1.00 1.00 0.00 0.00 0.00 3.6 268 Glue and Scott
1980
Britain 11.50 30.97 33.63 23.89 0.00 0.00 0.00 0.00 0.00 0.00 2.6 115 Leigh 2001a
France 0.00 0.00 23.00 62.00 15.00 0.00 0.00 0.00 0.00 0.00 3.9 80 Labitte 1951
France 6.00 9.00 35.00 46.00 19.00 7.00 1.00 0.00 0.00 0.00 3.67 123 Génot 2005
Switzerland 3.00 11.00 39.00 27.00 15.00 5.00 0.00 0.00 0.00 0.00 3.1 153 Juillard 1984
Netherlands 2.07 7.25 25.48 43.02 17.68 3.94 0.41 0.07 0.07 0.07 3.82 1448 Willems et al.
2004
Germany 0.2 2.6 11.8 37.7 32.8 10.7 4.2 0.00 0.00 0.00 4.1 126 Schönn 1986
Germany 5.00 9.00 31.00 41.00 12.00 2.00 0.00 0.00 0.00 0.00 3.5 269 Gassmann and
Bäumer 1993
Belgium 3.8 10.3 43.3 33 8 1.3 0 0.00 0.00 0.00 3.36 312 Smets, Huybrechts
(Flanders) and Cerulis
personal
communication
Belgium 5.60 18.30 36.80 32.10 6.40 0.60 0.10 0.00 0.00 0.00 3.18 1706 Bultot and Groupe
(Wallonia) noctua personal
communication
Netherlands 1.0 5.5 20.1 44.7 22.6 5.9 0.2 0.00 0.00 0.00 4.01 477 Van Harxen and
(Achterhoek) Stroeken
personal
communication
347

Belgium 0.8 5.50 29.70 40.60 20.30 2.30 0.80 0.00 0.00 0.00 3.85 123 Vanden Wyngaert
(Dijleland) 2005
 Chapter 8: Breeding Season

Table 8.12 Average clutch sizes of the Little Owl recorded across Europe

Mean
No. of clutch
Country Year clutches size Range Author

Britain 1939–1977 268 3.59 1–7 Glue and Scott 1980

Britain 1993–2000 115 2.65 1–4 Leigh 2001a
France 1918–1950 80 3.91 3–5 Labitte 1951
France 1984–2004 123 3.67 1–7 Génot 2005
Switzerland 1973–1980 153 3.12 1–6 Juillard 1984
Switzerland 1984–2000 189 3.70 – Meisser and Albrecht
2001
Germany 1974–1984 96 3.61 2–6 Exo 1987
Germany 1973–1988 265 4.42 1–8 Knötzsch in Schönn
et al. 1991
Germany 1960–1985 126 4.11 1–7 Schönn 1986
Germany 1971–1997 326 3.60 – Furrington 1998
Germany 1969–1988 161 4.13 2–7 Ullrich in Schönn et al.
1991
Germany 1974–1982 154 3.85 2–7 Illner in Schönn et al.
1991
Germany 1977–1985 153 3.65 2–5 Kimmel in Schönn
et al. 1991
Germany 1988–2001 812 3.71 – Keil unpublished
Germany 1978–1992 269 3.51 – Gassmann and
Baümer 1993
Germany 1974–1994 486 4.15 – Kämpfer-Lauenstein
and Lederer 1995
Hungary 1910–1976 89 5 3–8 Nemeth in Schönn
et al. 1991
Romania 1901–1975 21 5.24 4–7 Kohl in Schönn
et al. 1991
Russia 1989–2002 31 4.87 3–8 Il’yukh 2002
Czechoslovakia – 37 4.76 3–9 Hudec et al. in Schönn
et al 1991
Netherlands 1971–2022 17386 3.87 1–8 Van Harxen, Stroeken,
Boudewijn and van
Turnhout 2023
Netherlands 2003 17 3.33 2–4 Blanke 2003
Belgium 2000–2004 312 3.36 1–6 Smets, Huybrechts
(Flanders) and Cerulis personal
communication

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 8.9 Clutch Size

Table 8.12 (cont.)

Mean
No. of clutch
Country Year clutches size Range Author

Belgium 1999–2004 1706 3.18 1–7 Bultot and Groupe

(Wallonia) noctua personal
communication
Belgium 1999–2004 123 3.85 1–7 Vanden Wyngaert
(Dijleland) 2005

Clutch size is influenced by a number of factors, such as weather, food availability,

timing of laying, population density, age of female, geographic location, etc.. In Europe,
clutch size increases from west to east and decreases from south to north. Changes in
clutch size are thought to be related to climatic differences, with the owls’ preference
for semi-arid conditions (Exo 1992). In France, Génot (1992b) reported an increase
in clutch size in relation to the climatic shift from oceanic west to the south. Soler and
Soler (1992) hypothesized, in their study of latitudinal trends, that there is a significant
positive correlation between egg and clutch sizes and increasing latitude.
The average clutch size can vary substantially from year to year. Van Harxen and
Stroeken (personal communication) reported in their long-term study (1998–2021) that
the annual average clutch size ranged from 3.55 in 1991 to 4.49 in 2019, i.e., a
difference of almost one egg. They also found significant differences between individual
females (Table 8.13), e.g., 24 females that bred for five years each laid an average of
3.9 eggs.
At the level of the individual female, the differences were even greater, e.g., in four
years, female 016 laid nine eggs (2, 3, 2, 2) and female 252 laid 21 (6, 5, 4, 6).
Clutch size is affected by prey populations, especially small mammals, before the
laying phase. On average, females will lay 1.0–1.2 eggs more in the peak vole cycle years
than in low vole years (Exo 1992). The influence of rodents on clutch size has been
shown in captivity, where Little Owls easily produce large clutches (seven to eight eggs)
after optimal feeding during the prelaying period (Robiller and Robiller 1986). Large
clutches in the field were found in peak vole years before 1945 (Collinge 1922,
Uttendörfer 1939). Gassmann and Bäumer (1993) found a correlation between larger
clutch sizes and the amount of precipitation in March, before the egg-laying season.
Clutch size is shown to be density dependent. Pairs that breed in clusters lay fewer
eggs than pairs that nest in isolation. Bultot et al. (2001) found, in their study in
Wallonia (Belgium), that clutch sizes in clustered pairs laid, on average, 2.5 eggs,
whereas isolated pairs laid 3.
Clutch size is shown to be related to the egg-laying initiation date. Bultot et al.
(2001) and Finck (1989) both indicated that a larger clutch size was associated with
earlier laying dates. Van Harxen and Stroeken (personal communication) also found a

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 Chapter 8: Breeding Season

Table 8.13 Average clutch size per total years of active breeding

Breeding Ave. clutch No. of Min clutch Max clutch

years size females size size Range

1 3.98 264 1 7 6
2 4.1 125 2 7 5
3 4.12 69 2.33 6 3.67
4 3.95 35 2.25 5.25 3
5 3.9 24 2.2 4.8 2.6
6 3.98 15 2.83 5.17 2.34
7 3.98 8 3.14 4.57 1.43
8 3.77 7 3 4.38 1.38
9 3.87 5 3.67 4.41 0.74
10 4.8 1 0
14 3.64 1 0

Van Harxen and Stroeken personal communication

4.5

3.5
Average Clutch Size

2.5 y = –0.0359x + 4.3095

R² = 0.3674
2

1.5

0.5

0
0 5 10 15 20 25
Egg-Laying Initiation Day in April

Figure 8.13 Linear relationship between clutch initiation date and clutch size (1998–2021, n = 1075)
(Van Harxen and Stroeken personal communication).

positive relationship between the clutch initiation date and clutch size in the period
1998–2021. The earlier the first egg was laid, the larger the average clutch size
(Figure 8.13) (average clutch size = 4.3095 (on April 7) – 0.0359 eggs per day delay in
clutch initiation).

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 8.9 Clutch Size

4.20

4.10 y = –0.0123x2 + 0.0876x + 3.8472

R² = 0.7094
4.00

3.90
Average Clutch Size

3.80

3.70

3.60

3.50

3.40

3.30

3.20
1 2 3 4 5 6 7 8 9 10
60 55 56 56 55 42 27 18 11 7
Number of Breeding Seasons
Number of Females

Figure 8.14 Average clutch size per number of breeding seasons (second x-axis includes the number
of females) (n = 61 females, 390 clutches).

There are large differences in average clutch size between territories, mainly deter-
mined by the occupying pairs rather than the habitat. In 45 territories where the clutch
size had been determined at least 10 times, the average number of eggs ranged from 3.5
(14 years) to 4.63 (16 years). The difference is mainly accounted for by female 479, who
laid an average of 3.14 eggs per year over 7 years. Female 218 produced an average of
4 eggs per year in the same territory over three years. In territory 378, the good results
are mainly accounted for by female 317, who laid an average of 4.8 eggs per year over 10
years. The four other females produced an average of 4.3 eggs per year over 6 years.
Van Harxen and Stroeken (personal communication) found decreasing clutch size in
relation to the age of the female in those that were found to have bred at least five times
(Figure 8.14). On average, the highest clutch size was found between the third and fifth
breeding season (not necessarily corresponding with their exact age if found breeding
when unringed). In the first two years, clutch size was slightly less (3.91) compared to
the next three years (4.07). After the fifth breeding season, the average clutch size
dropped to 3.73. A female that bred for 14 consecutive years laid respectively 4, 3,
3 and 1 eggs in her last four years (4.0 on average in the 10 years before).
This corresponds to age-dependent reproductive success following a pattern where
reproductive success reaches an optimum in individuals of middle age. Bekaert (2006)
found the optimum number of nestlings with females aged 5–7 years. Breeding success
decreased again in older individuals. The clutch size reached an optimum for individ-
uals of 7 years. The number of fledglings had an optimum for females between 5 and
7 years old.

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 Chapter 8: Breeding Season

4.4
y = 0.0066x + 3.6706
R² = 0.1413
4.2

4
Average Clutch Size

3.8

3.6

3.4

3.2

3
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017
2018
2019
2020
2021
Year

Figure 8.15 Average clutch size per year 1976–2021 (n = 16 037, overall average 3.89) Source:
Meetnet Nestkaarten van Sovon Vogelonderzoek Nederland (Nienhuis et al. 2015).

Females whose exact age was known and who participated directly in reproduction
during the first breeding season (n = 66) had an average clutch size in their first season
that was 0.22 eggs higher than the long-term average for the population (4.26–4.02). Of
these, the 50 who had their first breeding season in the mouse-rich period of 2014–2021
did slightly better and had an average clutch size of 4.36 eggs, compared to 4.07 for the
entire population.
In a long-term analysis of the reproduction of the Little Owl in the Netherlands (Van
Harxen and Stroeken personal communication), clutch size gradually increased from
the mid-1970s onwards (Figure 8.15).
In the period 1976–1984, more than two-thirds of the data came from natural
cavities in trees in a limited area in the Betuwe and for the years 1984–1989 this was
still the case for just over one-quarter of the clutch sizes. After 1989, this applied on
average to just over 1% of the clutches. Clutches in natural burrows are not always easy
to control and possibly eggs were missed and the actual clutch size might have been
higher (J. van de Laar personal communication). From 2014 onwards, there were also a
series of good years (especially 2014 itself and 2019) that may have leveraged the trend.
The average clutch size per type of nesting site also shows that clutches in tree
cavities are, on average, somewhat smaller than those in buildings and nestboxes
(Table 8.14).
Just over 40% of clutches contained four eggs, clutches with three and five eggs were
fairly balanced with 25% and 20%, respectively (Figure 8.16). Together, these
accounted for 88% of the clutch sizes. Clutches with six eggs were not uncommon,
but even larger clutches (seven and eight eggs) occurred only sporadically. For clutches

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 8.9 Clutch Size

Table 8.14 Average clutch size and frequency per nest site type (1976–2021)

Type Average clutch size n

Nestbox 3.89 14 846

Cavity in building 3.82 211
Cavity in tree 3.62 390

Van Harxen and Stroeken personal communication

45 6747

40

35

30
3921
Percentage

25
3216
20

15

10
938
626
5 273
72 22
0
1 2 3 4 5 6 7 8
Clutch Size

Figure 8.16 Frequency distribution of clutch size in the Netherlands 1971–2021 (n = 15 815). Source:
Meetnet Nestkaarten van Sovon Vogelonderzoek Nederland (Nienhuis et al. 2015).

with one egg there is usually something incomplete and they have been abandoned
prematurely. This might also explain some of the two-egg clutches.
Eight clutches with nine eggs were found, three with 10 eggs, one 11-egg clutch, two
12-egg clutches and one clutch of 13 eggs. These were exceptional situations where
there may have been two females sharing the nestbox. This was confirmed for the 11-
egg clutch (Van Dooren and Hermsen 2019) and one of the nine-egg clutches (Grooters
and Grooters 2006). In two cases, the observer even suggested the possibility of three
females. Some of the seven- and eight-egg clutches might be related to double-
clutches too.
There are clear differences between bad and good micromammal years (Figure 8.17).
In bad years more clutches with two and three eggs, far fewer with five eggs and almost
none with more than five occur, in good years the reverse is observed, more clutches
with five and six and also with seven and eight eggs, but fewer with two and three eggs.
It is remarkable that the number of clutches with four eggs varies much less (40.2,
42 and 35.9) suggesting a four-egg clutch is optimal.

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 Chapter 8: Breeding Season

45

40

35

30
Percentage

25

20

15

10

0
1 2 3 4 5 6 7 8
Clutch Size
2003+2013 Poor mouse years All years 2014+2019 Rich mouse years

Figure 8.17 Frequency distribution of clutch size for all years n = 15 915, good (2014+2019) (n =
2287) and bad micromammal years (2003+2013) (n = 1197). Source: Meetnet Nestkaarten van Sovon
Vogelonderzoek Nederland (Nienhuis et al. 2015).

8.10 Replacement Clutches

In case the first clutch is lost, replacement clutches can be laid, especially when the first
clutch is early and prey resources favorable (Knötzsch 1978, Génot 2005).
In the Winterswijk area of the Netherlands, replacement clutches were recorded at
27 (8%) of 337 failed nests between 2001 and 2020 (Van Harxen and Stroeken 2022).
There were differences between years. Replacement clutches were recorded in 3 of 15
failed nests in 2014 (when voles and mice peaked), to none of 19 in 2016. Of 19 first
clutches, 15 failed before hatching and in four cases there were nestlings. Known causes
of failure in the first clutch were predation by a Stone Marten and a Squirrel, death of
the male/female, honeybees occupying the nestbox, eggs failing to hatch and disturb-
ance by humans. The mean laying date for the first egg in 16 replacement clutches was
May 24. Of 25 replacement clutches, 16 (64%) were successful and produced at least
one fledgling, while 9 (36%) failed for the second time.
On average, fewer eggs were found in 10 of 19 replacement clutches than in the first
clutches (3.7 in replacement versus 4.0 in first clutches); six replacement clutches had a
larger number of eggs than the first clutch and three had an equal number. The average
number of fledglings from the 19 nests was 1.7 for all nests and 2.7 fledglings for
successful nests. Of 21 nestlings weighed at the time of ringing, 13 young were a little
underweight and 8 were heavier than expected. Twelve were weighed again, just before
fledging. Each had gained weight, on average they were 6% heavier than expected.
In 19 of 27 replacement clutches the eggs were laid in the same nestbox, in 7 cases
the owls chose another nestbox in the same courtyard and once the owls moved 300 m

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 8.10 Replacement Clutches

to a neighboring territory. In 9 out of 12 cases the female that laid the first clutch and
the replacement clutch was the same individual, in 3 cases another female. In these
cases the initial female had died early, for unknown reasons.
Between 1984 and 2004, Génot (2005) recorded 8 replacements out of 64 failed
clutches (12.5%) in Voges du Nord (France). In two of the eight cases the owls chose a
different nestbox. Illner (in Schönn et al. 1991) reported on five replacement clutches in
Germany that contained slightly fewer eggs than the initial clutch: 3.4 versus 3.8 eggs.
Replacement clutches are uncommon, but rather successful. If there are sufficient food
resources to bring the female back into laying condition after the first clutch has been
lost, it is likely that there is also sufficient food to successfully raise young from re-
nesting attempts. As many prey items, such as beetles, larvae, caterpillers, earthworms
and young birds are less abundant later in spring, access to small mammals can affect
the outcomes. In 2020 in the Netherlands, all five eggs of a replacement clutch hatched.
The nestlings showed a decrease in weight and three of them died between days 4 and
18. The surviving two suddenly gained weight when, between day 32 and 39, the
meadow adjacent to the nestbox was mown: from 144 g to 185 g and from 142 g to
179 g, far above what would be normal at that age. A larder with four Common Voles
was even available, highly uncommon at this age.
2021 showed an unprecedented number of replacement broods in Winterswijk in
17 of the 49 failed clutches (34.7%). This “explosion” could partly be explained by the
fact that a relatively large number of nests failed as a result of the bad weather
conditions (cold, a lot of and prolonged rain) in May, while the feeding conditions
(i.e., available mice/voles) were good. Apparently, there was still enough prey when the
weather improved at the end of the month, with plenty of cockchafers available as they
appeared a few weeks later than usual due to the same bad weather conditions.
The average clutch size of the replacement clutches was 3.01 (n = 17), substantially
smaller than the average size of the first clutches (4.34, n = 126).
The average replacement clutch initiation date was May 28 (May 20–June 8, n = 10).
Compared to the average initiation for the first clutches in 2021 (Mean, Min-Max, n)
(April 13, March 25–May 12, n = 101), this was 45 days later.
Of the 17 replacement broods, 8 failed for the second time, 9 nests succeeded (53%),
yielding 20 fledglings (1  4, 3  3, 2  2 and 3  1), an average of 2.22 (n = 9) per
successful nest and 1.18 for all attempts (n = 17).
On the ringing day (average age 17 days), the 20 nestlings weighed, on average, 7%
more than expected. At the follow-up check (average age of 28 days), the average
condition had risen to 1.12 (16 young). Two loners were 26% and 31% heavier than
the expected value, respectively, weighing 178 g and 190.5 g at fledging.
In 14 of the 17 replacement broods, the female was identified. On 11 occasions it was
the same female (79%) and on three another (21%). Twice an alternative nestbox
was used for the replacement clutch, 28 and 110 m away from the first nest, probably
because of predation. The fact that the other clutches were laid in the same nestbox
might be due to the lack of alternative cavities.
Thanks to the nine replacement clutches, nesting success at the population level
increased from 64.7% to 71.2%, but still below the long-term average (74.2%, period

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 Chapter 8: Breeding Season

1998–2020). The average number of young per nest started rose from 2.14 to 2.29 due
to the replacement clutches, from below the long-term average (2.23, period
1998–2020, n = 2991) to above it. The condition of the young was 3% above that of
the first clutches and 7% above their expected weight. The large number of replace-
ments therefore also had a positive effect at a population level.

8.11 Incubation
Gavrin (1962), Bakaev in Sagitov (1990) and Glue and Scott (1980) reported that
incubation of the eggs began with the first egg laid. Exo (1983) and Ataev (1977) found
that incubation started after the laying of the second or third egg. In other cases,
incubation took place with the laying of the second to last egg (Illner in Glütz Von
Blotxheim and Bauer 1980, Robiller and Robiller 1986, Génot and Sturm 2001) or even
when the clutch was complete (Labitte 1951, Enehjelm 1969, Zav’yalov et al. 2000).
Illner (personal communication) believes that this is influenced by the clutch size, in
clutch sizes larger than two eggs, the brood patch is not fully formed until the later
stages of the clutch formation.
Camera observations in the Netherlands (Van Harxen et al. 2018a) have shown that
there is a gradual increase in incubation time. Although the female incubates for a short
time right from the first egg, it takes until the last egg is laid before she spends more
than 90% (22 hours) of the day on the eggs (Figure 8.18).
While the period of incubation has often been reported as 28 days, actual measure-
ments indicate variations. Exo (1983) found the incubation period to range between
27 and 33 days, Juillard (1984) reported 28 days and Ullrich (1973) recorded an
incubation period of 35 days. When Génot and Sturm (2001) studied the breeding
cycle of 25 captive Little Owls, incubation ranged between 18 and 29 days, an average
of 25 days. An average incubation period of 25 days was also found by Robiller and
Robiller (1986). The shortest period of incubation was 19.5 days, recorded for an owl in
captivity (König 1969).
Due to the gradually increasing number of hours that the female incubates, an exact
incubation duration is difficult to give. Calculated from the laying of the first egg, it
took an average of 31 days from the laying of the first egg in seven nests observed with
cameras, almost 25 days from the last egg and just over 20 days from the 90% limit
before the first egg hatched (Table 8.15). If the incomplete days at the beginning and
at the end are taken into account the average incubation duration is 25.2 days (Van
Harxen et al. 2018b).
Incubation breaks occur during the entire incubation period up to more than 20 per
day, with a combined duration ranging from 0:54 to 2:34 in seven observations
(Table 8.16).
Abdusalyamov (1971) suggested that both sexes participate in incubation, while
Gavrin (1962) noted that participation of the male in incubation is not clear because
both birds always stayed near the nest. Camera observations at 23 nests have shown
that only the female incubates. In her absence, the male sporadically made an attempt

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 8.11 Incubation

(a)
1 24:00
0,92 22:00
0,83 20:00
0,75 18:00
0,67 16:00
Hours of incubaon

0,58 14:00
0,50 12:00
0,42 10:00
0,33 8:00
0,25 6:00
0,17 4:00
0,08 2:00
0,00 0:00
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31
–0.083333
Days as of initiation of incubation
Egg Laying
(b) (c) (d)

(e) (f) (g)

Figure 8.18 Incubation times per day from eight camera observations. Black bars indicate egg-laying.
The horizontal black line indicates the 22-hour limit (Van Harxen, Stroeken and Sterringa personal
communication), (a) 2015 (b) 2016 (c) 2017 (d) 2018 (e) 2019 (f) 2020 (g) 2021.

to take a seat on the eggs. As soon as the females entered, however, he immediately gave
up his place. These kinds of attempts never lasted more than a few minutes (Van
Harxen and Stroeken personal communication).
In years of low prey resources the male struggles to provide enough food to the
breeding female. In cases of shortages she will leave the nest more often, leading to an
increase in egg deaths (Leigh 2001a).
The data from the camera observations were too limited to establish a solid relation-
ship between the number of incubation breaks, their duration, the prey supply and the
hatching of the eggs. However, it is striking that the relatively small number of breaks in
Dg17 was accompanied by a high prey supply. The female received 1.3 kg of prey
biomass during the laying and incubation period, including 66 mice. The number of
breaks in Hx18, on the other hand, does not seem to be related to the prey supply in the

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 Chapter 8: Breeding Season

Table 8.15 Number of incubation days for seven camera-observed breeding seasons

Number of incubation days

Clutch size Laying date First egg Last egg 22-hour limit Average

Hx15 5 Apr 25 31 23 21 25.0

Hx16 4 Apr 20 31 24 20 25.0
Hx17 4 Apr 14 32 24 20 25.3
Hx18 4 Apr 11 30 24 19 24.3
Hx19 3 Mar 31 30 25 19 24.7
Hx20 3 Mar 30 31 25 20 25.3
Dg17 3 Apr 01 31 27 23 27.0
Average 3.7 Apr 10 30.9 24.6 20.3 25.2

Van Harxen, Stroeken and Sterringa personal communication.

Table 8.16 Incubation breaks in seven camera-observed breeding seasons

Hx15 Hx16 Hx17 Hx18 Dg17 Dg18-I Dg18-II Ave.

Total number 329 390 318 297 156 331 219 291
of breaks
Average 15.7 19.5 15.9 15.6 6.8 11.8 8.4 13.4
number of
breaks per
day
Average time 01:23 01:28 01:11 01:35 00:54 01:15 02:34 01:28
per break
per day

Van Harxen, Stroeken and Sterringa personal communication.

other Hx locations, but to Dg17. The large number of breaks in Hx18, and their long
duration in combination with the low supply, did not lead to egg mortality (Table 8.17).

8.12 Hatching
Glütz Von Blotzheim and Bauer (1980) mentioned that eggs hatched at about one-day
intervals. In an aviary, Exo (1983) recorded that of nine clutches (containing 15 eggs),
two eggs hatched simultaneously and in three cases three eggs hatched during an 18-
hour period. Also in an aviary, Génot and Sturm (2001) found that of 22 clutches, 4 had
simultaneous hatching and that the eggs in the remaining clutches hatched over an

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 8.12 Hatching

Table 8.17 Number of incubation breaks and prey supply for five camera-observed
breeding seasons

Hx15 Hx16 Hx17 Hx18 Dg17

Total number of breaks 329 390 318 297 156

Total time during breaks 29:14 29:28 23:59 30:11 20:58
Clutch size 5 4 4 4 3
Hatching 4 3 4 4 3
Prey supply (g) 983 1070 1191 527 1353

Van Harxen, Stroeken and Sterringa personal communication.

Table 8.18 Hatching interval for 12 camera-observed breeding seasons

Clutch No. of Second Third Fourth First–last

size hatchlings interval interval interval interval

Hw05 4 4 1:00 0:16 3:15 4:31

Hv07 4 3 3:42 14:06 17:48
Hv09 4 4 9:45 4:59 0:58 15:42
Hv10 4 2 0:01 24:01
Hv11 4 2 18:04 18:04
Hx15 5 4 ? – 3:45 13:43
Hx16 4 3 3:15 7:32 10:47
Hx17 4 4 5:49 4:02 4:54 14:45
Dg17 3 3 6:08 5:26 11:34
Hx18 4 4 3:59 1:43 20:26 26:08
HX19 3 2 2:41 2:41
Hx20 3 3 1:29 6:20 7:49

Average interval 13:57

Extremes due to unhatched eggs in between in bold.

Van Harxen, Stroeken and Sterringa personal communication.

average of 1.2 days (range 1–4 days). Ataev (1977) found that chicks hatched over a
three-day period.
Van Harxen and Stroeken (2018) found an average period between the hatching of
the first and the last young (Table 8.18) ranging from 2:41 hours (two young) to 26:08
hours (four young) for 14 clutches that were under camera observation in the period
2005–2020 (five different females). In nests with four hatched eggs, it took an average of
15 hours before the last young hatched. In the four nests with three hatched eggs it
took an average of 12 hours and in the four nests with two hatched eggs just over

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 Chapter 8: Breeding Season

12 hours. In the nests with two hatched eggs, the average was strongly influenced by
the fact that in total only 8 of the 16 eggs hatched and that it seemed that in at least two
cases it was the last eggs laid that hatched. Leaving these out, it took an average of 3:13
hours for both eggs to hatch.
On average, it took 7:15 hours for the second egg to hatch, 5:33 for the third and
6:39 for the fourth. If we leave the extremes due to unhatched eggs in between aside,
i.e., second egg HV11, third egg Hv07 and fourth egg Hx18, it took 6:10 hours for the
second, 5:33 hours for the third and 3:13 hours for the fourth egg.
Van Harxen and Stroeken (2001) recorded the hatching of a Little Owl as follows:
“On May 26 14:45 we found two owlets and two eggs in one of our monitored nest sites.
The young owls were only a few hours old as their down was still damp. The weights of
the owlets were 10.8 g and 10.6 g. One egg, which had not hatched, was emitting a
peeping sound, and the owlet inside was breaking out of the egg with its beak and egg
tooth. The weight of this egg was 12.7 g. Whilst we were marking the other two young
for individual recognition at our next visit, the egg broke in the middle. The owlet was
pushing the eggshell back with its head, trying to free itself, resulting in the egg
splitting into two pieces. First the owlet’s head became more visible and then the owlet
released itself from the eggshell. The whole process took approximately two minutes,
after which the owlet was exhausted, and lay down motionless for a few seconds. The
newly hatched owlet weighed 10.1 g and the eggshell 1.7 g.”
The average weight of newly hatched owlets has been reported as 11 g and 12 g by
Mikkola (1983) and Schönn et al. (1991), respectively.
Hatching success is defined as the proportion of eggs that hatch divided by the total
number of eggs laid. The hatching success from Germany has been reported in several
studies (Table 8.19). Schönn (1986) reported that 62% (258 of 416) of the eggs recorded
in his study nests hatched. Similarly, Exo (in Schönn et al. 1991) found a hatching
success of 67% (180 of 269 eggs), Knötzsch (in Schönn et al. 1991) reported a hatching
success of 79% (925 of 1171 eggs) and H. Keil (personal communication) noted 86.5%
(2609 of 3016 eggs). In Switzerland, Meisser and Albrecht (2001) reported a hatching
success of 81% (566 of 700 eggs) and Juillard (1984) found the rate to be 71% (339 of
418 eggs). Génot (2005) reported a hatching success of 48% (223 of 452 eggs) in France.
J. Bultot (personal communication) found a hatching success rate of 83% (6265 of 7536
eggs) in Belgium for 1989 to 2004. Smets et al. (personal communication) found a
hatching success rate of 80% (1035 of 1284 eggs) in Belgium.

8.13 Egg Failure

The reasons why eggs do not hatch are diverse and range from being unfertilized to the
death of nearly mature embryos. Out of 372 unhatched eggs examined in the period
2014–2021, Stroeken and Van Harxen (personal communication) (Table 8.20) found
167 embryos in 459 eggs (36.4%) at all stages of development. This does not necessarily
mean that the other eggs were unfertilized. With the naked eye, an embryo is only
visible from the third day of life (Van den Burg 2017) and in many eggs the rotting
process had already progressed so far that the contents could no longer be examined.

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Table 8.19 Hatching success of the Little Owl across Europe

Eggs (clutch Hatched Fledged

Country number) (%) (%) Failed (%) Author

Britain 477 (156) 269 (56.4) 234 (49) 243 (51) Glue and Scott 1980
Britain 305 (120) 265 (86.9) 253 (83) 52 (17) Leigh 2001
France 452 (123) 223 (49.3) 201 (46.6) 229 (50.7) Génot 2005
Switzerland 478 (153) 339 (71) 279 (58.4) 199 (41.6) Juillard 1984
Switzerland 700 (189) 566 (81) 358 (51) 342 (49) Meisser and Albrecht 2001
Belgium 984 (257) 689 (77) 585 (65.4) 309 (34.6) Bultot et al. 2001
Germany 1171 (265) 925 (79) 706 (60.3) 465 (39.7) Knötzsch in Schönn et al. 1991
Germany 416 (102) 258 (62) 201 (48.3) 215 (51.7) Schönn 1986
Germany 3016 (812) 2609 (86.5) 2238 (74.2) 778 (25.8) Keil unpublished
Netherlands 1886 (471) 1473 (78,1) 1073 (56.9) 813 (43.1) Stroeken and van Harxen 2003
Belgium (Flanders) 1044 (312) 836 (80.1) 614 (58.8) 430 (41.2) Smets, Huybrechts and Cerulis personal communication
Belgium (Wallonia) 7536 (2346) 6265 (83,1) 5476 (72.6) 2063 (27.4) J. Bultot and Groupe noctua personal communication
Belgium (Dijleland) 473 (123) 344 (72.7) 273 (58) 200 (42) Vanden Wyngaert 2005
361
 Chapter 8: Breeding Season

Table 8.20 Breakdown of unhatched eggs (n = 459) and malformed embryos

(n = 167) 2014–2021

Unhatched All
eggs 2014 2015 2016 2017 2018 2019 2020 2021 years

Eggs 30 43 50 55 55 64 75 87 459
examined
Number of 11 21 10 21 22 16 28 38 167
eggs with
visible
embryo
Percentage 36.7 48.8 20.0 38.2 40.0 25.0 37.3 43.7 36.4
of eggs
with visible
embryo
Embryos
Total 11 21 10 21 22 16 28 38 167
number of
embryos
Number of 0 5 4 6 9 8 1 7 40
embryos
with visible
defect(s)
Percentage 0.0 23.8 40.0 28.6 40.9 50.0 3.6 18.4 24.0
with visible
defect(s)
Percentage 0.0 11.6 8.0 10.9 16.4 12.5 1.3 8.0 8.7
with visible
defect(s)
as part of
the total
number of
eggs
examined

Van Harxen and Stroeken personal communication.

Of 167 embryos, 40 (24%) were found to show an apparent abnormality (Stroeken and
Van Harxen personal communication): a lowered beak, crossed legs, spread leg, open
abdominal wall, extended neck, absence of brain, etc. Some embryos had multiple
handicaps: a combination of crossed legs, down-folded beak and too long wings, open
abdominal wall or missing leg. Many of these abnormalities are due to nutritional
deficiencies, i.e., lack of essential amino acids (Van den Burg 2017). Of the total of

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 8.14 Prolonged Incubation

459 eggs examined, 8.7% were found to contain an embryo with abnormalities
(Table 8.20).
Eggs can also not hatch due to overheating (Beersma and Beersma 2000). Embryos
that have been exposed to this show a striking red color. It also occasionally happens
that an embryo lies incorrectly in the egg so that the air chamber cannot be accessed.
Often this is a result of insufficiently turning the egg, typical for species that do not
immediately start incubation at the first egg. Flea droppings can sometimes cover an egg
in such a way that gas exchange becomes difficult or even impossible (Van den Burg
2017). Van Harxen et al. (2017) classified 44 unhatched eggs in 2015 based on the
extent to which they were smeared with flea droppings. Four eggs were so smeared that
gas exchange seemed impossible and for nine others the contamination was such that
gas exchange seemed at least difficult, altogether 22.7% of the eggs. Only 13 eggs
(29.5%) were completely free of flea droppings. In the years that followed, no such
degree of contamination was detected again.
In addition to infertility, loss of or abandonment by the adults, problem with the
nest structure or other complications, eggs are frequently lost to predators. The list of
predators of eggs includes: Magpie Pica pica, Fox Vulpes vulpes, Ermine Mustela erminea,
Hedgehog Erinaceus europaeus (Glue and Scott 1980), Jackdaw Corvus monedula (Staats
Von Wacquant-Geozelles 1890), Brown Rat Rattus norvegicus, Squirrel Sciurus vulgaris
and Dormouse Glis glis (Juillard 1984); the Stone Marten Martes foina is a particularly
effective predator of eggs or young of the Little Owl, especially in nestboxes without
protection systems (Furrington 1979, Schwarzenberg 1981, Schönn 1986, Génot
1992c).

8.14 Prolonged Incubation

Out of a total of 1553 complete (first) clutches of which the hatching success was
known, Van Harxen and Stroeken (personal communication) found 266 clutches where
none of the eggs hatched (17.1%). In an unknown number of cases, the eggs did not
hatch because they were not fertilized or because the embryos had died.
In such cases, the female often incubates the eggs (much) longer than usual. It is
then only approximately possible to determine how long the brood has been incubated
because the clutch start (and derived from this the breeding start) cannot be calculated
on the basis of the age of the young and it is also not known exactly when the female
stopped incubating. Schönn et al. (1991) reported “taube Eier” (unfertilized eggs) that
were incubated for 44 to 49 days. It is unclear whether these were individual eggs or
complete clutches.
Van Harxen et al. (2023) observed an incubation time of 60 days by taking into
account the first date with female and eggs, April 28, and the last date with female and
eggs, June 17, and respectively 67 days by taking June 24 into account, with only eggs
and no female observed, controlling for an average clutch initiation date that year of
April 14 and assuming that the first visit took place halfway through the incubation
period, and three to four days passed until the first visit without a female.

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 Chapter 8: Breeding Season

In 2021, the eggs did not hatch in a camera nestbox at Beleef de Lente. Because this
nestbox was under camera surveillance 24 hours a day from mid-February to the end of
July, the entire process from the laying of the first egg to the final abandonment of the
eggs could be followed (Figure 8.19). After the normal incubation period had expired
the eggs did not hatch. Nevertheless, the female continued to incubate relentlessly,
averaging just over 23 hours a day, i.e., until May 27, or 31 days after the eggs were due
to hatch (phase 3, prolonged incubation phase). Calculated from the moment she
started incubating, the female had been on the eggs for 56 days, only interrupted by
short trips, with a total duration of an average of less than an hour per day, i.e., more
than double the normal incubation time (Van Harxen et al. 2018a).
The incubation period of this female could be divided into four phases: egg laying (8
days), normal incubation phase (25 days), extended incubation phase (31 days) and
tapering phase (10 days). She spent an average of more than 23 hours per day in the
nestbox in both the incubation phase and the extended incubation phase. Only in the
last phase did the incubation time gradually decrease from more than 21 hours to zero.
As of day 66 the female was no longer constantly sitting on the eggs when she was in
the nestbox. An increasing part of the time she spent standing or sitting next to the
eggs, taking care of her feathers or resting (Figure 8.20).
On May 31 (day 68) the female stayed away for the first time for a whole night; that was
also the day on which she stood or sat next to the eggs for most of the time during the day.
That day she only incubated for seven hours. In the following days, the incubation time
gradually decreased. Six days later (day 75, June 6) no incubation took place for the first
time for a whole day. Although she still spent a short time on the eggs on days 76 and 78
(eight and seven minutes, respectively), day 74 was considered the last day of incubation.
The whole process – from the first egg being laid to stopping incubation – had then taken
two and a half times the normal duration. All this time, the male had continued to deliver
prey. The reason for the nonhatching remains unclear. The three eggs were all found to
contain a dead embryo, which died at different times during their development.

8.15 Number of Nestlings

The number of nestlings per brood is mostly expressed as the number on the ringing
day. In a long-term study in the Netherlands, this varied per year from 2.07 in 1976 to
3.63 in 2019, with a tendency to increase over the years (Figure 8.21).
There was much variation between years in different areas, as shown in Table 8.21,
showing that the number of nestlings differ not only per year but also per region and a
good year in one region does not automatically mean a good year in another region,
suggesting that local differences play an important role. Figure 8.22 illustrates the
geographical distribution of the ringed clutches.
The smaller average of nestlings in the western part of the Netherlands might be
related to habitat quality, as these are also the regions with lower densities and isolated
populations. The maturity of conservation activities might have had an impact locally,
causing extra variation.

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24:00
23:00
22:00
21:00
20:00
19:00
18:00
17:00
16:00
Hours by Female in Nestbox

15:00
14:00
13:00
12:00
11:00
10:00
9:00
8:00
7:00
6:00
5:00
4:00
3:00
2:00
1:00

-23 -20 -17 -14 -11 -8 -5 -2 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49 52 55 58 61 64 67 70 73 0 0

Days in Relation to Egg Initiation

Figure 8.19 Presence of the female in the nestbox (in hours per day) from March 1, 2021 to June 11, 2021 with egg-laying phase, normal incubation phase,
extended incubation phase and tapering phase indicated.
 Chapter 8: Breeding Season

0:00
23:00
22:00
21:00
20:00
19:00
18:00
17:00
Hours of female in nestbox

16:00
15:00
14:00
13:00
12:00
11:00
10:00
9:00
8:00
7:00
6:00
5:00
4:00
3:00
2:00
1:00
0:00
29/05/2021

30/05/2021

31/05/2021

1/06/2021

2/06/2021

3/06/2021

4/06/2021

5/06/2021

6/06/2021

7/06/2021

8/06/2021

9/06/2021

10/06/2021

11/06/2021
Sitting on the eggs Presence in the nestbox
Date

Figure 8.20 The residence time of the female on the eggs (incubation) in relation to the total presence
in the nestbox in the tapering phase from day 66 (May 29) to 79 (June 11).

4
Average number of nestling at ringing date

y = 0.0056x + 3.1164
R² = 0.0503
3.5

2.5
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
2016
2017
2018
2019
2020
2021

Year

Figure 8.21 Evolution of the average number of nestlings on the ringing date 1996–2021 in the
Netherlands (n = 17 824, mean = 3.17) (Van Harxen and Stroeken personal communication).

At almost 60%, broods with three or four nestlings were most common. Less than
2% had six or more nestlings (Figure 8.23).
Van Harxen and Stroeken (personal communication) found a clear relationship
between the abundance of voles and mice found in their long-term study in the eastern
part of the Netherlands and the frequency distribution of nestlings on the ringing date
in the complete Dutch dataset. In good years, there were more broods with five and six
nestlings and in bad years more with two and three (Figure 8.24).

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Table 8.21 Average number of nestlings on the ringing date per year with at least 10 records per location in the Netherlands (n = 5100, average
41 per location per year)

Lat. Long. 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 Ave.

Gaanderen e.o. 5156’49” 621’24” 3.54 3.06 2.68 2.97 2.98 3.48 2.97 3.45 3.09 3.35 3.16 2.91 3.90 3.40 3.26 3.21
Zuidoost- 5159’18” 636’01” 3.42 3.28 2.86 2.82 3.00 3.40 2.95 3.40 2.55 3.41 3.10 3.33 3.67 3.09 3.28 3.17
Achterhoek
Doesburg 5200’93” 607’01” 3.40 2.93 2.91 3.25 3.50 3.27 2.57 2.86 3.08 3.17 3.00 3.19 3.70 3.02 3.13
NW-Veluwe 5214’98 603’58” 3.65 3.45 3.07 3.29 2.83 3.36 2.71 3.63 3.94 3.16 3.10 3.08 3.27
Hellendoorn 5224’12 629’05” 3.82 3.70 3.55 3.04 3.10 2.79 3.58 2.83 3.34 3.47 3.30 3.46 3.37 3.02 3.31
Land van Cuyk 5130’36” 556’37” 4.18 3.22 3.69 3.93 3.88 3.00 3.64 3.19 3.41 3.30 3.32 3.95 3.35 3.50 3.54
Rijssen 5219’14” 631’16” 3.36 3.63 3.68 3.73 3.52 3.11 2.57 3.74 3.15 3.20 3.51 3.74 4.06 3.11 3.44 3.44
Zak van Zuid- 5125’02” 349’82” 2.88 2.38 2.00 2.53 2.88 2.47 2.19 3.40 2.57 2.60 2.87 2.62 3.13 2.61 2.45 2.64
Beveland
Warnsveld 5208’82” 615’09” 3.82 3.33 2.89 3.70 2.74 3.73 2.96 3.21 3.32 2.86 3.97 3.09 3.18 3.29
Heeten 5220’04” 616’65” 3.61 3.19 4.17 3.04 3.89 3.44 3.74 4.24 3.76 3.24 3.63
Neede 5208’54” 635’43” 3.26 3.07 3.15 2.87 2.84 3.21 3.10 3.76 3.27 3.64 3.75 3.27
Twisk 5244’59” 503’66 2.73 2.76 2.91 2.77 2.79
Wognum 5241’63” 507’02” 2.94 2.83 2.83 3.32 3.05 2.99
Udenhout 5135’74” 511’34” 3.12 3.20 3.68 3.43 4.64 3.28 3.77 3.59
Echt 5116’16” 556’43” 2.78 3.38 3.22 3.25 3.13 4.05 4.12 3.58 3.55 3.45
Betuwe 5157’08” 535’83” 2.79 2.70 2.81 2.45 4.00 2.65 2.64 3.13 3.00 2.54 3.04 2.92 3.00 3.15 2.85 2.87
Average 3.29 3.25 3.08 3.13 3.22 3.27 2.78 3.53 3.05 3.19 3.27 3.24 3.71 3.21 3.18 3.23
Min 2.79 2.38 2.00 2.45 2.83 2.47 2.19 2.86 2.55 2.54 2.87 2.62 2.76 2.61 2.45 2.64
Max 3.65 4.18 3.82 3.73 4.00 3.88 3.19 4.17 3.94 3.89 3.68 4.05 4.64 3.76 3.77 3.63
Variance 0.86 1.80 1.82 1.28 1.17 1.41 1.00 1.31 1.39 1.35 0.81 1.43 1.88 1.15 1.33 0.99
367
 Chapter 8: Breeding Season

Figure 8.22 Geographical distribution of the locations on which the average number of nestlings on
the ringing date was calculated.

8.16 Nest Success

Van Harxen and Stroeken (personal communication) showed that nesting success
(defined as the percentage of nests in which at least one juvenile fledged) varied
considerably between years, ranging from 51.2% in 2004 to 80.4% in 2013
(Table 8.22). On average, 72.1% of the nests are successful (1078 out of 1496).
The cause of failure could not be determined unambiguously in most cases, but the
loss of one of the partners, (egg) predation (by Squirrels, Weasels and Stone Martens),
unfertilized eggs, dead embryos and (human) disturbance turned out to be common

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 8.16 Nest Success

35
5328
30 5170

25
Percentage

20 3245

15
1843 1916
10

5
303
17 1
0
1 2 3 4 5 6 7 8
Number of Ringed Young per Nest

Figure 8.23 Frequency distribution number of young on the ringing date in the Netherlands
1971–2021 (n = 17 823). Source: Meetnet Nestkaarten van Sovon Vogelonderzoek Nederland
(Nienhuis et al. 2015).

40

35

30

25
Percentage

20

15

10

0
1 2 3 4 5 6 7 8
2003+2013 Poor mice/vole years All years 2014+2019 Rich mice/vole years
Number of Ringed Young per Nest

Figure 8.24 Frequency distribution of average number of young per nest on the ringing date for
all years (n = 17 613, mean = 3.17), for bad micromammal years (2003+2013) (n = 1164, mean =
2.82) and for good micromammal years (2014+2019) (n = 2155, mean = 3.58). Source: Meetnet
Nestkaarten van Sovon Vogelonderzoek Nederland (Nienhuis et al. 2015).

suspected causes. The low success in 2004 could at least partly be due to the extremely
low number of mice and voles. In 2015 a flea plague caused above-average failure and in
2021 bad weather in May was responsible for a considerable number of failed nests.
As shown in Table 8.19, the percentage of nest losses ranged from 17% in Britain
(Leigh 2001a) to 72% in France (Génot 2005). The loss of eggs and young from nests

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 Chapter 8: Breeding Season

Table 8.22 Percentage of successful nests in Zuidoost-Achterhoek (the Netherlands)

1998–2021

Year No. of nests Successful Failed Percentage successful

1998 44 38 6 86.4
1999 44 33 11 75.0
2000 41 29 12 70.7
2001 42 33 9 78.6
2002 49 35 14 71.4
2003 38 28 10 73.7
2004 41 21 20 51.2
2005 47 29 18 61.7
2006 46 33 13 71.7
2007 44 31 13 70.5
2008 45 31 14 68.9
2009 45 30 15 66.7
2010 46 34 12 73.9
2011 48 34 14 70.8
2012 54 39 15 72.2
2013 51 41 10 80.4
2014 68 53 15 77.9
2015 73 48 25 65.8
2016 79 60 19 75.9
2017 96 68 28 70.8
2018 99 77 22 77.8
2019 98 74 24 75.5
2020 123 93 30 75.6
2021 135 86 49 63.7
All years 1496 1078 418 72.1

Van Harxen and Stroeken personal communication.

can be attributed to many causes. Juillard (1984) reported losses of 31 complete clutches
to the following: 16 consisted of infertile eggs, 8 were predated, 2 were abandoned and
the cause for 5 others was unknown.
Of 48 nest failures reported by Schönn (1986), 15 were lost to predation, 4 were
infertile, 3 were abandoned, 2 were lost to flooding, 2 to human disturbance, 2 to the
nest falling, 1 due to a broken nest tree, 1 due to ants and 18 unknown.
Of 52 nest failures reported by Glue and Scott (1980), 28 were due to humans taking
the eggs or young, destroying the nest or shooting the adults, 8 were lost to predation,
and 16 were lost to other causes.

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 8.17 Egg Success

8.17 Egg Success

During 1998–2021, Stroeken and Van Harxen (personal communication) investigated
the percentage of the eggs that yielded a fledgling. This egg success averaged 55.95%,
but varied considerably between years ranging between 41.1% in 2004 and 64.6% in
2016. Remarkable is the relatively low recruitment success in the years 2002–2011 and
the high egg success of 2013 (Figure 8.25).
For the period 2013–2021, the causes of the difference between the number of eggs
laid and the number of fledged young were examined (Table 8.23). On average, 15.5%
of the eggs did not hatch (min 10.5% in 2013 and max 22.3% in 2015). The high
percentage in 2015 was probably caused by the fact that many eggs were covered with
flea droppings to such an extent that it made it difficult for the eggs to “breathe.” The
percentage of nestlings found dead (long-term average 3.3%) was also extremely high
that year at 10.1%, and below 0.8% in 2016. It mainly concerned young in the first
week of life who suffered badly from the flea infestation that year. On average, 15.9%
(min 11.6% in 2013 and max 21.7% in 2021) of the eggs were no longer present on the
second nestbox visit. Probably the majority of these eggs hatched, but the young died
before the second visit to the nest and were fed to the remaining nestlings. Some of the
eggs might have been eaten by the female or fallen victim to predation. For 2021 in
particular, eggs will have hatched, but the young could not be fed sufficiently due to
the bad weather in May and died fairly quickly after hatching. Finally, young that were
no longer present on the next visit to the nest, but not laid dead, were classified.

10.00
Percentage point difference in egg success relative to the average of 55.95%

5.00

0.00

-5.00

-10.00

-15.00
1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021
Year

Figure 8.25 Egg success 1998–2021, as deviation from the average (55.95%) (Van Harxen and
Stroeken personal communication).

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372

Table 8.23 Egg success statistics 2013–2021

2013 2014 2015 2016 2017 2018 2019 2020 2021 Average

N % N % N % N % N % N % N % N % N % N %

Eggs 181 228 247 260 286 343 409 427 535 2916
Unhatched 19 10.5 35 15.4 55 22.3 47 18.1 45 15.7 48 14.0 66 16.1 57 13.3 72 13.5 444 15.2
Eggs 21 11.6 31 13.6 40 16.2 36 13.8 40 14.0 41 12.0 65 15.9 75 17.6 114 21.3 463 15.9
“disappeared”
Nestlings dead 14 7.7 8 3.5 25 10.1 2 0.8 15 5.2 8 2.3 7 1.7 13 3.0 6 1.1 98 3.4
Nestlings 17 9.4 9 3.9 17 6.9 7 2.7 9 3.1 26 7.6 18 4.4 25 5.9 71 13.3 199 6.8
“disappeared”
Fledglings/egg 110 60.8 145 63.6 110 44.5 168 64.6 177 61.9 220 64.1 253 61.9 257 60.2 272 50.8 1712 58.7
success

Van Harxen and Stroeken personal communication.

 8.18 Condition

The most plausible explanation is that the majority died between visits and were fed to
the remaining young, removed from the nestbox by one of the parents or fledged too
early. The low egg success for 2015 can probably be largely attributed to the flea
infestation that year and for 2021 the weather conditions will have played a
decisive role.

8.18 Condition
In addition to known parameters such as clutch size, nest success and number of
fledged young, the condition of nestlings is also a useful tool to assess the reproduction
result. Stroeken and Van Harxen (2021) analyzed the condition of 3366 nestlings in
1069 nests in the period 1998–2021 by comparing the body weight measured on the
ringing day with the expected weight for the age and expressing this as the condition
index (CI). A CI of 1.0 means that the nestling weighs as expected, youngsters with a CI
higher than 1.0 are heavier than expected and a CI lower than 1.0 lighter
than expected.
They found that the average CI varied significantly from year to year, ranging from
0.95 in 2006 to 1.08 in 2019 (Figure 8.26). Calculated at day 19 (the average ringing
day), this meant that the youngsters in 2019 were on average almost 17 g heavier and
were a week ahead of those from 2006 in terms of weight (expected weight on day 19:
129.4 g, 2006: 122.9 g and 2019: 139.8 g).

1.09
Average condition index (CI) of nestlings at ringing date

1.07

1.05

1.03

1.01

0.99

0.97

0.95
2017
2018
2019

2021
2016
2013
2014
2010
2011
2006
2007
2008
2005
2003
2000

2002
1998
1999

2001

2004

2009

2012

2015

2020

Year

Figure 8.26 Average condition index (CI) of nestlings at ringing date per year, 1998–2021 (Stroeken
and Van Harxen 2022).

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 Chapter 8: Breeding Season

Average condion index (CI) of nestlings at ringing date

1.08

1.06

1.04

1.02

0.98

0.96

0.94

0.92

0.9
1 2 3 4 5 6 7
Brood size

Figure 8.27 Average condition index (CI) of nestlings at the ringing date per brood size (1069 nests,
4519 nestlings, 1998–2021) (Stroeken and Van Harxen 2022).

The variation in the CI was largely explained by the number of micromammals

(corrected for the number of nests visited in the first 10 days of the young period) in
larders (see Chapter 6, Diet).
The average condition decreased with the number of nestlings per brood. The
average CI only fell below 1.0 in nests with six or seven young (Figure 8.27), illustrating
that raising six young in a good condition proved to be a difficult task.
Micromammals appeared to be an important driver, both with regard to the number
of young and their condition. In good micromammal years, the condition of nests
where up to five young fledged was usually well above the expected value. In years
when there are few micromammals available, the parents usually only succeeded in
raising one fledged young to a condition at or above the expected value. In nests where
two or more young fledged, the condition was often below the expected value
(Figure 8.28).
Figure 8.29 illustrates that the CI is linearly related to the number of nestlings per
brood. Clutch sizes are well linked to available prey, yielding more and better-fed
nestlings at ringing age with larger broods.

8.19 Fledging Success

Table 8.24 summarizes the data collected in the main long-term studies across Europe.
Results shown in the table indicate the highest average number of fledged young per
breeding pair is 2.75 in Württemberg near Ludwigsburg, Germany (H. Keil personal
communication) and 2.66 near Lake Constance, Germany (Knötzsch in Schönn et al.
1991). The lowest average number of fledged young per breeding pair was 1.34 in

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 8.19 Fledging Success

1.1
Average condion index (CI) of nestlings at ringing date

1.08

1.06

1.04

1.02

0.98

0.96

0.94

0.92

0.9
1 2 3 4 5 6 7
Brood size
Bad vole years Good vole years

Figure 8.28 Average condition index (CI) of nestlings at ringing date per brood size in bad (yellow)
and good (brown) micromammal years (x-axis, number of young; y-axis, condition index (Stroeken
and Van Harxen 2022).

1.1
Average condition index (CI) of nestlings at ringing date

1.08
y = 0.0892x + 0.8136
1.06 R² = 0.5702

1.04

1.02
y = 0.0911x + 0.7309
1 R² = 0.5639

0.98

0.96

0.94
1.5 2 2.5 3 3.5 4
Average number of nestlings on ringing date for successful and all nests

Successful nests All nests Linear (successful nests) Linear (all nests)

Figure 8.29 Linear relationship between average condition index (CI) and average number of
nestlings on the ringing day for successful nests and for all nests (1998–2001, n broods = 1032)
(Stroeken and Van Harxen 2022).

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376

Table 8.24 Fledging success of Little Owl across Europe

Clutch size Brood size at hatching Brood size at fledging

Hatchling/ Fledging/
Egg/BP Hatchling/BP SBP Fledging/BP SBP

Country Mean n Mean n Mean n Mean n Mean n Author (Period of study)

Germany
Leipzig, Dresden 4.11 126 2.35 111 3.48 75 1.87 167 2.84 110 Schönn 1986 (1960–1985)
North Rhineland, 3.61 96 2.52 81 3.24 63 1.89 89 2.75 61 Exo 1987 (1974–1984)
Kleve
North Wurtemberg 3.6 326 2.4 326 3.3 235 Furrington 1998 (1971–1997)
Wurtemberg, 3.71 812 3.21 812 2.75 812 3.39 659 Keil unpublished (1988–2001)
Ludwigsburg
Lake of Constance 4.42 265 3.49 265 2.66 265 Knötzsch in Schönn et al. 1991
(1973–1988)
Wurtemberg, 4.12 141 2.78 203 3.78 149 2.35 203 3.34 143 Ullrich in Schönn et al. 1991
Göppingen (1969–1988)
Northrhine- 3.51 269 2.66 272 3.19 409 Gassmann and Bäumer 1993
Westphalia (1978–1992)
Mid-Westphalia 4.15 482 2.99 486 2.28 486 Kämpfer-Lauenstein and Lederer
1995 (1974–1994)
Voreifel 3.85 268 3.09 254 3.71 210 1.34 412 2.78 189 Zens 2005 (1987–1997)
Switzerland
Ajoie 3.12 153 2.22 153 2.78 122 1.82 153 Juillard 1984 (1973–1980)
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Bale 3.75 76 2.98 76 4.72 48 1.78 76 2.83 48 Baur in Krischer 1990 (1981–1990)
Genève 3.70 189 2.99 189 3.50 116 2.22 161 3.09 116 Meisser and Albrecht 2001
(1994–2000)
Britain
Nationally 3.59 268 2.40 241 Glue and Scott 1980 (1939–1975)
Cheshire 2.54 120 2.13 120 2.3 115 2.2 120 2.1 115 Leigh 2001a (1993–2000)
France
Nationally 4.24 94 3.62 53 3.70 30 2.84 237 2.96 218 Genot 1992d (1957–1990)
Northern Vosges 3.67 123 1.86 123 1.79 185 2.98 111 Genot 2005 (1984–2004)
Netherlands
Zuidoost- 4.00 471 3.13 471 3.77 348 2.25 471 3.08 348 Stroeken and Van Harxen 2003
Achterhoek (1986–2003)
Belgium
Flanders (Vlaams 3.36 312 2.68 312 3.13 267 1.97 312 2.3 267 Smets, Huybrechts and Cerulis
Brabant) personal communication
(2000–2004)
Wallonia 3.2 2346 2.7 2346 2.33 2346 J. Bultot and Groupe Noctua
personal communication
(1989–2004)
Flanders (Dijleland) 3.85 123 2.8 123 2.22 123 Vanden Wyngaert 2005
(1999–2004)

BP = breeding pairs, SBP = successful breeding pairs

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Germany in Voreifel (Zens 2005), in Switzerland 1.78 near Basel (Baur in Krischer 1990)
and 1.82 in Ajoie (Juillard 1984), 1.79 in Vosges du Nord, France (Génot 2005) and 1.87
in East Germany (Schönn 1986). The low fledging success obtained near Basel in
Switzerland (Baur in Krischer 1990) correlates with the disappearance of the species
in that region (Luder and Stange 2001). It is almost the same situation in Ajoie
(Switzerland) where Juillard (1984) recorded 1.82 fledged young per breeding pair with
just a few isolated pairs remaining. In addition to these reports, other data on fledged
young per breeding pair include: 2.84 (n = 39) in Czechoslovakia (Danko 1985a, b,
1986, 1989), 2.51 (n = 201) in Mediumrhine (Ingendahl and Tersteegen 1992) and 2.91
(n = 97) in Kaiserstuhl, Germany (C. Stange personal communication), 1.91 (n = 17) in
Austria (Ille 1996) and 1.6 (n = 12) in Russia (Khokhlov 1992).
In Germany, the average fledging success was reported to be better in nestboxes (2.47
fledged young per breeding pair (n = 76)) than in natural hollows (2.20 fledged young
per breeding pair (n = 10)) (Veit 1988) and Zens (2005) recorded 5.1 young on average
per female (n = 30).
The number of fledged young per successful breeding pair increases from west to east
in Europe (Génot 1992c) and within France too (Génot 1992c). Breeding success, in
particular the relation between the number of fledged young and clutch size, depends
on the condition of the female before incubation and therefore with feeding conditions
(Gassmann and Bäumer 1993). The authors found a higher fledging success than in
other German regions due to the good food situation in western Germany.
Climatic factors also influence fledging success. Precipitation in March seems to
improve food availability (earthworms) and thus the mass of the female in April.
While higher rainfall in March and April resulted in higher breeding success, high
rainfall in May and June, especially when the nestlings are younger than two to three
weeks, is correlated with high mortality of the young and a lower fledging success
(Lederer and Kämpfer-Lauenstein 1996), probably due to the moist environment in
the nestboxes (Gassmann and Bäumer 1993). In Belgium too, for each 100 l/m² of extra
precipitation in May and June, the number of fledglings decreased by 0.6 fledglings per
nest (Bultot et al. 2001).
Fledglings tend to leave the nest before being able to fly. They mostly jump and try
to climb back (Figure 8.30) along the tree in which the nest is situated. Returning
jumped nestlings to a nestbox does not keep the bird inside. Within a very short period
it fledges again. As of that moment both the female and the male start feeding fledlings
outside the nestbox (Figure 8.31). After leaving the nestbox, the fledglings remain
together and wait to be fed by the parents. They start developing some hunting skills
by observing possible prey (Figure 8.32).
Van Harxen and Stroeken (personal communication) showed that determining
breeding success as the number of young at ringing age over-estimates the real fledging
success. They compared the number of young reaching an age of at least 30 days (after
which they are supposed to start fledging) with the number of young on the ringing day
in 1035 nests between 1998 and 2021 (Table 8.25).
The difference between ringing and fledging success was calculated at 5.02% (3.14
versus 2.99). Out of 3284 nestlings ringed, 3163 survived until fledging, i.e., 121 died

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 8.19 Fledging Success

Figure 8.30 Little Owl fledglings try re-entering the nestbox that they fledged from by climbing the
nest tree.

Figure 8.31 Little Owl parents feeding fledglings on the ground in front of the nestbox.

(4.6%) and after ringing 1.45% of all nests failed. Partial loss of young in all nests was
3.68%. For successful nests only, the partial loss of young was 2.44% (79 of 3242).
The difference between ringing and fledging success between years was considerable,
ranging from 0% in 2021 to 26.3% in 2000 (including failed nests).

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Figure 8.32 Little Owl fledglings clustering together and observing possible prey.

To obtain comparable results in the future across studies, it is of the utmost import-
ance to visit a nestbox after the young have reached the age of 30 days and register the
age at the time of each observation to allow correct interpretation later.

8.20 Mortality
In a breeding dynamics study involving 15 nestboxes monitored over 8 years, Leigh
(2001a) showed that the highest mortality occurred at the egg stage, with 13% (40 of
305) of the eggs not hatching. The suggested causes for this were chilling of eggs, sperm
viability, experience of the female and possibly pesticide pollutants. The mortality rate
for owlets was significantly lower at 4.5% (12 of 265 hatched eggs). Death of the young
owls was caused by chilling, cainism and starvation. Most of the owlets that died did so
within five days of hatching (90%). High mortality levels in eggs and owlets correlated
with years when prey populations were low. In years when both micromammal and
invertebrate populations were low, the mortalities of both eggs and owlets were highest.
Field observations suggest that after hunting earthworms under wet conditions, the
adults transfer a lot of moisture into the nest site and onto the young, and this cool
dampness may increase the mortality risk. More detail is given in Chapter 10 on the
causes of natural and human-related mortality, and the impact of mortality on the
population regulation. In Germany, Zens (2005) recorded a total mortality rate of 50%

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Table 8.25 Over-estimations in breeding success numbers of theLittle Owl in Achterhoek (Netherlands) with or without taking mortality
after ringing into account (1998–2001)

Incl. failed nests Excl. failed nests

Number of nests 1035 Min Max SD Number of nests 1020 Min Max SD
Average ringed 3.14 Average ringed 3.14
Average fledged 2.99 Average fledged 3.04
Difference 0.15 Difference 0.10
Percentage overestimation 5.02 0 26.3 5.7 Percentage overestimation 3.40 0 13.5 3.5

Ringed 3284 Ringed 3242

Fledged 3163 Fledged 3163
Number of young lost 121 Number of young lost 79
Percentage loss 3.68 0 20.65 4.6 Percentage loss 2.44 0 12.5 3.1

Failed nests 15
Percentage failed nests 1.45 0 6.9 2.4

Van Harxen and Stroeken personal communication.

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(n = 412); of these deaths, 73% concerned owlets and 27% eggs. The mortality due to
predators reached 69.3%.
Of 51 mortality cases of young in Germany, 18 were caused by falling from the nest,
12 from predation, 2 from human destruction, 6 from loss of parents, 3 from disease,
2 from flooding and 8 were unknown (Schönn 1986). Of 60 mortality cases in
Switzerland, 9 were from predation, 20 from cainism, 6 from human destruction,
17 from the loss of parents, 1 from disease and 7 were unknown (Juillard 1984). Of
54 mortality cases in France, 7 were from predation, 7 from flooding, 3 from cainism,
1 from falling from the nest, 18 were unknown and 18 disappeared from the nest,
probably as victims of predation (Génot unpublished data). Mortality due to trampling
from cattle was noted by Juillard (1984). All eggs in 9 of 37 nests in Flanders did not
hatch, while two hatched clutches were predated by Stone Martens (Vanden Wyngaert
2005).

8.21 Cainism
Cainism was recorded in France involving 3 of 57 owlets (Génot 1992c), and in
Germany (Ullrich 1973, Knötzsch 1978, Exo 1983) where Zens (2005) recorded 33 of
332 owlets. Juillard (1984) noted that losses in a declining population due to cainism in
the nests was 33% (20 of 60 young). The victim of cainism is probably the youngest
owlet, eaten by the oldest (Ullrich 1973, Exo 1983, Génot unpublished data). Some
young were reported to feed on the carcass of a dead adult in a nest (Mills 1981). Van
Harxen and Stroeken (personal communication) found no evidence for cainism. Several
camera observations revealed that though dead nestlings up to a certain age were fed by
the female to the surviving siblings, in all these cases they died of natural causes
(probably starvation). They also found evidence of dead nestlings (still in their natal
down) lying between dead voles in the corner of the nestbox.

8.22 Lifetime Reproductive Success

One of the most important research priorities in owls is the acquisition of demographic
data and the development of good demographic models. To obtain good models input
variables are crucial. Age- or stage-specific estimates of survival and fecundity are
needed to parameterize population projection matrices with different distinct age or
stage classes, and to project future population trajectories (Noon and Franklin 2002).
Estimates of survival rates for Little Owls can be based on intensive capture–re-capture
studies and can yield age pyramids for populations. Fecundity rates can be estimated by
determining the reproductive output of birds of known age. See Chapter 10 on
Population Regulation.
Van Harxen and Stroeken (2020a) reported Life Reproductive Success (LRS) in
18 females with known year of birth, and first year and last year of breeding
(Table 8.26). Two females that bred for six years produced 17 and 15 fledglings (average

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 8.22 Lifetime Reproductive Success

Table 8.26 Females for which the entire Lifetime Reproductive Success could
be determined

Year of First Last Number of Number of

Female birth year year broods fledglings Average

3604090 2004 2005 2010 6 17 2.83

3689781 2011 2012 2017 6 15 2.50
3809482 2016 2017 2019 3 15 5.00
3771450 2015 2016 2017 2 6 3.00
3809556 2017 2018 2019 2 6 3.00
3769855 2015 2016 2016 1 5 5.00
3801687 2016 2017 2018 2 5 2.50
3709952 2011 2012 2012 1 4 4.00
3779812 2015 2016 2017 2 4 2.00
3827095 2018 2019 2019 1 4 4.00
3560100 2000 2001 2001 1 3 3.00
3763155 2014 2015 2015 1 3 3.00
3771370 2014 2015 2016 2 3 1.50
3801767 2016 2017 2017 1 3 3.00
4183716 1991 1992 1992 1 0 0.00
3666601 2009 2010 2010 1 0 0.00
3680342 2010 2011 2012 2 0 0.00
3826644 2018 2019 2019 1 0 0.00
All 18 females 36 93 2.58

Van Harxen and Stroeken personal communication.

2.83 and 2.50 per year). A female that bred for 3 years produced 15 fledglings, on
average five per year.
They found the same variation when comparing females for which no information
on year of birth was known, or for which one or more years of reproductive success was
missing. The 10 most successful females produced between 21 and 29 fledglings in 6–12
brood attempts (Table 8.27). The average number was highest in a female that produced
29 fledglings in six years: 4.83 per year.
A Lorenz curve is best suited to visualize the percentage of females as a function of
the percentage of reproduction (Figure 8.33). The top 10 most productive females
accounted for 7% of the number of fledged young (212 individuals); the top 5% of
the most productive females accounted for 21% of the offspring. The top 10% yielded
34% of the fledglings. Half of all fledglings were produced by 18% of the females. Half of
the females produced 86% of all fledglings. A third of all females produced 72% of
all fledglings.
They also found females with bad or even no reproduction success at all, e.g., one
female that laid nine eggs in four consecutive years did not succeed in producing any

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Table 8.27 Top 10 most productive females

Female Breeding Total Average number of fledglings

identification attemps fledglings per brood

3547290 6 29 4.83
3572852 9 28 3.11
3243212 12 28 2.33
3757618 6 25 4.17
3710019 8 25 3.13
3625901 7 24 3.43
3633134 9 23 2.56
3633132 9 22 2.44
3710033 6 21 3.50
3599233 8 21 2.63

Van Harxen and Stroeken personal communication.

120%

100%

80%
Percentage Fledlings

60%

40%

20%

0%
0% 20% 40% 60% 80% 100% 120%
Percentage Females

Figure 8.33 Lorenz curve of percentage of females as a function of percentage of fledglings

(n = 590 females producing 3035 fledglings (1987–2021).

fledglings at all. Another female produced exactly one fledgling in five years (0.2
per year).
Table 8.28 illustrates the lifetime reproduction from a long-term population study in
Belgium by J. Bultot (personal communication).

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 8.23 Extra-Pair Copulation

Table 8.28 Lifetime reproduction in Belgium

Average Total

Percentage
Individual Clutch nestlings/ Age of
female size Nestlings Fledgelings Eggs Nestlings eggs Years female

E142605 2.25 1.75 1.5 9 7 78 4 7

E196139 2.50 2.00 1.75 10 8 80 4 8
E196057 4.25 4.25 3 17 17 100 4 9
E183378 3.4 3.4 2.8 17 17 100 5 7
E183126 4 4 3.75 16 16 100 4 8
E196286 3.75 3.25 2 15 13 87 4 7
E214458 3.8 2.2 2.2 19 11 58 5 8
E229875 3 2.5 2 12 10 83 4 7

J. Bultot personal communication.

8.23 Extra-Pair Copulation

Despite some populations where high population densities have occured and plenty of
prey resources have been available, so far no extra-pair copulations have been found.
Müller et al. (2001) investigated the mating behavior of a population of Little Owls in
Westphalia, Germany, by taking blood samples from 53 nestlings, produced by
16 breeding pairs. Using multilocus DNA fingerprinting, the parentage of the young
owls from each nest could be established, indicating no successful extra-pair copula-
tions in this population (Müller et al. 2001). Additional studies conducted in high-
density populations should provide further insights into the relevance and impacts of
extra-pair copulations.
In a study in Cheshire, England, one record of bigamy was recorded in 10 years. This
involved two females each nesting approximately 100 m apart, one in a nestbox and
the other in a natural cavity. The timing of the breeding efforts was spread so that the
male could provide food for both nest sites, and both nest sites did not hit their peak
food consumption at the same time. R. Leigh (personal communication) also noted that
the male was caching food in the nests, which provided him some rest if the weather
prevented foraging for food. Illner (personal communication) has observed two females
incubating in the same nest on two occasions during his 25 years of studying over
1000 nest sites. Berghmans (2001c) recorded nine owlets in one nest site being cared for
by two females. The instances of bigamy mainly seem to be females that nest near to
each other or lay in the same nest site, incubate together and share parental care for the
owlets. However, in these instances the ability to incubate the eggs and brood the
young are the main influence on survival of the eggs or owlets.

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 Chapter 9: Behavior

Chapter Summary
New media such as internet-connected cameras in nestboxes can yield infrared images
in the dark leading to new insights and knowledge. Substantial new information has
become available from the groundbreaking webcam project Vogelbescherming
Nederland (BirdLife Netherlands) Beleef de Lente (www.beleefdelente.nl) that started
in 2007 and continues to this day. Volunteers selected video clips revealing remarkable
undiscovered behavior over 14 entire breeding seasons, day and night, from courtship
to egg laying to the fledging of the young.
Due to simpler and cheaper technology, more and more people have decided to
install such cameras in their nestboxes, which is expected to yield even more new
knowledge in the future, opening unprecedented opportunities for citizen science.
This chapter is complemented by a lot of information from the long-term research of
Van Harxen and Stroeken (personal communication) (from 1986) in their study area in
the southeast Achterhoek in the Netherlands.

9.1 Body Movements

The Little Owl can easily be observed during the day, especially when perched in
exposed locations like chimneys and power lines. In the Netherlands, nestboxes
designed with special balconies offer ideal roosting opportunities for the owls and allow
verification of nestbox occupation (P. Beersma personal communication). Other owls
prefer to roost in cavities by day. For this purpose it is advised to install two or three
nestboxes per territory to allow males to roost in a different cavity from breeding
females. During day roosting, most owls lean against one of the walls of the cavity
(Scherzinger 1980). While most owls roost relatively low to the ground (<10 m), radio-
marked owls have been found to roost in trees at higher altitudes.
Notable postures of Little Owls, also shown by other owls and raptors, are the cryptic
posture to minimize attacks by flying enemies, defensive turning against enemies at close
range and covering behavior to hide prey. The cryptic posture is a narrow, rigid, alert
upright stance, adopted as soon as the bird detects any kind of alien sound or disturb-
ance. Approaching persons or predators are located with short, nervous, horizontal
(Figure 9.1) and vertical (Figure 9.2) circular movements of the owl’s head, with the

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 9.1 Body Movements

Figure 9.1 Little Owl circular head movement.

Figure 9.2 Little Owl vertical head movement.

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 Chapter 9: Behavior

body in an upright position or with bended knees. On arousal, the owls alternate
bending their knees with extreme upright stretching of the body. This behavior gives
the impression that the owl is jumping (Heinroth and Heinroth 1926, Hubl 1952,
Scherzinger 1971a, Busse 1983). Flicking of the tail, wingtips and entire wings is
frequent among aroused and nervous owls (Scherzinger 1980).

9.2 Locomotion
Little Owls stroll or run when hunting for earthworms and other invertebrate prey.
Before flying off, the owls regularly repeatedly trample where they are. When pursuing
voles, owls prefer to walk, jump and run. Running also occurs when the owls are going
to cache some prey or are escaping danger. In contrast to most other owls, the flight of
the Little Owl is audible, because the distal radii of the outermost primaries are not
serrated (Sick 1937) and because the relationship between body mass and wing surface
(i.e., wing loading) is rather high (Schönn et al. 1991). The species mostly flies very low
to the ground (e.g., 1–10 m). During the day the Little Owl flies straight for short
distances, and crosses longer stretches in an undulating way, like a woodpecker.
When departing from a perch, the owl first drops to gain speed, flies and then lands
in an upward arc, reflecting a U-shaped flight pattern (Schönn et al. 1991). Flights can
be interrupted suddenly when the owl enters shelter or cover. Owls hunt from low
perches, and then capture prey after a straight flight or pounce (Baumgart 1980, Brands
1980, Scherzinger 1980, Busse 1983, Ille 1983, Mikkola 1983). When hunting, hovering
and gliding flight can also be observed (Figure 9.3).
The flying altitude has long been under-estimated. Eick (2003) observed Little Owls
via telemetry flying above high-tension power lines, crossing steep valleys and urban
areas at altitudes ranging from to 20–30 m and even higher. While it is easier to see owls
flying low above the ground, observations of owls flying higher gave important insights
into dispersal, migration and the degree of population isolation. Data suggest that rivers
and forest edges do not influence the owls’ choice of directional movements. Even
relatively broad valleys (e.g., Neckar and Enz valleys in Germany) are crossed. Telemetry
research has confirmed that Little Owls fly above – rather than through – large wooded
areas. Large wooded and urban areas have not been found to be barriers to the dispersal/
movements of the species.

9.3 Roosting and Sleeping

Sleeping birds are often almost invisible due to their excellent camouflage colors. The
feathers are fluffed up, the head is sunk into the body, causing all the white features of
the bird to be hidden. A leg is often pulled into the belly feathers with closed toes
(Scherzinger 1980). The eyes are closed by lifting up the lower-eye-lids (Heinroth 1926).
They often yawn when waking up (Figure 9.4).
Little Owls sleep or rest in places providing immediate cover and with open sight
and escape routes, e.g., tree crowns, shrubbery, attics and holes in walls. During bad

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 9.3 Roosting and Sleeping

Figure 9.3 Food transport and hunting behavior of Little Owl.

weather, Little Owls mostly use holes for resting (Exo 1987). During winter, owls
regularly spend the daytime in the stables of domestic livestock, hay barns and holes
in trees. The avoidance of cold temperatures plays a crucial role in the choice of the
roosts (Schönn 1986). Roosts are situated within the territory of the owl and are used by
the same birds for many years. These favorite locations are easy to detect through the
white chalk stripes of their excrement and pellets.

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 Chapter 9: Behavior

Figure 9.3 (cont.)

9.4 Preening
Optimally cared for plumage and well-maintained beak and claws are essential in
hunting and survival, therefore Little Owls spend a lot of time on these activities.
When preening, the owls partly or completely close their eyes by lifting their lower
eyelids (Heinroth and Heinroth 1926). The head feathers are cleaned as they lay on the

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 9.4 Preening

Figure 9.3 (cont.)

head, while the larger body, wing and tail feathers are cleaned one by one. The
uropygial gland is regularly drained for oil. The back, rump and base of the tail are
reached and preened over the shoulder and slightly lowered wing, under-tail feathers
are reached from beneath the wing. The primaries, secondaries and tail feathers are
pulled through the beak one by one, starting at the base. The base of the quill is
intensively nibbled. Legs and toes are mostly cleaned by stretching the entire leg while
keeping the foot closed. The toes are nibbled individually or pulled through the beak.
Little Owls only use their talons in grooming when they scratch the back of the head,
the ears and the facial region (Brands 1980, Scherzinger 1980). The head is also cleaned
by rubbing with the wing or shoulder. The beak is cleaned by rubbing it on branches,
especially after food intake (Scherzinger 1980). Incubating females tend to rub their
beak against one of the walls of the breeding cavity (Brands 1980).
Occasionally they “shower” by sitting or standing in the rain with wings and tail
spread out (Figure 9.5). Strong winds are not avoided. It is plausible that dust, sand and
pests are removed. Sunbathing, which Little Owls like to do, may also play a role in the
care of plumage to fight feather parasites. Sand and rain also help to remove parasites.
Parasites on the head are combated by making rapid back and forth movements with a
foot (scratching). The beak is cleaned by rubbing it along a branch. On one occasion, a
young shoot of the pear tree in which the nestbox was placed was observed to be
completely removed in two sessions by cutting it into pieces with the beak.
Parallel stretching of a wing, leg and tail on one side of the body often preceeds
preening activities and is also associated with the intent to fly (Schönn et al. 1991) or to
leave the daytime roost. Sometimes just one wing is stretched, sometimes both wings
are stretched towards the front and upwards, until both wings almost meet above the
head. Simultaneous stretching of both legs has also been observed.

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 Chapter 9: Behavior

Figure 9.4 Little Owl yawning when waking up.

9.5 Nest-Site Competitors

In Western and Central Europe, Jackdaws Corvus monedula and Stock Doves Columba
oenas are nesting-site competitors, since both species breed in cavities of a similar size.
Where cavities, both natural, in trees, and in buildings, are scarce, fighting takes place,
e.g., in pollard willows and buildings. Both species occupy nestboxes for Little Owls,
especially if there is no partition in them that obstructs direct view to the outside. Stock
Doves can be particularly tenacious once they have selected a nest site at a certain place
and try a second, third or even fourth time if a previous clutch is lost due to fights. They
defend themselves against Little Owls by tilting to one side and tapping firmly with the
free wing, sometimes intimidating the owls. Real fights can lead to feathers flying
around. Jackdaws are often numerically in the majority because several pairs try to
conquer the same cavity at the same time. They are not even afraid of attacking and
chasing a Little Owl from a branch and going after it. Both Jackdaws and doves are at a
disadvantage because Little Owls sleep in the nestbox during the day from February
onwards, sometimes even earlier, and can defend from the inside in the event of a
disturbance. The owls are not afraid to come out, choose a position in front of the
entrance of the nestbox and defend it with threatening behavior. If Jackdaws become
too intrusive, the owl may opt for a tactical retreat and flee back into the nestbox. Both

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 9.5 Nest-Site Competitors

Figure 9.5 Little Owl taking a rain shower.

species can also perform dive flights in an attempt to chase each other away. Such fights
can last for weeks, even after egg-laying.
Woodpeckers are sometimes intimidated or even attacked near the nest
(Figure 9.6)
Squirrels Sciurus vulgaris and Dormice Elyomis quercinus (Jacobs 2008) frequently
occupy Little Owl nestboxes, as a place to sleep in the winter months or to raise young.
It is unclear whether these really compete for nestboxes, or only move in if the owls
have no interest. The latter is probably the case when it comes to species such as the
Wood Mouse Apodemus sylvaticus and the Common Starling Sturnus vulgaris, which are
potential prey. This certainly applies to Great Tits Parus major and Collared Redstarts
Phoenicurus phoenicurus that are also found to breed in Little Owl nestboxes. Wood
Mice, which are excellent climbers, sometimes use a nestbox in winter as a sleeping
place and for storing nuts and dried fruit. If the stock is large, the nestbox may become
unsuitable for owls. This also applies if Squirrels, Starlings or Jackdaws have filled the
nestbox with nesting material in the absence of the owls. Often the entrance is blocked,
so that the owls no longer have access.

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 Chapter 9: Behavior

Figure 9.6 Little Owl intimidating Great Spotted Woodpecker Dendrocopos major.

Nests of bees Anthophilia spp. and wasps Vespidae, in particular the European hornet
Vespa crabro, are regularly found in nestboxes, especially after the young have fledged. If
such a nestbox is not cleaned before the breeding season, it significantly reduces the
chance that an owl will move in again. Bees and wasps can also be a nuisance. Once it
was shown with video footage that bees irritated a female who had small young at the
time, so much that she left the nestbox at half past eight in the morning to return only

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at the beginning of the evening, after the bees had left. In the meantime, the three
young had died (Van Harxen and Stroeken 2005).

9.6 Enemies
Naef-Daenzer et al.(2016) showed that Little Owls are not top predators but rather meso-
predators. They found that 78% of the mortality among tagged adult birds and fledgling
young was due to predation, with hardly any distinction between adult and young
animals. Foxes Vulpes vulpes, Common Buzzards Buteo buteo and Beech Martens Martes
foina turned out to be the most important predators in the study area around Ludwigsburg
(Germany). In the same study area Michel et al. (2016) found that the presence of Little
Owls within 150 m of a forest was greatly limited by the presence of Tawny Owls Strix aluco.
They argued that the negative effect of Tawny Owls was due to the avoidance of potentially
suitable habitat by intimidation rather than direct predation. However, research by Hessels
and Van Harxen (2020c) in Winterswijk (the Netherlands) showed that both species can
occur in high densities right next to each other, with Little Owl territories located less than
200 m from the edge of a forest in which Tawny Owls were observed. Both territorial calls
were regularly heard simultaneously.
Nevertheless, young Little Owls in particular regularly fall victim to Tawny Owls.
Van Harxen (2021) describes a situation in which a 72-day-old fledged young was
caught by a Tawny Owl in front of the nestbox. Sparrowhawks Accipiter nisus are
reported to regularly manage to surprise Little Owls by various observers, confirmed
by the discovery of “plucking places” in the vicinity of the nestbox. Mikkola (1983)
mentions Eagle Owls Bubo bubo, Long-eared owls Asio otus and Barn Owls Tyto alba as
predators and of the birds of prey especially the Goshawk (Accipiter gentilis). Dogs and
cats are also regularly reported as predators. Van Harxen and Stroeken (personal com-
munication) were once given the rings of four of five Little Owls ringed by them and
caught by a feral dog shortly after fledging.
In addition to being a nest-site competitor, squirrels also predate on eggs, but
whether the female owl is chased off the nest or this is during a breeding break is not
clear. Usually, loss of the entire clutch occurs; Van Harxen and Stroeken (personal
communication) observed a nest where all five eggs were consumed.
If Little Owls are given the chance, they defend themselves fiercely against predators
and other invaders. Several times camera recordings have shown that Beech Martens
that tried to enter a nestbox were attacked by both owls by diving flights (Van Harxen
and Stroeken 2021). Barn and even Tawny Owls are also attacked by Little Owls if seated
in front of the nestbox.
Several observers have reported feral cats trying to escape the attacks of Little Owls
by walking close to a wall or hedge; others hardly dare to go outside during fledging
time (Van Harxen and Stroeken personal communication).
Exo- and endoparasites can also cause an unusually high mortality among eggs and
small young, in 2015 as a result of a flea population explosion (Van Harxen et al. 2017).

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As a result of the wet and mild weather, many flea pupae had survived the winter and fed
on the blood of the breeding female at the brood patch, which is an excellent place for
this. The eggs were severely smeared by flea droppings and later with the blood of
newborn young, some of which weakened to such an extent that they succumbed
to anemia.
In the east of the Netherlands, the Beech Marten is known as the most important
enemy. Despite adjustments to the nestbox to try to prevent them from entering, the
predation of eggs and young can sometimes be significant. Regularly, bitten young,
and also adult animals with bitten necks, are reported in the nestbox. Nestboxes are
also regularly used as roosts while leaving characteristic latrines and unusual food resi-
dues (gingerbread, goose eggs, chicken heads). Visscher (personal communication) found
the head of a Great Cormorant Phalacrocorax carbo in a nestbox. Pine Martens Martes
martes have also been found to occupy nestboxes (Ruud Leblanc personal
communication).

Pine Marten Martes martes

Little Owls are frequent victims of the mobbing behavior of small passerines (Curio
1963). Owls will either ignore (Scherzinger 1970) or react to the passerines by looking
for cover (van Wacquant Geozelles 1890) or rarely grabbing at a mobbing bird with its
talons (Haverschmidt 1946). The cryptic posture adopted by the owls helps to avoid
being mobbed (Scherzinger 1971b). Using live owls as bait, the mobbing behavior of
passerines towards Little Owls has been exploited as a technique to capture passerines
in Italy (Schaaf 2005). The presence of the male often can be deduced from mobbing
passerines (Van Harxen and Stroeken personal communication).

9.7 Social Behavior

Little Owls maintain a mating bond throughout their lives and often remain in close
proximity to each other outside the breeding season. In addition, they regularly
spend the day sleeping together and resting. Each pair differs in the moment that
the female allows the male to join her in the nestbox. At the camera location in
Harreveld, the male only stayed in the nestbox during the day before egg-laying,

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 9.8 Territorial Behavior

while the male in Henxel spent a large part of the day in the nestbox from mid-
November. The female had already moved into the nestbox in August, but did not
allow the male to join her for the first few months (Van Harxen and Stroeken
personal communication).
Usually the male leaves the nestbox in the afternoon to report again at dusk. He does
not enter the nestbox but calls his partner. Often it takes a few minutes for her to leave.
In front of the nestbox, most of the time some affections are exchanged and sometimes
they mate. A little later, both disappear into the darkness. At the end of the night, the
female returns first; often it is still dark. As soon as she is inside, her wing and tail
feathers are thoroughly brushed, occasionally tapping the uropygial gland to collect
some oil. Shortly after sunrise, the male appears in front of the nestbox for a while
before entering. The day is spent sleeping and resting, taking care of the plumage and
the mutual exchange of affections. If a mouse happens to be in stock, both eat from it.
In the course of the afternoon, the male disappears outside, after which the pattern of
the previous day is repeated.
If the female is sitting on eggs or with nestlings later in the season and there is a
disturbance, she is warned by the male with some short alarm cries and leaves the
nestbox at once.
Part of the annual recurring actions is the choice of nesting place. Even if there is
only one suitable cavity that is used annually, the ritual is performed. This consists of
the male trying to seduce his partner to enter the nestbox while expressing all kinds of
excitement sounds. Once inside the male throws himself on his belly and digs out
a nest bowl underneath him with his legs. The nestbox litter sometimes causes such
a dust cloud that a clear view is obstructed. When the female takes over the action, the
mission is successful and the male leaves. The ritual repeats itself several times up to
egg-laying.

9.8 Territorial Behavior

Male Little Owls mainly delimit their territory through territorial calls. Spontaneously,
this happens most often in spring when dusk falls and they leave their roost to go
hunting. If there are intruders or strangers, this call, described as “ghuk,” often turns
into an excited “miaw,” increasing in pitch and frequency. On some evenings the
neighbors join in and in densely populated areas hooting is heard everywhere.
Neighbors recognize each other by their voices. Hardouin et al. (2006) found that
neighbors reacted significantly less often to each other than to intruders, a behavior
that is used during playback.
Some males react fiercely and aggressively, others behave more timidly. A few fly
towards the alleged intruder, sit on a pole or low branch, shout hoarsely and even
continue to do so when the light of a flashlight is pointed at them. Exo (1987) has
described attacks by males on dummies.
The willingness to react in densely populated areas is higher than where the density
is low. This means that in sparsely populated areas it is often difficult to determine the

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precise number of territories and the density is therefore often under-estimated.

Sometimes it is over-estimated because males sometimes follow the observer.
Rarely, it has been observed that an intruder is actively chased away by the territory
holder, helped by the female.
At fledging, territorial behavior takes place again. Male fledglings collect information
about occupied places by uttering the territory call and engaging in vocal duels or
physical fighting.

9.9 Division of Tasks

Little Owls have a pronounced division of tasks during the breeding period in which
the female incubates, keeps the young warm and feeds them, and the male takes care of
prey supply. When the young are about 10 days old the female goes hunting. The male
brings prey directly to the young if the female is not present. Sometimes, however, he
calls the female to hand over the prey. If in that phase the female falls away, the nest
usually fails. A case was observed in 2017 where the male tried to take over brood care.
After the female was killed by a Pine Marten while defending the nest, the male tried to
take care of the five-day-old young. The combination of keeping them warm, feeding
them and hunting turned out not to be feasible and two days later all the young had
died. It is remarkable that the male tried to tear pieces of a mouse and feed them to
the young, behavior only known from females. Initially, the nestlings did not accept
the food, but after a few failed attempts, they actually managed to take in some pieces
(Van Harxen 2021).
The same male made several attempts to sit on the eggs in the absence of his (new)
female in the following years. He stopped immediately when his partner entered the
nestbox and walked towards her. No attempt lasted more than a minute. When in
2021 the eggs did not hatch and the female gave up after 74 days of incubation, the
male tried several times to take a seat on the eggs. The longest attempt lasted 15
minutes. Similar to the female, the male brought the eggs under him by gently sliding
them under his chest and belly with his beak (Van Harxen et al. 2023).
In the nestling phase, the male is only welcome in the nest to deliver prey. When
struggling to get outside, the female forces him to do so by maneuvering him with her
head towards the exit. When he tries to eat from a mouse that is in the larder and
intended for the nestlings, she resolutely takes it away from him. Outside the nestling
period, she does allow him to eat stored mice.

9.10 Parents and Young

9.10.1 Communication in the Egg
About a day before the eggs hatch, the young make high-pitched squeaks from inside
the egg. For the parents, this is the signal that the birth is imminent. Especially for the
male, it is also a sign to start building up prey stock in a larder. If there are enough

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 9.10 Parents and Young

micromammals available, the supply is increased. The female responds by making

babbling sounds as she bends over the eggs.

9.10.2 Hatching
Camera observations have revealed that the female does not help with hatching.
After hatching, the female removes the shell residues by placing them at a safe distance
from the young with her beak. Just like the eggs, the young are kept warm
continuously.

9.10.3 Growing Up
In the first 10 days, before the eyes are open, the young are completely dependent on
their mother. During this period she keeps the young under her as much as possible to
prevent cooling. The young are not yet able to maintain their heat balance themselves
and cluster together in a heat pyramid when the female leaves the nestbox, preventing
heat loss as much as possible.
Young are preferably fed every one to two hours, about 15 times daily, if there is
enough prey in stock. If not, the feedings are limited to the moment that prey is
brought in by the male. Only around 10% of fresh prey suitable for the first days is
supplied during the day, and the number of feeds during the day is limited to a few or
even none.
As of the age of 14 days, the young try to leave the nest bowl and explore the nestbox.
Initially, the female tries to prevent this by pushing them back to the nest bowl, but soon
she gives up her attempts. The young still rest and sleep in the nesting bowl. In addition,
the female manages to keep them under her less and less. Eventually she gives up and
rests standing or sitting next to the young. She now hunts more and more, especially at
night, and therefore spends less time in the nestbox. Between days 15 and 17 she stays
outside the nestbox all night for the first time and the young are on their own.
Although feeding seems to depend on the young that presents first, females also
consciously choose a specific nestling and pass begging young at the front of the line to
give the prey to the young at the rear. Because the young do not differ very much in size
from each other and are therefore not easily recognizable individually, preference has
been shown to be a function of the yellowness of the beak (Avilés and Parejo 2013).
The young are regularly caressed by the female, by gently moving her beak between
their head feathers while expressing all kinds of babbling sounds. In this way, the male
also shows his affection, albeit less frequently.

9.10.4 Fledging
Once fledged, both parents, but especially the female, try to entice the young to stay
outside the nestbox permanently by offering prey outside the nest. This is done step by
step. As a first step, prey is delivered to an owlet that sticks its head out of the cavity.
After that, prey is kept in sight of the young, but not immediately released. Next the

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prey is kept in sight outside the nest, but only released after the young have left the
nest. In the end, most prey transfers take place outside of the nestbox. Sometimes prey
is not released immediately. Instead the female flies away, taking the prey with her and
tempting the young to follow. Even after all owlets have fledged, some take a seat in
front of the nestbox whereupon the female sometimes violently chases it away.

9.11 Handling Prey

Micromammals can be eaten in one go, but also gradually. A piece that is not immedi-
ately eaten is placed in a corner of the nestbox for later consumption. It can take hours
until the entire prey is consumed, e.g., on 5 March 2022 in the camera nestbox in
Lichtenvoorde. The female entered the nestbox at 6:41 am with a vole. Apparently she
had already eaten parts of it outside. She put it in the corner and went to sleep. At 7:10
the male entered, and discovered the mouse at 7:17. He picked it up with his beak and
held it with his right leg. He pulled out part of the intestines and offered the rest to his
partner at 7:21. She showed no interest, so he placed the residual mouse in the corner to
the right of the entrance. At 9.33 he moved the mouse from the corner to the left of the
entrance without eating it. At 10.41 he left the nestbox. At 12.25 the female picked up
the remainder of the mouse from the corner and ate it all.
Devouring an entire mouse takes very little time. Starting at the head, it takes about
10 seconds before only the abdomen with the tail of the mouse is visible and less than
20 seconds later that is also gone, totalling barely half a minute.
If a mouse is partially eaten the throat is often first. With the trachea, strings of
intestines are pulled loose and eaten, along with the head. Depending on hunger, a
larger or smaller piece of the mouse is consumed. The tail always disappears last. During
nestbox visits, micromammals can therefore be found at all stages of consumption,
from completely intact to only the posterior part of the lower body with the
tail. Occasionally only the tail remains. If many rodents are supplied in the nestling
phase, they are put in stock in their entirety. Feeding is then not done prey by prey, but
parts of several are consumed at the same time. A larger supply therefore always consists
of a combination of intact and partly eaten prey at all stages of consumption. On
warmer days, they can be found in all stages of decomposition, with some full
of maggots.

9.11.1 Earthworms
Earthworms Lumbricidae spp. are often fed to the young alive. They are then slurped
inside like a strand of spaghetti. But they are also eaten in pieces where the owl holds
the worm with one leg while cutting off pieces with its beak (Figure 9.7). They are
always consumed immediately and never put in stock. Sometimes they are handled
carelessly and they fall out of the beak when transferred to the female or the young.
Often they are not picked up and dry out at the bottom of the nestbox or at its entrance.
Remnants of earthworms consist of sand and the hard chitin brushes (chaetae).

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 9.11 Handling Prey

Figure 9.7 A Little Owl catching an earthworm.

9.11.2 Cockchafers
Cockchafers Melolontha melolontha are stripped of both shields (elytra) with the beak
before they are eaten. If that happens in the nestbox, these are easy to find, even a year
later. The rest of the beetle is consumed in its entirety. The hard chitin parts of the legs
and antennae are found in pellets or pulverized. Occasionally, if plenty of cockchafers
are caught (Figure 9.8) they may be stored for a short time, sometimes in a corner of the
nestbox, sometimes at the front. Van Harxen and Stroeken (personal communication)
once counted a pile of more than 25 fairly intact cockchafers at the entrance of
a nestbox.

9.11.3 Birds
Birds are partially plucked before they are eaten or fed to the young. This is not very
efficient because it takes a lot of time and the result is often poor. Wings and legs of
larger species sometimes remain in the nest. For small birds it is sometimes not even
worth the effort to pluck them. Hungry young can eat a headless Great Tit or Sparrow
inside as a whole, probably to prevent it being taking away by their siblings. It can take
up to half an hour for such prey to be eaten in its entirety.

9.11.4 Frogs
Frogs are regularly dehydrated and found more or less intact in the nestbox litter. Van
Harxen and Stroeken (personal communication) once found seven dried frogs Rana
spp., some with the head still attached, others with only the abdomen. The impression
is that frogs are caught more often than eaten. Frogs are sometimes still alive when they

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Figure 9.8 A Little Owl catching cockchafer.

are brought into the nestbox. Van Harxen and Stroeken (2011b) described a situation in
which, when pushing aside a wooden cover plate of a nest in a tree cavity, a frog took
the opportunity to escape. If the young are hungry, they can eat a frog in its entirety in
a matter of seconds. The legs disappear last.

9.11.5 Moles
Moles are rarely caught and almost never eaten in their entirety. The dense skin and
large front legs are not easy to digest. If they are eaten, the soft tissue in the abdomen is
consumed first. The skin is left untouched. As prey residue, the head with front legs and
an empty skin is sometimes found. Occasionally, an uneaten, dried mole disappears in
its entirety into the nestbox litter.

9.11.6 Insects and Other Small Prey

Insects, larvae and caterpillars are eaten in their entirety. This usually also applies to
moths. In the nestbox in the second half of the nestling period, caterpillars of moths
that come to pupate are sometimes found, e.g., the Gypsy moth Lymantria dispar. It is
unclear whether they are also eaten. Flies that enter the nestbox to lay eggs in a rotting
mouse are sometimes plucked from the wall of the nestbox. This also applies to maggots
that have started eating a mouse. In years with many micromammals and therefore
many rotting remains, maggots can be fed in large numbers. In 2018, in a camera
nestbox 108 maggots were fed on 17 out of 21 days, often several in a row, with a
maximum of 14 in a day (Van Harxen and Stroeken personal communication). Since
only the female spends substantial time with the young, she is the only one who grabs
and feeds maggots.

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 9.12 Two Females Nesting Together

Figure 9.8 (cont.)

9.12 Two Females Nesting Together

There are known examples of two females being found together in a nestbox, incubat-
ing a generally above-average number of eggs (Stroeken and Van Harxen 2018). The fact
that the eggs occasionally hatch proves that in these cases at least one male must have
been involved. Stroeken and Van Harxen (2018) once found six young in a nestbox
with two females which could clearly be divided into two groups that differed in age by
a few days. The same occurred in a nest in Brabant where all 11 eggs hatched and
10 young successfully fledged (Van Dooren and Hermsen 2019).
Usually a situation with two females lasts only one season, but in several cases it was
found that it can also occur over several seasons. In the years 2016–2019, Visscher
(2022) found two females that were found nesting together in a nestbox for four years
in a row. Together they laid 25 eggs (6, 7, 6 and 6) during that period, from which a total
of four young fledged (2 times 2).
The reason for this behavior is difficult to determine. Van Harxen (2021) suggested
that the second female is a floater who can still participate in reproduction in this way
and also increases her chances of taking over the place of the first female if it falls away.
The advantage for the first female is hard to assess since in most cases these broods do
not produce young, as the eggs do not hatch (Stroeken and Van Harxen 2018). There
seems to be a certain tolerance towards a strange owl in the territory anyhow. Van
Harxen (2021) reported that camera images regularly showed that a third owl was
present in the territory and sometimes even entered the nestbox, which was apparently
tolerated. Once it was even observed that a strange owl with a caterpillar in its beak
entered a nestbox where a female was currently nesting. The latter was clearly not

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 Chapter 9: Behavior

pleased and reacted fiercely with beak snaps and hissing sounds and the intruder
quickly left. The gender of the intruder remained unknown.

9.13 Behavior Towards Humans

As an opportunistic species Little Owls often reside in the vicinity of humans and are
adapted to human presence. It has been observed for newcomers that they show them-
selves less often and fly away earlier on approach than a former resident owl. The latter is
often easy to approach and does not care much about the presence of the resident as long
as attention is not specifically focused on it. The following year, the newly established
owl is generally more tolerant. However, individual differences are also observed.
Several landowners have reported that the owls just stayed in their favorite place, if
they walked past or underneath them. Even if they were working in the yard or garden,
the owls sometimes watched them with interest. With strangers in the yard, the owls
reacted sooner. Apparently, the owls can recognize people and assess whether they pose
a danger or not. Feral dogs and cats are also tolerated if experience has shown that they
don’t need to be feared. Others are sometimes fiercely attacked.
When opening the nestbox, some adult birds lay flat on the eggs or young and allow
themselves to be caught without any problem. Others leave the eggs and try to make
themselves invisible by standing elongated with the head forward in a corner. Very
occasionally they defend themselves with the claws and beak. It is often the same individ-
uals that exhibit this behavior; more frequently females than males. Camera images have
shown that after a disturbance to read the ring and/or measure and weigh, peace usually
returns after a few minutes and that the female takes a seat on the eggs or young again. Very
rarely will owls leave the nestbox after inspection for a short time. Owls that breed under
the roof tiles of a shed sometimes crawl away on approach; others sit tight on the eggs and
do not move even if a hand is slid underneath them to feel whether the eggs have hatched.
During the check, most owls undergo all manipulations without aggression. If they
feel that the hand around them is slackening, they sometimes try to wrestle themselves
free. The legs often show a gripping reflex, but they rarely really defend themselves. The
manipulation nevertheless causes stress, which can be deduced from the fact that many
owls defecate at some point, some when taking them out of the nestbox, others only
when they return again; most, however, during measuring and weighing. It is therefore
important to keep manipulation as short as possible and to quickly put the owls back.
The clutch is rarely abandoned after a nestbox inspection, but this can never be
completely ruled out.
Juvenile Little Owls that have been taken out of the nest to ring them also behave in
a relatively relaxed manner, although it can differ per individual. A few behave aggres-
sively by shouting loudly and making grasping movements with the beak and claws.
Snapping with the beak and defecating betrays at least some excitement. During
weighing, most of them stay relatively quiet in the weighing bowl, without any attempt
to escape. Young older than 25 days sometimes try to escape by quickly walking, or at a
later age, flying away. In general, ringing is endured relatively quietly.

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 Chapter 10: Population Regulation

Chapter Summary
A wide range of factors regulate Little Owl populations. These factors are typically
categorized as environmental (external) and demographic (intrinsic) factors (Newton
1998). This chapter has been written in two main sections following this categorization
theme. Little Owls have been shown to be directly and indirectly affected by habitat
loss, vehicle collisions, limited availability of nest and roosting sites, pesticides (i.e.,
secondary poisoning) and heavy metals, entrapment in anthropogenic structures (i.e.,
hollow metal power poles and chimneys, and drowning in water troughs), predators
and weather. They have also been shown to be susceptible to parasites, diseases and
injuries. While the Little Owl has co-evolved with a few of these (e.g., weather, preda-
tors, diseases, parasites), anthropogenic activities have substantially altered the land-
scape within which Little Owls exist(ed). When the population grows and owl densities
become higher, density-dependent processes take place and serve to stabilize the popu-
lation. In a metapopulation context, as populations become increasingly small,
immigration helps to support them, extending the survival time of these population
clusters. We summarize evidence along these lines from northern France, the
Netherlands and Denmark.
With the primary backdrop of habitat alteration, other factors (e.g., collisions with
vehicles, lack of suitable nest sites, reductions in prey due to pesticides) become additive
in reducing owl populations, and at some point in the decline the fragmented popula-
tions become susceptible to stochastic events (such as extreme weather events, e.g.,
several consecutive severe winters or poor nesting years) and individual populations
become extirpated from a given region. Unless the population decline is arrested and
reversed, the population will be unable to maintain itself over a 100-year timeframe
(the accepted timeframe for modeling trajectories of most wildlife populations) and is
doomed in the “extinction vortex.” This is what is currently happening to Little Owl
populations in Denmark (Pertoldi et al. 2012), Poland, Czech Republic and Slovakia
(Šálek and Schröpfer 2008).
The Little Owl is a very sedentary bird and will generally frequent the areas close to
its natal area. Dispersal begins when the young are 12–16 weeks of age and lasts up to at
least the end of October. Occasionally some owls do embark on long-distance move-
ments. Mean home-range sizes are 14–120 ha. Home-range and territory sizes depend
on the sex and age of the birds, population density, season and habitat. Density might

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 Chapter 10: Population Regulation

Figure 10.1 Locations of ringing and recovery (n = 108 444 observations) as reported in the EURING
Data Bank ring-recovery database (du Feu et al. 2016).

influence the reproductive success of the Little Owl, its habitat selection and its disper-
sal behavior. More and more Little Owl populations function as metapopulations
because of the fragmentation of the countryside.
For Little Owls, there is a critical need for demographic information from juvenile
owls (specifically on dispersal and survival data derived from radio-telemetry
studies) and from adults (specifically on survival and reproduction associated with
long-term mark–re-capture data on populations). The mating system hypothesis,
which predicts that the sex that establishes the territory should disperse shorter dis-
tances, was studied using the entire historical dataset of EURING, kindly made
available by its member ringing schemes. The dataset used contained 108 444 -
observations of ringing, re-capture and recovery of 59 743 unique ringed birds. The
locations of ringing and recovery of all observations are illustrated in Figure 10.1.
Ringing data thus far indicate that for live re-captures, Little Owls ringed as owlets
and then recovered at least one year later dispersed an average straight-line distance
of 14.69 km for females (n = 1303), 6.47 km for males (n = 460) and 11.61 km for
birds of unknown sex (n = 5475). For birds found dead at least one year later, the
figures were 9.07 km for females (n = 88), 6.94 km for males (n = 33) and 11.59 km for
owls of unknown sex (n = 2182). Birds ringed as adults and then later re-captured
alive dispersed an average straight line distance of 2.33 km for females (n = 2057),
2.45 km for males (n = 388) and 2.42 km for birds of unknown sex (n = 2815). For
those found dead at least one year later, the average disperal distances were 5.56 km
for females (n = 373), 5.59 km for males (n = 91) and 8.65 km for birds of unknown
sex (n = 721).
Nearly all of the demographic data we have on Little Owls to date come from studies
conducted in Europe. We strongly encourage and support researchers to undertake
projects focused on the demographic aspects of Little Owls in the Middle Eastern and
Asian portions of its range.

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 10.2 Environmental Limiting Factors

10.1 Introduction to Population Regulation

In his book on population limitation in birds, Newton (1998) stated that to understand
what limits bird populations within the habitats they occupy, we have to separate the
external (environmental) factors and intrinsic (demographic) features that these exter-
nal factors affect. External limiting factors include resources (e.g., food supplies, nesting
cavities), competing species and natural enemies (predators, parasites). Any one of these
factors can be considered limiting if it prevents a population from increasing or causes it
to decline. Particular populations may be affected by more than one, perhaps all, of
these different factors, but often one factor emerges with over-riding importance at any
one time. It is this factor that must be alleviated if we are to achieve an increase in
breeding numbers. Intrinsic (demographic) features include the rates of births and
deaths, immigration and emigration, the net effects of which mediate the influence
of external factors on local population trends. Both extrinsic and intrinsic factors can be
considered to “cause” population changes, the former as ultimate factors and the latter
as proximate ones. Thus, within suitable habitat, a population might be said to decline
because of food shortages (the ultimate cause) or because of the resulting mortality (the
proximate cause). Some extrinsic factors act in a direct density-dependent manner,
affecting a greater proportion of individuals as their numbers rise, while other extrinsic
factors act in a density-independent manner, sometimes causing big changes in
numbers, regardless of density. Severe winter weather might act in this way. A review
of factors involved in the population limitation in Holarctic owls was carried out by
Newton (2002).
The main challenge in understanding local and global population trends is to isolate
and quantify the different limiting factors. This can be done through experimentation,
by manipulating the available resources (adding food, adding extra breeding cavities) or
the impact of predators (using special predator-proof nestboxes or removal of predators,
adding fake predators like plastic snakes (Parejo and Avilés 2011)), or by collecting
various types of circumstantial evidence. Obviously, experiments are more focused
because some factors can be controlled for. In this chapter, we give an overview of
some of the factors that limit or influence Little Owl populations. We first consider
environmental factors, followed by demographic factors.

10.2 Environmental Limiting Factors

The main environmental resources on which Little Owls depend are food and nesting
sites. Other external factors that can reduce owl numbers include predators and
parasites, climate change and extreme weather, and human activity (e.g., habitat
alteration, disturbance, pesticide use).

10.2.1 Human Influence

In terms of human influences, the primary impacts on Little Owls come from changes
in human land use, persecution, application of agrochemicals, vehicle traffic, drowning

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 Chapter 10: Population Regulation

and electrocution. Each of these influences serve to limit the breeding output and limit
the health and sustainability of populations.
Van’t Hoff (2001) provides a good overview of the situation facing a Little Owl
population in response to large-scale landscape changes and loss of nest sites in the
Dutch region of Groningen. He found that the population declined markedly and
became much more fragmented. As in other places of Europe, a strong decline has been
noticed in Slovenia in recent years, attributed mainly to loss of suitable habitats, i.e.,
traditional orchards, and the use of pesticides in large quantities (Vogrin 2001). Other
examples of severe declines come from the Lower Savinja valley where the species was
common and abundant about 50 years ago (Dolinar 1951). Today it is extirpated in this
region, probably due to an increase in intensive agriculture (hop fields) that has
radically changed the valley. Denmark (Andersen et al. 2015), the Czech Republic,
Slovakia (Chrenkova et al. 2017) and Poland face structural population declines.
Changes in agricultural practices and land use are the main factors responsible for the
gradual decrease of the species in Spain. Urban development is a real threat, especially
in the eastern coastal semi-arid habitats and in the Basque countryside of the Atlantic
coast (northern Spain) (Martínez and Zuberogoitia 2004a). The reduction in dry-farmed
tree crops, in particular the carob tree (Ceratonia siliqua), has caused severe population
declines on the eastern coast. Pesticides diminish prey availability, land redistribution
reduces suitable sites for nesting or hunting, and the reduction of hedges and trees
along roads increases the risk of collisions with vehicles (Olea 1997, Blas García and
Muñoz Gallego 2003). Likewise, the enlargement of forested areas, especially in the Pre-
Pyrenees, has lead to a reduction in suitable habitats.
The most important causes of death, as reported by the EURING Data Bank (du Feu
et al. 2016) for 10 234 recovered dead birds are 27% road casualties, 8% drowning, 6%
predated by an animal, 2% shot, and railway casualty, hit glass, injured and hit man-
made structure, 1% each. Other minor causes account for 4% of all reported casualties
(Figure 10.2).
The most important period of death as reported by EURING Data Bank (du Feu et al.
2016) on 10 167 recovered dead birds is given in Figure 10.3. The principal period of
mortality is postfledging, May–September, accounting for 68% of casualties; June and
July account for 40% of all cases.
Recoveries of marked owls hold long-term, large-scale information on survival and
causes of mortality, but are prone to bias towards dead recoveries and casualties in areas
where potential finders are present. Thus, accounting for circumstance-related recovery
probabilities is crucial in statistical approaches. For the Little Owl, raw ring recovery
data suggest a strong human-related impact on survival. Naef-Daenzer and Grüebler
(2016) analyzed the proportions of the main causes of death using a large sample of
radio-tracked birds as a reference. They compared ring recoveries in southern Germany
collected in 1950–2012 (n = 465 dead of 2007 recoveries of 30 623 ringed birds) with
data from a radio-tracking study in the same region in 2009–2012 (n = 177 dead
recoveries of 377 tagged individuals). Two assumptions used in multistate ring recovery
modeling were unrealistic. First, not all dispatched rings remained available to potential
finders. Instead, 34% of tracked birds were displaced to sites where rings were

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 10.2 Environmental Limiting Factors

41 Railway casualty 44 Hit glass

50 Injured
1% 1%
2 Ring only found 1%
45 Hit man-made structure
1% 1%
10 Shot
76 Poor condition
2%
CAUSES OF DEATH (N = 10234)
3%
3 Leg & ring only found
4%
1 Bird found
22 Other
31%
4%

46 Entered building
5%

27 At nestbox/artificial site
5%

60 Taken by animal
6%

49 Drowned
8%
40 Road casualty
27%

Figure 10.2 Breakdown of circumstances of dead bird as reported in the EURING Data Bank ring
recovery database (du Feu et al. 2016).

2500

2000
Number of casualties

1500

1000

500

Month
Figure 10.3 Breakdown of month of death, as reported in EURING Data Bank ring recovery database
(du Feu et al. 2016) for all casualties, road-kills and drowning.

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 Chapter 10: Population Regulation

irretrievable, resulting in biased estimates of recovery probability. Second, the numbers

of irretrievable rings were disproportional, with 48% from predated birds and 5% from
human-induced mortality. Consequently, the sample of rings from which recoveries
were drawn differed from the sample of dispatched rings. Accounting for these biases in
a multistate model, Naef-Daenzer and Grüebler (2016) estimated the frequencies of the
main causes of mortality to be 45% for predation, 20% for casualties due to traffic and
buildings and 34% for all other causes. In radio-tracked birds, predation was even more
dominant (76%). Integrating mark–re-capture data and telemetry observations allowed
the detection and quantification of so-far unknown recovery bias and resulted in
improved estimates of key population parameters. According to Naef-Daenzer and
Grüebler the demography of Little Owls likely depends mainly on predator–prey rela-
tionships rather than on human-induced deaths.

10.2.1.1 EFFECTS OF TRAFFIC

The Little Owl suffers heavy losses to vehicle collisions on roads, which is perceived to
be the most important cause of human-induced mortality in Europe (Knötzsch 1978,
Exo and Hennes 1980). Among 15 bird species, the Little Owl was the second most
common victim of road traffic (n = 150) among birds brought to wildlife hospitals in
Belgium between 1987 and 1997 (Rodts 1994). Illner (1992) suggested that Little Owl
deaths due to vehicle collisions increase when roads transect villages. This correlates
with the owls’ relationship with man, as they routinely nest in or near human dwell-
ings in rural landscapes. Baudvin (1997) found that the rate of vehicle collisions with
Barn Owls and Long-eared Owls in France was based on the physical structure of the
roadbed (i.e., many fewer owls were killed where the roadway surface was set below that
of the surrounding terrain) and structure of roadside vegetation (fewer owls in areas
with taller natural or shrubby vegetation that limited access to voles).
It is difficult to know the exact proportion of deaths caused by traffic because deaths
from most of the natural causes (e.g., starvation, predation) go unobserved. Road
casualties may predominate because these are the owls that are more readily detected
by people. In a Belgian sample, collisions with vehicles was the cause of 34% of the total
Little Owl deaths (Bultot et al. 2001). Vehicle collisions represented 40% of known
mortality causes in eastern France (n = 28) (Génot 1991) and 36% (n = 53) in western
France (Clec’h 2001a). Clec’h (2001b) showed that 68% of breeding sites are situated
more than 1 km from high-traffic roads and speculated the reason to be because all owls
living closer had been killed. Of 89 Little Owl casualties in Belgium, 65% were birds less
than one year old and 35% were adults (Bultot et al. 2001). These data are consistent
with the mortality of Barn Owls in Britain, where Newton et al. (1997) found that 45%
of 1101 owls died from collisions with vehicles, and that 76% and 24% of these owls
were juveniles and adults, respectively.
Gomes et al. (2009) identified locations where roadkills were more frequent (i.e.,
road fatality hotspots) and factors that influenced the occurrence of such fatalities to
allow mitigation measures to be delineated. They used road fatality data collected from
311 km of roads in southern Portugal over a two-year period to compare the perform-
ance of five hotspot identification methods: binary logistic regression (BLR), ecological

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 10.2 Environmental Limiting Factors

niche factor analysis (ENFA), Kernel density estimation, nearest-neighbor hierarchical

clustering (NNHC) and Malo’s method. BLR and ENFA modeling were also used to
recognize roadkill deterministic factors. The results suggested that Malo’s method
should be preferred for hotspot identification. The main factors driving owl road-
killings were those associated with good habitat conditions for species occurrence and
specific conditions that promoted hunting behavior near roads. Based on these factors,
several mitigation measures were recommended.
Between 1995 and 2017, Bozó and Csatho (2017) carried out surveys and collected
road-kill data (n = 64) on the Little Owl in Battonya town and Kevermes village in
southern Békés county (Hungary). The number of casualties of the species increased
unambiguously over the 23 years studied. Most of the road-killed Little Owls (53
individuals, 82.81% of the total) were found in summer (June–August). In Battonya,
51 road-killed individuals (79.69%) were noted in the outer areas and 13 specimens
(20.31%) in the town. Their results highlighted that vehicle traffic is an important
mortality factor for the population.
Several studies have suggested that species mortality rates are positively related to local
population abundances. Because owls have shown both high road mortality rates and road
avoidance behaviors, we hypothesize that road-kill likelihood is not always directly linked
to their occurrence. Grilo et al. (2014) examined the relationship between the likelihood of
species occurrence in the vicinity of major roads and the road-kill risk for Barn Owls Tyto
alba, Tawny Owls Strix aluco and Little Owls Athene noctua. Their findings showed that
changes in movement patterns were probably the main behavioral mechanism that
threatened owls in roaded landscapes. The high likelihood of the occurrence of Little
Owls combined with low mortality rate suggested avoidance of road crossings. Although
it was clear that within species there was variation according to age- and territory-holding
status, closely related species had different sensitivities to roads as a function of their
varying food preferences. Grilo et al. (2014) believed that linking species distribution with
mortality risk may be more effective in conservation efforts and suggested measures should
be applied to reduce prey availability close to roads and in road verges.
The most significant mortality on roads has been found to be related to the fact that
Little Owls are vulnerable to vehicle deaths due to their habit of flying low from post to
post or hedge to hedge in pursuit of prey. Little Owls spend a great deal of time foraging
for food on the ground, which also makes them vulnerable to vehicles. When the owls
forage on the ground, they become dazzled by the headlights of vehicles, tend to freeze
and then cannot escape. The Little Owl can sometimes be surprised, feeding on the
remains of an animal killed by traffic (Van de Velde and Mannaert 1980).
Increasing road traffic, faster cars, a higher density of major road networks and,
perhaps, the physical structure of the roadbed and adjacent vegetation all serve to
increase vehicle-related mortality in the owl population. This perspective is supported
by comparisons of data from several countries. A study in Britain by Glue (1971a)
showed that 12% (24 of 198) of the owls died from vehicle collisions; in Germany,
Schönn (1986) found that road traffic was the cause of 23% (19 of 82) of the mortalities
there. Génot (1991) analyzed 751 Little Owls recovered in France for the cause of death;
52% of these owls were road traffic victims. A report commissioned by the Dutch

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 Chapter 10: Population Regulation

Department of Public Works on road traffic mortality among birds reviewed all records
of road traffic deaths and ringing recoveries for the period 1961–1990 (Van den Tempel
1993). The Little Owl topped the list of bird mortalities with 382 cases reported; 181
(47.4%) of these were ringed birds. The importance of this finding was that, based on
ringing data, nearly half of the owls in this study were killed by vehicle collisions.
Mortality due to railways is less well described in the literature, presumably because
the tracks carry less traffic than roads and are less frequented by people able to pick up
carcasses. Spencer (1965) showed that of 116 avian casualties due to railways, 41 were
owls and only 1 was a Little Owl. Génot (1991) found that of 751 dead Little Owls in
France, only one case of a railway casualty and three cases of plane casualties were
recorded. In former East Germany, three cases of railway casualties were recorded out of
98 mortalities (Schönn 1986).
The increasing road networks in Portugal threaten ecosystems by direct effects such
as increased mortality due to collision with vehicles and by various indirect effects
leading to road avoidance. Silva et al. (2012) censused Tawny Owls and Little Owls in
2005, 2007 and 2009 in a rural landscape in southern Portugal in order to study the
effects of roads and habitat characteristics on Tawny Owl density and Little Owl
presence. The presence of both owl species in the 70 census locations was coherent
between years. Their results showed that Tawny Owl density near major roads was
lower, with the negative effects extending possibly up to 2 km. The probability of Little
Owl presence was also negatively affected by proximity to major roads. The negative
effects of roads were significant, even considering habitat preferences and spatial auto-
correlation, which had the most marked effect on the density or presence of both owls.
The reduced occupancy by Tawny Owls and Little Owls of habitats near major roads
may be caused by several factors, including increased mortality, disturbance caused by
high traffic density and increased fragmentation. Traffic noise in particular may affect
intra-specific communication and hunting efficiency. Consequently, habitat near roads
may represent lower-quality territories for owls in southern Portugal (Silva et al. 2012).
Most studies have shown road deaths occurring mainly in July–August, which
suggests a link with postfledging dispersal. For example, an analysis of 418 deaths over
a three-year period in Spain showed the road mortality peak to occur in August
(Hernandez 1988).
The most important period for death by road kills, as reported in the EURING Data
Bank (du Feu et al. 2016) on 2745 recovered dead birds, is given in Figure 10.4. The
principal period of mortality by road kills (27% of total recoveries) is the dispersal
period June–July–August, accounting for 57% of casualties.

10.2.1.2 EFFECTS OF POISONING

Little Owls take a relatively wide range of small insect and rodent prey, which makes
them vulnerable to the effects of a wider variety of poisons.

Rodenticides, Anthelmentics and Organochlorines

Rodenticides are readily available and are used to poison and control rodents that
inhabit human dwellings or eat and contaminate foodstuffs that are destined for

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 10.2 Environmental Limiting Factors

800

700

600
Number of Road Kills

500

400

300

200

100

Month
Figure 10.4 Breakdown of month of death as reported in the EURING Data Bank ring recovery
database (du Feu et al. 2016) for road-kills.

human consumption. Little Owls rarely eat carrion; therefore the first-generation
rodenticides that killed the rodents outright rarely affect the owls directly. However,
most current rodenticides (which inhibit blood coagulation) are ingested by rodents
and carried for several days prior to death. In terms of LD50 values (lethal dose for 50%
of a sample, expressed as mg/kg body weight), the new chemicals are roughly 100–1000
times more toxic than the first generation (i.e., warfarin) rodenticides. It is the combin-
ation of greater toxicity and greater persistence that gives the potential for secondary
poisoning of rodent predators (Newton et al. 1997). While the rodents are alive (but
sick), they are available to the owls (perhaps more so than healthy rodents) and when
caught and eaten, the poison within the rodent is transferred to the owls. Beersma and
Beersma (2001) reported that owls affected by rodenticide poisoning suffer internal
bleeding from organs and blood vessels. This becomes visible in the acute stages, when
joints bleed, and the owls seem emaciated, even during times of high prey populations.
Eventually the owls die, the speed of which is denoted by the amount of rodenticide
accumulated from the prey.
Veterinary anthelmintics are used widely in the countryside to protect horses, cattle
and sheep against helminth fauna. The most used vermifuge is ivermectin, which
accumulates in the food chain through sustained bolus treatment (medicine given in
a pill or capsule) to grazing cattle and is toxic to Dung Beetles (Lumaret 1993). Cows are
typically given these medicines through their feed, and dung beetles pick up this
insecticide through ingestion of the cow dung. Little Owls consume both target and
nontarget species for these pesticides, ingesting an accumulation of the pesticide from
the prey. The effects on Little Owl biology are marked; the ability to use its senses for

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 Chapter 10: Population Regulation

foraging is affected and pesticide accumulation affects the breeding performance of

the owls.
While eggshell thinning is a result of an accumulation of organochlorines and
polychlorinated biphenyls (PCBs) in the metabolism of the owls, little information
is available from the national monitoring programs on the effect of organochlorines
and PCBs on Little Owls. In France, Génot et al. (1995) undertook a study on
organochlorines found in 12 dead Little Owls and 40 unhatched eggs. When they
compared their results to earlier studies in Europe, concentrations were found to
be lower than they were 15 years previously, and much lower than they were 30 years
ago. More specifically, dieldrin levels in Britain were reported at 6.5 ppm (Cramp
1963) and 0.1 ppm (n = 7) (Walker et al. 1967), while levels in France were found
to be 0.03 ppm (n = 4) (Tables 10.1 and 10.2; Génot et al. 1995). The concentrations
found in Little Owls were still significantly higher than for forest owls (Ravussin et al.
1990).
In Table 10.3, we offer results from a number of studies that have examined organo-
chlorine and PCB levels in Little Owls across Europe. In the Netherlands, Van den Brink
et al. (2003) showed that PCBs as well as DDE bioaccumulate from soil to worms and
from worms to owls. Mean concentrations and ranges in uropygial oil, expressed in
ppm, were 2.0 (0.8–6.2) for PCBs and 1.9 (0.8–17.9) for DDE. PCB levels in a Dutch
contaminated area were elevated and PCB patterns indicate hepatic enzyme induction
in owls. The levels of PCBs possibly combined with other dioxin-like compounds
may pose a risk to Little Owls in the Rhine floodplain.
Recently, brominated diphenylethers (BDEs), organochlorinated pesticides (OCPs)
and PCBs were measured in 40 Little Owl eggs in Belgium (Jaspers et al. 2005). The
major organochlorine pollutants detected were PCBs (median 2.6 ppm lipid, range
0.786–3.204 ppm lipid). BDEs were measurable in all eggs, although concentrations
were low compared to levels of persistent organic pollutants. The results revealed that
concentrations of most persistent organic pollutants were significantly higher in eggs
from deserted nests in comparison to addled eggs, even though there was no difference
in eggshell thickness between deserted and addled eggs (Jaspers et al. 2005).

Heavy Metals
Table 10.4 compares the levels of heavy metals in Little Owls across Europe. The effect
of industrial polution of watercourses that then carry the contamination and deposit it
on floodplains have been studied in detail in the Netherlands by Groen et al. (2000).
The Little Owl was an excellent study species, due to its widely varying diet, allowing
different poison absorption rates to be analyzed at different trophic levels (Figure 10.5).
The study examined the breeding biology of Little Owls in two areas (one contaminated
and an uncontaminated control) to compare breeding output. The results were inter-
esting, in that the output of fledged birds from the contaminated area was dramatically
lower than the uncontaminated control.
For Jongbloed et al. (1996), the Little Owl is particularly susceptible to the uptake of
cadmium, through the food chain of soil to worm, and worm to owl. This perspective
was also confirmed by Van den Brink et al. (2003) who considered that cadmium poses

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Table 10.1 Concentration of organochlorines and PCBs in owls in France expressed in ppm (mg/kg fresh weight)

Origin of sampled owls HCB HCH total HE D DDE TDE DDT PCB

Moselle Rahling – 0.002 0.001 – 0.005 – – 0.075

Bas-Rhin Ormersviller < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 <1
Bas-Rhin Weyer < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 <1
Bas-Rhin Weyer < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 <1
Nord Wallers – – 0.021 0.003 0.343 0.022 0.02 0.05
Nord Petite-Forêt – 0.002 0.059 0.002 0.012 – – 0.122
Nord Rumegies 0.029 0.002 0.016 0.002 0.028 – – 0.028
Lozère Drigas – – – – 0.005 – – 0.027
Lozère Drigas – 0.002 – – 0.01 – – 0.026
Lozère Drigas – – – – 0.005 – – 0.022
Lozère Aven Armand – – 0.01 0.11 0.4 – – –
Lozère Aven Armand – – – – 0.004 – – 0.011

After Génot et al. 1995.

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Table 10.2 Concentrations of organochlorines, PCBs and lead in eggs in France expressed in ppm (mg/kg fresh weight)

Year No. of eggs HCB HCH total HE DDE TDE DDT PCB Pb

1987 4 0.038 0.018 0.003 0.059 < 0.001 < 0.001 0.200
1988 4 0.013 0.003 0.008 0.080 < 0.001 < 0.001 0.420 0.043
1988 6 0.012 0.018 0.019 0.062 < 0.001 < 0.001 0.400 0.056
1989 2 < 0.001 < 0.001 < 0.001 < 0.001 0.48 < 0.001 1.28
1989 2 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 3.209
1989 5 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 0.958
1989 5 < 0.001 < 0.001 7.116
1991 7 < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 < 0.01 5.101
1992 5 < 0.01 < 0.005 < 0.01 0.214 0.214 < 0.05 5.284

After Génot et al. 1995.

 10.2 Environmental Limiting Factors

Table 10.3 Comparison of levels of organochlorines and PCB in Little Owls

across Europe

Author (Study No. of

Country period) eggs HCB DDE HE PCB

Belgium Joiris and Delbecke 4 0.01 0.11 0.7

1981
(1969–1974)
Netherlands Fuchs and Thissen 102 0.02–0.27 3–10 0.02–0.06 0.4–09
1981
(1972–1977)
Switzerland Juillard 1984 112 0.12 0.80 0.04 1.45
(1976–1981)
Germany Bednarek et al. 0.56 0.26 0.09 1.2
1975
(1972–1973)
Hahn 1984 23 0.049 0.344 0.026 0.647
(1981–1982)
France Génot 1992a 35 0.01 0.03 0.005 0.46
(1987–1991)

Mean and range of concentrations are expressed in ppm (mg/kg) in comparison to fresh
weight.

a particular risk when the owl feeds exclusively on earthworms over a long period
of time.
In eastern France, the mean concentration of lead in Little Owl eggs was 2.75 ppm
(n = 6) (Génot et al. 1995). This level was more than 10 times higher than the level of
0.2 ppm (n = 18) reported by Juillard et al. (1978) for eggs from Switzerland. As the Little
Owl is linked with prey in more open habitats, it appears much more likely to pick up
lead pollutants than forest owls. For example, Ravussin et al. (1990) found the mean
concentration of lead levels in eggs of the Tengmalm’s Owl to be 0.07 ppm (n = 25).
Except for some samples, the contamination of eggs with heavy metals was low in
France (Génot et al. 1995).
In southeastern Spain, the exposure of raptors to lead (Buzzard, Kestrel, Eagle Owl
and Little Owl) was recorded through samples of liver, kidney, brain, blood and bone.
Relationships were found between the size and age of the birds, proximity to areas of
human activity and lead concentrations in tissues. The larger the bird, the higher the
lead concentration in all tissues. Adults accumulated more lead than nestlings and
immature birds, except for lead in the bones which was lower for adults, possibly due
to the removal of lead from circulation and deposition in ossifying bone. The lead
distribution pattern revealed that bone was the principle part of the body for

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Table 10.4 Comparison of the levels of heavy metals in Little Owls across Europe.
Lead (Pb) Cadmium (Cd) Mercury (Hg) Chromium (Cr)

Country Author Weight n Liver Muscle Brain Feather Liver Muscle Brain Feather Liver Muscle Feather Liver Brain Feather

France Génot et al. 1995 Fresh 8 0.46 0.057 0.382 0.045 0.139 0.029
Italy Battaglia et al. Dry 38 0.35 0.10 2 0.05 0.003 0.05
(personal
communication)
Italy Zaccaroni et al. Fresh 41 0.20 0.057 0.110 0.054 0.288 0.057
2003 11 0.232 0.070 0.105 0.061 0.297 0.076
Netherlands Van den Brink et al. Dry 4 <40 <2.3 17 27
2003

Mean of concentration expressed in ppm (mg/kg dry or fresh weight according to the study).
 10.2 Environmental Limiting Factors

Figure 10.5 Accumulation rates of heavy metal poisons (cadmium) absorbed into the food chain
consumed by Little Owls (after Groen et al. 2000).

accumulation (900–6000 ppm, dry weight), followed by the kidney (46–375 ppm, dry
weight), the liver (35–315 ppm, dry weight) and the brain (14–149 ppm, dry weight).
The correlation between lead concentrations in soft tissues and in blood were high
(Garcia-Fernandez et al. 1997).
A review of biomonitoring of metals and metalloids in the Little Owl by Carneiro
et al. (2016) gives an overview of relevant publications from Portugal and Spain.
Cadmium was found in blood, liver, kidney, brain and bone. Lead was found in blood,
liver, kidney, brain and bone. Arsenic and zinc were found in liver and mercury in
feathers (García-Fernández et al. 1995, 1996, 1997, Pérez-López et al. 2008, Lourenço
et al. 2011).

10.2.1.3 OTHER ANTHROPOGENIC CAUSES

Drowning
Most drowning cases are of soiled females who have endured a breeding season without
bathing, and young birds that are leaving their natal territory. These owls become
immersed in cattle water troughs, which have steep slippery sides, their feathers
become waterlogged, they cannot recover and drown. In a French inquiry about
mortality causes between 1980 and 1990, Génot (1991) found that 21 of 751 (2.8%)
owl deaths were due to drowning. In the French region of Ile-de-France, drowning
represented 20% of the mortality among young owls (Lecomte 1995). Of 53 cases of
mortality in western France, 4 (7.5%) were due to drowning (Clec’h 2001a). In Belgium,
drowning represented 9.6% of 89 mortalities (Bultot et al. 2001).
The most important period of death by drowning as reported by the EURING Data
Bank (du Feu et al. 2016) on 667 recovered dead birds (8% of total recoveries) is given in
Figure 10.6. The principal period of mortality by drowning is the dispersal period, June–
July–August, accounting for 73% of casualties by drowning.

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250

200
Number of drownings

150

100

50

0
J

Month

Figure 10.6 Breakdown of month of death as reported in the EURING Data Bank ring recovery
database (du Feu et al. 2016) for drowning (n = 667).

Electrocution
One of the most common views that people have of Little Owls is of them sitting on top
of a telephone or electricity pole. Reports of deaths resulting from electrocution are
surprisingly rare. However, a report from a local birdwatcher in Cyprus reported a small
owl hanging by its feet from overhead electricity supply wires; it was assumed that the
owl had come into contact with two or more wires as it perched.
In an analysis of ringing recovery data on Little Owls in France from 1927–1990,
P. Raevel (personal communication) noted that 1.9% of the recoveries were of owls
recovered as a result of electrocution by power lines. This low rate of electrocution-
related recoveries classified the species as not sensitive to the risk of power-
line mortality.

Shooting and Other Persecution

Like most birds of prey, the Little Owl has been the subject of a high degree of
persecution, particularly in the late eighteenth century and early to mid-nineteenth
century. The Little Owl’s diet was not fully researched, and the suspicion was that the
main component of its diet was game birds. Many methods of persecution were used,
including shooting, pole or tunnel trapping (Hewson 1972), poisoning and egg
pricking, to name a few. Hibbert-Ware (1938a) undertook a survey on the diet of the
Little Owl and disproved the gamekeeper’s and hunting fraternity’s perspective of the
owl. In England and Wales it was noticeable that during World Wars I and II, when
most gamekeepers were serving their country, the population exploded and colonized
new areas, unhindered by the controls of persecution.

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 10.2 Environmental Limiting Factors

Even though laws have protected the Little Owl in the UK since the 1950s, persecu-
tion continues today in some places. Diurnal activities make the owl an easy target for
unscrupulous shooters. Likewise, the species is still killed in southern Italy where
24 Little Owls were reported shot in 1974–1975 and 9 in 1981 and 1983 (Falcone
1987). Similarly, in north Caucasia the Little Owl has been shot by local inhabitants
in all seasons and especially in the breeding period (Afanasova and Khokhlov 1995). In
the Ural region of Russia, shepherds rob the nests of Little Owls that breed in farms and
summer camps of cattle (Karyakin 1998). It seems likely that part of this indiscriminate
killing may have some roots in cultural beliefs (see Chapter 1).
The EURING Data Bank (du Feu et al. 2016) has reported only 156 shot birds (2% of
total recoveries).

Trapping in Hollow Metal Telephone Poles

In France, hollow metal telephone poles throughout the countryside constitute a trap
for many birds that are looking for cavities to nest or roost in. The hollow poles are
structured such that once the birds enter the hollow posts (which are open at the top)
they are unable to escape. In western France, eight Little Owls were found dead in
32 telephone poles (Clec’h 1993). In the region of Ile-de-France, 13 Little Owls were
found dead in 305 telephone poles (Savry personal communication). In the beginning
of the 1970s some 3 500 000 such telephone poles were put up in France (Clec’h 1993).
From the analysis of local records, we can imagine that these poles could very likely
cause the local extirpation of populations already suppressed due to other factors.

Trapping in Chimneys
As Little Owls are cavity nesters, and often live in proximity to villages and towns,
many Little Owls visit chimneys. Once inside, like the hollow metal telephone poles,
they are unable to fly/climb the steep walls, become trapped and die. Zvářal (2002)
believes that dying in chimneys and shafts is the most important mortality factor in the
Czech Republic. Of the overall mortality rates, this cause of death has been documented
in a number of areas: 13% (7 of 53) in western France (Clec’h 2001a), 7% (2 of 31) in
eastern France (Génot 2005), 19.5% (19 of 98) in eastern Germany (Schönn 1986) and
4% in Britain (Glue 1971a).

Land-Use Changes
The loss of habitat that has taken place from the 1960s until now is mainly responsible
for the decline of Little Owl populations in all of western Europe. The land-use changes
reflect the conversion of meadows into crop fields, drainage of wet meadows, cutting of
isolated trees (e.g., willows and oaks) or hedges, transformation of traditional orchards
into “fruit tree crops” (also called “low-stem orchards”) and the general shift to indus-
trial farming, with its associated large fields without perches, nesting sites or hunting
grounds, and with the use of pesticides. In addition, areas that were previously man-
aged in a pastoral or agricultural context have been encroached upon with the devel-
opment of big cities and intermediate-sized towns, highways, railways, roads, airports,
and industrial and business areas. The loss of habitat has been measured in a number of

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 Chapter 10: Population Regulation

study areas in Germany. In Schleswig-Holstein, the area of pasturelands was reduced by

20% between 1960 and 1980 (Ziesemer 1981). These former pasturelands were trans-
formed into arable fields, thus becoming less suitable for the Little Owl. In Northrhine-
Westphalia, on a study area of 4316 ha, the proportion of the landscape as grassland
decreased by 34% between 1919 and 1991 (Kämpfer-Lauenstein and Lederer 1995). In
Baden-Württemberg, between 1970 and 1973, 16 693 fruit trees (mainly apple and pear,
which offer the best holes for the Little Owl) were cut (Ullrich 1975a). In Westphalia,
Ilner (1981) reported that the decrease in Little Owl numbers was due to the loss of nest
sites and food resources from the conversion of a mosaic landscape composed of
meadows, pastures, hedges, orchards and field edges.
In Switzerland, Juillard (1989) explained that the decrease in the Little Owl popula-
tion between 1950 and 1985 was mainly due to the destruction of favorable biotopes, in
particular, the felling of several million fruit trees. This came from organized campaigns
to restructure the orchards in order to obtain fruit-tree crops.
A smaller-scale study was conducted in Northrhine-Westphalia (Germany) where
Loske and Loske (1981) noted the following transformation of an area of about 1 km²:
7% in crops in 1970 and 62% in 1980, 545 pollarded willows in 1970 and 271 in 1980.
Survey efforts conducted during this time found four calling Little Owl males in
1970 and only one in 1980.
Urban development is also a threat to Little Owl populations. An evaluation of land-
use plans in three municipalities in Northrhine-Westphalia, where 115 breeding pairs
of Little Owl were present, predicted the loss of 8% to 50% of the breeding pairs (Breuer
1998). The importance of a “green system” including mown meadows and orchards for
the species has been illustrated through the size of the home range of the Little Owl,
i.e., 20.3 ha in the “green system” and 74.8 ha in the “arable system” (Orf 2001).
In France, transformation of the rural landscape has been substantial. Four areas
(totaling approximately 26 000 km2) in western France lost 174 000 km of hedges
between 1962 and 1990. For all of France, there were some 48 million apple trees
in 1929 and 11 million in 1990. A substantial portion (20%) of this loss in apple
orchards occurred during the period 1981–1990. Also, significantly, 25% of the area
as meadows was converted to other land uses between 1970 and 1995 (Pointereau and
Bazile 1995).
In eastern France, the last available habitats for the Little Owl remain around villages
in “green belts,” which include orchards with old fruit trees, meadows, pastures and
gardens. Between villages, the former landscape has been modified by industrial agri-
culture, re-allocation of land in 1970–1990 (where people exchanged land to obtain
larger contiguous parcels, mainly for larger-scale agricultural purposes) and the recent
conversion of meadows into arable land resulting from European agricultural policies.
So, outside of the green belts containing habitat close to the villages, the landscape has
become unfavorable for Little Owls. Pairs become more and more isolated with frag-
mentation of their former habitat and the local populations are weak and very suscep-
tible to stochastic environmental disturbances, such as the storm on December 26,
1999, when several thousand fruit trees were blown down (Génot personal
observation).

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 10.2 Environmental Limiting Factors

In Austria the changes in landscapes, notably the increase in area of arable fields,
decrease of grasslands, and the reduction in habitat heterogeneity explained the large
decline in Little Owl. The patch size of agricultural areas and the insect prey density and
diversity are the important factors in maintaining populations (Ille and Grinschgl
2001). The enlargement of the size of parcels to more than 20 ha is followed by a
decrease in both the number of beetle species and number of individuals (Deveaux in
Lefeuvre 1981).
The decline of the Little Owl is often due to a synergy between different factors. In
the Northern Vosges (France) a combination of adverse climate (winters of 1985–1987),
predation, road traffic, loss of orchards and grasslands, fragmentation of good habitat
were the primary drivers behind the reduction in owl numbers (Génot 1992c).
Similarly, in the Basel region of Switzerland, the decline was due to predation by
Stone Martens, climatic conditions (in particular rainfall during feeding of nestlings),
lack of prey in hay fields, road traffic and a lack of connectivity to other surrounding
populations (that could offer demographic support) (Luder and Stange 2001).
Land-use changes in farmland rank among frequently discussed negative factors
causing the population decline of the Little Owl in Central Europe. Poprach et al.
(2017) analyzed land-use changes at nest sites of the Little Owl in the South-
Moravian region (Czech Republic) between the years 1976–1977 and 2014. In both
periods studied, the most important land-use type within 500 m of the Little Owl nest
sites was arable land (66.94–62.25%), followed by built-up areas (19.97–22.41%), while
other land-use types made up less than 5%. The proportion of the particular land-use
type did not change significantly between the years 1976–1977 and 2014. The most
important change was the decrease in the area of arable land by 4.69% and that of
orchards and gardens by 1.99%, while the surface area of built-up areas increased
slightly by 2.45% and that of meadows and pastures by 1.5%. The analysis showed
that at the known nest sites of the Little Owl in the South-Moravian region (Czech
Republic), there were no significant changes in the proportion of the particular land-use
types within 500 m from the nests between the years 1976–1977 and 2014. Based on
these results, Poprach et al. (2017) concluded that land-use changes in their study area
were not important as an influencing factor for the observed decline of the Little Owl.
The impact of climate change is expected to be mixed, depending on the region.
Desertification will have a negative impact, while milder winters will promote other
crops that might have a positive impact. In the steppe region of the former Soviet
Union, the ongoing desertification process, as seen in Dagestan, is a threat to the Little
Owl (Vilkov 2003).
Ilanloo et al. (2020) reported distance to forests, distance to human settlements and
distance to agricultural lands as the most important determinants of Little Owl distri-
bution in Iran. Human migration from rural areas to cities and availability of many
vacant human settlements surrounded by traditional and semi-modern agricultural
landscapes produce suitable habitats for the species in Iran.
While the majority of the data in the above discussion on habitat loss and modifica-
tion has come from Europe, it is reasonable to expect that similar types (and impacts
from) alterations in land use are occurring in other parts of the owls’ range.

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10.2.2 Food
The prey resources available to owls vary from year to year. Small mammal prey (e.g.,
voles) typically follow cycles with peaks of abundance every three to five years. In years
of scarcity, the breeding performance and survival rates of both adults and first-year
birds may be low, causing a population decline (Taylor 1994, Van Veen and Kirk 2000).
Resident owls respond to prey cycles in two ways, firstly by switching to other prey and
secondly by producing fewer eggs, or in some cases none at all (Newton 1998, 2002).
On average, female Little Owls lay 1.0–1.2 more eggs in good vole years than in poor
ones (Exo 1992).
The Little Owl’s ability to forage for a wide variety of small prey will aid it when one
of its prey species becomes scarce, enabling it to compensate to some extent. However,
in years when small mammal populations crash, the alternative prey, invertebrates or
birds, often provide less energy reward per unit of hunting effort.
Juvenile mortality tends to rise as the young gain independence from their parents,
and dispersal from natal areas tends to be longer in years when prey are scarce
than abundant.
Many farmland bird species have declined markedly in Europe in recent decades
because of changes in agricultural practice. To unravel specific causes of the decline of
the Little Owl in Denmark, Thorup et al. (2010) investigated the likely reasons for the
population decline in Denmark by identifying patterns of local extinction (5  5 km
grid cells) and estimating demographic parameters affecting local survival, focusing on
changes over time and their relationship to habitat characteristics. The distribution of
the Little Owl in Denmark contracted considerably between 1972–1974 and
1993–1996. The extent of contraction varied across the country, and the only habitat
correlation was that local disappearance was associated with smaller amounts of agri-
cultural land. Analyses of ring recovery data suggested a constant annual adult survival
rate of 61% from 1920 to 2002, which is similar to estimates from countries with stable
populations. First-year annual survival rates were much lower than values previously
reported. From the 1970s into the twenty-first century, the mean number of fledglings
declined from around three to less than two young per territory, but the decline in
clutch size was considerably less. Reproductive parameters were higher closer to habitat
types known to be important foraging habitats for Little Owls, and were also positively
correlated with the amount of seasonally changing land cover (mostly farmland) within
a 1 km radius around nests, as well as temperatures before and during the breeding
season. Experimental food supplementation to breeding pairs increased the proportion
of eggs that resulted in fledged young from 27% to 79%, supporting the hypothesis that
the main proximate reason underlying the ongoing population decline is reduced
productivity induced by energetic constraints after egg-laying. Danish conservation
efforts currently target enhancement of food availability during the breeding season.

Food Supplementation
In Little Owls, energy supply during growth is a major factor in the physical condition
and survival prospects of the fledglings. Differential quantity and quality of fledglings

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 10.2 Environmental Limiting Factors

produced under varying nestling food supplies are likely to affect the number of
offspring that are recruited into the breeding population. To better understand the
underlying mechanisms and associated consequences, a partial cross-fostering and food
supplementation experiment was undertaken by Perrig et al. (2014). They estimated the
effect of variation in food supply during growth on nestling survival and fledgling
phenotypic traits for 56 broods. Partial cross-fostering allowed only some nestlings
from the same family to be supplemented. Thus, intrinsic factors acting on survival
and phenotypic development could be statistically controlled for. Two synchronous
broods were denoted as partner broods. One of the two was randomly assigned to
experimental treatment (i.e., food supplementation). Depending on brood size, one
or two nestlings of similar age and body weight were exchanged among partner broods,
keeping brood sizes constant. Survival to fledging was much higher in food-
supplemented nestlings (98.6%) than in control nestlings (82.4%). Furthermore, sup-
plemented nestlings were, on average, 8.9 g heavier and were more likely to develop
subcutaneous fat deposits (99.4% versus 73.7% for treatment and control nestlings,
respectively). Supplemented nestlings also had, on average, longer wings than control
nestlings, but tarsi and culmen did not differ significantly. Furthermore, experimentally
supplemented fledglings struggled more when handled and emerged sooner from tonic
immobility than control fledglings. The irises of supplemented fledglings were less
intensely colored. The experimentally induced changes in nestling development prob-
ably affected individual performance beyond fledging. Nestlings from orchard-
dominated habitats were larger than those from habitats with mainly arable land. As
nestling food supply is largely determined by natural food availability, Perrig et al.
(2014) concluded that habitat quality affects Little Owl productivity and offspring
quality, and ultimately, population dynamics.
Since the population of Little Owls in Denmark is close to extinction Jacobsen et al.
(2016) undertook a food supplementation experiment to counter food limitation
during the breeding season. Efforts to improve breeding success include providing
breeding pairs with supplementary food and attempts to improve foraging habitats by
creating short grass areas near the nests. In addition to increasing the reproductive
output, feeding and habitat management may allow parents to work less hard, thus
improving their future reproductive value. Jacobsen et al. (2016) studied the working
efforts of five radio-tagged Little Owl pairs in years of absence and presence of food
provisioning, and/or access to short-sward vegetation areas near to the nest. They
quantified movement as the minimum flight distance per hour (MFD), using the mean
distance from the nest (DN) as a supplementary index. Under unmanipulated condi-
tions, males had higher MFD and DN than females. If provided with food and/or areas
with short-sward vegetation, males, but not females, reduced their MFD and DN
significantly. If MFD was adjusted for DN (the two measures correlated positively), both
sexes reduced their DN-adjusted MFD in response to food provisioning but not to
habitat provisioning. Food provisioning therefore had similar proximate effects on
the foraging effort of males and females, whereas provisioning of short-sward habitats
had an indirect effect on male but not female MFD, because of decreased commuting
distances between the nest and foraging sites. The results of Jacobsen et al. (2016)

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indicated that food provisioning not only leads to increased reproductive output, but
also to decreased work effort, which in turn may improve adult survival.

10.2.3 Nest and Roost Sites

A lack of natural cavities has been shown to limit Little Owl populations (Exo 1983), as
no new sites are available for birds to take up as first-time breeders or to relocate from
damaged nests. The Little Owl has high nest-site fidelity, making the problem worse,
should any damage occur to their nest.
The provision of additional nest sites provides support for the Little Owls and
accommodates other cavity-nesting species (who compete with Little Owls for the same
cavities). We much prefer the planting and long-term protection of native tree species
that cost-effectively provide natural cavities for owls, as well as other ecological bene-
fits. As a short-term remedy for the shortage of these cavities, nestboxes can be used. In
Chapter 11 on Management and Conservation we give an overview of nestbox cam-
paigns that have enhanced Little Owl populations.
In the steppe area of Portugal, the number of alternative suitable cavities emerged as
the main variable explaining nest-site selection, with the owls showing a preference for
stone piles containing larger-sized stones (Tomé et al. 2004).
The use of artificial nestboxes to bolster populations of Little Owls has increased
significantly, but spatial variation of nestbox occupancy rates and breeding success
within a nestbox population has been little studied. In a case study with 798 Little
Owl nestboxes established in central Germany, Gottschalk et al. (2011) studied the
dependence of occupancy rate and breeding success on the characteristics of the
surrounding habitat. The analysis focused on two aspects of general concern for
nestbox management: whether nestboxes were occupied for breeding or were left
unoccupied, and whether Little Owls had different reproductive rates, depending on
the location of the nestbox. A high resolution (1  1 m grid cells) land-use map was
used to analyze species–habitat relationships, and Generalized Linear Mixed Models
were used to predict suitable nestbox locations. During the period 2004–2006, 544
(68%) of the nestboxes were never occupied, 144 (18%) housed birds with low breeding
success and only 108 (14%) housed pairs that produced more than 2.35 nestlings
annually, a reproductive rate thought necessary for population stability. Nestboxes
were more likely to be occupied if they were located near orchards, at lower altitude,
and in areas with higher densities of fields and less forest. Higher breeding success was
associated with fallow fields and field margins, and with greater distance to roads and
forests. The results of Gottschalk et al. (2011) suggested that the efficiency of nestbox
programs could be substantially increased if unoccupied nestboxes were relocated to
sites where occupancy is more likely, and if unproductive nestboxes were relocated to
locations that would enhance breeding success.
Bock et al. (2013) studied roost-site selection throughout the year in order to collect
information about year-round habitat requirements at different life-history stages and
to get more insight into the roosting behavior of the owls. The roost-site selection was
studied by repeated daytime location of 24 adult and 75 juvenile radio-tagged

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 10.2 Environmental Limiting Factors

individuals from July to November. The owls preferred sheltered roost sites such as tree
cavities with multiple entrances. They increasingly used sheltered sites from summer to
winter and preferentially used sheltered roost sites with low ambient temperatures.
Juveniles used significantly less sheltered sites during dispersal than before and after-
wards, and used less sheltered sites than adults within their home range. The survival
probability of birds roosting frequently at exposed sites was reduced. Roost-site selec-
tion is probably driven by the two mechanisms of predator avoidance and thermore-
gulation, and the costs of natal dispersal may include increased predation threat and
higher energy expenditure for thermoregulation (Bock et al. 2013). Adequate roost sites,
such as multientrance tree cavities and extra nestboxes are important for Little Owls
and the habitat quality can be improved by manipulating their availability.

10.2.4 Predation
The predation rates on Little Owl nests in nestboxes with and without anti-predator
devices were monitored from 2000 to 2004 by J. Bultot (personal communication). The
overall predation rate on 1004 active nestboxes without anti-predator devices was
9.45%. No predation occurred on 509 nests (predation rate of zero (0)) equipped with
anti-predator devices. Thus, with the addition of anti-predator devices, the overall rate
of predation went down to 6.3% on 1513 active nests. Blanke (2005) studied five
research areas in the Netherlands, reporting a predation rate of 10% on 20 active nests
in Ruinen, 80% on 5 active nests in Geleen, 3% on 305 active nests in Zuidoost-
Achterhoek, 5% on 433 active nests in Doesburg, and 2% on 69 active nests in Raalte.
Vanden Wyngaert (2005) reported the predation rate by Stone Martens to be 5% on
37 active nests in Dijleland, Belgium. Eick (2003) radio-tracked 10 juvenile Little Owls
of which 5 (50%) were killed by predators (Goshawk, Fox, Tawny Owl or Goshawk,
unknown mammal and a Sparrowhawk). These owls were tracked 1, 1, 2, 48 and 101
days before they were predated.

Beech or Stone Marten Martes fuina

Mammals
The small size of the Little Owl means that it has many possible predators among
mammals and birds of prey. For mammals, the Stone Marten Martes foina is recognized
as an important predator of the Little Owl, as stressed by many authors (Knötzsch 1978,

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Ullrich 1980, Hahn 1984, Schönn 1986, Veit 1988, Krischer 1990, Haase 1993, Peter
1999, Luder and Stange 2001). Starting in the 1970s, ornithologists have been building
nestboxes designed to protect Little Owls from the Stone Marten (Schwarzenberg 1970,
Furrington 1979, Marié and Leysen 2001). The impact of marten predation on the eggs
of nesting Little Owls has been quantified in Germany and France. In Germany, Illner
(1979) found the predation rate on eggs to be 10–20%. In France, Génot (in Van
Nieuwenhuyse et al. 2008) found a predation rate of 28% (61 of 218) on unhatched
eggs. The Stone Marten is a generalist predator and will eat eggs as well as young or
adult owls. Martens are an important predator of Little Owls as they are agile climbers,
will search all of the tree holes and other cavities, and eat whatever small prey they find
within. Visits to holes and nestboxes by martens are revealed by their fecal droppings
found inside the boxes. Like others in the Mustelidae family, martens are territorial, and
by marking the places they visit, they also scent-mark the cavity against other martens.
Predation by Stone Martens probably does not depend on Little Owl densities, because
during foraging it visits the same cavities again and again, even after a clutch has been
found. Little Owls faithful to their nest even after a failure might become easy victims to
marten predation. The Stone Marten benefits from human activities, and is expanding
in numbers in cities as well as in the countryside (Van den Berghe 1998). Even if the
Little Owl is not the main prey of the Stone Marten, it can be considered an important
food source (Bergerud 1984).
In Portugal, the presence of two predators (the Stone Marten and the Common
Genet Genetta genetta) emerged as the main factor linked to nest-site selection by
Little Owls in woodland areas (Tomé et al. 2004). In the Volga Region of Russia,
Frolov et al. (2001) considered that the Little Owl suffers from predation from feral cats
and Stone Martens, whose numbers have increased in anthropogenic landscapes. While
predation by Stone Martens may threaten some isolated and small Little Owl popula-
tions, as Illner (1988) said, in the long term, predation has probably contributed little, if
anything, to the population decrease in comparison to the reduction in food supply.
Other mammals can destroy the eggs or chicks, including the Stoat Mustela erminea,
Hedgehog Erinaceus europaeus (Glue and Scott 1980), Brown Rat Rattus norvegicus (Glue
and Scott 1980, Coppée et al. 1995) or can kill the adults, such as the feral cat (Glue and
Scott 1980, Haase 1993), Fox Vulpes vulpes (Knötzsch 1978) and domestic dog (Glue
1971b, Juillard 1984). We have even witnessed trampling by cows as a cause of mortal-
ity among young that had fallen (or jumped) from the nest. During a re-inforcement
experiment in France, of 35 Little Owls born in captivity and equipped with radio-
transmitters at release, 22 were found dead within one day. In some cases the reason
was known: fox (2), cat (1), bird of prey (2), stoat or weasel (1), unknown carnivorous
mammal (2) (Génot and Sturm 2003). Through radio-tracking of Little Owls, Eick
(2003) found one juvenile Little Owl captured by a fox and one by an unidentified
mammal.

Birds
Interactions between top predators and mesopredators of the same guild often result in
habitat segregation restricting interactions to shared habitat edges. Although negative

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 10.2 Environmental Limiting Factors

edge effects are recognized as important spatial patterns in the ecology of fragmented
landscapes, the underlying mechanisms of predator–prey interactions resulting in
negative edge effects remain unknown. To disentangle top-down effects of intra-guild
predators and bottom-up effects of shared resources on mesopredator spatial distribu-
tion, Michel et al. (2016) recorded the occurrence of Tawny Owls in forests and their
prey, the Little Owl, in adjacent open areas over two years across 687 km² in southern
Germany. They developed a new, asymmetrical dynamic two-species occupancy model
investigating spatial interactions while accounting for imperfect detection. Little Owl
occupancy was strongly reduced within 150 m of forests, but only in the presence of
Tawny Owls. Analysis of over 30 000 telemetry locations of 275 Little Owls showed that
Little Owls strongly avoided areas closer than 150 m from the forest during range use,
suggesting that the negative edge effect is due to forest edge avoidance rather than
direct predation. Potential confounding mechanisms such as food depletion or habitat
avoidance at forest edges can be ruled out. Thus, top-down effects caused by avoidance
of intra-guild top predators shape the spatial distribution of mesopredators such as the
Little Owl. While habitat complexity mitigates multitrophic interactions within habi-
tats, it is expected to re-inforce multitrophic interactions between habitats, potentially
leading to the suppression of mesopredators from suitable habitats.
Little Owls are also found in the diet of birds of prey, as shown by pellet analyses.
Among the birds of prey, the Eagle Owl Bubo bubo seems to be the main predator
(Blondel and Badan 1976, Herrera and Hiraldo 1976, Mikkola 1976, Orsini 1985,
Simeonov 1988, Martinez et al. 1992, Jay 1993, Karyakin and Kozlov 1999), followed
by the Tawny Owl Strix aluco (Schnurre 1940, Staton 1947, Gunston 1948, März and
Weglau 1957, Mikkola 1976, Baumgart 1980, Schönn 1980, Ullrich 1980, Zuberogoitia
and Campos 1997, Eick 2003). Peaceful co-habitation is possible during the breeding
season (Rusch 1988) and might be due to the fact that the female does not hunt during
that period (I. Newton, personal communication). When Little Owl calls are broadcast
and a Tawny Owl answers, Little Owls remain silent (Schönn et al. 1991). The avoid-
ance of forest and forest edges was recognized as a response to the presence of the
Tawny Owl (Zabala et al. 2006, Michel et al. 2016).
While observed much less frequently, other birds of prey can kill Little Owls: Barn
Owl Tyto alba (Castellucci and Zavalloni 1989, Bonvicini and Maino 1993, Sgorlon
2005), Long-eared Owl Asio otus (Glue 1972, Mikkola 1976), Peregrine Falco peregrinus
(Formon 1969, Mikkola 1976, Schönn et al. 1991), Buzzard Buteo buteo (Christie 1931,
Mikkola 1983), Goshawk Accipiter gentilis (Mikkola 1976, Nore 1977, Schönn 1986, Eick
2003), Sparrowhawk Accipiter nisus (Nore 1979, Mikkola 1983), Red Kite Milvus milvus
(Mikkola 1983, Schönn et al. 1991, Eick 2003), Tawny Eagle Aquila rapax, Rough-legged
Buzzard Buteo lagopus, Booted Eagle Hieraaetus pennatus, Lanner Falco biarmicus
(Mikkola 1983), Marsh Harrier Circus aeruginosus, Black Kite Milvus migrans, Imperial
Eagle Aquila heliaca and Long-legged Buzzard Buteo rufinus (Gavrin 1962). In the desert
habitat of Kazakhstan without shelters, the Little Owl can become locally extirpated in
winter by Long-legged Buzzards and large falcons (Osmolovskaya 1953).
Besides birds of prey, a case of egg predation by a Magpie Pica pica was recorded (Glue
and Scott 1980). Magpies may also mob the owls (Ritzel and Wulf 1978).

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 Chapter 10: Population Regulation

Snakes
Parejo and Avilés (2011) studied whether three different Mediterranean cavity-nesting
bird species assess predation risk before reproduction and how this information influ-
ences their choice of breeding territory and subsequent reproductive strategies. To do
so, they experimentally increased predation risk by presenting realistic models of the
Spanish Montpellier Snake Malpolon monspessulanus at nest sites. Snakes constitute the
most important group of nest predators for open-nesting birds (reviewed in
Weatherhead and Blouin-Demers 2004), and evidence suggests that snake predation
is also an important determinant of nest failure for cavity-nesting birds (Hensley and
Smith 1986; Neal et al. 1993). The Spanish Montpellier Snake is the most abundant
snake in the Mediterranean area (Pleguezuelos 2003), and detailed studies have revealed
a high frequency of avian eggs and chicks in its diet (Monrós 1997).

10.2.5 Parasites and Disease

A complete list of 67 species of parasites of the Little Owl was given by Juillard (1984)
and Schönn et al. (1991). Additional insights into parasites were offered in the works of
Borgsteede et al. (2003), Sanmartin et al. (2004) and Ferrer et al. (2004). The last authors
showed that the Little Owl had the highest level of infestation by parasitic helminth in
the digestive tract of six species of owls in Spain, likely due to its generalist diet. The
main types of parasites include representatives from the following groups: Protozoa
(Falgellata), Sporozoa (Coccidia and Haemosporidia), Metazoa (Plathelminthes,
Trematoda and Cestoda; Nemathelminthes, Nematoda and Acanthocephala), Arachnida
(Acarina), Insecta (Mallophaga, Diptera and Siphonaptera). Another acarine species,
Ornithogastia ariadnae, found in Ukraine can be added to the list (Gushcha 1982).
Tomé et al. (2005) studied the relationships between the occurrence of blood
parasites and host traits in wild Little Owls in the Mediterranean habitats of southern
Portugal. Of 39 owls captured between February and August 1999, 16 (41.0%) were
infected with Leucocytozoon ziemanni. One individual was infected with Trypanosoma sp.
and another with a microfilaria. Age was the only variable influencing the prevalence of
L. ziemanni in a multivariate logistic regression model comparing infected and non-
infected owls. Prevalence was much higher in adults (82.4%, n = 17) than juveniles
(9.1%, n = 22). This difference was probably due to the scarcity of appropriate vectors
(black flies, Diptera, Siimulidae) in the area at the time of fledging. Univariate analysis
also suggested a trend for higher prevalence among females than males and a difference
between bill lengths of infected (shorter-billed) and uninfected (longer-billed) adult
owls. Korpimäki et al. (1995, 2002) studied the important relationships between blood
parasite loads, nest defense and reproductive performance in the Tengmalm’s Owl and
Eurasian Kestrel. Because reproduction and immune defenses are both thought to be
energetically costly, the birds may face a trade-off between allocation of resources to
reproduction and immunity. The authors found that defense against blood parasites is
indeed costly to the owls and kestrels (e.g., reduced clutch size, less vigorous defense of
offspring by male owls) and that costs may vary with the sex of the host species and
with prevailing environmental conditions. Little Owls are subject to a very wide range

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 10.2 Environmental Limiting Factors

of parasites (both internal and external). The implications from the studies described
above are that blood parasite levels have a very real impact on raptors, and through
additional investigations, new insights into the breeding ecology of Little Owls can
be ascertained.
During his study, Juillard (1984) found seven species of ectoparasites, mainly on
nestlings. Depending on the nature of the ectoparasites, they are typically found
around the eyes and bill, on the eyelids, on the body and in the feathers. One of the
ectoparasites found by Juillard in Switzerland, the dipteran Carnus hemapterus, was also
identified in Germany as a parasite on nestlings of many other birds, including the
Peregrine Falco peregrinus, Barn Owl Tyto alba, Kestrel Falco tinnunculus, Starling Sturnus
vulgaris, Great Tit Parus major, Stock Dove Columba oenas, Jackdaw Corvus monedula and
Carrion Crow Corvus corone (Walter and Hudde 1987). Walter and Hudde showed that
this ectoparasite did not reduce the breeding success. Two endoparasites have been
found in the intestine and in the liver of the Little Owl (Juillard 1984). Besides the
parasites, two commensal species of Lepidoptera, Trichophagatapetziella and Monopis
laevigella have been found in the pellets of a Little Owl in France (Courtois 1988).
Van Harxen et al. (2017) studied the occurrence of parasites in nestboxes during the
breeding season by classifying the degree of spots on the eggs caused by the excrement of
parasites. They compared nestboxes that are annually cleaned and others where the debris
remains in the nestbox. A classification scheme was published allowing scoring of the
amount of spots per egg in order to study the impact of cleaning nestboxes on parasitic
loads. Three research areas were compared, i.e., one with no cleaning of nestboxes, one with
partial cleaning before and during the breeding season and one with all boxes completely
cleaned before the breeding season. When no cleaning took place the percentages of
infected nests (34.8%) and eggs (34.5%) were significantly larger than the control areas
with partial cleaning (9.1%) and complete cleaning (8.5%) of the nest. The infection rate of
the eggs was, respectively, 6.8% and 7.1%. When no cleaning of the nestboxes took place
there was a significant difference in parasitic load when the nestbox was not occupied in the
preceeding year. Parasite larvae and eggs are killed by low winter temperatures and limited
humidity, which is typical for abandoned nestboxes, in contrast to occupied ones.
Regarding diseases, a list of infections is given by Schönn et al. (1991). Among the
main nonspecific diseases able to affect Little Owls are: (1) Newcastle infection (Schoop
et al. 1955), a viral infection that appears when the birds have limited food supplies, (2)
Hepatosplenitis Infectiosa Strigum, which can cause intestinal inflammation in Little
Owls (Schönn 1986), (3) bacterial infections such as salmonellosis, tuberculosis and
pseudomonasis and (4) fungal diseases such as aspergillosis. Finally, Streptococcus and
Staphylococcus infections affect Little Owls in captivity (Schönn et al. 1991). Bultot et al.
(2001) noted that Belgian bird-care centers have recorded 23% of incoming Little Owls
as being victims of disease.
A Little Owl was found in January 2021 with a tick, Ixodida, on its head in northern
Flanders (Figure 10.7) (De Taey and Aerts, Uilenwerkgroep Waasland personal
communication).
In addition to parasites and diseases that are much more prevalent, a few anomalies
or anatomical malformations have been reported. Juillard (1984) found a nestling with

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 Chapter 10: Population Regulation

Figure 10.7 Tick as exoparasite of a Little Owl.

a crossed bill, which was probably lethal because the young could not feed itself,
another with a necrotized leg due to a thorn under the ring of the bird. This one died
despite removal of the thorn. A third anomaly is very rare and is called “iris colobome”
(Reisinger 1926a, Juillard 1981). It is a tear of the iris provoked either during embryonic
development or due to a crash in a fully-fledged bird. During field studies, two other
cases of injuries to the iris were recorded, one in France (Génot unpublished data) and
another in Belgium (J. Bultot personal communication).

10.2.6 Weather
10.2.6.1 RAIN
Weather has been shown to influence the breeding performance of Little Owls
(Knötzsch 1988). For example, in the period leading up to egg-laying, rainy weather
conditions can hinder and sometimes prevent hunting, affecting the breeding condi-
tion of adult birds, and delaying the start of breeding. Bultot et al. (2001), highlighted
the constraints that weather puts on later breeding performance by showing that a
higher rate of owlet mortality was related to the amount of rainfall during May
and June.
At Lake Constance, Germany, Knötzsch (1988) considered that if the number of
cavities was sufficient, the only limiting factor for a population was the climate; this
also implied that food supplies were influenced by the prevailing weather. Larger
amounts of precipitation in March seem to result in improved food availability (earth-
worms) and thus higher weights of adult owls. In contrast, higher rainfall in May and
June, especially when the nestlings were younger than two to three weeks, reduced
fledging success, as a result of the moist environment in cavities or nestboxes (Knötzsch
1978, Exo 1983, Gassmann et al. 1994, Lederer and Kämpfer-Lauenstein 1996).

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 10.2 Environmental Limiting Factors

However, the reduced fledging success could also be due to reduced hunting success
for the adults at this time. Fledging success in Belgium (Bultot et al. 2001) showed a
negative association with cumulative precipitation in May and June. For every 100 l/m2
extra precipitation in this period, 0.6 fewer young were fledged per active nest.
Temporal variation arising from factors such as weather has been found to have a
direct impact in species–habitat relationships (Sunde et al. 2014). The radio-tagging of
Little Owls allowed the study of habitat use and selection (use/availability) of foraging
and how this varied as a function of weather, season and availability. Use of the two most
frequent land-cover types, gardens/buildings and cultivated fields, varied more than
threefold as a simple function of season and weather through linear effects of wind and
quadratic effects of temperature. Even when controlling for the temporal context, both
land-cover types were used more evenly than predicted from variation in availability
(functional response in habitat selection). Use of two other land-cover categories (pas-
tures and moist areas) increased linearly with temperature and was proportional to their
availability. The study showed that habitat selection by generalist foragers may be highly
dependent on temporal variables such as weather, probably because such foragers switch
between weather-dependent feeding opportunities offered by different land-cover types.
An opportunistic foraging strategy in a landscape with erratically appearing feeding
opportunities in different land-cover types may possibly also explain decreasing selection
of the two most frequently used land-cover types with increasing availability.

10.2.6.2 SNOW
Cold and snowy winters cause substantial losses in Little Owls, as was seen in
Switzerland in 1951 (Büchi 1952), in Denmark in 1956 (Poulsen 1957), in England in
1962–1963 (Dobinson and Richards 1964) and more recently in France in 1985–1987
(Génot 1990b) and Germany (Kämpfer-Lauenstein and Lederer 1995). The annual
mortality patterns of adult and first-year Little Owls in Germany indicate a peak in
mortality in January–February (Exo and Hennes 1980). Little Owls have more fat
reserves (15.1% of body weight) than Barn Owls (Piechocki 1960) and hence more
capacity to cope with periods of cold weather (Helbig 1981, Schönn 1986). During
severe or prolonged cold weather or snow events, the owls cannot feed themselves and
die of starvation or exposure (Poulsen 1957, Géroudet 1964). In France, Génot (personal
observation) found a dead Little Owl in January 1997 after three weeks of snow; the owl
weighed 110 g compared to a normal weight of 160–250 g.
After cold winters, as in 1939–1942 in Germany and 1894–1895 in Switzerland,
Peitzmeier (1952) and Luginbühl (1908), respectively, saw that Little Owl populations
declined and took about 10 years to recover. In the Kaliningrad Region, the number of
Little Owls decreased considerably after a severe winter (Grishanov 1994). In France,
after two severe winters in 1985 and 1986 having 25–26 days with 10–15 cm of snow,
the population decreased by almost 30% (Génot 1992c). In Germany, after the winters
of 1978–1979 and 1985–1986, 30% and 38% of the occupied sites were abandoned,
respectively (Schönn 1986).
The Tibetan Little Owls, Athene noctua ludlowi use the burrows of Himalayan
Marmots Marmota himalayana. An unusual winter, with exceptional amounts of snow,

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 Chapter 10: Population Regulation

flooded the burrows one spring, killing the majority of the marmots. The owls survived
this event by moving to lower elevations. Upon their return to the area, the Little Owls
did not breed in the marmot burrows, but rather looked for shelter in them, and
appeared to use the burrows that were occupied by marmots. One could speculate that
perhaps this was because the occupied burrows were warmer, or because the burrows
were not so full of small rocks and sediment caused by the flooding (van Orden and
Paklina 2003).
Hard winters that inhibit foraging activity are a major cause of both first-year and
adult mortality. This is reflected across the species range by the higher winter mortality
in countries that tend to have lower temperatures and longer periods of snow cover.
Snow cover presents problems for ground-feeding predatory birds, which cannot gain
access to their prey underneath (Schröpfer 1996).

10.2.7 Inter-specific Relationships

Aggressive interactions exist between the Little Owl and some other species. As shown
by Zuberogoitia et al. (2005) among European owls, most attacks were aimed at Little
Owls and the main aggressors were the Barn Owl and Tawny Owl. While the Barn Owl
may have been noted as being a potential predator of Little Owls (Lord and Ainsworth
1945), in captivity the species can get along peacefully (Festetics 1952), and in nature
the two species can live in the same building (Reynaud personal communication) and
even breed, as in Slovenia (Denac et al. 2002). Magpies Pica pica mob the Little Owl
(Ritzel and Wulf 1978), and the Eurasian Kestrel Falco tinninculus competes for nest sites
(Bonin and Strenna 1986) or prey (Moreau 1947). As noted in Chapter 2 on History and
Traditions, the mobbing behavior of small birds on Little Owls (and the human desire
to catch these small birds) was one of the main reasons that young owls were taken
from nests and raised in captivity for hunting for nearly 2000 years. In Italy, the
mobbing by small birds was concentrated within the breeding season (e.g., Swallow
Hirundo rustica and House Martin Delichon urbica), although some species displayed
mobbing behavior in autumn and winter (e.g., Tree Sparrow Passer montanus, Italian
Sparrow Passer italiae, Geat Tit Parus major, Blue Tit Parus caeruleus, Greenfinch Carduelis
chloris, Goldfinch Carduelis carduelis, Serin Serinus serinus, Stonechat Saxicola torquata
and Meadow Pipit Anthus pratensis; Casalini 2005). Due to either alternative and readily
available food sources, or fear of injury, some tolerance is observed between the Little
Owl and some animals which could be predators or competitors. Thus, Little Owls can
breed close (less than 50 m) to Tawny owls (Rusch 1988) or in the same nestbox as a
squirrel Sciurus vulgaris (Palmer 1989) or a Jackdaw Corvus monedula (Berghmans
2001d). Little Owls were able to breed in a big nestbox already occupied by Starlings
Sturnus vulgaris (Henrioux 1980). These events do not need to have biological signifi-
cance. Nest-site sharing may be a symptom of a shortage of suitable sites. A last example
of living together was in Italy, where a Little Owl shared daytime use of a winter roost of
the Long-eared Owl Asio otus (Mastrorilli 1999).
In food webs, top predators often feed at more than one trophic level. Top predators
may feed on primary prey but also on mesopredators that are the usual predators of

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 10.3 Demographic Limiting Factors

primary prey. Parejo et al. (2018) studied the cavity-nesting bird community of Little
Owls, Scops Owls and European Rollers in southern Spain as mesopredators in relation
to the Eagle Owl Bubo bubo and Long-Eared Owl Asio otus as top predators and several
smaller prey species that nested in similar cavities to the primary prey, e.g., Great Tit
Parus major and Spotless Starling Sturnus unicolor. Killing of mesopredators is sometimes
important enough to impact their demography in favor of competing species.
Mesopredators and primary prey can compete for nest holes. In a crossed design
Parejo et al. (2018) manipulated social information on a threat at the patch scale by
broadcasting calls of Little Owls indicating different levels of danger. They additionally
modified the availability of holes at the patch scale to manipulate competition for nest
sites. The effects on the structure of this cavity-nesting bird community, in which all
the species competed for nest sites but where some species shared predators with the
Little Owl and others were prey of the owl, were analyzed. Competitors sharing top
predators with the callers and prey of the callers interpreted alarm and nonalarm calls
differently. Competitors preferentially chose alarm and nonalarm patches over control
patches to breed, while prey selected alarm patches, taking into account the stress level
of their predators. In contrast, competition for nest sites affected the habitat selection of
prey species more than that of competitors of the callers. Parejo et al. (2018) provided
support for a differential value of alarm calls and competition for nest sites for different
receivers, related to niche overlapping among callers and eavesdroppers, therefore
calling attention to possible cascading effects by the use of information in this Little
Owl community.
Social information use between nest-cavity competitors can benefit fitness. Melanins
are pigments responsible for most intra-specific color variation in vertebrates, which is
often associated with changes in behavior. Parejo and Avilés (2020) explored whether
melanism is related to individual propensity to use social information in the color
polymorphic of Scops Owl Otus scops. They manipulated social information on
predation risk at nests by broadcasting calls of Little Owls and found that owlets of
brownish females exposed to alarm calls had lower levels of natural antibodies than
those of grayish females. In parallel, they found changes in parental behavior contin-
gent on coloration, because when exposed to the high-risk treatment of alarm calls of
Little Owls, brownish females returned to nests earlier than grayish females, and owlets
raised by brownish females were fed with smaller prey than those raised by grayish
ones. These results provide support for the role of melanins on the propensity to use
social information, which may help to explain the maintenance of melanin-based color
polymophisms wherever social environments are available.

10.3 Demographic Limiting Factors

The following discussion reflects a review of studies that contain the type of demo-
graphic data and analysis techniques that are needed to help answer fundamental
questions on population dynamics in Little Owls. Detailed long-term studies from
different geographical and climatic regions are required to resolve whether the observed

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 Chapter 10: Population Regulation

population decrease in Little Owls across Europe is due to poor reproduction or

enhanced mortality (Exo 1992).
Demographic factors reflect the number of births and deaths within a given local
population, the immigration of owls from other areas and the emigration of owls from
the local population. We first look at the factors that regulate births and deaths within
the local population and then zoom in on dispersal and movements of the birds. Some
density-dependent influences might be occurring (e.g., smaller clutch sizes from owls
within higher densities, later breeding from owls in lower densities). Therefore, we
describe the densities that are observed throughout the distribution area and then focus
in on the effects that this might have on local populations. The connectivity of local
small population clusters has been found to be increasingly important in the conser-
vation biology of wildlife species. In relation to this we further discuss meta-
populations of Little Owls and the impact this type of structure has on the species.

10.3.1 Births and Deaths

A local population that is considered isolated from the outside can be seen as the sum of
births and deaths. To keep things simple, first we will consider here that the local
population is “closed,” i.e., there is no immigration into or emigration from the
local population.
In stable populations, reproduction balances mortality. Mortality studies on birds
involve some kind of marking (usually ringing) and subsequent re-capture or re-
sighting. As owls are mostly nocturnal and reclusive, recovery rates of ringed owls are
invariably low, as few people re-sight, re-capture or recover dead ringed owls. From the
Finnish Bird Ringing Centre data, Saurola (2002, p. 43) reported that of 198 887 individ-
uals of 10 species of owls ringed between 1913 and 1999, some 25 492 (12.8%) were
recovered. This level of ringing and recovery represents a truly significant amount of
volunteer and professional effort. For Little Owls, the recovery rate was 2.1% for
755 ringed owls (Furrington 1998), 2.8% for 541 ringed owls (Juillard 1984), 3.9% for
616 ringed owls (Knötzsch 1985) and 2.5% for 435 ringed owls (Génot 2005).
Based on their work in western Germany and the Netherlands, Exo and Hennes
(1980) found mortality rates in adult owls to be 35% and for young owls 70%. From this
data, they estimated that each pair of Little Owls must produce 2.34 fledgings each year
for reproduction to balance mortality. Figures derived from ring recoveries can over- or
under-estimate the reproduction rate; for example, if the annual mortality rate for adult
and juvenile owls was only 5 percentage points lower (30% for adults and 65% for
juveniles, respectively) then each pair must produce only 1.7 fledged young per year to
compensate for mortality (Exo 1992). In Germany, the mortality for owls in their
second year of life was 30% (Mohr 1990). Mortality of male owls in their first year of
life was 27% and for females 30%; for owls between their second and sixth year of life,
the mortality rate was 26% for males and 27% for females (Knötzsch 1988). Exo (1992)
reported that extraordinarily high mortality occurs in severe, snowy winters. In France
the annual mortality rate was 85% for owls in their first year of life and 36% for adults
(Letty et al. 2001a). In Switzerland, the first-year mortality was 74%, and second year

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 10.3 Demographic Limiting Factors

64% (Glutz Von Blotzheim and Bauer 1980). In Bönnigheim (Landkreis Ludwigsburg,
Germany) 11 juveniles were followed by telemetry after dispersal and suffered a mor-
tality rate of 63% (Eick 2003).
Glue (1973) reported two main periods of seasonal mortality in Little Owls in
England: July–August for the young and May–June for adults. In France, Génot (1991)
found seasonal mortality peaks in July–September for the young and April–June for the
adults. In Germany, the peak mortality was in June–July for adults, with a peak in adult
mortality occurring just after fledging, probably because of the energetic bottleneck of
rearing young and molting (Exo 1992). In Britain, Glue (2002) analyzed ringing recov-
ery data and showed that most juvenile birds died from starvation in June–July, which
correlates to the time when the young owls gain independence from their parents.
The following data reflect longevity records for ringed Little “Methuselah” Owls. The
oldest Dutch bird was seen until its fifteenth year (Stroeken and van Harxen 2003d).
The oldest Belgian owl was found to have lived for 12 years and seven months in
Flanders (Vercauteren 1989) and in Wallonia an owl lived into its thirteenth year
(J. Bultot personal communication). The oldest ringed owl from Germany was reported
to be 15 years and seven months of age (Rydzewski 1978). Peter (1999) reported a
German owl that lived to be 11 years old and Sill and Ullrich (2005) a female of 12 years
old. Another old German owl was nine years six months (Furrington 1988), as was the
oldest French owl (Génot 2005).
In the Netherlands, Stam (2003) studied an owl population located east of Arnhem
and was able to record the specifics on 152 ringed female Little Owls during 1983–2002.
He found 54 females that bred once, 32 twice, 23 three times, 14 four times, 10 five
times, 7 six times, 7 seven times, 4 eight times and 3 nine times. The life expectancy of
these breeding Little Owl females averaged 2.8 years (all females bred at least once),
whilst in Germany, Zens (2005) recorded a life expectancy of 3.7 years.
Stroeken and van Harxen (2003b) followed a population with the following age
structure in 2003: ten females were in their second year, seven in their third year, five
in their fourth year, two in their fifth year, three in their sixth year, four in their seventh
year and one in its eleventh year. The life expectancy of these females averaged four
years. Stroeken and van Harxen (2003b) also reported the age structure for male owls as:
three males were in their second year, one in its third year, five in their fourth year, two
in their fifth year and one in its seventh year, indicating a life expectancy averaging 3.8
years. The ages are reported here are minimums, since most birds were ringed when
they were found for the first time in a nestbox.
The age diversity of ringed and recovered Little Owls, as reported by the EURING
Data Bank (du Feu et al. 2016) on 28 626 recovered birds, is given in Table 10.5. The
observed lifespan for birds ringed as adults but with age remaining unknown should be
incremented by at least one year before ringing.
The birds ringed as pulli entail the most complete view. Pulli that were still observed
alive on their last observation and that survived at least their first year were on average
1388 days old for females (n = 1303) and 1316 days old for males (n = 460). Pulli that
were observed dead on their last observation were on average 265 days old for females
(n = 17) and 170 days old for males (n = 26) if they died before one year old. When they

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Table 10.5 Breakdown of ages in days of dead and recovered Little Owl based upon the EURING Data Bank ring recovery database (du Feu et al. 2016).
Data is grouped by sex (determined a posteriori in case of pulli), ringed as nestling or as adult, either alive or dead at the last observation within or beyond
one year after ringing

Found Lifespan in days

Dead Time
Percentile
Ringed or after
Sex as Alive ringing N Mean Min Max SD Range Q25 Median Q75 99.99 99.95

Female Adult Alive First year 634 228.85 1 364 139.83 363 62 308 354 364 364
Male Adult Alive First year 195 208.97 1 364 136.97 363 70 256 343 364 364
Female Pullus Alive First year 321 338.18 51 364 35.09 313 333 346 356 364 364
Male Pullus Alive First year 141 294.87 2 364 73.14 362 270 318 347 364 364
Female Adult Alive First year 2057 1255 365 5286 809.83 4921 711 1087 1772 5286 5112
Male Adult Alive > 1 year 388 1092.55 365 4457 766.30 4092 538 760 1470.5 4457 4457
Female Pullus Alive > 1 year 1303 1388.27 365 4973 859.14 4608 723 1099 1821 4973 4973
Male Pullus Alive > 1 year 460 1316.13 365 3996 807.45 3631 697 1078 1782.5 3996 3996
Unknown Adult Alive > 1 year 2815.00 1118.75 365 4464 687.00 4099 655 940 1464 4464 4376
Unknown Adult Alive First year 1404.00 184.41 1 364 137.52 363 48 164 340 364 364
Unknown Adult Dead > 1 year 721.00 1245.29 365 4483 816.71 4118 627 989 1725 4483 4483
Unknown Adult Dead First year 722.00 109.68 1 364 107.05 363 19 74 180 364 364
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Female Adult Dead First year 150 169.41 5 364 119.76 359 48 150 287 364 364
Male Adult Dead First year 55 149.76 2 357 110.44 355 65 119 202 357 357
Female Pullus Dead First year 17 264.76 3 363 122.77 324 149 342 353 363 363
Male Pullus Dead First year 26 170.23 1 363 133.35 353 59 152 281 363 363
Female Adult Dead > 1 year 373 1356.76 366 4417 876.81 4051 707 1123 1824 4417 4417
Male Adult Dead > 1 year 92 1327.34 375 3470 700.14 3095 820 1176 1670 3470 3470
Female Pullus Dead > 1 year 88 1831.39 365 5112 1010.14 4747 1094.5 1514.5 2538.5 5112 5112
Male Pullus Dead > 1 year 33 1500.58 366 3951 929.66 3585 731 1437 1995 3951 3951
Unknown Pullus Alive > 1 year 5475 1260.22 365 5130 800.53 4765 690 1061 1700 5130 5086
Unknown Pullus Alive First year 3254 221.67 1 364 121.50 363 111 261 334 364 364
Unknown Pullus Dead First year 5720 104.65 1 364 99.85 363 31 62 145 364 364
Unknown Pullus Dead > 1 year 2182 1183.60 365 5056 870.49 4691 534 849 1495 5056 4914

28 626
Data are grouped by sex, either alive or dead at the last observation. and within or over one year after ringing.
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survived the first year, the average lifespan was 1831 days for females (n = 88) and
1501 days for males (n = 33). The split between the different observation dates and the
circumstances are crucial for meaningful comparison.

10.3.2 Juvenile Dispersal and Adult Movements

Earlier in this chapter, we described a “closed” population as one in which there is no
immigration or emigration. In wild populations of Little Owls, this simplified situation
is less frequently the case, and the dispersal of juveniles and movements of adults play a
significant role in the demographic (and genetic) connectivity between/among popu-
lations. Dispersal is the term that applies to a young owl as it leaves its natal territory to
mature, and to search out and settle into a territory of its own. A “successful” dispersing
owl is one who moves from his/her natal territory, disperses and lives long enough to
become part of a mated pair. Movements reflect the shifts that adults make, for example,
a female may breed in one location one year, and may move to a different territory for
the next breeding season.
The distance moved by dispersing male and female owls has provided key infor-
mation on aspects of population dynamics and metapopulation analyses (for examples
of juvenile dispersal studies, see Forsman et al. 1996, Ganey and Block 2005). Further,
the juvenile survival rate (simply the inverse of the mortality rate) has been shown to be
a key driver in the dynamics of other owl populations such as the California Spotted
Owl (Strix occidentalis occidentalis) (Franklin et al. 2004), Northern Spotted Owl
(S. o. caurina) (Forsman et al. 1996) and Mexican Spotted Owl (S. o. lucida) (Ganey
et al. 2004).
In some owl species (e.g., Northern Spotted Owls (Forsman et al. 1996), Tengmalm’s
Owls (Korpimäki 1988), Lanyu Scops Owls Otus elegans (Severinghaus 2002) and as will
be shown for Little Owls below) females disperse nearly twice as far as males. The
ecological basis for this appears to be the different breeding strategies of the sexes. In
the context of perpetuating the species, in order to be successful, males need to have a
territory and so will select one of the first vacant territories they come across, with even
a low-quality territory being better than no territory at all. In order for the female to be
successful, she must produce young, so she will “shop around” for a territory that
contains a mate, food and nest resources that will allow her to be successful. The
mating system hypothesis, which predicts that the sex that establishes the territory
should disperse shorter distances, remains tenable as an explanation for female-biased
dispersal in Little Owls.
Unlike the more northerly Asio and Surnia owls that embark on long-distance
migrations or movements, Little Owls are considered relatively sedentary, and will
generally settle in areas close to their natal origin. This was noticed when the Little
Owl colonized Britain in the late 1800s to early 1900s. It did so in small incremental
movements establishing new clusters of nests, and then moving out from the core
populations (Marchant et al. 1990).
Life is a balance, and within the context of temperate, Mediterranean and related
interior climates, the combination of their knowledge of local foraging habitat

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 10.3 Demographic Limiting Factors

combined with their varying prey capture techniques allows Little Owls to survive
within a limited range, and so do not have to migrate to search out areas with higher
prey densities. The home range of territorial Little Owls is pretty much fixed in size
throughout the year; however, in some cases the birds will range a little further in the
winter months due to the hardships of finding adequate prey. In some portions of their
range, Little Owls are less sedentary than elsewhere. For example, in Kazakhstan the
Little Owl is considered to be nomadic in winter, leaving the central part of the country
and migrating southwards (Gavrin 1962).

10.3.2.1 JUVENILE DISPERSAL CHARACTERISTICS

We examined data from 13 dispersal studies (Table 10.6) and offer a brief summary of
each study in this section. Eleven of these studies involved recoveries of ringed birds
and one study used radio-telemetry. Dispersal begins when the owls are 12–16 weeks of
age (Eick 2003) and lasts through at least the end of October. Most birds do not settle at
once. The earliest settlement was observed at the end of September. Young birds
disperse in random directions, and most settle within 20 km of their birthplace.
While a small proportion of ringing recoveries were from birds dispersing over 50 km,
some studies indicated that 4–9% of the juvenile owls will disperse over 100 km, and
distances of 182, 190, 220, 230, 270, 297 and 600 km have been recorded.
In typical ringing studies, researchers capture and ring the juvenile and (ideally)
all of the adult owls within a given area. Ringing continues in subsequent years, with
the researchers ringing the new offspring and re-capturing the owls that remain as
members of breeding pairs, or the juveniles that have since been recruited into the
study site’s breeding population. Unless the study area is large, few of the owls that
move off the study are recovered (as there are simply fewer people looking for owls
in those locations), thus establishing a recognized bias in the recovery rates of ringed
birds. With ringing programs, it is particularly challenging to determine what pro-
portion of the owls emigrate (move out) of the population, become nonterritorial
“floaters,” or simply die. Radio-telemetry studies like that of Eick (2003) are badly
needed for better determining the dispersal distances and survival rates of juvenile
Little Owls.
Given very large sample sizes (e.g., Saurola 2002), ringing studies do offer some
information about owl dispersal and movements (i.e., straight-line distance moved
and longevity) and are particularly valuable for investigating within-population
dynamics (e.g., reproduction and survival rates for known adults). In the paragraphs
that follow, we provide short summaries of the juvenile dispersal and adult movement
studies that have been conducted thus far on Little Owls.
Glue (2002) examined the British Ringing Data on 57 owls that had been ringed as
juveniles and later recovered. He found that the median straight-line dispersal distance
of these owls was 7 km (range 0–92 km, n = 57). In 1962, one bird ringed as a nestling in
Northumberland, northern England, was found dead later that year in Musselburgh,
Lothian, Scotland, a distance of approximately 35 km.
Birds from the introduced British population dispersed further on average than the
sympatric Tawny Owl. The sample sizes and distance dispersed (by distance category)

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Table 10.6 Juvenile dispersal studies – straight-line distances between natal site and recovery during the following breeding season

Total
Study young Males Females Results

Glue 2002 57 – – Glue found that the median straight-line dispersal distance of these
owls was 7 km. However, he found that there was a regional
difference between the owls originating in northern and southern
England. Median recovery distances for birds ringed (pulli and
adults) in the northwest (15 km, n = 4) and northeast (7.5 km,
P5–P95 0–92 km, n = 6) exceeded those of the southwest (2 km,
n = 111) and southeast (1 km, n = 171) regions
Exo and Hennes 1980 48 – – From November–March (inclusive) of their first winter, the median
dispersal distance for 30 owls in Germany was 15 km, and for
18 owls from the Netherlands, the median recovery distance
category was 10–19 km. Overall, about 55% of these juvenile
owls settled within 10 km of their birthplace and only 9%
dispersed distances over 100 km
Ullrich 1980 21 16 4 For 21 owls ringed in their nests as nestlings, 18 were recovered
0–16 km (average 6.6 km) from their birthplace, and one each at
22 km, 55 km and 190 km. The 20 owls that dispersed less than
55 km involved 16 males and 4 females, suggesting females were
more likely to move away although sex ratio at ringing was
unknown)
Kämpfer and Lederer 1988 130 – – In their Central Westphalia study area, juveniles settled on average
4.1 km away from their birthplace
Gassmann and Baumer 1993 29 9 13 In a 15 year-population study in Northrhine-Westphalia, the distance
of settlement of first-year owls was 5.7 km (n = 29), females
(6.4 km, n = 13) moved further than males (3.9 km n = 9)
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Veit 1988 11 – – In Hesse, young moved an average of 19 km, ranging from 2 km to

48 km
Juillard 1984 20 – – In Switzerland, young moved 0.5–16 km (average 5.8 km, n = 20)
Bultot et al. 2001 26 – – In Wallonia (Belgium), the distance of settlement of first-year owls in
two research areas with plenty of vacant nestboxes averaged
2.4 km in Ransart (min = 1.2 km, max = 4.4 km, SD = 1.2 km,
n = 9) and 1.2 km in Neufville (min = 0 m, max = 3.2 km,
SD = 0.9 km, n = 17)
Génot 2005 33 12 11 In Vosges du Nord (east France) juveniles disperse on average
6.5 km (n = 33) their birthplace, max 25 km. Males 5.6 km (n = 12)
females 9 km (n = 11)
Stastny et al 1996 – – – 50% of birds found no further than 10 km from the place of ringing,
only 4% found further than 100 km
Knötzsch 1988 127 68 59 In the Lake Constance study area between 1976 and 1987, the
dispersal of first-year owls averaged 4.5 km for males (n = 68) and
4 km for females (n = 59)
Eick 2003 (short-term radio- 11 – – In Germany (Bönnigheim, Landkreis Ludwigsburg) 11 juveniles
telemetry study) were followed by telemetry during their dispersal over 31 nights.
The birds moved on average 9.8 km and maximally 41 km per
night; nearly all juveniles came back to their birthplace after
exploratory movements in all directions after a few days
Desmarets and Braeckman 373 150 223 Analysis of Belgian ring recovery and re-capture data. Females
(in prep.) dispersed statistically significantly further than males (8.4 km,
n = 168 versus 2.6 km, n = 90). A higher percentage of female
pulli (75%) left their natal territory than males (60%)
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for 174 owls ringed as nestlings or May–July juveniles and recovered from August
onwards were: 111 dispersed 0–10 km, 30 dispersed 11–20 km, 21 dispersed 21–50
km, 7 dispersed 51–100 km and 5 dispersed over 100 km (maximum distance was 150
km) (British Trust for Ornithology data).
A juvenile ringed in September was recovered two years later 175 km north-
northeast of its ringing site. A full-grown bird ringed in December was recovered
16 months later (April) some 110 km northeast of its ringing site. Most of those ringed
as adults were recovered within 10 km (usually locally), but several were found up to
45 km away. One adult female was ringed in May in Dorset, and was recovered
13 months later in Hereford, having moved an impressive 182 km to the north-
northwest. Stragglers (none of them ringed) have crossed the Irish Sea to the Isle of
Man and eastern Ireland (British Trust for Ornithology data). Continental birds also
disperse relatively short distances. In their review of German and Dutch ringing data,
Exo and Hennes (1980) found that juvenile dispersal began in August. From November–
March (inclusive) of their first winter, the median dispersal distance for 30 owls in
Germany was 15 km, and 18 owls from the Netherlands were recovered in the distance
category of 10–19 km. Overall, about 55% of these juvenile owls settled within 10 km of
their birthplace and only 9% dispersed distances over 100 km.
Owls ringed after attaining breeding age (i.e., April onwards) were recovered a
median distance of 7.5 km (n = 53 owls) and a distance category of 0–9 km (n = 25
owls) in Germany and the Netherlands, respectively. Of these adult owls, 74% were
recovered within 10 km of their original ringing site (Exo and Hennes 1980).
In a population study conducted in North-Württemberg, Germany, Ullrich (1980)
found first-year birds to disperse 0.5–220 km. Most of the owls dispersed under 40 km,
and there were three cases in which juveniles that dispersed in autumn later returned to
near their natal area, one coming from 36 km away.
As found in other owl species, recoveries of siblings showed different within-brood
dispersal characteristics (i.e., siblings dispersed different distances and in different
directions). Ullrich (1980) reported on dispersal aspects for pairs of siblings from three
nests: (1) 3 km west, 36 km northeast, (2) 3 km west-southwest, 38 km north and (3)
3 km northeast, 190 km southwest. Different young from the same clutch dispersed in
random directions.
For 21 owls ringed at their nests as nestlings, 18 were recovered 0–16 km (average 6.6
km) from their birthplace, and one each at 22 km, 55 km and 190 km. Here, the 20 owls
that dispersed less than 55 km included 16 males and 4 females (Ullrich 1980).
In a population study in Central Westphalia, juveniles settled an average of 4.1 km
away from their birthplace (n = 130). Some 75% of the first-year owls trapped after
November 1 were re-captured later within the breeding area (Kämpfer and Lederer
1988).
A population study around Lake Constance between 1976 and 1987 found that the
dispersal of first-year owls averaged 4.5 km for males (n = 68) and 4 km for females (n =
59) (Knötzsch 1988).
In a 15-year population study in Northrhine-Westphalia, the dispersal distance and
settlement of first-year owls was 5.7 km (n = 29), with females (6.4 km, n = 13) moving

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 10.3 Demographic Limiting Factors

further than males (3.9 km, n = 9). For pairs of young (siblings) from six nests, one bird
was found in a different direction and at a significantly greater distance than the other
(Gassmann and Bäumer 1993). In Hesse, young moved an average of 19 km (n = 11)
with a range of 2–48 km (Veit 1988).
The dispersal distance and settlement of first-year owls in an 11-year population
study in Northrhine-Westphalia was 5.17 km (n = 88), with females (6.8 km, n = 34)
moving further than males (3.8 km, n = 51). Of 153 owls born, only 19 bred in the study
area (Zens 2005).
In Switzerland, young owls dispersed an average of 5.8 km (range = 0.5–16 km,
n = 20) (Juillard 1984).
In a population study in Wallonia (Belgium) (Bultot et al. 2001), the dispersal
distance and settlement of first-year owls in two research areas with plenty of vacant
nestboxes averaged 2.4 km in Ransart (min = 1.2 km, max = 4.4 km, n = 9) and 1.2 km in
Neufville (min = 0 km, max = 3.2 km, n = 17). Here, owls appeared to show a preference
for dispersing in a southerly and easterly direction. No relationship was found between
the population density and the emigration distance, possibly because of the lack of re-
captures outside the research area.
In Vosges du Nord (eastern France) juveniles dispersed on average 6.5 km (max =
25 km, n = 33) from their birthplace, females (9 km, n = 11) moving further than the
males (5.6 km, n = 12) (Génot 2005).
In the Czech Republic, 50% of the ringed birds were found within 10 km of the place
of ringing, and only 4% were found further than 100 km away (Stastny et al. 1996).
In Belgium (Desmarets and Braeckman in prep.), male pulli (n = 90/150: 60%)
dispersed an average of 2.6 km (max distance 18 km), while female pulli (n = 168/
223: 75%) dispersed 8.4 km (max distance 150 km). Of all birds that were ringed as
pulli, 30.8% did not disperse, 40% of all males, 24.6% of all females. Males dispersed
significantly shorter distances than females when ringed at the nest. The same holds for
birds ringed as adults, but with shorter average distances (5/170 males (2.9%) showed
an average dispersion of 0.27 km, 32/456 (7%) females dispersed on average 1.13 km).
Occasionally some owls do embark on long-distance movements. The longest recov-
eries and direction of travel of ringed birds are as follows: Mittel-Franken, Germany
(June nestling) to Bas-Rhin, France (January), 230 km west-southwest; Hessen,
Germany (October) to Halle, Germany (April), 270 km northeast; Dresden, Germany
(July, full-grown) to Austria (February), 297 km south; two North-Württemberg nest-
lings (June) were found in October in Switzerland (220 km southwest) and Zielona
Gura, Poland (600 km northeast) (Glutz Von Blotzheim and Bauer 1980, Furrington
1998). Such erratic movements as these are consistent with aspects of vagrancy to
Helgoland and southern Fenno-Scandia.
Eick (2003) conducted a radio-telemetry study on juvenile Little Owls during and
after dispersal in Germany (Bönnigheim, Landkreis Ludwigsburg). He tracked 11 juven-
iles with telemetry over 31 nights (Eick 2003). On average, the birds moved 9.8 km and
maximally 41 km per night (Table 10.7). The movements showed no systematic direc-
tion. Such random dispersal behavior is typical for this rather sedentary owl species
(Bauer 1987, Berthold 2000). The juveniles tended to re-visit their natal site at regular

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Table 10.7 Exploratory excursions of juvenile Little Owls in Germany

Excursion Total distance Average distance per Max distance per

duration (km) night (km) night (km)

3 nights 15 7.5 7.5

3 nights 20 7 8.5
5 nights 35 7 12.5
5 nights 76 15.2 41
>2 nights >25 - 25
2 nights 4 2 2.5
5 nights >80 16 25
3 nights >25 8.3 12
2 nights 25 12.5 19
31 nights >30.5 9.8 41

After Eick 2003.

intervals between exploratory excursions of three to five days (Eick 2003). This behavior
of exploratory excursions followed by a return to the nest was also observed in radio-
marked Northern Spotted Owls prior to their formal dispersal from the natal territory
(Miller 1989).
Zens (2005) followed 11 juvenile Little Owls with radio-transmitters in Germany
(Mechernicher Voreifel). In July and August, juveniles flew around their nest site up
to 300 m. In September, they flew further during the night, up to 1.25 km from their
birthplace. During one night they moved from 560–690 m to 980–3100 m. By this time
54.5% of the juveniles were already dead or missing. At the end of September or
beginning of October, the juveniles left their birthplace.
The use of radio-telemetry in juvenile survival studies has only been conducted on
small numbers of Little Owls. Due to the high mortality rate of juveniles, one has to
equip sufficient numbers of birds with radio-transmitters to be able to track a sufficient
sample of them from dispersal through to ultimate pairing and nesting. The critical
advantage provided by radio-telemetry studies over regular ringing efforts is in that the
“outcomes” of individual owls become known.
The historical EURING data on Little Owl direction of dispersal (n = 19 239)
(Table 10.8) supports the idea of indifference, with a small negative tendency to the
north (21%) in contrast to east (26%), south (26%) and west (27%). This obviously will
depend on high-density populations and their respective locations in the country.
Ringing areas near borders might be biased due to lower cross-border recovery com-
pared to national programs. A breakdown of the data as a function of national ringing
schemes shows associations between directions and schemes. Western European owls
from the Netherlands (NLA), UK (GBT), Belgium (BLB) and Germany (DEW) are pos-
itioned in the middle of the direction labels, meaning they have similar counts for all
directions. Other ringing schemes, however, have more associations with some specific

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Table 10.8 Juvenile dispersal distances in km of dead and recovered Little Owls based upon the EURING Data Bank ring recovery database

Distance in km
Found
Percentile
Ringed Dead or Time after
Sex as Alive ringing n Mean Min Max SD Range Q25 Median Q75 99.99 99.95

Juvenile dispersal distances ≥ 0 km

Female Pullus Alive >1 year 1303 14.69 0 317 24.84 317 4 7 15 317 317
Female Pullus Alive First year 321 16.80 0 341 32.01 341 3 7 17 341 341
Female Pullus Dead >1 year 88 9.07 0 86 13.48 86 3 4.5 11 86 86
Female Pullus Dead First year 17 5.53 0 18 4.69 18 3 4 5 18 18
Male Pullus Alive >1 year 460 6.47 0 118 12.31 118 2 3 6 118 118
Male Pullus Alive First year 141 7.28 0 104 13.63 104 2 3 5 104 104
Male Pullus Dead First year 26 12.81 0 166 33.66 166 0 2.5 5 166 166
Male Pullus Dead >1 year 33 6.94 0 76 14.19 76 2 3 5 76 76
Unknown Pullus Alive >1 year 5475 11.61 0 758 29.83 758 3 5 10 758 603
Unknown Pullus Alive First year 3254 9.34 0 424 25.56 424 0 3 8 424 292
Unknown Pullus Dead First year 5720 5.25 0 1401 26.59 1401 0 0 3 1401 443
Unknown Pullus Dead >1 year 2182 11.59 0 319 23.82 319 2 4 11 319 319
Juvenile dispersal distances > 0 km
Female Pullus Alive >1 year 1206 15.73 2 317 25.52 315 4 8 16 317 317
Female Pullus Alive First year 291 18.53 2 341 33.15 339 4 8 18 341 341
Female Pullus Dead >1 year 75 10.33 2 86 14.22 84 3 6 12 86 86
Female Pullus Dead First year 15 6.27 2 18 4.50 16 4 5 7 18 18
Male Pullus Alive >1 year 392 7.49 2 118 13.06 116 3 4 7 118 118
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448

Table 10.8 (cont.)

Distance in km
Found
Percentile
Ringed Dead or Time after
Sex as Alive ringing n Mean Min Max SD Range Q25 Median Q75 99.99 99.95

Male Pullus Alive First year 117 8.72 2 104 14.54 102 3 4 6 104 104
Male Pullus Dead First year 19 17.53 2 166 38.55 164 2 3 17 166 166
Male Pullus Dead >1 year 25 8.72 2 76 15.96 74 3 3 6 76 76
Unknown Pullus Alive >1 year 4685 13.43 2 758 31.86 756 3 6 12 758 603
Unknown Pullus Alive First year 2324 13.03 2 424 29.44 422 3 5 12 424 292
Unknown Pullus Dead First year 2540 11.77 2 1401 38.93 1399 2 4 9 1401 511
Unknown Pullus Dead First year 1711 14.54 2 319 25.72 317 3 7 14 319 319
du Feu et al. 2016. Data are grouped by sex, either alive or dead at the last observation. and within or over one year after ringing.
 10.3 Demographic Limiting Factors

Row Coords.
0.00899 Col. Coords.

Figure 10.8 Associations between EURING ringing schemes and principal directions of dispersal
through correspondence analysis of counts of Little Owls based upon the EURING Data Bank ring
recovery database (du Feu et al. 2016) (n = 19 239 individual Little Owls). BLB – Belgium/Brussels;
CZP – Czech Republic/Praha; DEH – Germany/Hiddensee; DER – Germany/Radolzell; DEW – Germany/
Wilhelmshaven; DKC – Denmark/Copenhagen; DKS – Denmark/Skagen Bird Observatory and Ringing
Station; ESA – Spain/Sevillia; ESI – Spain/San Sebastian; FRP – France/Paris; GBT – United Kingdom;
British Trust for Ornithology – HES – Switzerland/Sempach; HGB – Hungary/Budapest; IAB – Italy/
Bologna; NLA – Netherlands/Arnhem.

directions (Figure 10.8), e.g., Switzerland (HES) is directed more eastward, Italy (IAB)
more to the west, Germany (DER – Radolfzell and DEH – Hiddensee) to the north, and
Spain (ESA – San Sebastian) to the south and (ESI – Madrid) to the north.
Birds ringed as pulli systematically differ in dispersal distance after more than one
year of age according to the sex, i.e., females always dispersed further than males, when
found alive or dead. Females that were still alive on their ultimate observation had an
average dispersal of 14.69 km (median = 7 km, n = 1303), while for males it was 6.47 km
(median = 3 km, n = 460). When found dead, a similar difference was found for females
9.07 km (median = 4.5 km, n = 88) and for males 6.94 km (median = 3 km, n = 33).
When observed alive within a year after birth and ringing, females dispersed on average
16.80 km (median = 7 km, n = 321) and males 7.28 km (median = 3 km, n = 141). For
dead birds the situation was reversed, with 5.53 km for females (median = 4 km, n = 17)
and 12.81 km for males (median = 2.5 km, n = 26), probably due to low sample sizes
that can lead to an unrepresentative discovery of dead individuals.

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 Chapter 10: Population Regulation

10.3.2.2 ADULT MOVEMENTS

Relatively few data exist to show the movements of birds of breeding age. We examined
data from eight studies and a few anecdotal notes regarding the movements of adults,
and offer a summary of those here. Those available confirm field observations that the
great majority stay within territories through the autumn and winter, but reveal travel
up to 45 km, and that longer movements away from their previous nest site occur
rarely. Some owls return to their previous nests the following year. Again, we caution
readers in recognizing that these studies involved ringed owls, and of the inherent
recovery bias that ringing studies are subject to.
Exo and Hennes (1980) recorded the movements of owls ringed after attaining
breeding age (i.e., April onwards). These owls were recovered a median distance of
7.5 km (n = 53 owls) and in the median distance category of 0–9 km (n = 25 owls) from
their natal sites in Germany and the Netherlands, respectively. Of these adult owls, 74%
were recovered within 10 km of their original ringing site.
In their Central Westphalia study area, adult movements away from their last nest
sites differed according to sex, with males moving 2.3 km and females 6.3 km (n = 137).
Among adults, 10.2% of males and 1.8% of females showed no site fidelity; they settled
1.7 km away from their former sites (Kämpfer and Lederer 1988).
In a Northrhine-Westphalia study area, adult males showed a higher fidelity
(1.37 km, n = 11) than females (2.2 km, n = 19) (Zens 2005).
In Switzerland, adults moved an average of 1.6 km (range 0.4–4.7 km, n = 12)
(Juillard 1984). In Vosges du Nord (eastern France) two adults were recorded as moving
12 and 32 km (Génot 2001). Ullrich (1980), Fuchs (1986) and Eick (2003) also reported
re-captures of birds that successfully bred somewhere, then moved up to 36 km to breed
elsewhere and subsequently returned to the original breeding place the year after.
Nightly movements of adults of 1–1.5 km up to 3–3.5 km were reported by Exo
(1989), especially in the fertile period. This might indicate the willingness of adults to
solicit extra-pair copulations.
A median dispersal distance of just 2 km for Little Owls ringed and recovered,
including all age classes, provides factual evidence for a strongly sedentary species,
generally site-faithful, but undertaking limited postnatal movements spanning autumn
and winter (Glue 2002).
Movements of adult owls are in random directions (Table 10.9), sometimes at higher
elevations and do not appear to follow linear landscape structures, e.g., canals, power
lines. Eick (2003) even observed Little Owls crossing larger forests. The final settlement
of these owls was relatively late and might have been related to finding a partner.
Settlement is a dynamic process and is driven by two factors, i.e., birds want to settle
in the close vicinity of existing population clusters as much as possible and adult birds
want to push conspecifics out of their active territory to avoid competition (Génot and
Van Nieuwenhuyse 2002). This dynamic process is confirmed by juvenile displacement
behavior. It should be stressed that presumably isolated populations might be less
so than previously assumed. Little Owls disperse at flying altitudes of at least
20–30 m and sometimes even higher, avoiding physical barriers such as forest and
cities (Eick 2003).

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Table 10.9 Adult dispersal distances in km of dead and recovered Little Owls based upon the EURING Data Bank ring recovery database

Distance in km
Found Time
Percentile
Ringed Dead or after
Sex as Alive ringing n Mean Min Max SD Range Q25 Median Q75 99.99 99.95

Adult dispersal distances ≥ 0 km

Female Adult Alive >1 year 2057 2.33 0 179 8.87 179 0 0 2 179 113
Female Adult Alive First year 634 1.40 0 117 5.77 117 0 0 0 117 117
Female Adult Dead >1 year 373 5.56 0 442 26.92 442 0 2 3 442 442
Female Adult Dead First year 150 5.02 0 178 19.68 178 0 0 3 178 178
Male Adult Alive >1 year 388 2.45 0 112 10.13 112 0 0 2 112 112
Male Adult Alive First year 195 0.97 0 25 2.73 25 0 0 0 25 25
Male Adult Dead First year 55 3.40 0 59 9.50 59 0 0 3 59 59
Male Adult Dead >1 year 92 5.59 0 86 13.33 86 0 2 3.5 86 86
Unknown Adult Alive >1 year 2815 2.42 0 319 13.82 319 0 0 0 319 266
Unknown Adult Alive First year 1404 2.66 0 285 14.73 285 0 0 0 285 285
Unknown Adult Dead >1 year 721 8.65 0 332 25.93 332 0 2 6 332 332
Unknown Adult Dead First year 722 8.11 0 262 24.94 262 0 2 6 262 262
Adult dispersal distances > 0 km
Female Adult Alive >1 year 498 7.15 2 113 13.28 111 2 3 6 113 113
Female Adult Alive First year 141 6.04 2 117 10.96 115 2 3 6 117 117
Female Adult Dead >1 year 139 13.21 2 442 42.96 440 2 3 8 442 442
Female Adult Dead First year 61 12.08 2 178 29.59 176 2 4 7 178 178
Male Adult Alive >1 year 110 6.55 2 92 14.60 90 2 3 4 92 92
Male Adult Alive First year 43 4.35 2 25 4.40 23 2 3 5 25 25
451

Male Adult Dead First year 21 6.10 2 29 7.89 27 2 3 5 29 29

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452

Table 10.9 (cont.)

Distance in km
Found Time
Percentile
Ringed Dead or after
Sex as Alive ringing n Mean Min Max SD Range Q25 Median Q75 99.99 99.95

Male Adult Dead >1 year 32 12.31 2 86 19.62 84 3 5 9 86 86

Unknown Adult Alive >1 year 474 12.05 2 319 30.75 317 2 3 7 319 319
Unknown Adult Alive First year 280 13.15 2 285 30.85 283 2 4 11 285 285
Unknown Adult Dead > 1 year 321 18.56 2 332 36.36 330 3 6 15 332 332
Unknown Adult Dead First year 385 15.20 2 262 32.56 260 3 6 13 262 262
du Feu et al. 2016. Data are grouped by sex, either alive or dead at the last observation within or over one year after ringing as an adult bird (real age
unknown).
 10.3 Demographic Limiting Factors

Birds ringed as adults systematically differed in dispersal distance of age within one
year after ringing according to the sex, i.e., females always dispersed further than males,
when found alive or dead. Females that were still alive on their ultimate observation
had an average dispersal of 1.4 km (n = 634), while for males it was 0.97 km (n = 195).
When found dead, a similar difference was found for females, 5.02 km (n = 150) and for
males, 3.40 km (n = 55). When observed alive at least a year after birth and ringing,
females dispersed around the same distances on average, 2.33 km (n = 2057), as males
2.45 km (n = 388). For dead birds the situation was similar, with 5.56 km for females
(n = 373) and 5.59 km for males (n = 92). So, for the breeding dispersal of adults, the
differentiation between adult birds is no longer relevant.

10.3.3 Immigration
The dispersal of juveniles has a substantial effect on immigration. However, estimating
and modeling immigration is a real challenge. In the past, several methods have been
developed to estimate immigration rate, but they either require strong assumptions or
combine in a piecewise manner the results from separate analyses. In most methods, the
effects of covariates cannot be modeled formally. Abadi et al. (2010) developed a Bayesian
integrated population model which combines capture–re-capture data, population
counts and information on reproductive success into a single model that estimates and
models immigration rate, while directly assessing the impact of environmental covari-
ates. They assessed parameter identifiability by comparing posterior distributions of
immigration rates under varying priors, and illustrated the application of the model with
long-term demographic data of a Little Owl population from southern Germany.
Demographic data were collected near Göppingen, where nestboxes were installed and
the population monitored from 1978 to 2003. Each year all nestboxes were checked
several times and reproductive success (n = 353 breeding attempts) was noted, nestlings
were marked with rings (n = 659) and captured and marked adults that were in the
nestboxes at the time of the checks (n = 73) were recorded. Capture–re-capture data, data
on reproductive success and population size were available. A two-level index of vole
abundance (high, low) was recorded each spring, based on the number of vole hills and
holes. Abadi et al. (2010) assessed the impact of environmental covariates on immigra-
tion. The resulting posterior distributions were insensitive to different prior distributions
and dominated by the observed data, indicating that the immigration rate was identifi-
able. Average yearly immigration into the Little Owl population was 0.293 (95% CI
0.183–0.418), which means that ca. 0.3 females per resident female entered the popula-
tion every year. The immigration rate tended to increase with increasing abundance of
voles. The means to estimate and model immigration is an important step towards a
better understanding of the dynamics of geographically open populations of Little Owls.
The demographic estimates obtained from the developed integrated population model
facilitate population diagnoses and can be used to assess population viability. The
structural flexibility of the model can constitute a useful tool for conservation.
Besides stochastic and deterministic factors, the spatial extent and the isolation of a
local population may have an impact on the contribution of the different demographic

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 Chapter 10: Population Regulation

components. Using long-term demographic data, Schaub et al. (2006) performed retro-
spective population analyses of four Little Owl populations (Basel, Göppingen, Genève
and Friedrichshafen) with differential spatial extent and degree of isolation to assess the
contribution of demographic rates to the variation of the growth rate of each local
population and to the difference of the growth rate among populations. Schaub et al.
(2006) used life table response experiments (LTREs) for the retrospective perturbation
analyses, which can be an accurate way to identify the demographic parameters rele-
vant for population changes. LTREs estimate the contribution of a demographic par-
ameter to population change from the sensitivity of population growth rate to changes
in the demographic parameter and on the magnitude of this change using a simple
female-based model that assumes that Little Owls start to reproduce at an age of one
year and that the reproductive performance and survival of reproductive individuals
does not change with age. The number of breeding females in the year t+1 is given by
the sum of the number of females that were already reproducing in year t and that have
survived and not emigrated in year t+1, the number of young females produced in the
year t that were locally recruited in year t+1, and the number of female immigrants.
In all populations, variation in fecundity contributed least, and variation in adult
survival contributed most, to the variation in the growth rate in three of the four
populations. Between-population comparisons revealed that differences mainly
stemmed from differences in immigration and local juvenile survival. The relative
importance of immigration to the growth rate tended to decrease with increasing
spatial extent and the isolation of the local population. None of the four local popula-
tions was self-sustainable. Because the local populations exported and imported indi-
viduals, they could be considered as open recruitment systems in which some of the
recruited breeding birds were not produced locally. The spatial extent and the degree of
isolation of a local population has an impact on local population dynamics; hence
these factors need to be considered in studies about local population dynamics and for
deriving conservation measures.
A specific aim of the study by Schaub et al. (2006) was to understand the demo-
graphic reasons for the two local population extinctions (Basel and Friedrichshafen).
The decline until 1995 was due to a strong decrease in local juvenile survival, resulting
in lower local recruitment that could not be halted by increasing adult survival. Hence,
there was a decline in the gain component, rather than an increase in the loss compon-
ent that caused the population to decline. A possible environmental change that could
have caused this decline was a decrease in food availability due to changes in farming
practices. This was evidenced by declining body mass of the chicks at the time of
marking. In Basel, the demographic parameters were not very variable and had no
trends over time. Consequently the growth rate remained fairly constant, but as it
was less than one the population declined asymptotically. The equilibrium of gain
and loss had already been disturbed before the study started, and it was difficult to
infer which ones had been involved. However, Schaub et al. (2006) could compare
demographic parameters with those from the other populations, taking into account
that the spatial extent of the populations differed. Fecundity was in the range of the
other populations and did not contribute to the difference in growth rate, suggesting

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 10.3 Demographic Limiting Factors

that extinction was not related to a decrease in fecundity. The spatial extent of the Basel
population was the smallest, thus Schaub et al. expected the estimates and the contri-
butions of local juvenile and adult survivals to the difference in the growth rate among
populations to be lower, and the estimate and the contribution of immigration to the
difference in the growth rate to be higher, than in the other populations. While these
expectations were true for local juvenile survival, they were not true for local adult
survival and particularly not for immigration. This suggested that the main demo-
graphic reasons for the extinction of the Little Owl population in Basel were declines
in immigration and adult survival rates that had already happened before the study
started in 1978. Extinctions of many local populations in the vicinity of the focus
population due to changes in farming practices (Luder and Stange 2001) were probably
the reason for insufficient immigration, and predation of Stone Martens on breeding
females may have contributed to low survival (Luder and Stange 2001). This compara-
tive analysis by Schaub et al. showed that the demographic mechanisms resulting in
local extinction differed, despite the reason for both population changes being
farming practices.

10.3.4 Genetic Variability

Loss of genetic variability can be an extra determining factor in declining Little Owl
populations (Pertoldi et al. 2012). Pertoldi et al. assessed if the dramatic decrease in
population size during recent decades in Denmark could be observed at the genetic
level and the degree of genetic relationship between the surviving owl population in
Northern and Western Jutland and the (now extinct) population on the isle of Funen.
Their research helped to determine whether the remaining population was part of one
large population that had declined, or if genetically distinct population segments had
previously been present. These issues were addressed using data obtained by analysis of
microsatellite DNA from museum samples of owls from Jutland and the isle of Funen,
covering the period from 1918 to 2000, and from more recent samples collected from
the owl population (2001–2010). The Danish Little Owl population serves as a good
example for analyzing genetic patterns of a declining and decomposing population.
The genetic variability was compared with samples from a southwestern German
population with an overlapping set of genetic markers. The number of alleles in five
Danish populations were lower than the German population, indicating a lower genetic
variability, which may be caused by recent population declines resulting in genetic
bottlenecks, a historical low effective population size due to its relative peninsular
isolation and dramatic weather events in the past leading to dramatic reductions in
numbers, as in the 1940s, when the species disappeared from the Danish isle of Lolland
and in Sweden.

10.3.5 Offspring Sex Ratio Bias

Two general mechanisms can lead to biased offspring sex ratios in different ways and in
various periods of reproduction: sex-biased survival or parental sex allocation. Tschumi

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 Chapter 10: Population Regulation

et al. (2019) examined nestling sex ratio patterns between hatching and fledging, sexual
size dimorphism and factors affecting nestling survival using growth and survival data
from 846 individual Little Owl nestlings with known sex from 307 broods in Germany,
the Netherlands and Denmark. Nestling sex ratio was female-biased, mainly due to a
significant female bias in the first-hatched chicks. Females showed a higher body
weight than male nestlings at ringing and the body weight of nestlings decreased with
hatching sequence. Nestling survival was higher in females (Φ = 0.91) than in males
(Φ = 0.85), and survival rates were positively related to body mass and negatively to
brood size. Although the observed lower survival of males can cause an overall female-
biased sex ratio, the sex dimorphism and survival patterns found by Tschumi et al. are
unlikely to explain the conspicuous sex ratio pattern with a female bias in the first-
hatched nestlings and the increase in female bias across the season. These results point
towards interacting mechanisms of parental sex allocation strategies and sex-specific
survival. As the female bias was allocated to the first rank that is most likely to survive,
the female bias will increase under suboptimal breeding conditions. Hence, Tschumi
et al. (2019) suggested that under suboptimal ecological conditions, higher investment
into females is adaptive in Little Owls.
Van Harxen and Stroeken (2021) continued their research in the Winterswijk region
in the years 2014–2017. They found no significant differences between the proportion
of female and male nestlings, not over the entire dataset and not based on only those
nests where all eggs hatched and all hatched young were sampled. Although the ratio
(proportion of females) fluctuated in both datasets, the differences were not significant
on an annual basis nor over the total of all the years. An analysis based on the ranking
order over all years also revealed no significant differences. In addition, female nestlings
were not significantly heavier than their male siblings.
The results that Tschumi et al. (2019) found for their research area in 2013 were thus
confirmed by the results from the 2014–2017 period. It is plausible that the suboptimal
conditions under which the German and Danish Little Owls grew up in comparison
with their Dutch counterparts mattered. Despite some possible methodical difference
in the age determination, the German nestlings weighed 115.3 g on ringing day (day
20) on average and the Dutch were substantially heavier around the same day, i.e.,
137.2 g on average, which is a difference of almost 19%. This confirms that the overall
food conditions in the Winterswijk study area were substantially better during the
study period. The micromammal index was significantly higher for the years
1998–2012 in each separate study year. The survival from hatching to ringing was
0.842 in the German study versus 0.903 in Winterswijk.
During the nestling phase, male nestlings were on average as heavy as their female
siblings, in contrast to the German study where the female nestlings were on average
2.6% heavier. Gender-linked mortality due to size differences was therefore not
observed in Winterswijk. The relatively small weight differences between adult male
and female nestlings (7–8%) in Winterswijk would not feature a difference in gender
ratio under normal circumstances. The main causes for a bias in the ratio in the
Winterswijk study area seemed to be absent. Under favorable food conditions, the sex
ratio does not deviate from a 50:50 ratio.

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 10.3 Demographic Limiting Factors

10.3.6 Capture–Re-capture analysis (MARK)

An analysis of capture–re-capture data for three research areas in Flanders
(Zuiderkempen (1991–2005), Dijleland (1998–2005), Hageland (1992-2005)), one in
Wallonia (1989–2005) and one in the Netherlands (Achterhoek) (1986–2005)) yielded
8550 capture–re-capture observations collected during 86 man-years of conservation
and data collection on 5649 Little Owls within diverse anthropogenic habitats and
confirmed that reproductive success, just as survival, is influenced by a combination of
extrinsic (environmental factors) and intrinsic factors (individual quality) (Bekaert
2006). Four different habitat typologies were distinguished: small-scale landscapes,
built-up areas, and field- and pasture-dominated.
In addition to density and social parameters, the reproduction of Little Owls was
tested for age dependency, with clutch sizes and number of young increasing with
increasing age to a maximum, after which reproductive success in older
individuals declined.

10.3.6.1 REPRODUCTION AND SURVIVAL

In contrast to the number of chicks that hatched and fledged, clutch size significantly
differed between the study areas (χ2 = 28.91, DF = 3, p <0.0001, n = 830) where Little
Owls in Wallonia and the Hageland lay fewer eggs than individuals in the Achterhoek
and Dijleland. Breeding success as the percentage of the clutch that yielded fledglings
varied between 72.6% in Wallonia and 57.2% in the Netherlands (χ2 = 20.46, DF = 3,
p <0.0001, n = 802). Little Owl populations were stable (Hageland, the Netherlands and
Wallonia) or showed slight growth (Dijleland and Zuiderkempen). The survival was
smallest in the Netherlands, where survival was time dependent, and greatest in
the Zuiderkempen.

10.3.6.2 LANDSCAPE DEPENDENCY

Clutch size (χ2 = 12.14, DF = 3, p <0.007, n = 633) and breeding success (χ2 = 20.46,
DF = 3, p <0.0001, n = 802) differed significantly between the different habitat typolo-
gies. Little Owls occupying built-up areas had the highest breeding success. Individuals
breeding in small-scale landscapes had the lowest breeding success. Clutch sizes within
meadow-dominated habitats were the largest, in built-up areas the smallest. The opti-
mal number of eggs was found in nesting sites with 0.5–2 ha of built-up area (χ2 = 14.39,
DF = 3, p <0.0024, n = 633).
Bekaert (2006) found that the reproductive success and survival within a certain
habitat type was only slightly influenced by the landscape composition. Reproduction
models showed that the quantity of built-up areas had a negative impact on clutch size.
This impact was compensated by higher breeding success (number of fledglings/
number of eggs) within these habitats so that the absolute number of fledged young
was not significantly lower than in other habitats.
Based upon survival and reproductive success, as an indicator of habitat quality, all
habitat typologies were of the same quality, probably due to the high degree of land-
scape heterogeneity in the respective research areas, since most long-term conservation
is done in rather optimal habitats. The probability that a particular habitat is of good

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 Chapter 10: Population Regulation

quality is therefore mainly determined by other factors. Both the weather and the
density as extrinsic, and individual quality (age, social variables) as intrinsic factors,
each affect the reproductive success of the Little Owl.

10.3.6.3 AGE DEPENDENCY

Age-dependent reproductive success has already been described for many animal
species (Cichon 2003). The relationship between age and reproductive success follows
a pattern in birds where reproductive success reaches an optimum in individuals of
middle age (Ryder 1980).
The breeding success and the number of fledglings significantly depends on age (χ2 =
13.75, DF = 4, p = 0.0085, n = 827).
An optimum is reached in individuals with an age of 5–6 years. Breeding success
decreases again in older individuals. These results are also obtained by estimating a
quadratic function (F = 0.0062, DF = 14, p = 0.0062, n = 5) (Figure 10.9).
Figure 10.10 gives the distribution of breeding females in the different research areas.

10.3.6.4 WEATHER DEPENDENCY

Population growth is also influenced by the winter temperature. From 1987–1996 the
population was not stable. Fluctuations in population growth paralleled fluctuations in
winter temperature. The population growth experienced a minimum in 1991 after a
severe winter with an average daytime temperature of 0.8 C, and a maximum in
1993 after a mild winter in 1992 with an average daytime temperature of 5.8 C.
A mild winter has a positive effect on the available food during the following breeding
season, with a direct impact on the number of fledglings. The rise in population growth,
the year after a mild winter, may possibly be the result of the optimal conditions of the
previous year.

4.5
y = –0.0122x2 + 0.1734x + 3.2545
4 R² = 0.542
Number of eggs, young, fledglings

3.5
y = –0.0144x2 + 0.15x + 2.6207
R² = 0.4866
3
Eggs
2.5 Young
Fledglings
2 Poly. (Eggs)
y = –0.021x2 + 0.2144x + 1.9198
R² = 0.6622 Poly. (Young)
1.5 Poly. (Fledglings)

0.5

0
0 2 4 6 8 10 12 14 16
Age (years)

Figure 10.9 Influence of age on the reproduction of the Little Owl. The clutch size reaches an
optimum in individuals of 7 years. The number of fledglings has an optimum in individuals between
5 and 6 years old.

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 10.3 Demographic Limiting Factors

In contrast to survival, population growth within the different habitat typologies is

time dependent. After eight years, the population growth within the different habitat
typologies converges to 1, resulting in a stable population. In the early years, large
fluctuations in population growth can be observed largely caused by changes in winter
temperature. The impact of a severe winter on population growth depends on the
habitat typology. Individuals within built-up areas will experience a greater negative
influence from a severe winter than individuals in meadow-dominated or small-scale
landscapes, where a severe winter apparently has no impact on population growth.
Based on the buffering effect against extreme weather conditions and the associated
impact on population growth, small-scale landscapes are of higher quality than built-up
habitats. The strong negative impact of winter temperature on population growth
within built-up areas may have to do with very low recruitment. Just like the number
of fledglings, the survival of individuals is influenced by winter temperature (F1.16 =
41.58, DF = 17, p = 0.005, R2 = 0.54). Figure 10.11 shows the linear relationship between
winter temperature and survival.

10.3.6.5 INFLUENCE OF SOCIAL PARAMETERS

Population growth is the result of both immigration and the settlement of local recruits.
However, the vast majority of ringed pulli are not re-captured. These individuals leave
the local population or breed in natural cavities. The share of local recruits relative to

6
The Netherlands
Wallonia
Zuiderkempen
5
Hageland
Dijleland

4
Age (years)

0
1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004
Year

Figure 10.10 Age distribution of breeding females. Comparison between the different research areas.

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 Chapter 10: Population Regulation

100
y = 0.0705x + 0.4423
90 R² = 0.3376
Survival (%)

80

70

60

50
3.5 4 4.5 5 5.5 6
Average Winter Temperature (°C)
Figure 10.11 Linear relationship between survival and winter temperature.

immigrants is therefore relatively small (18%). Breeding success is significantly greater

in females who change partners (χ2 = 9.76, DF = 1, p = 0.0018, n = 334). If a female ever
moves (χ2 = 6.85, DF = 1, p = 0.008, n = 828) or is single, this has a negative influence on
the total number of eggs laid (χ2 = 4.25, DF = 1, p = 0.05, n = 828).
Bekaert (2006) found that the number of local recruits did not increase with increas-
ing number of fledglings. Of the total number of local recruits, 43% were already
breeding the next year. For the other 57%, who were only found later as breeding
individuals, it was not known where these individuals were during the period between
ringing and re-capture. Compared to immigrants, local recruits in Flanders had a higher
reproductive success. Local recruits that breed immediately the first year after birth
within the same population have an energetic advantage compared to immigrants. This
advantage can lead to greater reproductive success. This may conflict with the age
dependence found where young individuals have a lower breeding success. The ener-
getic advantage seems to be more important than the age of individuals for
reproductive success.
There have been cases where young individuals first migrated to another population,
bred there for one to several years, and then returned to their original population.
Alternatively these local recruits might have bred in natural nesting sites without being
re-captured. Another possibility is that these individuals did not breed and were terri-
torial. These so-called floaters wait until a partner and nest location become available.
Within the Dutch population, ex-singles have a higher reproductive success com-
pared to annual breeders.
The occurrence of local recruits differed significantly between the different research
areas, with Wallonia and Hageland having more local recruits.
No significant difference was found for survival of local recruits (LRT test, χ2 = 16.4,
DF = 15, p = 0.15). In the Netherlands, single birds (0.81  0.02) had a significantly
higher survival rate (LRT test, χ2 = 4.32, DF = 1, p = 0.037), compared to breeding
individuals (0.67  0.01). For Flanders no significant differences were found.
Per year, the average nearest-neighbor distance and the proportion of singles correl-
ated positively. Finding a partner strongly depends on the density (F = 29.69, DF = 12,

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 10.3 Demographic Limiting Factors

0.6

y = 0.0005x –0.4095
0.5 R² = 0.8017
Concentration singles

0.4

0.3

0.2

0.1

0
1000 1100 1200 1300 1400 1500 1600 1700 1800 1900
Nearest-neighbor distance

Figure 10.12 Occurrences of singles as a function of density. Concentration singles = number of

singles/(number of singles + breeding individuals). The smaller the nearest-neighbor distance,
the higher the density.

p = 0.0002, R2 = 0.71, n = 13), the lower the density, the smaller the chance that a Little
Owl will find a partner (Figure 10.12).

10.3.6.6 POPULATION MODELING WITH MARK

In addition to survival and re-capture, population growth and recruitment per research
area can be calculated. MARK calculates population growth based on population par-
ameters that provide a sufficient amount of capture–re-capture data to be estimated.
Figure 10.13 provides an overview of the estimated parameters by analyzing capture–re-
capture data.

10.3.6.7 DEMOGRAPHIC PARAMETERS

Figure 10.14 shows survival, population growth, number of fledglings and winter
temperature. The harsh winter of 1990–1991 is associated with declining population
growth and survival. The number of fledglings continued its downward trend. The
following year (1992), featured a relatively mild winter leading to both higher survival
and number of fledglings.
Different hypotheses have been tested regarding the difference in survival, re-
capture rate and population growth within the different habitat typologies. The best
model (lowest AICc) is one where survival is time dependent, but does not differ
between different typologies, the re-capture rate is habitat dependent and the popula-
tion growth both varies over time and between habitats. Individuals within a habitat
typology characterized by built-up areas have the greatest chance of being re-captured
(0.66  0.030). Within small-scale landscapes, the chance that a individual is re-
captured becomes the smallest (0.56  0.031). There is a close relationship between

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Alive and
p re-sighted
Alive and
in sample
F 1–
p Alive but
Alive not re-sighted
1– Alive and
S F
left sample
Marked and
released alive Dead and
at start of year 1 r reported
–S
Dead
1–
r Dead but
not reported

Figure 10.13 Overview of parameters that can be estimated using MARK. S = survival,
1 – S = mortality, F = probability of a bird staying in the research area, 1 – F = probability of emigration,
r = probability that a bird dies and is found dead and retrieved, p = probability that a bird is still
alive, did not emigrate and is re-captured.

1.8 9

Number of fledglings and average winter temperature

Survival Population growth Winter temperature Fledglings

1.6 8

1.4 7

1.2 6
Population paramters

1 5

0.8 4

0.6 3

0.4 2

0.2 1

0 0
1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006
Year

Figure 10.14 Different demographic parameters (survival, population growth and reproduction
(number of fledglings)) and the average temperature in winter. Until 1998 survival, population growth
and reproductive success are linked to winter temperature, with all having a minimum in
1991 followed by a maximum in 1992.

population growth and winter temperature, especially in extreme weather conditions.

Immediately noticeable are the large fluctuations in population growth from 1987 to
1996. From 1997, the populations within the different typologies had a stable popula-
tion growth around 1. The impact of winter temperature on the population growth

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 10.3 Demographic Limiting Factors

within the different typologies was not the same in all research areas. In habitat
characterized by built-up areas, the population showed a declining population trend
after a hard winter. This trend was reflected to a lesser extent in the population within
habitats characterized by pasture. Within small-scale landscapes, a harsh winter had
little influence on the population and in habitats characterized by arable land, the
population declined less quickly than the year before.
A peak in population growth came after a year with a mild winter and was greater
within built-up areas followed by meadow-dominated habitats.

10.3.7 Home Range and Territory

A bird’s home range is defined as “the area that embraces all activities of a bird or pair
over a given time period.” Generally, a yearly home range is used, but sometimes there
is a seasonal area (e.g., breeding season area). A territory is defined as “the area around
the nest that is defended” according to Newton (1979). Finck (1990), on the other
hand, considers a territory to be “the defended area for food and nesting.” No matter
which definition is used, the best way to get home-range values is by attaching a small
radio-transmitter to the bird, allowing the movements and activity of the Little Owl to
be tracked. Generally, the transmitter is put in place through a backpack harness and
weighs 6–9 g, that is to say 2.5–4.5% of the body weight of the bird. Typically, the
main reason for assessing home-range size is to define the area within which the species
finds selected habitats and resources. While the backpack transmitter may seem an
inconvenience for the Little Owl, it has been shown that the transmitter does not
influence the bird’s breeding success. Exo (1987) and Zens (1992), respectively, reported
the following breeding success data: 2.5 fledglings/pair with transmitter (n = 4 pairs)
and 1.33 fledglings/pair without transmitter (n = 15 pairs); 1.5 fledglings/pair with
transmitter (n = 4 pairs) and 1.29 fledglings/pair without transmitter (n = 31pairs).
To accurately determine home-range size, a minimum of 30 locations per bird are
needed (Kenward 1987) over a specified timeframe (e.g., 12 months).
Home-range sizes are often quantified by use of the Minimum Convex Polygon
method. This is the area enclosed by a line that connects the outermost observations
(Figure 10.15). The Minimum Convex Polygon and Adapative Kernel are methods used
to assess the home-range dimensions of owls (and other species). In Table 10.10, we
summarize home-range sizes based on the Minimum Convex Polygon method. In
addition to the data presented in Table 10.11, Eick (2003) monitored three adults and
four juveniles during dispersal time from July to November (48–120 days of monitor-
ing). The available data on home-range sizes reflect efforts that were either data-
intensive for a given season (e.g., breeding season), or reflect a relatively low number
of locations gathered over a longer period. In order for annual home-range sizes to be
calculated, additional rigor in data collection is warranted. For most owl species, it is
recognized that an asymptote in radio-locations occurs with increasing home-range
size. That is to say, after about 100 or so radio-locations, the home-range sizes either do
not increase in area, or do so only minimally. It is important to note that for conser-
vation applications, radio-locations gathered over a full year (or more), for both male

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 Chapter 10: Population Regulation

and female members of a pair, are the most useful, as the area involved most accurately
represents the area actually used by a pair of owls. Thus, ideally, researchers should
gather locations on radio-marked owls at least twice a week (during the night when the
owls are active) for 12 months. Gathering only one location per night would minimize
problems associated with auto-correlation and the independence of data.

Table 10.10 Summary of Little Owl home-range sizes from different studies, as
calculated using the Minimum Convex Polygon or MCP method

Home range in ha

Little Owls Annual Density in Study

monitored mean Min Max pairs/km2 area in km2 Author Country

12 14.5 1 50 1.7 35 Exo 1987 Germany

19 14.6 2 107 1.6 29 Finck 1990 Germany
4 27.4 1 150 0.15 260 Zens 1992 Germany
8 31 5 107 0.09 437 Génot and France
Wilhelm
1993
15 51.3 6.5 137 0.5 - Orf 2001 Germany

The home-range values reflect the size of area within the MCP during the time the owl
was followed.

Figure 10.15 Examples of home ranges (after Exo 1989 and Génot and Wilhelm 1993).

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 10.3 Demographic Limiting Factors

Figure 10.15 (cont.)

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Table 10.11 Summary of Little Owl home-range sizes from different studies in Europe

Author and Time followed No. of Home range

study location Sex Age in months locations (MCP) in ha

Orf (2001) Male Adult 5 102 137.1

Main-Taunus Male Juvenile 6 74 44.7
(Germany) Male Adult 6 127 6.5
Male Adult 6 251 43.1
Male Adult 6 123 56
Female Adult 6 163 32
Male Juvenile 4.5 105 41.5
Male Adult 3.5 97 128.9
Female Adult 4.5 43 39.1
? Adult 2 25 11.2
Génot and Male Adult 9 180 50.2
Wilhelm Male Adult 2 65 13.1
(1993) Male Adult 3 50 15.6
Vosges du Nord Male Adult 1 30 46.5
(France) Female Adult 8 190 84.1
Female Adult 3 95 26.2
Female Adult 3 55 28.4
Female Adult 4 110 107.4
Zens (1992) Male Juvenile 10 25* 148.9
Voreifel Female Adult 6 14* 17.8
(Germany) Male Adult 7 18* 51.6
Female Adult 9 16* 73.8
Exo (1987) Male Adult 14 45* 41.5
Niederrhein Female Adult 10 30* 38.4
(Germany) Male Adult 14 29* 10.1
Female Juvenile 10 17* 9.6
Male Adult 10 14* 4.6
Female Adult 12 20* 3.1
Male Adult 12 26* 4.9
Female Adult 12 15* 2.5
Male Adult 8 19* 17.8
Female Adult 8 14* 20.7

When the number of locations is followed by an asterisk (*), it reflects the number of night-
time locations acquired during the first three or four hours of the Little Owls’ activity.

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 10.3 Demographic Limiting Factors

Finck (1990) and Grzywaczewski (2009) were the only authors to study the size of
the Little Owl territory using a decoy and a loud speaker. The territory size of 19 tracked
males ranged from 1–68 ha (mean 12.3 ha) and of 10 tracked males and one female
ranged from 1.8–95 ha (mean 19.9 ha). Finck showed that Little Owls defended their
territories all year, with the aggressiveness of the owls changing according to the
season. Grzywaczewski (2009) reported a difference according to the season with a
range of 2.5–65 ha (mean 18 ha) during the egg laying and incubation period, 1.9–95
ha (mean 28.2 ha) during the nestling period and 1.8–34 ha (mean 19.9 ha) for the
fledgling period. Grzywaczewski et al. (in prep.) compared the active range of 33 Little
Owls, 16 in agricultural and 17 in urban landscapes in 2002–2004 between April 15 and
August 12. The territory size was smaller in urban (2.15  0.89 ha, range 0.3–27.9 ha)
than in rural (6.08  1.29 ha, range 0.5–92.4 ha) landscapes. Urban territories of the
Little Owl are characterized by lower cover of grassland, tall crops, short crops, gardens
and orchards, as well as a higher cover of built-up areas than territories in rural
landscapes. Territory size in urban landscapes was negatively correlated with cover of
grassland, short crops and cover of buildings, whereas in rural landscapes territory size
was negatively correlated with season progression and positively by altitude above sea
level. The rate of territorial displays was similar between urban and rural territories;
however, birds differentially utilized various structures. In urban territories birds com-
monly used buildings, whereas in rural territories, birds used electric pylons and trees.
The rate of aggressive displays was similar between agricultural and urban sites.

10.3.7.1 USE OF THE HOME RANGE

As with many other owl and raptor species, breeding Little Owls have been found to
concentrate their activities in a portion of their overall home ranges. For example,
Génot and Wilhelm (1993) found that the owls they studied in France had a mean
home-range size of 31 ha, and spent 80% of their time in an area of 3.5–6 ha. Zens
(1992) found that owls concentrated their activities in an area of between 20% and 80%
of the home range and Exo (1987) found that owls concentrated their time in an area of
between 5% and 50% of the home range.
Eick (2003) studied some adults and dispersing juveniles during the breeding season.
The breeding area used by the adults was 20–30 ha. Predispersal juveniles used different
sized areas depending on their age: during the first week after leaving the nest, 1.5 ha
(n = 5 individuals); nine weeks after, 15.8 ha (n = 6 individuals) and 12 weeks after,
31 ha (n = 3 individuals). Note that the juveniles did not have a “home range” but
rather that the areas given here reflect their predispersal excursions. Exo (1987) showed
that 50% of Little Owl vocal activity was observed in a radius of less than 50 m around
the nest or roost. Orf (2001) found a mean activity radius (distance from the nest or
roost) of 78 m during courtship, 219 m during the breeding season and 181 m in winter.
For Zens (1992), the activity radius ranged from 60–120 m in June, the radius increased
to 170–300 m in September–November and to 420–530 m up to 1.5 km in winter. In
France, the mean activity radius was 430 m, depending on the sex and the season; it was
longer in winter but some distances were also long in July (up to 2.6 km).

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Figure 10.16 Lunar periodic changes in activity of a pair of Little Owls kept under natural light
conditions. (a) Daily activity in relation to moon phase, total from seven lunar cycles * –p < 0.05
(Wilcoxon–Wilcox test). (b) Example of variation in the daily activity total perch use over one moon cycle.

Exo (1989) determined the daily activity pattern. Activity peaks of about 1–2 hours
just after sunset and before sunrise, when light conditions of <1 lux were observed. Less
activity was recorded around midnight. Males and females showed the same activity
patterns, except in the breeding season (April/May–July). Their activity lasted between
5.5 and 9 hours per night, while during the breeding period the daytime activity was
longer than in the night.
Little Owls are more active during the full moon (Figure 10.16) phase but they move
less when it is raining and the wind speed is stronger than 3 Beaufort.
In France, hunting perches used by the Little Owls were in trees (82.2%), on fence
poles (13.7%), in hedges (3.9%) and in buildings (0.2%) (Génot and Wilhelm 1993).
Orf (2001) showed that Little Owls spent the daytime mainly in nestboxes (44.3%),
hedges or trees (38.5%), natural cavities (15.5%) and in other sites such as buildings
(1.7%). Of 39 dispersing juvenile owls, Eick (2003) found that 56% of the roosts were
in woodpiles, 23% in branches of trees, 8% in nestboxes, 5% in tree holes and 8%
in other places.

10.3.7.2 VARIATION IN HOME-RANGE AND TERRITORY SIZES

Home-range and territory sizes depend on different factors, e.g., the sex and the age of
the birds, the population density, the season and the structure of the habitat, and the
density and availability of prey.

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 10.3 Demographic Limiting Factors

Sex and Age of the Little Owls

Exo (1987) has monitored pairs of owls the longest (Table 10.11) (one pair over one year
and three pairs over 10 months), and all authors found Little Owl males to have home
ranges larger than those of females, except for certain males during the end of summer
and beginning of autumn (Exo 1987, Génot and Wilhelm 1993). Sunde et al. (2009)
followed 27 radio-tagged owls from 14 territories over two years in Denmark. Home-
range sizes were more than twice as large in January–March than during the rest of the
year, with no consistent differences between males and females. On an annual basis, a
pair spent on average half of their time (50% MCP) within an area of 2.6 ha (95% CI
1.5–4.5) and 90% of their time (90% MCP) within an area of 41 ha (range 27–64 ha).
The overlap of home ranges between males and females can be matched perfectly
(Exo 1987, Zens 1992) or can overlap by 38–94%, depending on the season (Génot and
Wilhelm 1993). During the breeding season, Eick (2003) found the range of adult
females was within the range of their partner for two pairs (males on average 31.7 ha,
females 21.9 ha). After independence of the young, both adults used different plots. The
territories of neighboring males did not overlap throughout the year (Exo 1992). Orf
(2001) found no significant relationship between the age of the Little Owl and the size
of the home range. Finck (1993) showed that new settlers defended larger territories
(15 ha, n = 5) than well-established Little Owls or long-term settlers (8.3 ha, n = 6).
While established owls are more knowledgeable about their territory due to their longer
occupation, other possibilities for the differences in home-range sizes are possible (e.g.,
quality of sites, food availability).
Sunde et al. (2009) studied home-range overlap between mates for 12 pairs in total,
9 pairs for the nonbreeding period, 6 pairs during the posthatching period. Mean mate
overlap was not significantly lower than 1 in any of the situations, showing that mates
generally shared one communal home range. Parents with dependent offspring
expressed the lowest level of range-sharing, with a mean-adjusted overlap of 85%.
They compared 348 inter-mate distances from 11 mate pairs, of which 37% were less
than 100 m, compared to 21% of randomized inter-mate distances. The observed inter-
mate distances were significantly less than expected from random. The proportion of
times mates were observed within 100 m of each other (a distance short enough to
facilitate intra-pair communication and mate guarding) varied across months, but was
not related to time before the start of incubation. Hence, no mate guarding was
apparent.

Density and Neighbors

No differences in territory size were detected between population densities of 0.75
calling males/km² and 2.25 calling males/km² (Finck 1990). Van Nieuwenhuyse et al.
(2008) examined the density data and average home-range size (as seen in Table 10.11),
and found no significant linear relationship between them. Sunde et al. (2009) reported
a substantial difference between pairs, with a tenfold difference in range size
between the smallest and largest home ranges. Pairs with neighbors nesting within
1.5 km of their own nest had two to three times larger home ranges than pairs with
no neighbors.

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Table 10.12 Comparison of the median areas (ha) used by Little Owls during
different seasons

Area used

Winter Courtship Breeding Summer

Nov–Feb Mar–Apr May–Jun Jul–Oct Author

4.2 4.7 3.4 7.2 Exo (1987)

26.7 15.2 21 9 Finck (1989)
48.2 – 12.9 15.6 Zens (1992)
14.9 11.9 4.6 4.5 Génot and Wilhelm (1993)
21.8 12.3 31.5 – Orf (2001)

Season
With the exception of data from Exo (1987) and Orf (2001), home-range sizes were
larger in winter than during the other periods (Table 10.12). Home ranges are substan-
tially larger in winter than during the courtship period, as behaviorally, the attachment
to the breeding cavity is looser than during courtship and breeding. The variation in
home-range size is likely also linked to the seasonal availability of prey. Orf (2001)
found the largest median home-range size in May and June, during the feeding of the
female and nestlings, and considered this to be a result of the owls’ need to move
further afield within a poor-quality hunting range. The seasonal variation in activity
distance and range size that peaked in January–March and dipped in summer in
Denmark (Sunde et al. 2009) was comparable to that found by Finck (1990) and
Zuberogoitia et al. (2007). The seasonal increase in range size during the cold season
might not necessarily be motivated by foraging decisions, since owls lowered their
activitiy distance from roosting sites on cold nights.

Roosts
Sunde et al. (2009) studied roost use by Little Owls in Denmark. In daylight the owls
roosted inside buildings or (less often) in trees/bushes <50 m from buildings. All owls
roosted in association with the buildings used for nesting, or occasionally in adjacent
buildings <300 m away. Half of all telemetry fixes of nocturnally active owls were
located within 125 m of the nest, 75% within 282 m and 90% within 569 m, the
maximum being 3070 m. The mean distance from nest/roosting sites varied between
territories, across months, in relation to breeding status in interaction with sex and as a
function of weather parameters. On a seasonal basis, mean nest distance was largest in
January–April and smallest in May–June. Outside the breeding season the distances
were similar for males and females. In males, nest distance did not vary significantly
with breeding status, whereas females stayed closer to the nest while attending young
compared to the nonbreeding season. A highly significant quadratic relation existed
between the distance and temperature, owls being observed most distant from the
nest at temperatures around 6 C. The distance was positively correlated with the

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 10.3 Demographic Limiting Factors

temperature in the coldest months of the year and negatively correlated in the warmest.
Additionally, the distance was on average 17% higher in dry nights compared to
rainy nights.

Structure of the Habitat

The structure of the habitat has a large influence on the size of the home range and territory.
In terms of habitat structure for Little Owls, we often think of agricultural land uses linked
to the seasonal development and height of the vegetation. Finck (1990) considered four
types of land use having an influence on Little Owl foraging as grassland (meadow and
pasture) with free access for hunting year round, barley and wheat fields which are unsuit-
able (due to their height) from early May to mid-July and from mid-May to mid-August,
respectively, corn fields which are unsuitable for hunting from July to September/October
and sugar-beet fields which are unsuitable from September to November.
Many authors have stressed the importance of grasslands in the home range of the Little
Owl. The species can survive with only 0.5–3 ha of low-cut grassland (Exo 1987, Zens 1992,
Eick 2003). Finck (1990) observed that throughout the year, grassland within the owl
home ranges was used proportionally more than its availability. This is because pastures
and meadows offer a continuous food supply almost year round. Further, in a study of the
Little Owl’s prey in France, Dewulf (personal communication) found that the timing of
grassland mowing had an impact on the availability of beetle prey: the diversity of beetles
was larger if the grass was mowed early, and smaller if mowed later in the season. This
observation was also supported by Grimm (1986) who recommended that grass be mowed
early (end of May) rather than grazed, to obtain the highest diversity of prey.
Access to ground-based prey is negatively correlated with vegetation height. Uncut
meadows could be a “reserve” of food that is simply not available to the owls because of
the vegetation height. Exo (1987) showed that all of the Little Owls studied used crop
fields. For example, they spent 20–40% of their time in the crop fields from August to
February/March. Several unpaired birds had larger areas of crop fields than grasslands in
their home ranges. Sometimes, grasslands have no perches for the owls to hunt from,
which might be a limiting factor for them. Eick (2003) recorded 769 locations of
juvenile Little Owls, and found 45% of their locations to be in mowed grassland and
26% in vineyards. Little Owls can use intensive vineyards, preferring fallows and areas
with short grass. For hunting, of 698 locations of foraging juveniles, 62% were found to
be in orchards, 20% in vineyards, 15% in gardens and 3% in grassland with high grass.
The preponderance of evidence suggests that Little Owls get their food supplies from
different areas within a large diversity of natural and pastoral landscapes having
meadows, pastures with variable grazing intensity, a limited area of crops (because of
pesticides issues) and plenty of edges between grasslands and other types of land use.
No consistent correlations were found by Sunde et al. (2009) between home-range
size and availability of preferred habitat types.

Landscape Heterogeneity and Breeding Success

The amount of high-quality habitat patches, their distribution and the resource acces-
sibility therein play a key role in regulating habitat effects on reproductive success.

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Heterogeneous habitats offer nonsubstitutable resources (e.g., nest sites and food) and
substitutable resources (e.g., different types of food) in close proximity, thereby facili-
tating landscape complementation and supplementation. To better understand how
spatial resource separation in homogeneous agricultural landscapes affects reproductive
success, Michel et al. (2017) investigated the relationships between farmland hetero-
geneity and Little Owl reproductive success, including potential indirect effects of the
heterogeneity-dependent home-range size on reproduction. Little Owl home ranges
were related to field heterogeneity in summer and to structural heterogeneity in winter.
Clutch size was correlated with the amount of food-rich habitat close to the nest,
irrespective of female home-range size, suggesting the importance of landscape com-
plementation. Nestling survival was positively correlated with male home-range size,
suggesting the importance of landscape supplementation. At the same time, fledgling
condition was negatively correlated with male home-range size. Michel et al. (2017)
stated that decreasing farmland heterogeneity constrains population productivity by
two processes: increasing separation of food resources from nest or roost sites results in
low landscape complementation, and reduction of alternative food resources limits
landscape supplementation. Their results suggest that structural heterogeneity affects
landscape complementation, whereas the heterogeneity and management of farmland
fields affect landscape supplementation. Thus, to what extent a reduction in the
heterogeneity within agricultural landscapes results in species-specific habitat degrad-
ation depends on the ecological processes (i.e., landscape complementation or supple-
mentation) that are affected.

10.3.8 Density Dependence

Some processes might be driven by local population density. The higher the density,
the fiercer the competition might be and the higher the impact on the breeding success
(Newton 1998). In Chapter 6, we gave an overview of the variability in observed
densities through the number of pairs/calling individuals per km² and nearest-neighbor
distances. Where habitats are not disturbed, most bird species remain relatively stable in
numbers over longer periods. Their breeding numbers fluctuate moderately from one
year to another, mostly between limits that are smaller compared to what their birth
and death rates would allow (Lack 1954).
It seems that relatively few Little Owl populations live in habitats not subject to
anthropogenic activities. Exo (1992) looked at the role of nest-site availability in
limiting population densities of Little Owls, to investigate how the provision of nest-
boxes could partially compensate for the loss of natural sites. The results illustrated that
the population density increased until the carrying capacity was met, then other
controlling factors become more prevalent, and after three to five years the growth rate
diminished. Similarly, evidence for a strong increase in population numbers after the
introduction of additional artificial nesting cavities has been illustrated in other loca-
tions (Van Nieuwenhuyse and Nollet 1991, Bultot et al. 2001, C. Stange personal
communication). On the other hand, in some populations that are more severely
impacted by anthropogenic factors, such as the Little Owl population in the

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 10.3 Demographic Limiting Factors

Northern Vosges (France), the addition of nestboxes has served to slow the rate of
population decline, and helped to support the few small population clusters that
remain (Génot 2001).
More detailed observations on density dependence were recorded by Bultot et al.
(2001), as part of the nestbox provisioning project in Wallonia, Belgium. By comparing
isolated pairs and clustered pairs, it was found that owls that nested as isolated pairs
produced an average of one more egg per breeding season than the owl pairs that nested
in clusters. However, for owl pairs that nested as part of clusters, density dependence was
found to affect the timing of egg-laying – the higher the owl density, the earlier the
laying date became. This might indicate that social aspects become important at higher
densities, reducing the number of eggs and/or changing the timing of egg-laying. Density
dependence in any demographic parameter, whether births or deaths, immigration or
emigration, can be caused by competition for resources, such as food, nest sites or territor-
ial space. Natural enemies can also be a cause if predators kill an increasing proportion of
prey individuals as prey density rises, or if parasites infect a greater proportion of host
individuals as host density rises. In any event, the lines of evidence from the provision of
nestboxes have helped to illuminate the onset of density-dependent regulatory affects.
Since the densities of the Little Owl population may themselves influence the Little
Owl in its reproductive success, habitat selection and dispersal behavior have been
studied together. In Flanders (Belgium), the vicinity of conspecifics in surrounding grid
cells proved to be a better predictor of occupation for a given grid cell than the
landscape composition (Van Nieuwenhuyse and Bekaert 2002). These results suggest
that correlations of habitat and demographics be examined and interpreted in light of
this density-dependent perspective. Van Horne (1983) identified social interactions
within wildlife populations as a potential habitat classification issue. In the structure
of some populations, dominant breeding animals excluded more numerous, submissive
or nonbreeding animals from the highest-quality habitat. Habitat classification based
solely on aural surveys may result in a model that identifies suboptimal habitat as
critical, and exclude optimal habitat (Van Horne 1983). Protecting only suboptimal
habitats would negatively influence the breeding success and overall stability of the
population (Christie and Van Woudenberg 1997).

10.3.9 Metapopulation
A metapopulation is a spatial distribution of distinct subpopulations, separated by large
distances or barriers and connected by dispersal movements. This patchy population
pattern changes over time (Opdam 1991, Hanski 1999). At high densities, the spatial
distribution pattern of Little Owls remains clustered, with the “saturation level” of owls
determined by the territorial behavior of the species, consisting of regularly spaced
calling males (Bretagnolles et al. 2001, Van Nieuwenhuyse et al. 2001e). At this point,
the variance in territoriality and subsequent increase in population density (Fuchs
1986) appears to stress the importance of social aspects over landscape conditions.
More and more, Little Owl populations in western Europe are functioning like
metapopulations (rather than a consistently interactive regional population), as the

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fragmentation of the habitat results in localized population declines and areas with very
low owl densities. In the Northern Vosges (France) Génot has studied a metapopulation
since 1984 (Génot 2001). It is composed of three clusters of owls (Figure 10.17), with
distances of 18 km between clusters A and B, 33 km between A and C, and 36 km
between B and C. These three subpopulations had about 10–15, 5–10 and 5–10 pairs of
Little Owls, respectively. A semi-barrier of deep forest separated the cluster C from A and
B. Arrows in Figure 10.17 show the movements of ringed birds between the different
clusters. The mean distance of young dispersal was 6 km (n = 15). Cluster D was
occupied in 1984–1987 but had since disappeared. A population viability analysis
applied to this metapopulation revealed a high exctinction risk for population sizes
below five breeding pairs (Letty et al. 2001a). A difference between the baseline model,
which indicated a continuous population decline, and the field data showed that some
birds from outside these population clusters immigrated into and supplemented the
local stock. In particular Little Owls from the Sarre region of Germany, and areas
adjacent to the Northern Vosges, flew across the area to supplement owls in cluster
C (see Eick 2003) and also clusters A and B (Figure 10.17). The sustainability of these
population clusters can therefore only be explained by a strong connection with
German populations, demonstrating some sink–source dynamics. Both small
local population sizes and low observed genetic structure can only be compensated

Figure 10.17 Subpopulations and immigration of Little Owls in the Northern Vosges, France (after
Génot 2001).

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 10.3 Demographic Limiting Factors

by a considerable immigration rate (more than two immigrants per year per
population cluster, representing 17% of the current local population) (Bouchy 2004).
In conservation biology theory, this degree of immigration support is called the “rescue
effect.”
The results from Eick (2003) show much larger dispersal distances of juveniles than
previously known. New insights into Little Owl dispersal activities through radio-
telemetry studies have shown that more dynamic juveniles undertake exploratory
excursions of three to five days in all directions up to 40 km in a single night before
coming back to their birthplace. They even cross large forests, flying at altitudes of
20–30 m (or more) above the ground. Little Owl dispersal behavior allows exchange of
individuals between population clusters previously considered to be isolated. In
Groningen (Netherlands), a similar metapopulation pattern had developed due to the
increasing isolation and fragmentation of the population (van’t Hoff 2001). In 2000,
three population clusters remained in Groningen: Oosterwijtwerd in the east with six
pairs, Oldehove in the west with five pairs and Peize (north-Drenthe) with 11 pairs
(J. van’t Hoff personal communication). The distances between the different clusters
were Oosterwijtwerd to Oldehove 27 km, Oosterwijtwerd to Peize 28 km, and
Oldehove to Peizenorth 13 km. The average nearest-neighbor distance between owl
pairs within each cluster was 1494 m (Oldehove), 1140 m (Oosterwijtwerd) and
763 m (Peize).
In Groningen, isolated pairs declined much faster than those in remaining popula-
tion clusters (Figure 10.18). The vicinity of the last population clusters in Groningen
seemed more attractive to owls than the actual quality of the habitats. Most recently
abandoned (and isolated) territories were situated in apparently optimal habitat, while

40

35 Isolated territories

30 Clustered territories
Number of Territories

25

20

15

10

0
1994 1995 1996 1997 1998 1999 2000 2001 2002 2003
Year

Figure 10.18 Trend in the number of breeding pairs or territories of Little Owls in Groningen,
between 1995 and 2002, for isolated and clustered pairs in Oosterwijtwerd and Oldehove
(Netherlands) (Van’t Hoff 2001).

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 Chapter 10: Population Regulation

the population clusters remained in suboptimal habitat. A key value of these vacant
“habitat isolates” lies in their role for a future recovering population.

10.3.10 Population Viability Analysis

When a bird population is facing extinction, ecologically artificial conservation actions
such as feeding and captive breeding may be considered as temporary efforts to rescue
the remaining birds until environmental conditions for a self-sustaining population
have been restored. Population viability analysis can be used to evaluate different
management options for endangered owl populations. Andersen et al. (2015) used
the program VORTEX to explore different management strategies using the Danish
Little Owl population, which is declining mainly due to lack of food during the
breeding. Four scenarios were run, simulating 25 years of population dynamics, i.e.,
the “do nothing” scenario, the captive breeding scenario, where individuals are supple-
mented to the population, the food supplementation or habitat improvement scenario
and the scenario combining captive breeding and food supplementation/habitat
improvements. For each scenario 1000 iterations were run with the literature-based
estimates given in Table 10.13. In scenarios where no management actions were taken
the population went extinct within 12 years. When supplementing individuals con-
tinuously, the population remained extant but the population size remained small. If
captive breeding was initiated without habitat improvements or artificial feeding, the
Danish population would likely become a population sink and never become self-
sustaining. Food supplementation/habitat improvements could restore the population,
although there must be capacity to secure food/habitat for a minimum of 100 individ-
uals to minimize genetic losses. A captive breeding program had to be maintained for
about 10 years in order to secure a viable wild population. By combining food supple-
mentation and the release of captive-bred individuals, the population had a chance of
being restored and become independent of human aid. Food supplementation must be
considered a temporary solution to keep the population alive while awaiting
habitat restoration.
It is increasingly important to estimate both the number of individuals it takes to
create a genetically viable population and the population trend. Andersen et al. (2017)
calibrated population viability analysis and habitat suitability modeling to estimate
population viability and future prospects of the Little Owl across Europe. Models were
made for Italy, the Balkans, Spain, Portugal and the Netherlands using the respective
parameters given in Table 10.13. The results showed a high risk of population declines
over the coming 100 years, especially towards the north of Europe, whereas populations
towards the southeastern part of Europe have a greater probability of persistence. In
order to be considered genetically viable, individual populations must count
1000–30 000 individuals. As Little Owl populations of several countries count
<30 000, and many isolated populations in northern Europe count <1000 individuals,
management actions resulting in exchange of individuals between populations or even
countries are probably necessary to prevent losing <1% genetic diversity over a 100-
year period. At a continental scale Anderson et al. (2017) suggest, with their habitat

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Table 10.13 Estimated parameters used in VORTEX simulations

Estimateb
a a
Parameter Estimate References IT, Balkan, ES, PT, NL Referencesb

Number of iterations 1000 10

Adult mortality (aged 1+) 32% (SD 10) Thorup et al. 2013 35%, 35%, 35%, 35%, Van Nieuwenhuyse et al.
36.7% 2008, Thorup et al. 2010
Juvenile mortality (aged 0–1) 85% (SD 10) Thorup et al. 2013 70%, 70%, 70%, 70%, Thorup et al. 2013
80%
Environmental correlations in 0.5 Default value 5 (adult), 10 (juvenile)
mortality rates
Mating structure Monogamous Van Nieuwenhuyse et al. Short-term Van Nieuwenhuyse
2008, Thorup et al. 2010 monogamous et al. 2008
Breeding age 1 year Thorup et al. 2010 1 year Thorup et al. 2010
Maximum age of reproduction 11 years Calculations based on 15 years Van Nieuwenhuyse et al.
formula in Lande 1988 2008
Density dependency No Van Nieuwenhuyse et al. Yes Van Nieuwenhuyse
2008 et al. 2008
Mean no. of fledglings/brood 2.12 Jacobsen 2006, Thorup et al. 4.64, 5.24, 4.4, 3.3, Jacobsen 2006, Thorup et al.
2010 3.78 2010
SD (mean no. fledglings) 1.15 Jacobsen 2006 1.0, 1.0, 1.0, 1.2, 1.0
Maximum no. of progeny 8 Jacobsen 2006 10, 7, 10, 5, 10 Van Nieuwenhuyse et al.
2008
Ratio of breeding pairs 66% Jacobsen 2006 85% Tomé et al. 2004, Jacobsen
successful in getting 2006
fledglings
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478

Table 10.13 (cont.)

Estimateb
a a
Parameter Estimate References IT, Balkan, ES, PT, NL Referencesb

Number of breeding attempts 1 Van Nieuwenhuyse et al.

per year 2008
Sex ratio at birth (male:female) 50:50 Thorup et al. 2010 50:50 Thorup et al. 2010
No. of males in breeding pool 100% Assume all attempt to breed 100% Assume all attempt to breed
No. of females in breeding 100% Assume all attempt to breed 100% Assume all attempt to breed
pool
Min age of dispersal 1 year Pedersen et al. 2013 1 year Pedersen et al. 2013
Max age of dispersal 11 years Calculations based on 3
formula in Lande 1988
Probability of dispersal 5% Sunde et al. 2009 5%
Dispersing sex Both Van Nieuwenhuyse et al. Both (70% survive
2008 dispersal)
Population size 120 indiviuals Eskildsen and Vikstrøm 2011 Variable in order to find
MVP
Catastrophe 1: Cold winter 5% occurrence; Poulsen 1940, Poulsen 1957 Rate of 5%; survival Poulsen 1940, Poulsen 1957
mortality 75% 75% of normal
of normal
Catastrophe 2: High rainfall Rate of 5%; Bultot et al. 2001, Van
reproduction 75% of Nieuwenhuyse et al. 2008
normal
Dispersal is not included in the “do nothing” and captive breeding scenarios.
a
Andersen et al. 2015
b
Andersen et al. 2017
 10.3 Demographic Limiting Factors

suitability analysis, that the Little Owl would be affected positively by increasing
temperatures and urban areas, whereas an increase in tree cover, annual rainfall,
grassland and sparsely vegetated areas would affect the presence of the owl negatively.
However, the low predictive power of the habitat suitability model suggests that habitat
suitability might be better explained at a smaller scale. Furthermore, in the simulations,
the European population was considered as a complex of inter-connected populations
and subpopulations of which some might persist as true metapopulations, hypothesiz-
ing the relevant use of the method for population estimation. The results might be
over-optimistic with regards to population size and overstate the role of dispersal and
genetic exchange between de facto populations.

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 Chapter 11: Management
and Conservation

Chapter Summary
We open this chapter with a brief overview of the status of and threats to the Little Owl.
We then offer a conservation strategy for the owl that involves five critical success
factors: Knowledge, Limiting Factors, Evolution of Landscape Conditions, Legislation and
Policies, and People. Thereafter, we describe four main drivers to implement this strategy,
focused on Monitoring, Management, Standardized Methodologies and Data Management.
The long-term conservation of the Little Owl is complicated, as the species is largely
linked to an agriculturally dominated landscape. The condition of this landscape can
change rapidly and significantly due to human demographics, changes in policies and
management. The conservation strategy described in this chapter requires a multiscale,
multidisciplinary approach, with collaboration between different stakeholders (conser-
vationists, scientists, different authorities, farmers) and additional research into the
ecology of the species (Chapter 12 Research Priorities). This strategy must be applied
at different levels: local, regional, national and international. We encourage people
involved in this conservation strategy to work broadly, openly and to freely co-ordinate
on issues, data and management efforts that will benefit the broader array of species and
environments of which the Little Owl is a part.

11.1 Conservation Status and Threats

In Europe, the Little Owl is considered to have a threat status of Least Concern (IUCN).
The species is protected by the EU Birds Directive (Directive 2009/147/EC of the European
Parliament and of the Council, of 30 November 2009, on the Conservation of Wild Birds), by
the Bern Convention (Convention on the Conservation of European Wildlife and Natural
Habitats) and the EU regulation on the trade of fauna and flora (Commission Regulation
(EU) No 1320/2014, of 1 December 2014, amending Council Regulation (EC) No 338/97 on
the Protection of Species of Wild Fauna and Flora by Regulating Trade Therein).
The status of the population at the EU level was evaluated at the species level; this
was based on reports delivered by Member States under Article 12 of the Birds Directive

Supplementary resources are available online at www.cambridge.org/littleowl. These include

appendices as well as tables which previously appeared in the first edition (prefixed with ‘S’).

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 11.2 Conservation Strategy for the Little Owl

(European Environment Agency 2021). The European Union (EU28) Red List assess-
ments are based principally on these official data reported by EU Member States. All
Member States are requested to monitor bird species and send a report every six years.
This information includes population sizes and trends (short and long term) for breeding
and wintering populations, as well as pressures and threats for Special Protection Area
trigger species. In addition, population status and trends are assessed at the EU level.
The reporting period 2013–2018 data was used by the European Environment Agency
(2021). Detailed information about reporting under Article 12 of the Habitats Directive
can be obtained from www.eionet.europa.eu/etcs/etc-bd/activities/reporting/article-12/.
The reported data as well as the assessments of status and trends at the EU level can be
visualized using the web tool http://bd.eionet.europa.eu/article12/. The reporting period
2006–2012 data were used by BirdLife International (2015) and Keller et al. (2020).
The threats to the Little Owl are mainly due to changes in human land-use practices.
Some threats are becoming more prominent across the European range of the Little
Owl, such as the reduction in the amount of tree lines, the deterioration of high-stem
orchards and the increase in the area of subsidized maize. The Little Owl is also
negatively impacted by collisions with vehicles on roadways and by pesticides that
reduce invertebrate prey.
Populations can disappear because of low density, lack of immigration, and
fragmentation of habitat and populations (Génot 1992c, van’t Hoff 2001). More and
more Little Owls near villages in central Europe are threatened by building activities
without consideration of nature conservation in land-use plans (Breuer 1998). In Iran,
human migration towards cities offers many opportunities for the species in abandoned
rural dwellings (Ilanloo et al. 2020). Climate change is expected to influence the
distribution range of the species. Throughout the rest of the breeding range, little is
known about the current population levels, threats or long-term trends.

11.2 Conservation Strategy for the Little Owl

We propose a conservation strategy featuring five critical success factors: Knowledge,
Understanding of Limiting Factors, Evolution of Landscape Conditions, Legislation and
Policies, and People. We will first characterize the components and follow with a review
of the drivers that are essential in implementing these components.

11.2.1 Five Critical Success Factors for the

Conservation Strategy
11.2.1.1 KNOWLEDGE
One of the first steps in developing a practical and cost-effective conservation program
is to identify the current status of knowledge on the species and define the gaps in the
key scientific aspects (Tucker and Evans 1997). We will update the body of knowledge
offered by Van Nieuwenhuyse, Génot and Johnson (2008).
To assist in examining the current state of knowledge (and knowledge gaps) on the
species, we offer a summary of the Little Owl literature, through time, by geography and

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 Chapter 11: Management and Conservation

by subject area. Importantly, we are making the Bibliography of the Little Owl Athene noctua
available online (www.cambridge.org/littleowl). The timeline of the Bibliography is for
publications from 1495 through to early 2020. This Bibliography (n = 3444 references)
represents all of the Little Owl literature of which we are currently aware; we welcome
any additions to this literature base, and apologize for any missed citations.
Most of the references appear after the 1960s. The relative number of publications
since this time is 2951, and 2558 of these come from the period 1980–2020
(Figure 11.1). Most of the publications from 1880–1940 come from Britain, and are
associated with the status of the Little Owl after its introduction there at the end of the
nineteenth century.
Regarding the geographic distribution of the Little Owl literature, a total of 3045 titles
have been assigned to geographic areas (Figure 11.2), with 2805 of the references concerning
Europe. This can be explained partly by the distribution area of the species, but mostly by a
readily accessible European literature base. Western Europe (France, Germany, Belgium, the
Netherlands, Britain, Switzerland) accounts for 2526 references, Mediterranean Europe
(Spain, Italy, Greece, Crete, Cyprus) 133, with eastern Europe (former Soviet Union,
Poland, Czech Republic, Hungary, Romania) reflecting 401 references (Figure 11.3).

DECADES
1880–1900, 76, 2% 1910, 83, 2%
<1880, 45, 1%
2020, 89, 3% 1920, 67, 2%
1930, 77, 2%
1940, 66, 2%
2010, 679, 20%
1950, 74, 2%

1960, 132, 4%

1970, 261, 8%

2000, 745, 22% 1980, 482, 14%

1990, 563, 16%

Figure 11.1 Temporal distribution of the Little Owl literature (1495–2022).

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 11.2 Conservation Strategy for the Little Owl

CONTINENTS
Africa, 55, 2% Asia, 71, 2%
Middle East, 36, 1%
World, 18, 1%

Europe, 2865, 94%

Figure 11.2 Geographic distribution of the Little Owl literature across the range of the species.

TOP12
Czech Republic, 66,
2% Hungary, 64, 2%
Poland, 72, 3%
Spain, 92, 4%
Germany, 642, 25%
Russia, 93, 4%

Belgium, 128, 5%

Switzerland, 175,
7%

Italy, 204, 8%

France, 447, 18%

Netherlands, 270,
11% United Kingdom,
273, 11%

Figure 11.3 Geographic distribution of the Little Owl literature in Europe.

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 Chapter 11: Management and Conservation

M
G
T
C
P
C
R V
B P
G
M I
0.05150

D
D C
T H D
R
B
C A
– N M
C M

– P
P

– D T
S

–
P
–
– – – – – – Row Coords.
0.11009 Col. Coords.

Figure 11.4 Topic–decade correspondence analysis of Little Owl literature associations.

The majority of publications come from countries where the species was historically
declining (France, Germany, Switzerland, Belgium, Netherlands, Austria, Luxembourg).
This is confirmed by the chronology of studies, whose 2951 references appear after
1960, which is the beginning of the Little Owl population decline across Europe.
A clear gradient can be observed in the correspondence analysis output of counts of
papers and the topics they handle versus the decade (Figure 11.4), from more descrip-
tive research on morphology (1960), behavior (1970), mortality, distribution and
threats (1980), to reproduction, nestboxes, population dynamics, densities and surveys
(1990), telemetry and conservation (2000), dispersal and climate (2010) towards intra-
guild interactions and genetics (2020).
As for topics in relation to geography, a gradient is shown in Figure 11.5 from
dominance of the UK (1960), Bulgaria and Romania (1970), Austria, Germany and
Luxembourg (1980), France, Greece and Russia (1990), China, Italy, Poland and the
Netherlands (2000), Slovakia, Belgium and Switzerland (2010) towards Denmark,
Hungary and Turkey (2020).
As for typical topics (Figure 11.6), diet is predominant in Slovakia, Algeria, Turkey
and the Balkans; Poland, Hungary and Slovenia typically address population estimates,
Belgium, the Netherlands, Switzerland and Germany are mainly focused on population

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 11.2 Conservation Strategy for the Little Owl

0.8

0.6

0.4

0.2

0.0
0.11188

–0.2

–0.4

–0.6

–0.8

–1.0

–1.2

–1.4
–1.2 –1.0 –0.8 –0.6 –0.4 –0.2 0.0 0.2 0.4 0.6 0.8 1.0 1.2 Row Coords.
0.16002 Col. Coords.

Figure 11.5 Country–decade correspondence analysis of Little Owl literature associations.

G
M

P T
S D

D
H Vocalization
P
B C
N A
M R
R
T
C
–
D
P D
P
B

–
– – – Row Coords.
Col. Coords.

Figure 11.6 Country–topic correspondence analysis of Little Owl literature associations.

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 Chapter 11: Management and Conservation

dynamics, breeding biology and conservation, France and the Czech Republic concen-
trate on surveys, mortality and distribution, while the UK is associated with behavior
and Italy and Spain with contamination, parasites and taxonomy.

11.2.1.2 LIMITING FACTORS

Essential criteria for suitable Little Owl habitat are year-round prey availability, prey
accessibility, vertical landscape structures with cavities (for nesting and roosting, with
multiple entrances) and limited predation pressure (see Chapter 6). We recommend
management activities that increase the availability and long-term replacement of
nesting cavities, provide better availability and accessibility to prey, and offer other
habitat conditions that serve to limit the impact of predators. Finally, before we can get
a clear picture of the relative importance of the different factors that influence the
population dynamics of the species (Exo 1992) in a given region, we need reference data
on these key resources and aspects.

Food Availability
Improving the amount of prey for the Little Owl can be done through different
management activities that increase habitat patch heterogeneity by reducing the aver-
age parcel size (ha). Grimm (1986) found larger and more numerous prey in a meadow
edge beside a brook than in grasslands. The trampling of vegetation by cattle yielded
more Carabidae beetles, especially near parcel edges (but trampling by cattle has other
potentially habitat-damaging consequences, such as a reduction in growth of cavity-
bearing trees). Tall vegetation is crucial for small mammals, with edges tending to have
larger diversities and densities (Dalbeck et al. 1999a,b). Planting shrubs, hedges and tree
lines serves to promote the abundance of insects and other prey.
If we want to study the food availability and its possible impact on owl population
numbers, we need more information on the prey choice of the owls, as well as the
variance of those prey in space and time. An international database of small-mammal
distributions and especially densities is needed, as is better data on the relative abun-
dance of earthworms Lumbricidae and key beetle species. Standardized measures for vole
abundancy are needed.

Food Accessibility
Being principally a perch-and-pounce hunter in southern regions (Fajardo et al. 1998,
Tomé 2011), the availability of perches is fundamental to Little Owls. A substantial
portion of perches are provided in anthropogenic landscapes by fences and fence poles.
Tempero-spatial rotation in mowing activities can create gradients between tall and
short vegetation, and offers favorable conditions for the owls to access prey. Hypothesis
testing that local habitat conditions can be optimized by a combination of artificial
perches and phased mowing to increase prey accessibility is crucial (similar to shrikes)
(see Yosef 1993, Van Nieuwenhuyse et al. 1995).

Nesting and Roosting Cavities

The provision of nestboxes or other cavities, while labor intensive, has been shown to
be a successful short-term solution to the lack of nesting and roosting cavities in

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 11.2 Conservation Strategy for the Little Owl

different regions (Schwarzenberg 1970, Knötzsch 1978, Loske 1978, Furrington 1979,
Schönn 1980, 1986, Schönn et al. 1991, Exo 1992, Haase 1993, Bultot et al. 2001, Bock
et al. 2013, Grüebler et al. 2013, 2014). Nestboxes have been readily accepted, even
when a minimal owl population remains in a given area. In some cases researchers have
found that nestboxes that were newly installed at the beginning of April were occupied
after only two weeks, indicating that nest cavities were a limiting factor for the floater
owls that were otherwise incapable of nesting (Bultot et al. 2001). Nesting and roosting
sites can be created for the short term by enhancing existing previously unsuitable
cavities, by modifying buildings to provide holes and crevices for owls, or by hanging
nestboxes. In the longer term, cavities can be offered through planting of new trees and
managing existing trees (fruit trees, pollard willows). New research dealing with the
characteristics of multientrance nestboxes and roosts, and experiments with newly
installed nestboxes just before the start of the breeding season might offer insights into
the shortage of nesting cavities and surplus of breeding pairs.

Natural Cavities
More suitable sites can be created by the enhancement of natural cavities. Cyclic
management of coppiced willows by volunteers offers the best guarantee for a stable-
amount of suitable sites. This method is preferred since the trees also offer opportunities
for prey species and can act as hunting perches too. Importantly, provision of these
types of nesting structures is the most natural and cost-effective.

Buildings
Across large parts of its breeding range, the Little Owl occupies buildings. Hence,
modifying barns, chapels and dilapidated structures by adding suitable holes in the
walls or under roofs is easy and sustainable. Furthermore, it also offers good opportun-
ities for positive interactions with people to help the species due to its association
with humans.

Nestboxes
We wish to stress that the last resort to increase the number of nest sites is through the
installation of nestboxes. Nestboxes should be avoided whenever possible since preda-
tors such as Stone Martens easily recognize the boxes and remember their locations
during their hunting rounds. This is especially the case when larger numbers of a
standardized type of nestbox are used (Kirchberger 1988). Importantly, nestboxes are
often not available for Little Owls due to use by other animals such as Starlings Sturnus
vulgaris, Tits Parus spp., Tree Sparrows Passer montanus, Dormice Glis glis, Garden
Dormice Eliomys quercinus, hornets, bees, wasps and bumble-bees, and House Mice
Mus musculus (Pitzer 1995). Thus, before installing any nestboxes, the initial population
should be determined through a standardized survey method. This will provide specific
insights into what extent the owl population may actually need newly offered nesting
places. Furthermore, a clear view of all available nestboxes in the study area should be
obtained and communicated during this inventory. Nestboxes that are not known to
researchers prior to efforts can influence the interpretation of results. When installing

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 Chapter 11: Management and Conservation

nestboxes, we advise to do so in both seemingly suitable and unsuitable habitats and to

study the difference in use by owls in both habitats.
Installing nestboxes should be seen as creating excellent opportunities for research.
Sampling of the species is made very easy through the use of nestboxes. We consider
follow-up monitoring of the response of owls to the addition of nestboxes as manda-
tory. It is important for researchers to understand that population trends produced by
nestbox studies alone are not always reliable (Illner 1990).

Predators
The acrobatic skills of Stone Martens have been illustrated by Marié and Leysen (2001)
and possible measures to avoid predation of Little Owls by martens were suggested. To
get a better assessment of the impact of predators, we are very supportive of co-
ordination with existing Stone Marten (Martes foina) monitoring projects. These types
of investigations/co-ordination efforts are important, given the population increase in
marten species in Europe (Van den Berghe 1998) and its possible impact on areas with
low populations of Little Owls.

11.2.1.3 EVOLUTION OF LANDSCAPE CONDITIONS

Across its global range, the Little Owl is closely associated with agricultural landscapes
(Tucker and Evans 1997). The impact of changes to what was likely a largely
pastoral landscape is unclear due to a lack of data from a historical baseline condition.
However, as illustrated by van’t Hoff (2001) in the Netherlands, this kind of data
is crucial in the study of the causes of population trends. The Flemish Little Owl
Project (Van Nieuwenhuyse, Leysen and Leysen 2001d) paid much attention to the
set-up of a referential database of soil, land-use and biological quality data, and roads,
rivers and related spatial landscape conditions. This reference database will be used to
follow changes in landscape in a digital and exploitable way, and made available.
Contacts and co-operation between conservation groups and official organizations,
e.g., Intra Eco Network Europe (IENE), need to be established to jointly and more
efficiently assess the impact of habitat fragmentation on birds by roads and other
societal infra-structure.
So far all habitat selection studies have been carried out in anthropogenic land-
scapes. New research on habitats is needed in natural areas like deserts (northern Africa)
and steppes (Kazakhstan) and at higher altitudes (Tibet, India).

11.2.1.4 LEGISLATION AND POLICIES

An excellent overview of the opportunities for conserving the wider environment, and
indirectly the Little Owl, through international and European legislation is given in
Tucker and Evans (1997). An important challenge that remains is to analyze these
different instruments in more detail and to translate the opportunities into a broader
international context for the conservation of the Little Owl and its habitat.
In Belgium, legislation on nature conservation has evolved from a species-protection
scheme for “useful” species towards a more territory-directed conservation strategy, e.g.,
Natura 2000 (http://europa.eu.int/comm/environment/nature/home.htm). In recent

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 11.2 Conservation Strategy for the Little Owl

years, more attention has been directed at species protection, again due to the formal
adoption of the Communication on a European Biodiversity Strategy, adopted on
February 4, 1998 (http://europa.eu.int/comm/environment/docum/9842sm.htm).
With this strategy, the EU re-inforced its role in finding solutions for biodiversity within
the framework of the United Nations’ Convention on Biological Diversity (also called
the Biodiversity Treaty, see www.ciesin.org/TG/PI/TREATY/bio.html).
In Flanders, the Little Owl is a protected species through the European Bird
Guideline 79/409 (Royal Decree of 9/9/1981 changed by decree of the Flemish
Government on November 20, 1985). Since 1998, the Decree on Nature Conservation
(Belgisch Staatsblad January 10, 1998) offers different opportunities for Little Owl
conservation. The fundamentals of the decree are species protection, territory-directed
conservation, general basic ecological quality and a focused tackling of species groups of
interest. The implementation of Little Owl conservation measures can be done through
management contracts as mentioned in European Union Ordinance 2078/92. With
these contracts, farmers can get paid to create or maintain certain habitat elements in
the landscape (e.g., wooded banks, pollard willows, ponds, field edges). These contracts
are essential for safeguarding viable breeding places for the Little Owl. Despite their
availability, there is no systematic use of management contracts. Additional instru-
ments in Flanders include financial subsidies for the planting and maintenance of
pollard willows, high-stem orchards, wooded banks and other small landscape elements
in most communities through the GNOPs (Gemeentelijk Natuur Ontwikkelings Plans,
or communal nature development plans). Despite some good and pragmatic initiatives,
there remains a huge gap in the use of economic and policy mechanisms and their
conservation impact on the species (Tucker and Evans 1997) in most parts of Europe,
let alone the rest of the Little Owls’ distributional range.
In Luxembourg, the Règlement Grand-Ducal of March 22, 2002 (Règlement de
biodiversité), regulating a set of measures for the conservation of biodiversity, was
made official on April 4, 2002. This law provides funds and the opportunity to take
measures for the protection of rare and/or endangered species. The costs of actions
especially helpful to the Little Owl, like nestboxes or planting fruit trees around villages
are 90% covered by the Ministry of Environment.
In Switzerland a conservation plan has been set up in the canton of Genève since
2012, with clear objectives, positive results and sustained follow-up (Meisser 2013,
2014, 2015) and later extended to the entire country (Meisser 2016, 2017, 2018,
2019, 2020, 2021, 2022) dealing with habitat improvements, co-ordination of local
conservation projects, follow-up of the effectiveness of the measures, bringing together
different stakeholders and solving conflicts, targeting communication and improving
understanding of the species.

11.2.1.5 PEOPLE
People are another key factor in any proposed Little Owl Conservation Strategy. Up to
75% of the European Little Owl population occurs in agricultural and grassland habitats
(Tucker and Evans 1997). Especially for this species, environmental protection and
biodiversity conservation should be an integral part of all uses of the environment,

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and of all policies of all socio-economic sectors (agriculture, forestry, tourism). In the
longer term, sustainable use of the environment will require substantial changes in
society’s use of natural resources, energy and transport. It is therefore essential that the
public is aware of the implications of the current and increasing intensity of its use of
the environment, and of the fact that everyone has a role to play in the conservation of
biodiversity in the wider environment (Tucker and Evans 1997). The Little Owl offers
an excellent model to implement this in Europe because of is positive emotional impact
on humans. We propose the species become a flagship for well-managed agricultural
landscapes in Europe.
Specific action plans should be made to educate and influence policy-makers, land-
owners, land-users, conservationists and children.
The International Little Owl Working Group (ILOWG) first emerged in 1999, with the
aims of increasing communication on Little Owl research, conservation and education
across Europe. A great deal of work had been undertaken by individuals, many of whom
began to develop nestbox schemes, tree management initiatives, research and monitor-
ing programs. A great deal of information had been generated by these programs, and
all that was needed was a mechanism for sharing the information, hence the concep-
tion of the ILOWG. A group developed to readily support communications to out-
posted owl workers so that they were able to tap into a mine of experience from the
ILOWG community. The success of this truly international family reflects a member-
ship of over 60 members from 11 countries. This took the level of exchange of infor-
mation and co-operative fieldwork up to a new level. Since then six symposiums have
been organized that involved national and international presentations, and discussions
regarding standardizing methodologies, generating great interest from across Europe,
and owl workers returned home full of ideas and energized about developing more
initiatives for Little Owls in their regions. Importantly, the participants were eager to
contribute to the larger program and to understand and share in the conservation
priorities for Little Owls across Europe.
The future of Little Owls is in our hands. Within the remit of fiscal agri-ecology
schemes we need to develop methods to inform farmers and landowners how to best
manage the land to benefit Little Owls.
We need to understand the populations and the causes of declines better, so we can
be more prescriptive with the conservation measures we apply.
The population data will serve to add strength to lobby for the inclusion of the Little
Owl as a priority agri-environment target species. This will help promote the needs of
the owl, and lead to the development of national action plans.
The role of the voluntary Little Owl fieldworker has never been more important.
Individually we need to continue to develop our conservation programs, advise farmers
and landowners, monitor nest sites, manage and plant more trees, educate and take the
owl to the people.
More international co-operation can be achieved by continuous sharing of experi-
ences, developing best practice methods, developing workshops on key monitoring
skills and by holding regular symposiums. The key to achieving these points is commu-
nication and sharing information, which is the foundation of the ILOWG.

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 11.2 Conservation Strategy for the Little Owl

This volume is a testimony to all voluntary fieldworkers and conservationists who

have freely given their time and energy; their reward, to see the owl that they love and
cherish continue to survive and flourish.

11.2.2 Implementing the Strategy: Four Drivers

The five components described above reflect the overarching initiatives that embody a
Little Owl Conservation Strategy. The four drivers that we propose in this section are
specific measures to implement the initiatives: Monitoring, Management, Standardizing
Methodologies and Data Management.

11.2.2.1 MONITORING
The overall goal of natural resource monitoring is to develop scientifically sound infor-
mation on the current status and long-term trends in the composition, structure and
function of ecosystems, and to determine how well current management practices are
sustaining those ecosystems. The purpose of the Little Owl Monitoring Plan is to assess
trends in owl populations and habitat. Monitoring data will be used to evaluate the
success of the various conservation measures in arresting the downward trends in some
owl populations and in assessing the habitat conditions necessary to support viable owl
populations throughout the range of the owl.
A survey (or inventory) is an extensive point-in-time effort to determine the location
or condition of a resource, including the presence, class, distribution and status of
plants, animals and abiotic components such as water, soils, landforms and climate.
Surveys provide information on the distribution, abundance, habitat associations and
primary ecological aspects of species – information required for developing sound
conservation strategies. Monitoring differs from surveys or inventories in adding the
dimension of time, and the general purpose of monitoring programs is to illuminate
population trends, identify specific aspects needed for conservation action and evaluate
the effectiveness of current management programs. Survey protocols and monitoring
programs are necessary to ensure that changes detected by fieldworkers are actually
occurring in nature and not simply a result of measurements taken by different people
or in slightly different ways.
Elzinga et al. (1998) defined monitoring as “The collection and analysis of repeated
observations or measurements to evaluate changes in condition and progress toward
meeting a management objective.” Monitoring objectives should include six compon-
ents to be completed (Elzinga et al. 1998): the indicator or “vital sign” to be monitored,
the location or geographical area, the attribute of the indicator to be measured (e.g.,
population size, density, percentage cover), the intended management action (increase,
decrease, maintain), the measurable state or degree of change for the attribute and the
timeframe. Monitoring data are most useful when the same methods are used to collect
data at the same locations over a long time period (e.g., more than 10–12 years). It is
important to note that cause-and-effect relationships usually cannot be demonstrated
with monitoring data, but monitoring data might suggest a cause-and-effect relation-
ship that can then be investigated with a research study. The key points in the

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definition of monitoring are that: (1) the same methods are used to take measurements
over time, (2) monitoring is done for a specific purpose, usually to determine progress
towards a management objective and (3) some action will be taken based on the results,
even if the action is to maintain the current management.

Monitoring the Species

In our Monitoring Plan, we propose the development of a network of some 30 Vital Sign
Demographic Monitoring Areas, where detailed mark–re-capture studies are employed.
The primary objective of demographic studies is to detect trends in the vital rates of the
species. More specifically, demographic analyses on owls employ mark–re-capture stud-
ies, typically on large identified areas, where repeated surveys are conducted to locate,
mark and re-observe or re-capture pairs of owls and their offspring. Many of the
demographic areas we are recommending are in operation now, and for these, we
suggest formalization of a more detailed and rigorous study design. For some areas
within the range of the species, this would mean the identification of new demographic
study areas. Rather than having a lot of study areas, we propose having fewer, but more
intensively investigated areas, within which some 50–100 pairs of owls (or habitat for
the same) are monitored routinely. Adult survival rates, fecundity, and population and
metapopulation trend analyses would be based upon the work conducted by Franklin
et al. (2004), Forsman et al. (1996) and Ganey et al. (2004). For countries without Vital
Sign demographic study areas, we recommend distributional surveys and
ecological studies.

Monitoring the Habitat

Van’t Hoff (2001) offered interesting insights from the historical data on landscape and
Little Owl distributions in Groningen, the Netherlands. Unfortunately few historical
landscape data are available in digital formats. Acquiring a digital version of the
landscape information (e.g., satellite images) at regular intervals of time (say, every 10
years) is extremely important because we can then monitor the changes in landscape
variables important to Little Owl populations. Van Nieuwenhuyse and Leysen (2001)
described some key principles for a long-term monitoring program. In relation to Little
Owl ecology, they identified which landscape elements cause habitat heterogeneity
(i.e., dependent and independent variables) and which habitat types should be moni-
tored and determined the relevant sample sizes to be acquired.
To obtain better insights into habitat selection, a multiscale approach is suggested to
more readily detect differing patterns that occur at different scales (Swindle et al. 1999,
Apolloni et al. 2018, Fattebert et al. 2018, Mayer et al. 2021). The multiscale approach to
the study of habitat selection is based on the conceptual framework suggested by
Johnson (1980). Consequently, general habitat selection can be regarded as a hierarch-
ical process, with owls determining what is an appropriate place for breeding at a small
scale, with foraging areas determined at a broader scale (see Laaksonen et al. 2004).
Knowledge should become available on the macro-scale of countries and regions, on
the meso-scale (a population cluster), at the individual home-range scale and at the
fine-scale (components within individual territories). The first projects following a

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 11.2 Conservation Strategy for the Little Owl

scaled approach were done in Flanders (northern Belgium) (Van Nieuwenhuyse et al.
2001c), in France (Ferrus et al. 2002) and in Spain (Martinez and Zuberogoitia 2004b),
Czech Republic (Šálek et al. 2016) and Switzerland (Apolloni et al. 2018). A detailed
monitoring scheme was documented for the Netherlands (SOVON 2002) with special
emphasis on the selection of a stratified sampling scheme.

Rationale of Monitoring Programs and Demographic Studies

Long-term demographic monitoring programs have been established for relatively few
owls. While some of these monitoring programs are explicitly tied to recovery plans or
conservation strategies, others are not. Importantly though, these studies have
advanced the science of conservation biology, have been influential in affecting land-
use changes and have provided some fundamental information on the ecology and
conservation of owls and their habitats. Further, these studies have provided an import-
ant springboard for furthering the science of wildlife species and have offered import-
ant insights into the broader health of the ecosystems these owls inhabit.
Some studies also include radio-telemetry to examine dispersal of juvenile and adult
owls. Many studies often begin through projects that are designed to illuminate the
basic ecology of the species, such as estimating owl numbers and densities, and identi-
fying habitat relationships. With demographic studies, the emphasis of such projects
shifts to estimating trends in reproduction and survival. The types of questions that
drive demographic studies include: “How many owls are needed to maintain viable
populations?,” “Are the populations really declining because of habitat loss?,”
“Are individual populations isolated from other populations?” and “How is the popu-
lation (or metapopulation) responding to our management efforts?” Demographic
information and estimates of habitat loss are often central issues in litigation
and procedural arguments levied by competing advocacy groups (e.g., environmental
and industry groups). In some situations, analyses of demographic data have been
used to demonstrate that proposed (or current) conservation plans are insufficient to
provide for the long-term welfare of the species. Finally, the science behind species
and ecosystem management, of which demographic analyses are an important part,
has become a biopolitical cornerstone in national and international conservation
policy.
Alongside demographic data come a large number of mathematical population
models that have been designed to investigate the hypothetical responses of owl
populations to different kinds of landscape management. The distinction between
population modeling and population analysis is that the former is an abstraction and
the latter is an objective assessment of empirical data. Population models are con-
structed by depicting, mathematically, the characteristics of a population and then
examining hypothetical population behavior under a variety of assumptions. They
can be simple or complex depending on the number of parameters used and their
purpose. On the other hand, population analyses are based on objective evaluation of
the life-history information of the owl derived by capturing, marking and re-sighting
the same birds over long periods of time. It is this latter aspect of data for population
analyses that our Little Owl Monitoring Plan is focused on acquiring.

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Goals of the Little Owl Monitoring Plan

The specific goals of the Little Owl Monitoring Plan are to:

• Identify status and trends in the health of the Little Owl population and its key
habitat components;

• Provide early warning signals for situations that require intervention;

• Suggest remedial treatments and frame research hypotheses;

• Determine compliance with laws and regulations;

• Provide reference data for comparison with more disturbed sites;

• Provide scientifically rigorous data to enable managers to make better informed

management and policy decisions.

Specific population-related and habitat monitoring questions that direct the Plan are
as follows.
Population Questions:

1. Is implementation of the various conservation measures and policies effective in

reversing the declining population trend and maintaining the historical geographic
distribution of the Little Owl?
2. What is the trend in rates of demographic performance (adult survival,
reproduction, turnover and the annual rate of change of owl populations)? Do these
trends support a conclusion that the conservation measures are working to achieve a
stable or increasing population?
3. Can the status and trends in Little Owl abundance and demographic performance
be inferred from the distribution and abundance of habitat?
4. Can the relation between owl occurrence and demographic performance be reliably
predicted given a set of habitat characteristics at the landscape scale?
5. How well do habitat-based models predict occurrence and demographic performance
of owls in different land allocations (e.g., parks, reserves and matrix lands)?

Habitat Questions:

1. Is Little Owl habitat being maintained and restored as prescribed under the various
conventions, plans and policies? This general question has two key components:
a. What is the trend in amount and changes in distribution of habitat? Questions
relevant to specific parameters and how they are changing through time include:
i. What is the structure and composition of habitat at a variety of spatial
scales (nest site, home range, landscape, regional population)?
ii. What proportion of the total landscape is owl habitat?
iii. What is the distribution of sizes of habitat patches?
iv. What is the distribution of distances (connectivity) among habitat patches?
v. What are the primary factors leading to loss and fragmentation of habitat?
b. What is the trend in the habitat that provides dispersal connectivity between
populations? Questions relevant to specific parameters and how they are changing
through time include:

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 11.2 Conservation Strategy for the Little Owl

i. What is the structure and composition of dispersal habitat at the home-

range and landscape scales?
ii. What proportion of the landscape represents dispersal habitat? How is it
changing through time?
iii. What are the primary factors leading to dispersal habitat changes at the
home-range, regional population and landscape scales?

A number of Little Owl study areas currently exist, and some of these are suitable to
become part of a network of formal demographic study areas (see Sampling Design
below). Most of the current study areas are in European countries; study areas in other
portions of the owl’s range would need to be developed. In this plan, the ongoing
population demographic monitoring would continue in selected areas. Data about the
abundance and demographic performance of owls would be combined with habitat
data from the demographic study areas to develop models to predict owl occurrence
and demographic performance. Sampling the population and habitat as described in
this plan will ensure efficient use of resources and consistency of the information
needed to conserve owl populations.
Concurrent with demographic data collection, range-wide habitat conditions would be
monitored to track changes in conditions by using owl habitat maps derived from the
national and international vegetation maps and imagery. This analysis would reflect
several spatial scales (i.e., nest site, home range, landscape). Landscape components, such
as area of pastureland, would be monitored on the basis of critical structural and manage-
ment activities that reflect ecological conditions for the owl. The assumption is that the
larger ecosystem will retain its ecological integrity to the extent that key habitat elements
and physical processes are sustained. Population data from the demographic studies and
habitat information from companion studies on vegetative characteristics at these sites
will provide the data needed to model patterns of Little Owl occurrence and demographic
performance to home range and landscape characteristics of the vegetation. Provided the
models are field tested and shown to be predictable at a level of uncertainty (associated risk)
mutually acceptable to scientists and decision-makers, emphasis could shift from mark–re-
capture studies to increased reliance on monitoring of owl habitat and use of predictive
models to indirectly estimate the occurrence and demographic performance of owls.

Development of Conceptual Models at the Home-Range and Landscape Scales

An initial step in the monitoring task is to select indicators that reflect the underlying
processes governing the dynamics of species populations. Indicator selection is facili-
tated by first developing a conceptual model (Noon 2003). The conceptual model
outlines the interconnections among ecosystem processes (key system components),
the structural and compositional attributes (resources) and how the condition of the
resources affects the owl population (see Van Nieuwenhuyse et al. 2001a). The model
should also indicate the pathways by which populations accommodate natural disturb-
ances and how they demonstrate resilience to such disturbances. A framework to guide
model development is to link ecosystem process and function to measurable compon-
ents of the resources (that is, structural and compositional attributes). Changes in

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 Chapter 11: Management and Conservation

resource value, in turn, can be used to make predictions of expected biological response.
Measurements of biological systems are affected by the scale of observation. To deter-
mine the appropriate scale(s) of indicator measurement, the temporal and spatial scales
at which processes operate and populations respond must be estimated (at least to a first
approximation) and clearly identified in the conceptual model. The indicators selected
for measurement reflect known or suspected cause–effect relationships to population
dynamics. A satisfactory model provides a justification, in terms of current ecological
principles and theory, for the indicator(s) selected for monitoring and how knowledge
of the status and trend of the indicator reflects underlying process, function and
population response.
On the basis of other demographic work done on owls (Forsman et al. 1996, Mulder
et al. 1999, Franklin et al. 2004, Ganey et al. 2004) and on the conservation biology
science upon which that work was based (e.g., Gaines et al. 1999, Noon 2003), selected
indicators should:

• Reflect fundamental population processes and changes in stressor levels (that is,
habitat change)

• Be representative of the state of the population

• Be measurable and quantifiable.

Further refinement identified indicators with the following properties:

• Their dynamics parallel the dynamics of the species populations

• They reflect a rapid and persistent response to change in the state of the
environment

• An accurate and precise estimation is possible (that is, a high signal-to-noise ratio)

• There is a high likelihood of detecting a change in the magnitude of the indicators

given a change in the state of the ecosystem

• They demonstrate a low level of natural variability, or additive variation, and

changes in their values can readily be distinguished from background variation

• Measurement costs are not prohibitive.

The conceptual models developed for the Little Owl focus on negative habitat change
(both loss of area and fragmentation) and positive habitat change (landscape manage-
ment as a function of food availability, improved breeding opportunities and better
security) as the prime determinants of the species population dynamics at this time.
We have developed models at two spatial scales. One model focuses on habitat change
(both stressor and driver) processes relevant at the home-range scale, a meaningful scale
in the context of the dynamics of individual pairs of owls (Figure 11.7). The other
model focuses on changes in habitat pattern at the landscape scale (Figure 11.8). At this
scale, habitat change affects the size, and thus the likelihood of persistence of local
populations, and the connectivity among populations.

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 11.2 Conservation Strategy for the Little Owl

For both conceptual models, possible indicators are in two broad categories – habitat
and populations. This was done: (1) to document the well-established relationship
between habitat change, at a variety of spatial scales, and the dynamics of owl popula-
tions and (2) to suggest a list of habitat variables that may serve as surrogate indicators
once habitat-based models are developed.

Selection of Vital Sign Indicators

Population and metapopulation studies on owls have indicated that the adult survival
rate is the primary indicator driving population dynamics. Franklin et al. (2004) stated
that for the California Spotted Owl, while the adult survival rate provided the baseline
indicator for the population trajectory, fecundity and recruitment provided the vari-
ability around that trajectory. For fecundity, demographic analyses focus on the
number of female young produced per adult female.
At the home-range scale (Figure 11.7), habitat stressors (–) and drivers (+) affect the birth
and death rates of individual owls. The consequences of habitat change at this scale can be
assessed directly by measuring various demographic rates, or indirectly by measuring
habitat attributes correlated with variation in these rates. Both approaches are appropriate,
but direct measures of population attributes are clearly less ambiguous. For the near term,
we recommend a continued focus on direct measures of population attributes. Specifically,
we propose that monitoring efforts focus on precise and accurate estimates of adult survival
rate by using capture–re-capture methods (see details in Forsman et al. 1996, Franklin et al.
2004, Ganey et al. 2004). We emphasize adult survival rate as the most relevant indicator of
population status because: (1) variation in this rate most affects changes in population
growth rate and (2) it is responsive to habitat change at the home-range scale.
The second conceptual model addresses the landscape scale, focusing on stressors (–)
and drivers (+) affecting local populations and the interactions among these populations
(Figure 11.8). Relevant population-level indicators at this scale are population size and
distribution. At this scale, habitat change affects not only dynamics of local pairs of owls
but also connectivity among local populations. Attributes such as habitat patch size,
shape, number and distribution determine the connectivity and stability of local popula-
tions. The population consequences of habitat change at the landscape scale are ultim-
ately the same as at the home-range scale, even though the mechanisms driving the
change are different. In practice though, landscape-scale habitat variables will probably
be of insufficient resolution to predict population response at the home-range scale.
The conceptual models document the relations between habitat dynamics and
population response at two spatial scales. Until generic habitat-based predictive models
are developed, we recommend monitoring at the population level by refining current
European-based studies and developing new demographic studies in northern African,
Middle Eastern and Asian regions. At the same time, we propose an extensive inter-
national research effort to extend and generalize local Swiss, German, Czech and
Danish predictive models relating population dynamics to habitat variation (e.g.,
Chrenková et al. 2012, Šálek et al. 2016, Michel et al. 2017, Apolloni et al. 2018,
Grüebler et al. 2018) at the home-range scale and the landscape scale. This parallel
research effort is necessary to link habitat conditions to Little Owl population

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Processes acting as stressors Effects on habitat of stressors

Human disturbance (human induced) Decrease in large, live, solitary trees

Intensive agriculture Decrease in large snags
Removal of hedges and old trees with cavities Decrease in pasture/grassland
Abandonment of agriculture and grazing Decrease in species and number of insects and small mammals
Permanent habitat loss - Decrease in high-stem orchards
Pasture/grassland conversion Increased distance between patches of pasture/grassland
Cessation of agricultural land use leading to less grazing Creation of hostile landscape matrix
Forestation Creation of larger, more homogeneous patches
Urbanization Increased levels of human disturbance
Human development
Shooting
Trapping
Unintentional traps (chimneys, ponds, water troughs, telephone poles

Habitat disturbance (natural)

Wind damage
Climatic influence on the species and on prey species
Predation
Inter-species competition for cavities and food
Flooding Loss of pasture/grassland
Parasites - Decline in amount of habitat
Postfledging kills from trampling by cows

Local management as a function of food availability Effects on habitat of drivers

Optimal mowing management
Optimal grazing Optimization of pastures/grassland
Managing residual plots Restoration of habitat quality
Planting hedgerows, woodbank, Benjes hedges Creation of diversity and heterogeneity in patches
Perch provision + Mitigation of human disturbance
Digging ponds Larger trees with more decadence
Reduction in habitat scale by adding fences Creation of natural nest sites
Reduction in habitat scale by planting hedges Increase of habitat heterogeneity
Increased edges
Local management as a function of breeding opportunities
Increase of roost availability
Increase of nesting cavity availability
Planting high-stem fruit trees More roosts(two exits)
Planting solitary trees + More nesting cavities
Improving buildings
Installing nestboxes

Local management as a function of more security

Optimizing barns and sheds
Improving quality of artificial cavities (two exits, ventilated, marten proof)
Reduction of drowning risk
Reduction of poisoning effects Less poisoning
Mitigation of traps + Less drowning
Less mortality
Local re-inforcement
Food supplementation
Re-introduction
Supplementation of owls

Figure 11.7 A conceptual model of the effects of natural and human-induced stressors ( ) and drivers
(+) on Little Owls at the home-range scale.

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 11.2 Conservation Strategy for the Little Owl

Effects on Little Owls Potential indicators

Decrease in the area of pasture/grassland Habitat

Loss of nest sites Distribution of patch size
- Decrease in habitat quality Declines in reproduction
Decreased prey abundance Decreased survival of juveniles
Secondary poisoning Average perimeter-to-area ratios for habitat patches
Area in pasture/grassland
Area in intensively managed agriculture (e.g., row crops)
Heterogeneity in patches
Area in old, high-stem orchards
Large tree abundance-distribution (e.g., pollard willows)
Tree species composition
Pesticides/heavy metal levels

Populations
Population size
Population distribution
Nest initiation (percentage population breeding)
Reproductive success
Creation of nest sites Territory occupancy rate
Increased habitat quality Territory turnover rate
Increased prey abundance Juvenile dispersal rate
+ Increase prey diversity Juvenile survival rate
Better food availability Adult survival rate
More optimal energetics of the owls Replacement speed of killed adults
Greater site fidelity Concentration of floaters

Habitat heterogeneity
Number of different crops
Average parcel area
Percentage short/tall vegetation
Number of cavities
Number of perches
Year-round optimal mix of vegetation heights
Less predation Year-round optimal mix of prey availability
+ Improved survival
Improved fecundity

Fine-scale resources (after Tschumi et al. 2020)

Permanent meadow area
Less impact of predators Land-use richness
+ Less mortality Low-intensity meadows
Occurrence of grazed areas
Different cutting regimes
Number of dry stone walls, stone piles, stacks of wood, brush
piles, hedges, summer houses, unmaintained buidings and
equipment shelter buildings
Total number of tree cavities >6 cm diameter and >20 cm depth
Occurrence of potential roosting sites in buildings, palette stacks
Small rodent index
Number of trees
Sampling date

Figure 11.7 (cont.)

dynamics. For the most part, landscape-scale models will be based on habitat attributes
estimated from remotely sensed data. Home-range-scale habitat models will rely on
low-level aerial photography and ground plots to estimate the relevant variables.
Habitat models applicable at the landscape scale will be more extensive but will predict
population status with less resolution than models applicable at the home-range scale.
Because landscape-scale models make simplifying assumptions about habitat quality at
the home-range scale, both types of predictive models are necessary.

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Processes acting as stressors Effects on habitat of stressors

Human disturbance (human induced) Decrease in the area of pasture/grassland

Intensive agriculture Large-scale declines in habitat quality
Enlargement of parcels due to land re-allotment schemes Loss of connectivity between habitat patches
Removal of hedges and old trees with cavities Increased distance between patches of pasture/grassland
Abandonment of agriculture and grazing - Creation of hostile landscape matrix
Road construction Creation of larger, more homogeneous patches
Monoculture Increased levels of human disturbance
Permanent habitat loss Reduction in landscape heterogeneity
Pasture/grassland conversion Increase of scale of the landscape
Forestation
Urbanization
Human development
Windmills
Shooting
Trapping
Unintentional traps (chimneys, ponds, water troughs, telephone poles.)
Gentrification of holiday villages
Horse(ification) of rural landscape

Habitat disturbance (natural)

Wind damage
Climatic influence on the species and on prey species -
Predation
Inter specific competition for cavities and food
Flooding
Desertification
Parasites Effects on habitat of drivers
Postfledging kills from trampling by cows
Optimization of pastures/grassland
Processes acting as drivers Restoration of habitat quality

Human geographical evolutions + Creation of diversity and heterogeneity in patches

Mitigation of human distrubance
Human migration to cities leading to abandoned dwellings
Reduction of scale of the landscape
Processes acting as drivers

Landscape management as a function of food availability

Optimal mowing management
Optimal grazing
Managing residual plots
Planting hedgerows, woodbanks, Benjes hedges More roosts (two exits)
Perch provision + More nesting cavities
Digging ponds
Reduction in landscape scale by adding fences
Reduction in landscape scale by planting hedges

Landscape management as a function of breeding opportunities

Increase of roost availability
Increase of nesting cavity availability
Planting high-stem fruit trees Less poisoning
Planting solitary trees + Less drowning
Improving buildings Less mortality
Installing nestboxes

Landscape management as a function of more security

Optimizing barns and sheds
Improving quality of artificial cavities (two exits, ventilated, marten proof) +
Reduction of drowning risk
Reduction of poisoning effects
Mitigation of traps

Population management
Food supplementation
Re-introduction
+
Supplementation of owls

Figure 11.8 A conceptual model of the effects of natural and human-induced stressors ( ) and drivers
(+) on Little Owls at the landscape scale.

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 11.2 Conservation Strategy for the Little Owl

Effects on Little Owls Potential indicators

Large-scale declines in habitat quality Habitat

Loss of connectivity between habitat patches Distribution of distances among habitat patches
Decreased rate of pair formation Distibution of patch size
- Declines in reproduction Decreased rate of pair formation
Decreased survival of juveniles Average perimeter-to-area ratios for habitat patches
Decrease dispersal success Area in pasture/grassland
Increased rates of nest failures Area in intensively managed agriculture (e.g., row crops)
Isolation of local populations Road density
Local populations subjected to demographic stochasticity Heterogeneity in patches
Decrease of gene flow
Deline in owl numbers

- Decline in the amount of suitable habitat

Isolation of remaining suitable habitat

Populations
Population size
Population distribution
Nest initiation (percentage population breeding)
Reproductive success
Territory occupancy rate
Territory turnover rate
Juvenile dispersal rate
Juvenile survival rate
Adult survival rate

+ Better food availability

More optimal energetics of the owls
Dispersal metrics
Replacement speed of killed adults
Concentration of floaters

Fine-scale resources (after Tschumi et al. 2020)

Less predation Permanent meadow area
Land-use richness
Improved survival
+ Improved dispersal
Low-intensity meadows
Occurrence of grazed areas
Improved fecundity Different cutting regimes
Number of dry stone walls, stone piles, stacks of wood, brush
piles, hedges, summer houses, unmaintained buidings and
equipment shelter buildings
Total number of tree cavities >6 cm diameter and >20 cm depth
Occurrence of potential roosting sites in buildings, palette stacks
Small rodent index
Number of trees
Less impact of predators
+ Less mortality
Sampling date

+ Re-inforcement of population numbers

Figure 11.8 (cont.)

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Sampling Design
Population monitoring. The capability of a monitoring method to provide data
about vital rates for individual owls was considered key to whether a method would meet
the indicator monitoring needs identified in the conceptual models. Three alternative
methods for monitoring owl populations have been evaluated. The first, based on mark–
re-capture methods, would provide detailed estimates of demographic rates and the
annual rate of population change (see Forsman et al. 1996, Franklin et al. 2004 for
methods and rationale). The second alternative method is based on repeated counts in
randomly selected census plots and would provide estimates of rates of population
change and rates of change in occupied habitat. The third method, density study areas,
would require a total count of individual territorial owls in an area of several km2 and
would provide estimates of the rate of population change and change in occupied habitat.
Mark–re-capture methods were used to monitor territorial Little Owl populations in
the Low Countries (Bekaert 2006). The study provided estimates of rates of survival,
reproduction and turnover, as well as detailed data about location of nests, roosts and
habitat conditions at nest sites. Data about survival and reproduction were used to
estimate the annual rate of population change and to investigate trends in reproduction.
Bart and Robson (1995) suggested that a random survey using the playback technique
(for Northern Spotted Owls) could be done for considerably less money over broader areas
than the labor-intensive mark–re-capture approach. The results could be used to estimate
rates of population change, but not to estimate rates of survival, reproduction or turnover.
The density-estimate study area approach has been used in parallel with mark–re-
capture projects for several years. The density-estimate method assumes a 100% survey
of a land unit of about 20 to 100 km². In each density study area, observers record the
number and location of all owls encountered in one or repeated surveys during a single
season. Density areas are re-surveyed annually. The results can be used to identify
substantial changes in the number of owls, but do not estimate rates of survival,
reproduction or turnover.
Based on previous analyses on Spotted Owls (e.g., Forsman et al. 1996), only the
mark–re-capture method within demographic study areas will satisfy the array of
indicator data needs. It is also the only method that will provide the option to pursue
predictive model development, which is a key option within the overall monitoring
strategy. Given the declining nature of Little Owl populations, we consider estimates of
the demographic rates to be key elements of a monitoring plan. We therefore propose
that the primary sampling method for monitoring should be mark–re-capture studies in
the demographic study areas.
An additional consideration for population monitoring is the use of an independent
estimate of population trend for comparison with the results from the demographic
studies. An additional, scientifically credible method would provide an independent,
empirical assessment of the estimated population trend to compare with observed
demographic parameters. For example, declines in survival and reproductive rates
discerned from demographic studies should be reflected in declines in number of owls
encountered during repeated surveys. We recognized both the random census plot and
density study area methods as having the potential to provide an independent estimate

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 11.2 Conservation Strategy for the Little Owl

of population trend. Considerable uncertainty exists, however, about the sample size of
survey units and the number of years of survey required to detect small changes in
population trend. If the rate of population change is small (say 1% per year), and owl
detection rates differ between years, many years would be required to detect a <5% rate
of change in the population. If the rate of change is large (>5% per year), however, the
method would probably detect a significant downward trend in only a few years.
Because of uncertainty about the statistical power of the census plot approach, we
recommend additional estimates of the statistical power of the method and its cost-
effectiveness. Likewise, further analysis should be conducted on the efficacy of the
density study areas to provide an independent estimate of population trend. Once
these additional analyses are completed, the adoption of a method to obtain an inde-
pendent estimate of population should be given further consideration.
Habitat Monitoring. The dynamic changes in vegetation structure and compos-
ition of grassland ecosystems in the Little Owl’s range reflect the underlying biotic and
physical driving forces, including intensive and pastoral agricultural practices and other
human-caused sources of disturbance. For this book, we have assumed that knowledge of
vegetation structure and composition (amount and distribution) does not yet allow
reliable prediction of Little Owl occurrence or the demographic performance of the owls.
Habitat monitoring for Little Owls will consist of two separate, but related,
initiatives.
One is tied directly to each of the demographic study areas. Patch-specific vegetation
classifications and habitat evaluations will be completed in each study area (e.g., see
Van Nieuwenhuyse et al. 2001c). Habitat assessments of demographic study areas will
be based on plot data, vegetation description information and standard aerial photog-
raphy to develop structure and composition attributes for owl habitats. These data will
be used with the population data to assess the relationships among varying demo-
graphic responses and varying habitat conditions at the nest-site, home-range and
landscape scales. These relations will form the basis of the (future) predictive models.
Accurate and timely monitoring of changes in the status and trends of vegetation
(habitat) should provide reliable early warning signals to eventual changes in the
population viability of Little Owls.
The second habitat assessment will estimate baseline, range-wide Little Owl habitat
conditions and track change in habitat conditions over time. The range-wide coverage
also will provide unique capabilities for spatial analysis, portrayal of geographic distri-
bution of habitat, and analysis of patch statistics. A good example is offered by Ilanloo
et al. (2020) in their study of habitats in Iran. A range-wide vegetation map is urgently
needed and will be developed from satellite imagery. This is a case where the current
analytical tools permit the assessment of the universe of data, thus eliminating the need
to stratify and sample the data, which would require statistical analysis to describe the
precision of the estimates and would be more expensive. A baseline for range-wide
habitat condition will be established and periodically re-assessed to describe habitat
condition and trend. This will allow tracking of the amount of nesting and foraging
habitat at a variety of spatial scales. It also will provide information on habitat condi-
tions in population connectivity areas. Continuous follow-up is needed, given the

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speed of climate-change-induced alterations of climatic circumstances in the global

Little Owl distribution area.
Habitat-based aspects of the monitoring plan offer several advantages:

• Habitat monitoring can build on existing inventory programs.

• Estimating the trends in vegetative structure and composition represents a

prospective, as opposed to retrospective, approach to ecological monitoring.

• Monitoring vegetation change is more cost-effective than directly monitoring

populations of all the possible species for which management agencies
are responsible.

• The baseline and subsequent vegetation maps will be applicable to other species, and
supportive of the broader array of conservation assessments and
biodiversity management.

Some limitations to a strictly habitat-based approach to monitoring include:

• Some unknown proportion of the variation in species population dynamics may not
be driven by changes in habitat amount and distribution (for example, population
fluctuations due to behavioral and prey-related influences).

• Changes in habitat may not predict population responses to other stressors (for
example, environmental toxins, changing environmental conditions and
competitive interactions).

• On the basis of the above two limitations, a strictly habitat-based monitoring

program may have limited ability to predict changes in species viability
and distribution.

The following variables are a minimum set of attributes thought to influence the
presence and abundance of Little Owls at the landscape scale and to be monitored
range-wide:

• Distribution and area of the pastureland/grassland corresponding to owl nesting and

foraging habitat;

• Frequency distributions, by area size class, of habitat patches;

• Frequency distributions of distances between habitat patches;

• Tall versus short vegetation ratios;

• Heterogeneity of landscapes.

These parameters may be further summarized at a variety of spatial scales including

land allocation, administrative unit, physiographic region and range of the Little Owl.
Predictive Modeling. At some future point, to make the transition from
population-based monitoring to a completely habitat-based program will require an
intensive period of habitat model development. We recognize this task to be primarily
one of research, taking into account that recent research looks extremely promising.
Now it is a matter of setting up international co-operation, scaling up methods that

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 11.2 Conservation Strategy for the Little Owl

prove useful, replication elsewhere, more experiments and more copy/paste in other
geographies. We envision that a completely habitat-based monitoring program must
identify those aspects of vegetation structure and composition that have the greatest
power and precision to predict the number, distribution and demographic performance
of owls at the landscape scale, as well as to explain the observed variation in demo-
graphic rates at a home-range scale. This task will require characterizing the vegetation
at a variety of spatial scales in the existing demographic study areas. The combination
of spatially referenced data from both the owl demographic studies and mapped
vegetation attributes provides the fundamental data for the model-building phase.
The degree to which these models explain the observed variation in owl distribution
and demographic performance will estimate the certainty with which habitat variation
predicts population performance and stability. Explained variation is thus a direct
measure of the confidence we have in habitat as an appropriate monitoring surrogate
for population performance. In addition, validating model predictions by independent
field surveys is essential; that is, the models will be used to predict owl population
response, which must then be verified by direct field measurement of owls from several
landscapes with different population levels. Validation testing will use the range-wide
habitat map derived from the regional or country-based vegetation maps. Once reliable
models are developed, existing habitat conditions across extensive landscapes can be
assessed and the expected owl occupancy, distribution or demographic performance
predicted, depending on which predictive level of monitoring is implemented. We have
made huge progress, but we still have a long way to go before we will be able to shift to a
habitat-only monitoring program at a global scale.

Development of a Network of Vital Sign Monitoring Areas

Vital Signs are key elements that indicate the health of the Little Owl population and
the key habitat components it depends on. Vital Signs can be any measurable feature of
the population or environment that provides insights into the state of the species. The
term is synonymous with “ecological indicator,” but use of the term and the analogy to
an individual’s health is helpful in explaining the need for monitoring to managers,
politicians and the public.
We propose the development of a network of some 30 monitoring areas across the
range of the Little Owl. The scientific framework for the conceptual approach for this
work is based upon Mulder et al. (1999), the National Park Monitoring Program for the
USA (http://science.nature.nps.gov/im/monitor/vsmTG.htm), Gaines et al. (1999), the
European Biodiversity Monitoring and Indicator Framework (www.strategyguide.org/
ebmf.html) and the Framework for Monitoring Biodiversity Change in Canada (www
.eman-rese.ca/eman/reports/publications/framework/context.html).
Our proposed network of monitoring areas reflects an array of study areas spread
across the major ecological regions (biomes) within the range of the Little Owl. Each
monitoring area should contain 50–100 pairs of Little Owls, or equivalent habitat in
areas where population restoration is planned. A network of such monitoring areas is
recommended because it is currently unrealistic to expect that all countries within the
range of the Little Owl will be able to adequately fund and consistently support

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monitoring at the optimum scale and intensity desired. A monitoring program requires
professional-level staff who can analyze and interpret data, prepare reports and provide
the information in a useable format to managers, scientists and other interested parties.
Thus, the compromise position is to identify a base network of monitoring areas, and to
develop a core program for the Little Owl. Thereafter, the participating parties in the
network can identify what specialists are needed, and allow each network component
to then leverage their core resources with other resources and partnerships to build a
monitoring program that can also be integrated with monitoring efforts for other
natural resources (e.g., Important Bird Areas, UNESCO World Heritage Sites, National
Parks, Wildlife Refuges, etc.).
The experimental design of the demographic study areas is crucial for estimating owl
birth and survival rates. This is important because the individual vital rate estimates
provide data about population indicators. These data will be used in the population
meta-analysis and as input to predictive models. The large, quality datasets associated
with existing Little Owl study areas provide a foundation for the monitoring program.
The majority of the potential Vital Sign demographic study areas were developed by
contacting researchers currently working in these areas, and discussing aspects of owl
densities, distribution, logistics and previous research efforts on the Little Owl.

Design of Methods within Each Vital Sign Demographic Monitoring Area

Sample Sizes and Protocols. Precise estimates from mark–re-capture studies require
large samples of marked owls. We recommend that sample sizes for individual demo-
graphic studies be 50 pairs of owls (100 individuals), with samples of 75 to 100 pairs
preferable. Demographic parameters monitored in mark–re-capture studies should include
survival and reproductive rates by age-group and sex. From these data, the annual rate of
population change can be estimated. Occupancy and turnover rates at historical owl sites
in the demographic study areas should also be monitored because these variables may
provide key information about the response of owls to variation in home-range and
landscape-scale features. We anticipate that mark–re-capture studies will employ ringing
as the primary means of tracking individual owl histories. The first initiatives with RFID
pigeon rings look promising to trace often inaccessible males entering nestboxes without
capturing them (Hameau and Millon 2022) Protocols for assessing survival, reproduction,
occupancy rates and turnover rates of owls will need to be consistently followed across all
study areas (for examples, see Forsman et al. 1996, Franklin et al. 2004).
A Technical Manual (in prep.) should be able to guarantee the needed harmoniza-
tion and standardization. A Survey Protocol has been developed to provide consistent
and cost-effective methods for monitoring demographic aspects of Little Owl popula-
tions. This protocol is now going through a formal peer-review process, and will be
published separately from this book.
Demographic Area Habitat Assessment. Habitat classification and mapping
will be completed for each of the demographic study areas. To date, the habitat
characterization has been conducted independently for each of the demographic study
areas. As a result, there are differences in the measured attributes, classification stand-
ards and resulting map products. Consistency of the vegetation classifications is import-
ant to the modeling that will follow, thus there is a need to resolve differences between

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 11.2 Conservation Strategy for the Little Owl

the methods. The resolution of these differences may involve some re-sampling and re-
classification of variables in some of the demographic study areas. The responsibility for
completing this will be assigned to the leaders of the respective demographic studies.
All re-assessment and new classification efforts will follow the consensus standards
developed during the resolution process.
The set of vegetation attributes that best characterize Spotted Owl habitat will differ by
physiographic region and study area. Investigators will therefore first identify the key
attributes in each demographic study area to be used to define habitat depending on
presumed habitat relations in that region. The landscape-scale habitat maps will be based
on attributes acquired from the regional vegetation map or specified for inclusion in
subsequent versions of the vegetation map. This approach is essential so that more general
predictive models can be built that apply outside the demographic study-area boundaries.
Range-Wide Habitat Assessment. The basic information needed for range-wide
monitoring of owl habitat is a set of map layers that collectively characterize Little Owl
nesting and foraging habitats. An overlay of map layers will allow the development of a
GIS-compatible database used to describe the amount and distribution of habitat in
relation to land allocations or other geographic areas of interest. Once developed, the
map would be updated periodically to track habitat change. Periodic updates of the map
layers in the near term will allow the estimation of changes in amount and distribution of
habitat over time resulting primarily from agricultural intensification and human devel-
opment. Changes in vegetation due to forest succession are not expected to provide any
significant changes in habitat condition for several decades. In the initial years of moni-
toring, detecting biologically significant changes in habitat condition will require periodic
inventory at 10-year intervals. The map would be re-compiled and habitat conditions re-
assessed in synchrony with the schedule for monitoring map product updates.

11.2.2.2 MANAGEMENT
Little Owl conservation in cultural landscapes is feasible and small-scale enough for
volunteers to play an important role. Some measures are good for Little Owl habitats in
and around gardens. Some of those measures physically benefit the species, e.g., the
installation of a nestbox, stacking wood as a perch or covering a water trough to prevent
owls from drowning. Other measures such as setting up a piece of waste land, grazing
donkeys or horses, making a branch wall or growing rye in a vegetable garden are
examples that indirectly help improving the habitat conditions for prey animals and
thus food availability. Favorable conditions can be obtained by omission of activities
too, such as not cleaning up a pile of stones, not breaking down an old shed or not
mowing part of the lawn every week. Here we offer suggestions and descriptions of how
to improve the overall survival conditions for Little Owls, largely based upon
Parmentier and van Paassen (2009). A distinction is made between measures to improve
the food situation, the breeding grounds and the safety of fledglings. Opportunities for
optimization are possible for grasslands, timber stands and others.

Habitat Bottlenecks
Since the 1960s, the number of Little Owls has declined sharply (Van Nieuwenhuyse
et al. 2008). The main cause of this decline is the deterioration and gradual

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disappearance of small-scale cultural landscapes after World War II. Many high-stem
fruit trees were replaced in the second half of the twentieth century by low-stem trees,
removing many potential breeding cavities. The maintenance of pollard trees was also
neglected, leading to damage during storms and eventually loss. Many old sheds were
demolished and many breeding cavities were made inaccessible due to renovation of
buildings. For the Little Owl, these developments meant decreasing nesting opportun-
ities in tree cavities (pollard willows, high-stem fruit trees) and (ancillary) buildings.
Clearing high-stem orchards, intensifying agricultural land use and clearing residual
plots also had negative consequences for prey availability, leading to sharp decreases in
the numbers of insects and voles. Low prey density necessitates that the owls make
longer food flights, while searching for prey for longer takes a lot of extra energy (Sunde
et al. 2009, Holsegård-Rasmussen et al. 2009, Staggenborg et al. 2017). There are
boundaries to this, which presumably in many areas have been exceeded, resulting in
poor breeding results. In addition, owls are crossing roads more often, which increases
the risk of death from traffic. Fortunately, there are also regions where the population is
more or less stable. Small-scale landscapes and locally less intensive land use offer the
Little Owl enough living space. Thanks to increasing interest, orchards are well main-
tained and are also restored locally. Nowadays pollard trees and other small landscape
elements are better maintained too. In these regions, Little Owl conservation has taken
place for a long time in the form of installing nestboxes. The combination of extra
nesting facilities, small-scale measures that promote the food situation and reduce
physical threats to Little Owls and interest in conservation among volunteers has
caused the decline to stand still in several regions (Habel et al. 2015, Meisser 2021).
The principal bottleneck for the Little Owl in western European agricultural landscapes
is the deteriorating food situation (Thorup et al. 2010, Šálek et al. 2016). Habitat improve-
ments are preferably undertaken at the scale of a local breeding population in at least five
territories and have higher success rates when close to existing territories. Since Little Owls
are resident birds, fledglings explore locations near their birthplace. The further away
from an occupied territory it is, the less chance that a fledgling will discover that spot.
The best results are achieved with planting and maintenance of adjacent landscape
elements. In addition, if landowners know the local circumstances it has a stimulating
effect. This approach also improves settlement in small-scale cultural landscapes.
The presence of nesting Little Owls indicates that the area is suitable to offer
opportunities to strengthen the population. Without existing pairs in the neighbor-
hood, however, support of local people is usually significantly lower, even with the co-
operation of critical farmers regarding their land. At a conservation site, there are
various tasks for local volunteers. It is therefore desirable to choose an area where a
Little Owl group already exists and a local birding group is active. It is recommended
that an area of between 1000 and 2000 ha is chosen. Within an area of this size, there is
a good chance that fledglings can find an unoccupied territory in the vicinity of their
birthplace. The further apart suitable sites are, the less likely it is that the owls will find
suitable habitat and survive winter. Large-scale agricultural areas are less suitable
because of the lack of sufficient old, cavity-rich pollard trees, hedges and tree rows,
which are essential in small-scale cultural landscapes.

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 11.2 Conservation Strategy for the Little Owl

Initiatives start by mapping existing nesting sites for Little Owls by collecting infor-
mation about the current and historical breeding sites in the area. A dotted map of the
breeding grounds giving an overview of the local distribution is an excellent baseline.
When the existing population is clustered within the area one should consider
concentrating on those subareas. Before starting the project, it is advisable to
conduct a stakeholder analysis amongst farmers, fieldworkers from agricultural associ-
ations, nature conservation volunteers, reserve managers, and local nature and
bird lovers.
Plans are needed to adapt an owl-friendly yard by first confirming whether there are
opportunities for Little Owls and what bottlenecks exist, and describing them in a
systematic way via a Little Owl territory scan. It is not only the yard with buildings,
vegetable garden, orchard, pastures, residual plots, surrounding fences or the access
road with rows of trees that are important. Plots with wood stands, crops in the vicinity
offering food, roosting and nesting places are also part of the territory. A Little Owl pair
typically uses an area with a radius of between 100 and 300 m around the (potential)
breeding site. The size of this territory depends on the local food supply. An initial
exploration is advised, together with landowners, in order to immediately gain insights
about land use and management of the various elements in and around the yard, the
land use in adjacent plots and the history of the yard and the buildings. A rough sketch
of the type of landscape, the presence of other yards, timber stands, plots, waterways
and roads, and the current management of the various elements and use of the plots
offers a good baseline. An aerial photo can bring an extra overview. Identification of the
nearest Little Owl territory and whether there is connecting habitat with the (prospect-
ive) territory, and the type and use of adjacent parcels is necessary. Find out who uses
these plots and ascertain how the yard and buildings are used and what condition they
are in. It is advisable to measure the length and surface area of the various elements.
State whether Little Owls have nested in and around the yard in the last decades. Also
inquire about recent changes with regard to the expansion of buildings, land use or
planting of trees. Make a systematic inventory of which elements the yard contains that
are important for the food supply and its quality through the breeding season and the
rest of the year. Identify bottlenecks with respect to food supply and opportunities for
improvement. Check for nesting opportunities in hollow trees, sheds and buildings, or
nestboxes. Conclude whether there is sufficient breeding opportunity and provide
solutions in case they are lacking. Shelter and roosts are important for adults and
juveniles, hence investigate the accessibility of buildings and the availability of trees
with hollows, woodpiles, perches and hollow tubes. Assess whether there are sufficient
hiding places and roosts and how these can be improved. Investigate which threats and
dangers there may be in and around the yard (roads, railways, yard work, pets, drinking
troughs, recreationists, pesticide use, etc.). Verify if disruptive activities take place in the
yard. Future plans can also be important in case the landowner wishes to demolish a
shed in which a local owl pair breeds. Conclude whether the hazards and threats have
been remedied or prevented and if not how this can be achieved. After collecting all the
data, draw conclusions about the suitability of the yard and the environment for Little
Owls and whether there are sufficient elements present providing nesting cavities, food

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and physical safety. Indicate how the habitat can be improved, keeping in mind that
not all missing elements are of equal importance.
A Little Owl management plan contains a map of the current situation of the yard
and its surroundings, and elements to be constructed or changed as part of the current
management. Highlight all changes, yielding a quick overview of the different elements
within the potential territory. It is advisable to estimate the cost and how much work
will need to be carried out. Think about who will perform the work and when it can be
carried out. This management does not necessarily have to be done by the resident. It is
conceivable that others undertake the work, such as a local volunteer group maintain-
ing high-stem fruit trees or pollard willows. Explain the draft design plan to the
landowner, identifying adjustments and consequences in terms of work and funding,
leading to a consensus and a definitive layout plan.

Measures to Improve Food Availability

Little Owls have very varied and opportunistic diet needs near their breeding place and
roosts. The reduction in food availability due to the loss of characteristic elements of
small-scale cultural landscapes is one of the main causes of the decline of Little Owl
populations (Šálek et al. 2016).
Figure 11.9 illustrates the different phases in managed landscapes in the Lower
Countries and visualizes the consequences of lack of management. Figure 11.10 illus-
trates the phases in the UK up to the optimal Little Owl habitat locally. Figure 11.11
illustrates how prestine Little Owl habitats in Portugal deteriorate due to modern
agricultural changes.
Many pollard trees, hawthorn hedgerows and high-stem fruit trees have disappeared.
In addition to more intensive land use, the surfaces of herbaceous grassland and fields
with wildflowers have declined significantly and many parcels merged through land re-
allocation projects, reducing the number of field borders (Šálek et al. 2016). In regions
where there are still reasonable numbers of Little Owls, grazed pastures, rough corners,
high-stem fruit trees, sheds, vegetable gardens, pools, hedges, woods and tree rows still
occur. It is important to maintain and restore small-scale cultural landscapes or restore
extensive land use in order to offer a very diverse habitat leading to a variety of food.
Five basic rules apply for a greater diversity and number of plant and animal species to
be realized, i.e., variation in space, variation in time, customization, short-term and
long-term wins. These rules should be used as the basis for a management plan. The
more variation in spatial management, the greater the availability of prey, the better
chance of good results.

Grassy Vegetation
Grasslands rich in wildflowers and prey are crucial. Herb-rich vegetation with an open
structure and a soil that is not too acidic and is rich in earthworms are preferred.
Earthworms and insect larvae live in abundance in and on the ground. Caterpillars,
larvae, beetles and other insects look for food there. Not only is the presence of prey
important, but it needs to be within reach too, necessitating places where the vegeta-
tion is shorter than 10 cm. Research (Grzywaczewski 2009) has shown that the presence

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 11.2 Conservation Strategy for the Little Owl

(a)

(b)

Figure 11.9 Effects of landscape management in function of Little Owls in the Lower Countries.
(a) Baseline without specific management. 1. Corn field; 2. Channelled water. (b) First initiatives
reducing the general landscape scale by planting willows, wood stands and introducing sheep for
grazing. 1. Grain field; 2. Benjes hedge; 3. Nestbox; 4. Vegetable garden; 5. Owl-friendly drinking

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(c)

(d)

Figure 11.9 (cont.) trough; 6. Young pollard willows; 7. Afforestation; 8. Solitary tree; 9. Free
running stream. (c) Tempero-spatial diversity in management, i.e., trimming pollard willows,

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 11.2 Conservation Strategy for the Little Owl

of grassland with short vegetation around the breeding site is very important. This is
especially true in years when voles are scarce and during the nestling period between
mid-May and the end of June. A grassland with an open wildflower-rich structure that is
attractive for Little Owls does not automatically have to be of botanically high quality.
A lawn or meadow under a high-stem orchard, a grassy residual plot or a grazed pasture
can also contribute to a good habitat, as long as there is a lot of variation. In order to
maintain or develop a wildflower-rich grassland, defertilization of the soil is key. The
more nutrient-rich the soil, the fewer wildflowers and the less prey animals. The
grassland should be fertilized as little as possible and mown as needed, with the cut
grass removed. Different grasslands, such as lawns, hay fields, grazed pastures and
verges offer plenty of prey variation.

Herb Borders
Many homes and farms in the countryside have a well-maintained, monotonous lawn
free of weeds. Except for earthworms, prey is scarce. By transforming the edges into a
wildflower border, the lawn rim attracts butterflies, insects and beetles, but also voles
find cover and food. Insects use the intensively mown part of the lawn to catch heat,
making short grassy vegetation as important as tall grass. Edges should be mown along
a hedge or ditch less frequently than the rest of the lawn in June, August and October
offering the weeds the chance to flower, set seed and spread. By disposing of the
clippings, this part of the lawn can be impoverished further. Manure should be avoided
as it stimulates the growth of grasses, suppressing weeds. Preferably native wild seeds
are used that can be collected from flowery roadsides or in neighboring wildflower-
rich meadows.

High-Stem Orchards
Grassland under a high-stem orchard is ideal for foraging as it is often grazed with sheep
or calves. The fallen fruit is an attractive food source for insects and livestock manure
attracts dung beetles. To avoid overgrowth and ensure the presence of short grass, it is
necessary to graze or mow the orchard. Extensive grazing with different animal species
is preferable to mowing because the vegetation height varies more due to grazing.
Grazing with sheep in combination with other animals yields the greatest variation.
Horses are less suitable in an orchard because they damage the trees, donkeys are
preferred because of their smaller size. Vegetation shorter than 10 cm provides less
cover for voles but allows hunting. Mow the first time preferably after the flowering
period in mid-June and a second time in September or early October. When possible,
mow in steps and not all at once or in a spatio-temporal mosaic form (Michel et al.

Figure 11.9 (cont.) different grazers, digging a pond, bordering fields with hedges. 1. Grain field
with wildflower border; 2. Benjes hedge; 3. Nestbox; 4. Vegetable garden; 5. Owl-friendly drinking
trough; 6. Periodic trimming of pollard willows; 7. Hay rack in hay field; 8. Pond; 9. Pollard
willows; 10. Solitary tree and woodpile; 11. Grazing cows. (d) Lack of maintenance causing further
closing of the landscape, less grazing and promoting circumstances for Tawny Owls. 1. Large forest;
2. Overgrown Benjes hedge; 3. Nestbox; 4. Overgrown vegetable garden; 5. Overgrown pond;
6. Untrimmed pollard willows; 7. Rampant undergrowth.

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(a)

(b)

(c)

(d)

Figure 11.10 Effects of landscape management on Little Owls in the UK. (a) Baseline without
specific management. 1. Lawn; 2. Slope; 3. Cattle field; 4. Broken wall. (b) First initiatives reducing the
general landscape scale by planting willows, restoring stone walls, and introducing hay fields with
hay stacks, a pond and nestboxes. 1. Hedge; 2. Rose rack; 3. Vegetable garden; 4. Sapling; 5. Restored

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 11.2 Conservation Strategy for the Little Owl

2017). Remove the grass cuttings. Let the edge grow so that a wildflower-rich border
develops. Mow the edge with the second mowing to prevent roughening. In southern
Germany and Switzerland high-stem orchards are called Streuobstwiesen (Henrichs and
Fischer 2006, Fiedler 2013).

Grazing
Little Owls prefer short, grazed areas to hunt (Grzywaczewski 2009). Horse or donkey
pastures are attractive as they are hardly fertilized and the vegetation is varied and rich
in structure due to the permanent, low-intensity grazing. Rough places with flowering
weeds, next to parts that have been grazed are typical in combination with horse
latrines of concentrated horse dung causing the vegetation to become rougher. The
flowers and horse manure attract a wide variety of insects and their larvae. Sheep or goat
grazing is attractive too, whether or not in combination with young stock as long as the
grazing is extensive. High livestock density is detrimental for all kinds of prey.

Residual Grassland Plots

Most farms have grassland plots that are harder to maintain or too wet, often oblique or
irregularly shaped along a watercourse, wood bank or forest edge. These are potentially
ideal places for prey and can be easily converted into rugged and varied vegetation with
varying heights. When properly managed, species-rich, grassy vegetation is obtained,
often rich in butterflies that search for food, reproduce and hibernate there. In late
summer, fall and winter voles find seeds offering the resident owls enough food all year
round. Setting up a residual plot on a grassland parcel usually means cleaning up and
cutting the vegetation short first. At places with differences in moisture balance and soil
types, more diversity is to be expected, with more kinds of plants.

Mowing
To maximize variation in vegetation height it is important to mow as often as neces-
sary, depending on the fertility of the bottom. If the vegetation grows quickly and
consists mainly of grasses and nettles, mowing at least twice a year is advised, before
blooming to prevent seeding and promoting exhaustion of the plants. After a few years,
the vegetation will grow slower and the frequency can be reduced to once or twice a
year, preferably in June/July and September/October. Species-rich vegetation with
limited crop production should be mown only once every two years in August. Ideal
is to mow in blocks or mosaics, i.e., one half in May/June and September, with the other
half offering spring and summer blooming opportunities. Prey can reproduce in the tall
vegetation and be caught in the short vegetation.

Figure 11.10 (cont.) wall. (c) Further scale reduction by planting high-stem orchards, tempero-
spatial diversity in management, i.e., trimming pollard willows, different grazers, planting solitary
trees. 1. Pollard willows; 2. Slope with wildflower; 3. Hay bales in hay field; 4. Nestboxes;
5. Vegetable garden; 6. Flower garden; 7. Hay rack; 8. Pollarded linden tree; 9. Pond; 10. Sapling;
11. Wood rack. (d) Final stage with grazing by horses alternating with hay field. 1. Trimmed pollard
willows; 2. Trimmed pollard linde; 3. Wood bank; 4. Solitary tree; 5. Horse pasture; 6. Woodpile; 7.
High-stem orchard.

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Figure 11.11 Effects of landscape changes in function of more industrialized agriculture in Portugal.
(a) Prestine Little Owl habitat with old olive trees and stone heaps. 1. Old barn with nesting
opportunities; 2. Quiet cliff; 3. Grazing donkey; 4. Grazing sheep; 5. Rock pile; 6. Dunghill; 7. Old tree
plantation; 8. Wild flowers. (b) Deterioriation of Little Owl habitat by restoration of buildings, turning
them into holiday homes, outdoor sports, removal of old olive trees and replacement by young low-
stem olive and almond trees as cash crops. 1. Modern house; 2. Cliff used by paragliders; 3. Paved
ground; 4. Vineyard; 5. Almond plantation.

Disposing of the clippings promotes defertilization of the soil and species richness of
the residual plots. When the vegetation mainly consists of grasses with a lot of rough
weeds, dispose of the clippings immediately. In case of species in less nutrient-rich soil,
let the clippings dry for a few days to give these plants the opportunity to seed.
However, always dispose of the clippings within a week. When the plot is isolated in
the landscape introduce flowery hay from a neighboring nature reserve or a seed
mixture of native wildflowers.

Cereal Fields
Instead of a grassy corner in the yard, the plot can (partly) be sown with cereals such as
rye or barley which voles and granivorous birds love, offering some more variation to
the diet of owls in cold periods. Favorable locations for a grain field in the yard are a
sunny spot adjacent to a thicket, a rough corner or heap of branches in the vegetable

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 11.2 Conservation Strategy for the Little Owl

garden. Using crop rotation, a different part of the vegetable garden can be used every
year for rye, barley or oats. The crop remains unharvested, the grains fall from the ears
to the ground spontaneously, benefiting prey all winter long.

Field Borders
More diversity in arable land can be introduced by constructing a wildflower border or
fallow strip, taking a strip out of production along the edge of the plot and sowing with
a wildflower mixture, omitting fertilization or giving room for spontaneous develop-
ment of the vegetation. In spring and summer, the flowers attract insects, butterflies
and beetles, which in turn attract insectivorous animals. In fall and winter, the
remaining seeds attract voles and birds alike. After harvesting the commercial crop,
the wildflower border and fallow strip still provide cover and food for various prey
species. Suitable locations are along wood banks, hedgerows, thickets or ditches. These
parts of the parcel are often less productive anyway, due to shadowing. Optimize the
seed mixture in such a way that there are flowering plants that bloom all summer long
by paying attention to the flowering period of the species. Strive for seeds of native
origin fitting in well with the natural presence of the species and preventing exotic
flora. The sowing period is around mid-April.
To prevent the development of too dense and too rough vegetation on the fallow
strip, it is desirable not to fertilize it. Mow in late summer between mid-July and mid-
September to avoid excessive crop growth. In this way the vegetation is sufficiently tall
again in winter to provide cover for birds and mammals. If the farmer wants to put the
fallow strip back into production, plowing is best as late as possible to maintain the
undergrowth of the vegetation as long as possible, so that prey can benefit from
it longer.

Hedgerows
Little Owls often hunt along plantations and wood stands, in and under hedges. Plenty
of prey animals such as voles, amphibians, birds, butterflies, caterpillars and other
insects live in hedges, woods, solitary trees or tree rows. Well-maintained timber stands
are therefore crucial within good habitat.
Hedges can be planted or sprout spontaneously, forming an attractive hunting
ground. If hedgerows are not pruned, they grow fully in height and width. Hedges
regularly consist of different types of shrubs such as Hawthorn Crataegus monogyna or
Blackthorn Prunus spinosa. Overgrown hedges can have a rich undergrowth of wild-
flowers. They are also very attractive for various bird species and butterflies because of
the breeding opportunity, food and shelter that hedgerows provide to them. Hedges
higher and wider than 4 m have the most natural value. Amphibians, insects and
mammals such as voles and shrews use hedges as shelter and as pathways. The most
suitable locations for planting hedgerows are places adjacent to other timber stands,
since connected hedges are richer in plant and animal species than solitary ones.
Pruned hedges are straight linear elements of shrubs planted closely together grow-
ing around lawns and vegetable gardens. They consist of shrubs of Hawthorn or
Blackthorn that are cut in a block shape providing shelter or nesting opportunities for

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birds. Hedgerows can be created by planting shrubs in a row 25 cm apart. Preferably

regional shrub species are used. In cases of more than one species, put at least five
shrubs of the same species next to each other.
Hedgerows need maintenance to remain vital. Without maintenance, a hedge hollows
and eventually turns into a wood bank or even a forest border. Prune the
hedge, depending on the growth rate, two to five times a year. If vitality is greatly
reduced, it is best to cut the hedge at 10–20 cm above the ground without cutting the
hedge straight down, but let it flare out slightly at the bottom to keep the
hedgerow denser.

Wood Banks
Wood banks are long narrow wood stands overgrown with shrubs and trees forming a
partition between plots. The most suitable locations are along field edges and roads,
paths or banks. The usual form of maintenance for linear plantings is coppice manage-
ment. Depending on the situation and the species, the wood bank is fully cut every
7–20 years. By removing part of the timber stand annually or every few years, the
work is split in time and space, promoting diversity. Wood banks are most
valuable when more than 10 m wide, especially when trees and shrubs are used.
Preferably plant shrubs on the edges and trees in the middle and in groups of the same
species. If cattle graze the parcel it is wise to place a grid to protect against damage while
still allowing the cattle to graze at the lower edge, leading to more variation in
the undergrowth.

Solitary Trees and Tree Rows

Solitary trees are freestanding, prominent trees in the yard or in a garden grassland plot.
Tree rows are organized as avenues, usually on both sides of a driveway. Solitary trees are
mainly oak, elm, beech, nut or chestnut. For avenues, linden, poplar or beech are
frequently used. Little Owls often use solitary trees and tree rows to perch or roost. In
solitary trees with cavities, they also breed. If fallen from the tree or on the ground,
scurrying owlets can climb back when threatened. Voles find food under solitary fruit
and nut trees. Because of the large amount of caterpillars on the leaves, they are also an
important food source for young Little Owls. The most suitable location for a solitary tree
is in front of or next to a house, preferably near an existing nesting site. Trees can also be
planted at prominent landscape points on or outside the yard, provided it is at a sufficient
distance from other trees, shrubs or buildings. When planting trees, use native, local
species or varieties. Trees are planted in the winter months from October to mid-March,
but not during frost. Do not plant a solitary tree with a large crown too close to existing
plants and buildings. Place one or two tree posts next to the tree and fix the tree here.
When planting a solitary tree in a grassland plot, place a grid or a tree basket around
it as protection against livestock. Attach the basket at a maximum of 10 cm above the
ground to allow grazing and mowing underneath. This has the additional advantage
that fledglings can crawl under it in case of danger and climb into the tree. Avoid
grazing horses or goats around unprotected solitary trees.

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 11.2 Conservation Strategy for the Little Owl

Other Foraging Optimizations

The presence of one or more woodpiles in the yard is favorable for prey, especially in fall
and winter, as they form a dry shelter for many prey animals. High piles of wood are
also used as vantage points and hiding places when fledglings scurry around on the
ground in yards where cats and dogs roam. An attractive woodpile functions as a food
source all year round. For the woodpile, firewood or planks are suitable, but also
branches that are cut during the maintenance of trees and bushes. The pile must be
at least 1.5 m high to serve as a vantage point.

Benjes Hedges
Benjes hedges or deadwood hedges (Benjes 1997) are hedges that are created by linear,
loose deposits of mainly thinner wood cuttings, such as branches and twigs, through
seeds or initial planting. Benjes hedges offer birds and other animals protection and
food, so that they can accelerate the sowing of trees with their droppings or
food depots.
The principle of the Benjes hedge is not to create hedges by planting them but let
them develop spontaneously through wind and seeds from the droppings of resting
birds. For this purpose, wood cuttings (branches, twigs, sticks) are mixed up, as a pile or
in strips, as a wall, loosely stacked or, better, simply tipped, which in turn serves to
protect the growing plants. The advantages of such a system are, on the one hand, its
low production costs, since it is not plants but seeds that are brought in naturally or
autochthonously, and branches are often available as waste. In addition, the loosely
stored dead wood provides immediate habitat for numerous species of birds (mainly
hedge-breeders), small mammals and insects.
The Benjes hedge proves to be useful by quickly and inexpensively making a contri-
bution to the network of biotopes in extensive, field-adjusted, intensively used agricul-
tural landscapes such as meadows and fields. Children and adolescents can also easily be
involved in nature conservation and environmental protection by building, observing
and monitoring such a hedge, so they function as vectors for environmental education.
Depending on the landscape, instead of the desired berry-bearing thorn bushes, tree
species can also be used, usually the wind-spread forest species, such as birch and
willow, and commonly also ash.
With a Benjes hedge the prunings of trees are neatly stacked lengthwise between rows
of posts. A branch wall is comparable to a pile of branches, but looks a lot neater. A fresh
wall has a height of 1–1.5 m. Over time plants will grow through the branches. After a few
years, bushes grow above the branches and take on the shape of a hedge. A branch wall
houses many prey animals, such as voles, toads, insects and birds because it provides
excellent food, shelter and a wintering place for them. Stack the branches lengthwise.
Branches come from the pruning, pollarding or felling of trees and shrubs. The thicker
branches should be laid at the bottom and thinner branches on top. Due to rotting
the branch wall slowly collapses unless it is annually supplemented. The most suitable
locations are places near the pruned trees and shrubs, which saves a lot of dragging. The
edge of an orchard is a good location, but also near a residual plot or rough patch.

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Improving Conditions for Voles

There are various options to improve circumstances for voles by ensuring extra food
and dry, sheltered wintering places. Healthy vole populations guarantee sufficient food
for the owls in every season. Taking measures to encourage a larger population must be
done cautiously to avoid them becoming a pest, e.g., by only helping vole populations
to overcome a difficult (winter) period. The easiest way to help voles in winter is to
spread out some straw in a quiet place in the yard and scatter some grain over it now
and then. This can be done in a residual plot, a pile of branches, a woodpile or the
corner of a shed, or by setting up an old-fashioned hay stack. The tripod can reach
about 2 m high. Replace the hay up to several times a year.
A vole pyramid of about 1.5 m in height can be built by using long branches as ribs
that can be placed together at the top. At the bottom attach a grid of iron or branches at
a height of about 30 cm. The grid is covered with pruned wood. Underneath some straw
or hay can be put and grains scattered for voles. A plastic tube can be placed into the
construction to add grain. It is important to offer dry shelter in fall and winter inside
woodpiles, branches or grass, preferably near main food sources such as under nut and
fruit trees, oak or chestnut. A corrugated sheet laid flat, a pile of stones, some pipes or
old roof tiles casually stacked on top of each other all make excellent hiding places. An
additional advantage of sufficient hiding possibilities is that in winter fewer voles will
look for shelter indoors.

Perches
Little Owls often hunt from a perch like a wooden fence post. But it can also be a low
tree branch, the roof of a shed or a fence offering a clear view of the surroundings.
A wide range of posts and other perches within a territory are desirable. On grassland
plots, Little Owls mainly use wooden fence posts with a height of at least 1.5 m as a
perch. Mobile perches can be used in mown parts, in relation to mowing management,
offering an ideal view of prey.

Ponds
Ponds have a great attraction for animals and therefore form a good hunting ground.
Amphibians like frogs, toads and salamanders, but also dragonflies, lay their eggs there.
Birds and mammals come to drink or bathe. All kinds of aquatic animals live in the
water. Species-rich riparian and water vegetation is also attractive to many butterflies or
(water) insects. In the past they served, among other things, as the drinking water
supply for cattle. If properly constructed and managed, ponds can accommodate a lot of
life. There are several suitable locations for digging a pool or pond so it is fed by seepage,
rainwater or groundwater on an impermeable layer, guaranteeing water is also present
during dry periods. In places with relatively nutrient-poor water, more species benefit
from the pool than when the water is nutrient-rich.
New ponds are best dug within 300–400 m of other pools, making it easier for
animals to discover such a pool compared to isolated ones. A pool in a sunny spot is
beneficial for the development of the eggs and larvae of amphibians and dragonflies.
The best location for a new pond is in a residual plot or at the edge of a lawn. The size

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 11.2 Conservation Strategy for the Little Owl

and depth of a newly dug pool depend on the animal species for which it is intended. It
is important for the Little Owl that the pool is attractive and suitable for prey such as
water beetles and frogs. The minimum surface area is about 50 m2, or a pool with a
diameter of about 8 m. The depth should be at least 80 cm to account for frost and also
depending on the water supply. If the bottom of the pool is below the groundwater
level, make the deepest point of the pool 1 m below that level. Apply an impermeable
layer like loam or clay if the soil is permeable and the bottom of the pool is above
groundwater level. This can be done using a combination of foil and a layer of soil of
20–30 cm. Bentonite mats are also excellent for making a pond bottom watertight.

Vegetable Gardens
In yards, gardens with both ornamental plants and vegetables and wildflowers are
usually present. The flower garden contains many plants and shrubs that are attractive
to butterflies. The butterflies lay their eggs on the host plants and the caterpillars eat the
leaves. What is good for butterflies is also positive for other insects. The vegetables and
wildflowers that are present in the vegetable garden ensure prey all year round. All this
invites the Little Owl to visit the vegetable or flower garden regularly during food
flights. By carefully selecting the composition of flowers and vegetables, flowers can
be present, with nectar and pollen, for much of the year. Native species generally
contain more nectar than cultivated plants. Many wild species also have great orna-
mental value for the garden. Native wildflowers are attractive not only to humans, but
also to countless animals. Nettles are an important food plant for caterpillars. Give
them a chance to grow where they do not cause any nuisance.

Measures to Improve Breeding and Roosting Opportunities

Another bottleneck is the lack of sufficient breeding and roosting opportunities. Some
Little Owls change nest sites in successive years probably as a reaction to parasites in the
nest (Van Harxen et al. 2017). Breeding cavities also disappear or become unsuitable.
Optimally there are several nesting opportunities available, allowing choice. Insufficient
housing can be remedied in the short term by improving existing cavities in trees and
buildings or by installing nestboxes. In order to ensure nesting opportunities in the
longer term, it is advisable to properly maintain the buildings and wood stands with
breeding cavities and if necessary plant new trees. High-stem fruit like apple, pear, plum,
cherry and nut trees are highly appreciated for breeding and roosting in their hollows.
Old apple, pear and nut trees often contain suitable breeding cavities from an age of
50 years and not only in the trunk, but also in thick side branches. Old fruit trees are
therefore very valuable and should be preserved and properly managed. It is recom-
mended to choose different heritage fruit varieties, as they also flourish without
pesticides. When choosing trees consider their suitability for soil type, and disease
resistance and pollination aspects, because not all species and varieties grow equally well
everywhere. When planting trees, space new trees sufficiently far apart and support the
trees with one or two tree poles. Preferably don’t use creosoted posts for this, but
environmentally friendly preserved posts or posts of oak, sweet chestnut or acacia wood.

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Fruit Trees
To maintain a fruit tree, it is very important to prune regularly. Optimal maintenance
for a maximum fruit yield is not always favorable for owls. Cavities are formed by
rotting branches that have been sawn off or blown away. The frequency of mainten-
ance therefore depends on the goal. It is important to first draw up a pruning plan
before choosing which trees to use primarily for the owl and which for the fruit. When
the fruit trees are maintained in favor of cavities, it is sufficient to prune every two
to three years. If fruit production comes first annual pruning is crucial. The cavities
that develop in the trees are mainly a result of the rotting of hackneyed branches. The
rotting process is therefore favorable for the Little Owl. Do not use an anti-rotting
agent on the cut surfaces. By keeping the branch stumps at 3–10 cm, cavities are
created earlier than when you cut the branches off completely. Keep old and dead
trees as long as possible as they often provide good cavities for roosting or breeding.
Plant a replacement tree right next to a dead tree. Remove trees that have died of
cancer. Alternatively, scratching out cavities inside old and rotten trees can yield
excellent cavities.

Pollard Trees
Pollard trees are typical for western European landscapes. The best known pollarded tree
species is the willow, but also alder, ash, oak, linden, poplar and chestnuts occur as a
pollarded tree. There are excellent breeding cavities in pollard trees. Cutting them
regularly prevents the pollard trees from falling during a storm or the head of the tree
from splitting, causing the hollow in the trunk to become useless for owls. The most
suitable locations for planting new pollard trees are along ditches and roadsides. The
roots re-inforce banks. Of all pollard trees, the pollard willow is the easiest to plant by
sawing a straight branch of shoot material with a length of 2.5–3 m. Cut the bottom
obliquely. Peel strips of bark from the bottom meter. Put the stake about 1 m deep in
the ground. Plant new pollard willows in sufficiently moist soil and at a distance of
4–8 m. The same can be done with poplars. For other types of pollard trees, rooted
young trees are planted. In very windy places, two or three tree poles are needed next to
the new pollard trees, secured with tree straps. Protect the new plantings against being
eaten by providing individual tree baskets. Another possibility is to place a grid 1 m
away from the row of trees. After some years the grid can be removed unless horses or
goats are used for grazing. For proper maintenance of pollard trees all the branches of
the tree should be removed once every four to six years; alder should be cut every five to
seven years and ash once every seven to ten years. It is recommended to check for any
burrows present before pollarding a row of trees. Do not truncate all the trees in the
same year in view of the temporary unavailability of hiding places.
To mitigate disturbance, perform the work in winter, preferably before the end of
the year. Prevent cavities from being crushed since there is a danger of them being
trampled during pollarding. If a pollard willow is rotting or has mold inside, it is
possible to create extra breeding places by scratching out cavities. However, larger
burrows are preferred, because these remain tidy for longer and give nestlings
more room to move. A narrow, curved corridor between the flight opening and the

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 11.2 Conservation Strategy for the Little Owl

actual breeding cavity is ideal. Between the sludge-covered bottom and the top of the
cavity there should be 15–20 cm of space. The round flight opening should have a
diameter of 7 cm.

Solitary Trees
Solitary trees with hollows either offer excellent breeding opportunities or are perfect
places to install a nestbox, especially if it has horizontal branches. Little Owls also often
use solitary trees as roosts and for foraging. The presence of one or more solitary trees in
the yard is therefore an enrichment of the territory. Consider planting one or two
solitary trees like walnut, oak, beech or chestnut when they are lacking. Natural cavities
with openings that are too large are uninteresting to owls when rainwater can flow into
the cavity or when too easily accessible for predators. This problem can be solved by
partially sealing the opening with a zinc or wooden plate.

Buildings
Instead of nesting in natural hollows in trees, Little Owls often use niches, crevices and
small spaces in sheds, stables or pens. Those buildings can stand in the yard, but also in
the fields such as milk sheds and shelters for cattle. In some regions Little Owls more
often breed in sheds than in trees (Šálek et al. 2016) opting for small spaces, e.g., under a
loose roof tile, in the space behind a windbreak, under the bottom row of tiles or under
a reed or corrugated roof. Cavities in walls due to missing bricks are also favored if the
space has an entrance opening of about 7 cm. In addition to breeding, cavities are used
to roost. Buildings must be accessible at all times. Make sure that during the breeding
season the parents can fly in and out, day and night, by leaving a window or door open,
if there is no other access. Be careful during restoration of a shed not to close off access
to potential breeding places. It may be worth checking out other options that keep the
shed accessible, like a concrete nestbox that can be installed in the wall.
In many yards there is a disused shed. By making existing spaces in buildings
accessible, additional housing options can be created. The opening can be round or
square, with a diameter of up to 7 cm, possibly via some roof tiles or corrugated iron,
giving the owls access to the space under the roof. Making an entry hole in the facade
takes more work, but the birds get easy access to the entire space at the attic. Such an
entrance opening should not be made on the south side of the roof. In case of poor
ventilation the space under that side of the roof can become too hot on sunny days.

Nestboxes
When the number of breeding cavities in trees and buildings is a limiting factor it is
recommended to install at least two nestboxes, one for nesting and one for roosting.
Little Owls like to use nestboxes to nest in, as long as they meet specific criteria
(Gottschalk et al. 2011). They should provide enough nesting space, be dark inside
and watertight. Adequate ventilation openings must be provided. The nestboxes must
be protected from rain and wind. There must be a potential exit for fledglings, prefer-
ably via a second entrance to the cavity.

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Even during the breeding season, boxes are used as roosts by the male. If other suitable
roosts are missing, extra nestboxes do no harm and can be a solution to competition from
Jackdaws. Furthermore it is advised to also install one or more nestboxes in the area or in
a neighboring yard for dispersing fledglings (Habel et al. 2015).
To improve the chances of breeding success, it is important to place nestboxes in the
right place (Gottschalk et al. 2011). The best place for a nestbox is in a quiet corner of
the yard with plenty of cover in the vicinity in the form of trees or accessible buildings.
Nestboxes can be placed in a solitary tree, against the facade of a building or behind an
opening in the facade, or in a barn. An indoor site has the advantage that the nestbox
lasts longer, because it is not exposed to the weather. Avoid busy places in case
fledglings inadvertently jump out of the nest and are easy prey for cats and dogs. Do
not hang nestboxes in full sun to avoid the temperature rising too high, causing the
embryos or young to die (Beersma and Beersma 2000, Van den Burg et al. 2003, Widmer
2011, Grüebler et al. 2014). When placed in a tree, hang the nestbox in the shade of the
foliage. Beware that the entrance opening is not covered by dense foliage. When
hanging the nestbox on a wall choose the shadow side. The entrance should not be
directed towards the most rainy wind direction to prevent possible rain getting inside.
To let the owls take advantage of the spring sun, complement the entrance with a
balcony in a southeast direction.
Little Owls are not picky about whether or not to accept a certain type of nestbox.
The diversity in natural nesting sites proves that they are not very critical. In practice,
therefore, various types of nestboxes are used. The classic model is long and sturdy and
measures about 80  25  25 cm. The interior contains two bulkheads with a round fly-
through opening directly behind the vestibule. These partitions form a kind of air lock,
behind which the dark breeding space is situated. In this way the nestbox is made less
attractive for nest competitors such as Jackdaws Corvus monedula and Stock Doves
Columba oenas. There is also a model equipped with an entrance and exit, through
which the owl can flee in the event of an unwanted visit from a predator like the Stone
Marten (Marié and Leysen 2001, Vanden Wijngaert 2017). Wood as the base material,
painted on the outside, is preferred because of its good moisture regulation. Multi-
entrance roosting places are preferable and should be developed (Bock et al. 2013).
The height at which the nestbox hangs above the ground is not important for the
owls. In practice a height of 2–4 m is sufficient but try not to make it too difficult for
inspections and make sure the nestbox is easily accessible, facilitating nestbox control.
Place the nestbox in a tree in such a way that it rests on a branch, allowing fledglings
that are not yet capable of flying to walk. If that is not possible, tie a thick cut branch
against the balcony so that the owlets still have the opportunity to move back and
forth. The back of the nestbox should be slightly lower than the front, preventing the
eggs rolling against the front and breaking when adults go in and out. Provide the
bottom with a layer of absorbent material, for example sawdust, peat or old wood chips,
in which the owl can scrape a hollow. Long straw or hay are unsuitable as ground cover.
Winter is the best time to install nestboxes, before the start of the courtship and
breeding season in February or March. Good maintenance is essential for the successful
use of nestboxes. Cleaning, maintenance and possible replacement of nestboxes should

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 11.2 Conservation Strategy for the Little Owl

always take place after the breeding season, unless there is an urgent reason to do so
sooner. When inspecting nestboxes annually, pay particular attention to whether
they are still watertight, the strength of the construction and the fixing points. Check
that the nestbox is also connected to a wire attached to the tree, preventing it from
falling from the tree if the screws or nails give way. Clean nestboxes annually after the
breeding season and check whether the nesting material threatens to block the exit.
Remove all branches from nestboxes that have been occupied by Jackdaws or Stock
Doves. Remove all debris and deposit a fresh layer of absorbent substrate at the bottom,
to reduce parasite survival (Van Harxen et al. 2017). Cleaning the nestbox during the
breeding season reduces the parasite pressure at that time. Nestboxes where no Little
Owls have nested in the previous year are free of parasites in the following breeding
season. Hence, cleaning nestboxes after the breeding season results in much fewer flea
droppings (and therefore fleas) and probably also far fewer parasites in nestlings. Small
nestlings mainly suffer from blood-sucking flies like Carnus hemapterus, larger ones are
more bothered by Louse Flies Hippoboscidae spp. For (comparative) research into the
presence of parasites it is therefore important to make a connection with the age of the
nestlings. In situations where a nest (box) cannot be cleaned, it is advised to install a
second box, so that the Little Owls have the opportunity to choose a clean box.
The most favorable time to inspect and clean the nestbox is from September to
November. From December on nestboxes are occupied.
To address the alarming situation of declining Little Owl populations in some areas,
conservation plans have been set up at an international level (Leigh 2001b), at national
levels in Austria (Kirchberger 1988), France (Génot et al. 1999, Lecomte et al. 2001), the
Netherlands (Plantinga 1999) and at regional levels in Flanders (Van Nieuwenhuyse
et al. 2001b), in Geneva (Meisser and Albrecht 2001), and in Groningen, the
Netherlands (van’t Hoff 1999). In many countries, actions are underway to protect
and manage Little Owl habitat, such as through the maintenance of pollard willows
(Loske 1978, Bultot 1996, Bultot et al. 2001), oak hedges (Meisser 1998, Meisser and
Albrecht 2001) and orchards (Harbodt and Pauritsch 1987, Grimm 1989, Juillard 1997).
Since 1970 (Schwarzenberg 1970), many nestboxes have been established, partly to
compensate for the lack of natural cavities and also for scientific study. To reduce
vehicle collisions on roads, special action on perches along roads has been found to
decrease mortality (Hernandez 1988). Optimization of existing territories by improve-
ment of prey abundance and accessibility through the rotation of grassland manage-
ment, and the placement of additional perches (similar to those for shrikes (Laniidae)
(Van Nieuwenhuyse et al. 1999)) have been suggested to help conserve remaining owl
populations. To improve postfledging survival, concrete or PVC tubes can be laid on
the ground below or near breeding cavities. These tubes are used by fledglings as shelter
and prevent them from getting trampled by cows (van Harxen and Stroeken 2003).
Schwarzenberg (1970) developed the original artificial nestbox, which is basically a
wooden pipe. Its use has spread widely in German-speaking countries. Such nestboxes
are made by means of narrow planks, held together by glue. The length of the pipe is
80–100 cm, it has an inner diameter of about 18 cm, and the diameter of the entry hole
is 6.5–8.0 cm. As a rule, the upper part of the pipe juts out somewhat overhead, creating

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a small overhang over the entry hole. In the rear part of the pipe a hole for checking and
cleaning of the nestbox is present. It is possible to make an analogous square nestbox of
boards or to use a piece of empty tree trunk. Usually nest pipes are protected against
penetration by Martens by installing two front walls at a distance of 7–8 cm with entry
holes at different sides. On the side opposite to the entry hole, ventilation holes with
diameters of 1.5–2 cm are made. Predators are thought to be unable to do two tight
turns in such a narrow space while the Little Owl has no entry problems. Pipes are
attached above or below horizontal branches of trees or under roofs of buildings. In the
Czech Republic the species has occupied plastic boxes shaped like cubes with sides of
25 or 30 cm (L. Oplustil personal communication). Bultot (1996) developed nestboxes
using recycled 12-bottle wine cases.
More and more nestboxes are equipped with either video or photographic tools to
automatically monitor the visiting frequency and the prey brought in by the parent
owls during the breeding season (Juillard 1984, Blache 2001). The observed plasticity in
the choice of nesting sites allows the possible development of the more diverse variants
of artificial nestboxes.

Measures to Improve Safety

By taking targeted measures or by omitting activities, the safety of both young and
adults in and around the yard can be mitigated. Fledglings leave the nest after 30–35
days. They cannot fly yet and scurry around helplessly, often on the ground. The
parents keep an eye on them, but can’t really protect them. Fledglings face many
threats, like cats, dogs, polecats and Stone Martens. In order to escape from pets and
predators, the juveniles, especially if they do not fly, should quickly find shelter. If the
nestbox hangs in a tree or on a wall, an exit opportunity is needed. If a good run-out
over tree branches is missing from a nestbox in a tree, branches or a plank can be
attached next to the flight hole. In addition, provide hiding places under or in the
vicinity of the nest, such as a pile of stones or tiles, grid posts or pallets. A firewood
store, a pile of branches or a pile of dry pipes are also good shelters, as are rabbit holes.
A sturdy potato box with 7 cm holes made just above the ground and completely
covered with branches also suffices as a hiding place. Dense vegetation can also provide
protection. For the fledglings, the presence of trees in the vicinity of the breeding cavity
is important. As soon as they can fly, at twilight they go to rest and take shelter in trees.
When a little older they stay there during the day. Cavities also offer the young extra
hiding options in bad weather conditions. Fledglings that have fallen out of the nest or
have left the nest too early can easily be put back in the nest, preventing them from
becoming victim to predators. Once they have flown out, it makes little sense to put
them back in, because then they will soon leave the nestbox again.

Barns
Fledglings born in a barn or stable can also be found on the ground. Offer them shelter
in the form of a pile of wood, boxes or other material that can give them extra
opportunities to hide. Make an entrance to the cavity as needed. Check that they can
get out on their own. Pets such as cats and dogs can take plenty of victims. Cats stalk the

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 11.2 Conservation Strategy for the Little Owl

young at night and at dawn while looking for food on the ground. A bell around
the cat’s neck can warn the owlets. Try to keep cats indoors at night as much as
possible during the fledgling period, preventing predation and also protecting the
food-seeking fledglings without hindering parents from hunting. A dog can injure
the owls internally or bite them to death when given the chance. Keep dogs inside in
the months of June and July and keep them away from the nesting site as much
as possible.

Hiding and Roosting Places

Adult Little Owls like to sit alone in sheltered places during the day, where they have a
good view of the surroundings. In calm weather they can be found on a woodpile, on
the ridge of the roof or in a hollow of a pollard tree. In fall and winter they prefer to sit
in a cavity or nestbox with an entrance to the southeast to enjoy the sun. They often
change places, and male and female have their own preferences (Bock 2011, Bock et al.
2013). During the breeding season, the male often uses a different cavity from the one
where the nest is located. It is important to offer variation in cavities in pollard, fruit or
solitary trees. Make accessible spaces in sheds and install at least one extra nestbox as a
roost. When pollard trees in the yard are used as shelter, consider not truncating all the
trees in the same season. Owls are often already in their breeding cavities in pollard
trees from February or March, so preferably do any work before the turn of the year.
Also be careful when clearing fallen trees, since owls may be hiding in them. They rely
on the protection of the cavity and do not always fly away when the tree is cut into
parts. They can be injured or killed during sawing. Sometimes they also remain in place
during transport of the sawn tree parts and become displaced. Check cavities carefully
before cutting a tree.

Unnatural Dangers
Car traffic is one of the main causes of death of Little Owls, low flying owls are often
victims. This can happen while using the verge as a hunting ground, but also when
crossing the road to bring prey to the nest. They often use fixed flight routes. Fledglings
can be killed if they search the road for insects that have collided with cars. This not
only happens on main roads, but also on quiet agricultural roads. When there is less
food available in the yard, roads become more important as hunting grounds. By
guaranteeing sufficient prey in the yard all year round, the necessity for the owls to
hunt or pick up insects by the roadside is reduced.
It seems an obvious solution to place the nestboxes as far as possible from roads. The
question, however, is whether it actually helps. Roadsides are places where many voles
occur and hence are an attractive but dangerous hunting ground, especially if there are
poles beside the road. Remove fence posts and other perches from the roadside if
possible. Make sure the vegetation there is somewhat roughened, making it less attract-
ive for hunting voles. If the fixed flight route to and from the nest crosses a road, the
birds risk being hit. Hence, force the birds to fly higher during the crossing by placing a
high branch wall or Benjes hedge. In the longer term, this can be achieved by growing a
tall hedge.

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Drowning Risk
Wells, cattle troughs, tubs, bathtubs, and even buckets and watering cans, which are all
open at the top, have steep walls and are filled with water or some other liquid, pose a
high risk of drowning, especially for fledglings or females that have their first bath after
a month of uninterrupted incubation. The birds fall in when they see something
interesting or when they want to drink water and can no longer escape. Since adults
also drown in this way, the danger exists all year round. There are several options for
avoiding drowning in a water bowl, in the first instance by preventing the owls from
falling in. To do this, cover the water container during the breeding season with
chicken wire or turn the trays over so that they cannot fill up. If the owls fall into a
water trough, they must be able to escape. Keep the water bowl full and put a shelf in it
or hang a cloth over the edge so they can climb out on something. Replace the plank in
time so that it does not sink from saturation. A second shelf diagonally against the wall
can allow the owl to escape on its own.

Poisoning
An invisible and unintentional, but deadly, danger to Little Owls is poison. This does
not only concern rodenticides, but also insecticides, which are often used in agriculture
to control harmful insects such as leatherbacks and grubs in the grassland or Colorado
Beetles in potato fields. Flies are also controlled with insecticides in stables. Owls are
harmed through eating contaminated prey, which easily happens, because the prey is
dazed or dead from the poison and is easy to catch (Beersma and Beersma 2001). The
poison accumulates in the owls’ bodies and is harmful for egg production and the
survival of the embryos, but ultimately also for the parent birds themselves. In addition
to this direct effect on the owls, the use of poison also has an indirect negative effect by
limiting local prey abundance.
Nuisance from voles can be caused by too much food being available to them. Store
attractive food such as chicken feed out of the reach of rodents. If there are too many
rodents do not use rodenticides but live traps, and freeze the killed voles to keep them
as supplemental feeding during food scarcity, after heavy snowfall. To avoid the risk of
poisoning the owls, insecticides should be avoided and if absolutely necessary replaced
by biological methods. Herbicides have no direct effect on Little Owls, insects and other
animals but do have an indirect effect. They cause wildflower richness to decrease
sharply, and as a result shelter and food sources for insects are lost, thereby reducing
the amount of insects and the owls’ food supply.

Traps
Little Owls are curious, inquisitive birds that like to look around in cavities and scurry
around. Especially after the disappearance of nest sites by demolishing, for example, a
shed or cutting down a breeding tree, the owls will look for a new nesting place and are
in danger of being trapped. Even when threatened, they crawl away into holes and can
get stuck. Chimney pipes and telephone poles are a known danger to the adults when
looking for a new nesting place or in cases of imminent danger, they can hide in them
and get trapped. Only if they are discovered and liberated in time will the owls survive.

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To avoid being trapped, it is best to make chimney pipes inaccessible by covering them
with a piece of gauze. An additional advantage is that the gauze blocks Jackdaws and
House Sparrows Passer domesticus. Besides chimney pipes, fruit nets also cause owls to
become easily entangled and die.

Food Supplementation
Food has been shown to be a crucial limiting factor for Little Owl populations.
Supplementing extra food to breeding pairs has been used experimentally, mainly in
Denmark, Switzerland and Germany.
Thorup et al. (2010) investigated the likely reasons for the population decline in
Denmark by identifying patterns of local extinction and estimating demographic par-
ameters affecting local survival, focusing on changes over time and their relationship to
habitat characteristics. Experimental food supplementation to breeding pairs increased
the proportion of eggs that resulted in fledged young from 27% to 79%, supporting the
hypothesis that the main proximate reason underlying the ongoing population decline
was reduced productivity induced by energetic constraints after egg-laying.
Conservation efforts in Denmark have been suggested that target enhancement of food
availability during the breeding season.
Perrig et al. (2014) studied the underlying mechanisms and associated consequences
of differential quantity and quality of fledglings produced under varying nestling food
supplies and the way they affect the number of offspring that recruit into the breeding
population (see Section 10.2.1 for details) and concluded that habitat quality affects
Little Owl productivity and offspring quality, and ultimately, population dynamics.
Jacobsen et al. (2016) studied the effect of food limitation during the breeding season
on the decline and almost extinction of Little Owls in Denmark (see Section 10.2.1 for
details). The results indicate that food provisioning not only led to increased repro-
ductive output, but also to decreased working effort, which in turn may improve
adult survival.
Perrig et al. (2017) studied postfledging survival and the energy supply to the
growing juveniles. Using a food supplementation experiment, they determined the
effect of variation in food supply on the survival of juvenile Little Owls from hatching
to two months post fledging. Experimental broods were food-supplemented for 36 days
during the nestling and the early postfledging period. The fate of 307 juveniles (95 of
them provided with extra food) was determined by nest monitoring and radio-
telemetry. In unsupplemented birds, the rates of survival measured at five-day intervals
were lowest during the nestling stage, remained low during the early postfledging stage
and steadily increased after about two weeks post fledging. Food supplementation
substantially increased nestling survival, but Perrig et al. (2017) detected no direct
treatment effect on postfledging survival. Instead, they found a strong indirect effect
of food supplementation, in that fledglings of good physical condition had markedly
higher chances of surviving the postfledging period compared with those in poor
condition. Experimental food supplementation increased survival over the first three
months from 45% to 64.6%. This suggests that energy reserves built up during the
nestling stage influence postfledging survival and ultimately parental reproductive

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output. The low nestling and postfledging survival shows that the early life-history
stages constitute a crucial bottleneck in the reproductive ecology of Little Owls. The
strong treatment effects on the number of independent offspring indicate that natural
variation in food supply is an important determinant of spatio-temporal patterns in
Little Owl demography.
Grüebler et al. (2018) experimentally increased the food intake of Little Owl nest-
lings in two habitat types differing in food availability and used unsupplemented
broods as controls. The food supplementation experiment allowed the effect of habitat
type to be disentangled from the effect of the nestlings’ food intake on parental
provisioning behavior. Camera traps recording each parental visit allowed visiting rates
and diet composition to be quantified by applying a hierarchical multinomial model
explicitly accounting for the observation process. Food supplementation caused
parents to switch to smaller food items and to increase visiting rates, resulting in similar
biomass brought to nestlings. Irrespective of the food supplementation, parents in low-
quality habitats fed 63% of the biomass compared to those in high-quality habitats.
Grüebler et al. (2018) found an increase in nestling survival rates in response to food
supplementation in low-quality habitats, but not in high-quality habitats. They showed
that habitat quality affects the biomass of prey delivered to the brood, whereas the
nutritional state of the brood affects prey selection or the foraging modes of the parents.
Reproductive output directly reflected habitat quality in terms of food availability,
identifying food as the main factor underlying differential reproduction within and
between populations.
Food supplementation as such was done in different ways and sometimes in com-
bination with partial cross-fostering, i.e., switch half of the nestlings of similar age
between two nests to control for different factors that are inherent to family units (e.g.,
genetic factors, common nest environment) and that can cause dependence of survival
rates among family members. Randomization of the treatment and partial cross-
fostering prior to experimental food supplementation partially controlled for such
effects.
Thorup et al. (2010) supplied two pairs during the 2005 breeding season and three
different pairs during 2006 with supplemental food (one to three dead domestic mice or
newly hatched chickens per day). Food was supplied daily from hatching until the
young fledged.
Perrig et al. (2014) conducted a partial cross-fostering and food supplementation
field experiment. Partial cross-fostering allowed supplementing only some nestlings
from the same family. Thus, intrinsic factors acting on survival and phenotypic devel-
opment could be statistically controlled for. Two synchronous broods were denoted as
partner broods. One of the two was randomly assigned to experimental treatment (i.e.,
food supplementation). Depending on brood size, one or two nestlings of similar age
and body weight were exchanged among partner broods, keeping brood sizes constant.
Exchange took place at ca. 14 days of age. Conservation concerns prohibited earlier
manipulation of nestlings. Food supplementation started after exchange. All broods
were visited every second day for 36 days. Thus, visits continued beyond fledging at
ca. 30 days of age, when nestlings leave the nestbox for the first time. After fledging,

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 11.2 Conservation Strategy for the Little Owl

juvenile Little Owls stay within close proximity of and regularly return to the nestbox.
During visits, a total of 480 g of dead laboratory mice per nestling were deposited inside
the nestbox of supplemented broods, 20 g per visit and nestling for the first six visits,
30 g per visit and nestling thereafter. Food supplementation represented a ca. 40%
increase in food compared with natural food supply (Juillard 1984).
In the study by Jacobsen et al. (2016), several pairs received supplementary food.
Supplemented food consisted of dead 1-day-old (30 g) poultry chicks (Gallus gallus
domesticus), from a commercial poultry hatchery, and laboratory mice (Mus musculus).
The food was provided from March until August every day or every second day at a rate
of approximately 90–210 g per nest per day.
Perrig et al. (2017) completed the data of Perrig et al. (2014) with additional broods
that were not cross-fostered (due to a lack of a simultaneous partner brood). A total
of 93 nests were included in the analysis, of which 31 were food-supplemented.
Supplementation was continued as for the nestling period, and mice were consumed
as quickly as they were in the nestling phase. Perrig et al. (2017) defined the
postfledging stage as the period from the age of 30 days to the onset of dispersal at
the age of 95 days.
Given the fact that supplemental food has a significant added value for nestling
survival, it can be considered as a valuable conservation measure, either by supplying
dead prey or by improving habitat conditions for prey, as illustrated above.

Re-introduction and Re-inforcement

As the species has disappeared or declined in many European regions, some experi-
ments with re-introduction (where the species had been extirpated and owls were
released) or supplementation (when the species was still present but extra owls were
released to increase the population) have occurred (Table 11.1). These efforts took place
mainly in Germany, France, Switzerland, the Czech Republic and Denmark.
The project in Jura Souabe, Germany (Mohr 1989) was started in 1990 with 15 Little
Owls hatched and raised in captivity and then released into the wild. This experiment
was a re-inforcement of a wild population that increased from two pairs in 1982 to 25
pairs in 1990 thanks to nestboxes (Mohr personal communication).
In Hesse the project was a re-introduction (Stahl 1982). The captively raised owls
were released during September 1980 and 1981. They were placed in nestboxes with a
metal balcony, the balcony helping the young birds to stay inside and acclimate to
their environment. In 1982, two released pairs bred in nature, producing nine
fledged young.
In Lower Saxony (Möller 1993), a re-introduction effort was started in 1989; five
Little Owls were released the first year and 10 birds released every year thereafter. The
young were released from an aviary containing young and adults from which only the
young could fly out. In 1992, seven pairs bred in nature, but with a low breeding success
of 1.57 fledging/breeding pair.
In Mecklenburg, the project was also a re-introduction (Bönsel 2001). A total of
42 Little Owls were released in 1996, 49 in 1997, 49 in 1998 and 31 in 1999. In 1999,
seven territories were surveyed and four breeding pairs were found.

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Table 11.1 Different projects of Little Owl re-inforcement and re-introduction in Europe

No. of owls Type of

released Year experiment Final result Region Country Author

79 1983–2002 Re-introduction 0 BP in Harz Germany E. Kartheuser (pers. comm.)

nature
102 1983–2002 Re-introduction ? Göttingen Germany F.-K. Schöttelndreier (pers.
comm.)
266 1987–2002 Re-inforcement 4 BP in Brandenburg Germany P. Haase (pers. comm.)
nature
109 1978–1990 Re-introduction 0 BP in Coburg Germany Leicht (1992)
nature
15 1990 Re-inforcement ? Jura Souabe Germany Mohr (1989)
? 1980–1981 Re-introduction 2 BP in Hesse Germany Stahl (1982)
nature (Limburg)
31 1987–1990 Re-introduction ? Hesse (Fulda) Germany K. Burbach (pers. comm.)
25 1989–1992 Re-introduction 7 BP in Lower Saxony Germany Möller (1993)
nature
? 1988–1999 Re-introduction 3 BP in Thuringe Germany F. and F. C. Robiller (pers.
nature comm.)
171 1996–1999 Re-introduction 4 BP in Mecklenburg Germany Bönsel (2001)
nature
92 1982–1986 Re-inforcement O BP in Bern-Freiburg Switzerland K. Robin (pers. comm.)
nature
85 1993–2002 Re-inforcement 1 BP in Alsace France Génot (2005)
nature

BP in nature means a breeding pair containing at least one released owl.

 11.2 Conservation Strategy for the Little Owl

In the Coburg region where the Little Owl was extirpated (Leicht 1992) 109 young
reared in captivity were released between 1978 and 1990 by differing methods. Some
owls were released directly in spring and fall, others were reared through the adoption
by wild owls of young born in an aviary and kept in contact with the parents in the
aviary. Ultimately, no breeding pairs were recorded in the region.
In Thüringen, in particular the Weimar region, Robiller and Robiller (1986, 1992)
conducted a re-introduction experiment from a complex of 16 aviaries built as veterin-
ary equipment and to study Little Owl behavior. The species disappeared around
1970 from the Weimar area. Thanks to the few adults released from captivity and the
wild birds probably attracted by the birds in the aviary, two pairs bred in the wild and
produced 15 fledglings between 1988 and 1992.
In the Göttingen region, from 1999–2003, 79 young Little Owls were ringed and
released in an area of about 200–300 km2 (Kartheuser personal communication). As of
2001, there were still no wild Little Owls forming a population, while some single birds
stayed for some months in that region. The habitat for Little Owls at the release area is
too isolated from the densely populated areas in Westphalia, more than 100 km away
(H. Illner personal communication). The climate of this area is cooler and there is on
average much more snow than in Westphalia. The number of owls released are thought
to have been too low and the optimal habitats too isolated and it remains unclear
whether the available food and nesting sites were sufficient.
In Brandenburg, and in Haveland in particular, an experiment began in 1987 to supple-
ment a declining population (Haase 1993). Between 1987 and 2002, a total of 266 birds were
released from captivity; 90 in spring (March–April) and 176 in fall (August–September). The
survival of the owls after release was better in fall than in spring. During this period, 9 birds
were recorded as breeding in nature and 15 died (of which 12 were killed and one wounded
by a collision with a vehicle). Different methods for release were used. A mobile aviary was
put in place, young staying inside for 7–10 days prior to release. The owls were fed with live
prey before the aviary was opened at twilight and kept open over several days. A little mobile
cage was put in place for release, the young stayed inside for 14–26 hours and were fed, and
the cage was opened at twilight. Young owls born in captivity were put in wild clutches of
the same age and not exceeding four to five young, and thereafter raised by the wild adults.
This method was limited due to the low number of Little Owls in nature.
In Sarre, 40 Little Owls were released between 2001 and 2003 and two females bred
in 2003 (Stelzl personal communication).
Most of these experiments had no real monitoring; we would encourage observers
involved with experimental studies to routinely provide status reports on these efforts,
so that we can collectively gain from specific insights.
Since 1993, a re-inforcement experiment has been undertaken in the Northern
Vosges Biosphere Reserve (France). This re-inforcement took place in a population that
was isolated due to habitat fragmentation, had weak breeding success and in which the
demographic viability had been analyzed (Letty et al. 2001b). The aim was to avoid
extinction due to inbreeding (Frankel and Soule 1981). This project is part of a program
of global activities aimed at protecting the Little Owl, in the framework of the national
conservation plan of the French Ministry of Environment (Lecomte et al. 2001).

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Table 11.2 Details of the numbers of Little Owls released in Alsace (France)

Number of birds released

Year 1993 1995 1996 1997 1998 1999 2000 2001 2002 Total

Nestbox with 5 6 4 2 17
balcony
Nestbox 12 17 8 13 9 59
Complete wild 3 4 2 9
nest

Total 8 10 6 2 12 17 8 13 9 85

Genetic fingerprinting was carried out to analyze the existence or absence of

genetic links between owls in different geographic regions in order to find out
whether these birds could be cross-bred in captivity. This was crucial to establish the
degree of relationship between the birds. The analysis revealed that genetic variability
was not unique for each region. Two unrelated birds from the same region could be
more distant (genetically speaking) than two birds from different regions. Even though
the sampling of nine individuals was restricted, the marked similarity coefficients
coming from owls in different localities indicated the existence of an important genetic
proximity between the individuals studied. Such values thus reflect both the absence
of great genetic diversity locally and the homogeneity of the similarity coefficients
obtained.
Young born in captivity were either released into nestboxes or placed in wild broods
in order to complement them. A total of 85 young were released from 1993–2002
(Table 11.2) in an area where a population of about 10 pairs was living. Of 35 birds
fitted with transmitters from 1995–2001, 22 were found dead shortly after being
released, 12 disappeared from the release site and have not been traced, and one was
followed for five months before disappearing.
Four young owls were released directly into nestboxes in the northern Netherlands
in the summer of 2001. While it probably results in higher mortality than aviary-raised
owls, this method was chosen, and one pair and two males were released. The female
died during the winter (killed on a road), but the three males stayed around the release
site for one year. Finally the three males disappeared too, having dispersed or died
(van’t Hoff personal communication).
Young placed in wild broods were readily adopted by the parent birds. The birds
born in captivity were of an equivalent age to those born in the wild. Extra prey was
supplied to facilitate feeding. One owlet, born in captivity in 1996 and placed in a wild
brood of only one chick to complement it, survived the very snowy winter of 1997. This

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 11.2 Conservation Strategy for the Little Owl

male paired with a wild female and together they produced two young in 1997 and four
in 1998 from a nestbox situated 600 m from the release site.
Despite staying for three to four weeks in balcony nestboxes, the Little Owls left
them once the metal balcony was opened and did not come back later. This is the
reason why birds were ultimately released from nestboxes having no balcony.
Amongst the known causes of death, Little Owls found dead after being released into
the wild were either victims of a predator or crushed by a tractor. The mortality rate
among the young born in captivity and released was high because the birds spent a lot
of time on the ground.
Due to the difficulties in producing young in captivity and the logistics of such
experiments, the decision was taken to stop re-inforcement and wait for the results of a
genetics study for more details about the structure of the population and to establish a
demographic and genetic viability model (Génot and Sturm 2003).
Crucial in repatriation programs using captive-bred individuals is the selection of
suitable habitat, one that meets the species’ biotic and abiotic needs through space and
time. However, securing adequate locations does not always mean that repatriated
individuals remain at the release site. Bušina (2021) provided a record of unusual
postrelease spatial behavior and erroneous habitat selection for a repatriation program
that was launched in the Pilsen region in the Czech Republic to counter the constant
decline in the size of the local population. During observations, the absence of forest
avoidance typical for central European Little Owls, along with underground
roosting, was observed. More specifically, one of the soft-released individuals settled
in a patch of dense forest establishing a home range equivalent in size to 3.54 ha.
Within its home range, underground cavities were regularly used for diurnal roosting.
Bušina (2021) was not able to determine whether this behavior was due to captive
breeding conditions.
Some movement experiments were carried out in the Netherlands by Jacobs (2003).
Manipulation of eggs, young and entire broods were undertaken with varying degrees
of success. The manipulation of eggs (addition or replacement of eggs) yielded the most
positive results, so long as the nesting place remained the same. Moving an entire
clutch, including the brooding female, did not work. The success of adding one or
more nestlings to another nest (after death of one of their parents) was most successful
when the difference between the nestlings of both clutches was not too large and if the
nestlings were at least one week in age. Artificial feeding of the pair with dead mice,
sparrows and one-day-old chicks helped the birds. Displacement of an entire clutch of
nestlings was too risky if they were less than 10 days old.
Mortality caused by predators has been described previously in many released indi-
viduals. To counter this Alonso et al. (2011) used prerelease training of juvenile owls to
avoid predation. Anti-predator training of juveniles was tested in a sample of recovered
owls raised in captivity in Brinzal Owl Rescue Center (Madrid, Spain). Nine owls were
conditioned during their development to a naturalized goshawk and a large live rat,
whose presence was paired to the owl’s alarm call. All nine owls and seven nontrained

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 Chapter 11: Management and Conservation

individuals were then released during the late summer and fall, and radio–tracked for
six weeks to test their survival. In total, 71.4% of the trained owls survived, while only
the 33.3% of the untrained group were alive at the end of week 6. The only cause of
death that was detected was predation. Anti-predator training, therefore, seems to be
beneficial in maximizing survival after the release of juvenile Little Owls.
A re-inforcement or re-introduction program can only succeed if some key criteria
are fulfilled. An analysis of the factors explaining the decline or the disappearance of
the species in the study area should first be conducted. Wild owls should not be used,
only birds reared in captivity or taken from care centers. The identification of good
habitats in the release area through a survey is key. In the case of re-inforcement, owls
should be able to establish a connection to an existing population in the release area.
Landowners, farmers and inhabitants living in the release area should be informed
about the initiative. Good logistic guidelines for rearing in captivity are given by Stahl
(1982), Robiller and Robiller (1992), Möller (1993), Génot et al. (2000). Radio-
transmitters should be used to monitor postrelease behavior. Enough individuals need
to be released per year (because of the high mortality rate). A local conservation
management plan for the habitat should be run at the same time as the re-introduction
or re-inforcement program. A DNA fingerprinting analysis of the wild and released owls
is needed to understand the genetic diversity of the former, and the viability of the new
population. For rearing in captivity one needs to guarantee a complete separation of the
breeding pairs, avoid using owls living in captivity for a long time due to the possibility
of abnormal behavior, have an outside aviary to allow training of the Little Owl to catch
living prey and different items (birds, rodents, earthworms, insects) and training of
predator avoidance actively.
The release must happen after the owls are 80 days old, either during their normal
dispersal (August–September) or the following spring. There are no comparative
studies from which to choose a method, but for the Tawny Owl, the owls become
less suited to release after longer periods of captivity compared to younger owls
(Meyer-Holzapfel and Räder in Schönn et al. 1991). The three main methods of
release are described as: young born in captivity and placed in wild broods (if the
age is the same), the exchange of sterile eggs in nature with fertile eggs from an aviary
and putting the young with their parents in a special aviary, the same one used for
rearing and placed at the release site. The aviary is left open until the young leave the
parents and the young are released directly by placing them in nestboxes located in
good habitats.
The released owls must be ringed. There is a lack of knowledge to establish the
optimal number of owls to release per year to be successful. But because of the high
mortality among the young, we consider that a minimum of 20–30 individuals per year
during each of 10 years is necessary with the last method of release.

11.2.2.3 STANDARDIZED METHODOLOGIES

Standardizing the methodologies for Little Owl studies involves using consistent ter-
minology, survey methods, demographic data and habitat analyses. Certainly, we are

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 11.2 Conservation Strategy for the Little Owl

not urging researchers and managers to suppress their innovative talents, but rather, we
feel we must standardize core aspects of our work in order for us to maximize the
benefits from our collective efforts. Part of the standardization process starts with the
use of common terminology; in support of this, we offer definitions for some frequently
used terms in the Supplementary Glossary.

Survey Methods
Survey protocols are detailed study plans that explain how data are to be collected,
managed and reported, and are a key component of quality assurance for natural
resource monitoring programs. There are typically three primary objectives for
surveying owls: (1) documentation of presence/absence, (2) enumeration (density,
absolute or relative abundance) or (3) population dynamics (size, distribution, trends
in abundance over time). In our survey protocol, we focus on surveying to determine
population dynamics. This is the most rigorous of the three objectives, and documen-
tation of owl presence and enumeration of owls can also be determined using this
protocol. While a census is defined as a complete count of every individual in a given
population (very rarely possible in wild populations), a survey is an estimate of the
number of animals in a given area (e.g., number of territorial pairs) or of the relative
frequency of encounters with animals (e.g., number per unit transect length or time
period). Repeated surveys can be used to estimate the trend of a population. Surveys
provide an efficient way of collecting information from a large number of sites and can
also help determine the overall distributional range of a species in different seasons. The
structure of surveys is intended to reduce observer bias, and surveys are best conducted
in a standardized way to ensure reliability and validity.
Thus, for owls, as it is truly unlikely that a complete count of all individuals could
realistically be determined for any wild population (due to methods, budgets or detect-
ability of the species), a survey is a more appropriate approach to estimating numbers,
frequencies and trends (see also Thompson et al. 1998).
Survey efforts are ultimately aimed at determining the total number of territorial owls
(paired and unpaired) present in a given area. Not all territorial owls readily respond to
playback, either because they are not within hearing range when playback surveys are
conducted, or because they are engaged in other more pressing behaviors (e.g.,
hunting). It requires energy from the owls to respond to territorial intruders, and the
owl must balance its energy expenditure in territorial defense against its other nightly
and seasonal activities.
The broadcasting of prerecorded conspecific vocalizations is a common technique
used to detect secretive and vocally active raptors (Fuller and Mosher 1981). Playback is
widely used to solicit responses from territorial owls, whose elusive and nocturnal
habits typically prevent visual detection (Bibby et al. 1992). Playback is the most
common method for surveying Little Owls.
Response Rate and Detection Rate. An important aspect of surveying using
playback is quantifying the playback efficiency (i.e., the response rate). In determining
the response rate, we are trying to determine how many of the owls holding territories
that can hear the playback actually respond to it. In order to determine the response

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 Chapter 11: Management and Conservation

rate, one needs to know the locations of the existing owls (prior to playback solicita-
tion), as determined through radio-telemetry, and then broadcast the playback and
record the number of responding owls. Thus, the response rate reflects the proportion
of owls that, when within hearing range of the broadcast calls, actually respond to the
playback. For example, if 5 out of 10 owls that were within the broadcast range of
playback recordings actually responded, this is a response rate of 50%. However, unless
the owls’ locations are first determined with radio-telemetry, observers employing
playback have no way of knowing whether surveys without responses reflect the
absence of owls, whether the territory holders were outside of hearing range or whether
owls were present and did not reply.
Playback is a method that increases the chances of finding the Little Owl relative to
spontaneous unstimulated observations (Exo and Hennes 1978). The method uses the
urge of the Little Owl to answer to territorial calls. Exo and Hennes indicated that in
favorable areas, 80% to 90% of the Little Owls present can be detected in one single
inventory. Research in Mediterranean areas has shown that in the February–April
period, the Little Owl effectively responds to simulated sound in 50% of cases, for
example in Italy (Centili 2001), and that three rounds of playback yield 87% of all
existing territories. Zuberogoitia et al. (2011) give an average detection rate of 45% for
one, 68% for two, 80% for three and 95% for four playback sessions. The circumstances
in these studies with other densities and mountainous landscapes were not really
comparable with the Flemish project (Binnemans et al. 2021). A better comparison is
the situation in Zeeuws-Vlaanderen where De Smet (2019) detected 73% of the popula-
tion in the first round, 93% in the second and 100% in the third.
The detection rate is a summary of the number of owls that were recorded across a
given area, e.g., (number of individuals heard)/(number of playback sessions per-
formed) (Galeotti 1989, Sará and Zanca 1989, Centili 2001). While easier to calculate,
these summaries invariably yield rather low rates of detection, as they cannot separate
owl absence from owl silence and incorporate surveys conducted in areas of unsuitable
habitat (no owls present to respond). The calculated detection rate is inversely propor-
tional to the number of playback stations surveyed with no owls responding.
An alternative way of determining the response rate is to establish survey call points
along a transect, and conduct multiple (e.g., six to ten) visits across the survey route,
conducting playback surveys at the survey points over a given timeframe (e.g., the
breeding season). During these surveys, the observer records the responding owls and
maps their locations. During each visit across the route, it is probable that some “new”
owls (those that did not respond, or were not within earshot of the broadcast during
previous visits) would be detected. Additional survey visits would be conducted until no
new owls were detected along the transect. Analysis of the data would indicate the
number of visits required until 90–95% of the territorial owls had been detected. As an
example, for the Northern Spotted Owl, a species considered to be a relatively good
responder, some six visits across the survey routes are required to effectively detect the
resident owls. Field tests of dectectability for the Powerful Owl (Australia) have indi-
cated that it may require up to 18 visits to survey stations involving listening and call
playback to detect 90% of the territorial owls within a given area (Wintle et al. 2005).

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 11.2 Conservation Strategy for the Little Owl

Similarly, eight visits to survey stations involving listening and call playback were
required to detect 90% of Sooty Owls (Tyto tenebricosa) (Wintle et al. 2005).
Further, the sex of the owl’s vocalizations being broadcast may make a difference. For
example, if you were broadcasting the vocalizations of a male owl (this is most appro-
priate, since males are the primary defenders of territorial boundaries), why would the
resident female owl respond aggressively to another male owl (and potential mate)?
Further, if you were to simultaneously broadcast the vocalizations of a pair of owls (i.e.,
male and female), a resident owl who was unpaired (i.e., a resident single) may well be
too intimidated to respond to two intruders. However, the broadcasting of female calls
(solely) may be a good way to efficiently locate nests (as females often respond aggres-
sively to other female owls). Field surveyors should be aware that sex, and behavioral
and seasonal aspects all have a role to play in the response rates of owls.
In developing a standardized survey protocol, we examined the array of methods
currently being used. In Table 11.3, we give an overview of the factors that influence the
efficiency of the broadcasting methods. Temperature does not appear to have much of
an effect, the weather usually does and is therefore used as a criterion for broadcasting.
Most researchers do not perform playback in conditions of rain or strong wind, as
reduced owl vocalizations have been observed during those times.
Centili (2001) examined the response rate of Little Owls to playback surveys at
known occupied sites. The overall response rate between February and July
was 49.6%; three playback visits to each site were necessary to locate 87% of the
resident owls. The response rate did not change significantly during these months,
consistent with the results of Zuberogoitia and Campos (1998) and Mastrorilli (1997)
(Table 11.4). There was no significant relationship between response rate and distance
from playback station to occupied sites that were within the broadcast reach. This
suggests that playback was as effective in eliciting an owl’s vocal reaction inside a
territory as was outside of the territory. Thirty-six percent of the replies came from
occupied sites over 450 m away from playback stations (max distance 1100 m). Since
a commonly used detection radius around playback stations is 400 m, beyond which
owl calls are considered inaudible, it is suggested that caution be used in incorporating
detection radii in survey schemes in order to minimize the risk of over-estimating
owl numbers.
Clewley et al. (2016) evaluated a playback survey methodology for Little Owls in the
UK to determine detectability and practicalities that could inform potential future
national monitoring efforts. Nocturnal playback surveys were carried out in an open
landscape in the east of England in spring 2015. Twelve 2  2 km survey squares were
each visited four times. Little Owl calls were broadcast from three locations within each
square along a linear transect and spaced 500 m apart. The responses of owls of any
species were recorded. Little Owl occupancy was later confirmed through existing nest
monitoring efforts. Little Owls were detected responding to playback in all survey
squares and were confirmed to be breeding in 8 of the 12. Detection and response rates
were greater when combining data from multiple broadcast locations. The likelihood of
Little Owls responding decreased with greater distance of broadcast location from
known nest site, but no relationships were identified with time of evening or the

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Table 11.3 Overview of the factors that influence the efficiency of Little Owl broadcasting methods

Day/ Census
Reference Efficiency Method Weather Time of year Temperature Distance Night period

Exo and Hennes 80–90% Percentage of known Influence Influence No influence – – End Feb–
(1978) owls answering mid-Apr
Centili (2001) 49.60% Percentage of known – Limited – No influence – Feb–Jul
owls answering influence
Zuberogoitia >100% No. individuals No influence No influence – – – Jan–Dec
and Campos heard/no.
(1998) playback sessions
Mastrorilli 28.20% No. individuals Influence Limited – – – Sep–May
(1997) heard/no. influence
playback sessions
Galeotti and 24.90% No. individuals – – – – – Unknown
Morimando heard/no.
(1991) playback sessions
Pirovano and 54.70% No. individuals – – – – – Unknown
Galeotti heard/no.
(1999) playback sessions
Navarro et al. 120% before No. individuals – – – – Influence Apr–May
(2005) dusk; 180% heard/no.
after dusk playback sessions
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Table 11.4 Overview of the probability of response to broadcasting by month of the year

Reference Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Centili (2001) Percentage – 47.00 52.00 69.00 40.00 53.00 38.00 – – – – –

known
owls
Stimuli – 17 21 13 20 31 21 – – – – –
Zuberogoitia Percentage 21.43 36.96 45.07 51.92 46.38 19.12 49.32 19.15 17.5 32.07 9.76 6.67
and Campos successful
(1998) stimuli
Stimuli 28 46 71 52 69 68 65 47 40 53 41 30
Mastrorilli Percentage 23.00 29.00 22.00 29.00 28.00 28.20 – – 31.00 33.00 35.00 24.00
(1997) succesful
stimuli
Stimuli 56 24 22 54 57 37.4 – – 29 33 17 45
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presence of other owl species. Playback methodology proved to be effective at increas-

ing the detectability of Little Owls.
During playback experiments, the distance from the surveyor to the call influences
the chances of detection across nocturnal species in farmland and woodland habitats.
Orlando et al. (2021) evaluated how distance affects surveyor detection capability,
expressed as the probability of hearing broadcast calls and of estimating their distances
correctly. They conducted a playback field experiment in farmland and woodland areas
within an agricultural landscape in winter and summer 2020. Vocalizations of five
species (Little Owl, Tawny Owl Strix aluco, Long-eared Owl Asio otus, Common
Nightingale Luscinia megarhynchos and Water Rail Rallus aquaticus) were broadcast at
various distances to a surveyor, who attempted to detect them and, if successful, to
classify them into predefined distance zones. Binomial GLMMs were used to estimate
detection probability as a function of distance, and the effects of habitat and season on
this relationship. The distance of the broadcast call from the surveyor had a significant
effect on detection probability in both habitats. In woodland, the probability of hearing
calls was significantly higher in winter, while estimating distances correctly was gener-
ally higher in summer. An increase in field experience improved the detection capabil-
ity, mainly in terms of distance estimation, for which errors were mostly over-estimates.
The probability of hearing calls and the distance estimation accuracy varied between
species. Most Little and Tawny Owl calls were detected within a 200 m radius of the
surveyor, while this was 100 m for the other species. For a multispecies community-
level study, playback surveys are thus likely to be most representative of a 200 m radius
surrounding the surveyor where the probability of detection is highest, while estimates
of distance from the observer are likely to be inaccurate in most cases.
To better deal with the issue of estimating owl numbers, we recommend the use of
triangulating on responding owls, to better discern and accurately map their locations.
In triangulation, an observer acquires two (or three) compass bearings on the respond-
ing owl (e.g., from points that are 100 m apart), and then plots both the observer’s
locations (on a detailed map or aerial photograph) as well as the compass bearings to
the located owl from those locations. This will serve to pinpoint the location of the
responding owl. Observers will need to be attentive, to ensure that the owl does not
move between the acquisition of the second and third compass bearings; if it does, then
a new series of bearings will be required to map its location.
Table 11.5 gives an overview of published survey methods, displaying an array of the
call types used, time of survey activities, playback sequences, number of visits across the
survey routes, inter-station distance and suggested periods for experiments. Important
differences exist between the current methods; some groups use rather stringent
responses (i.e., only “ghuk” calls) to accept a calling Little Owl as a territorial bird
(Bloem et al. 2001), while other methods accept all the different calls of the owls as
responses (Verwaerde et al. 1999). Hardouin (2002) showed that only the call of the
male has a territorial function, suggesting the use of the “ghuk” call during broadcasting
efforts. These slightly different survey programs lead to differences in the data collected,
frustrate interpretations and limit the utility of the acquired data. Clearly, a shared goal
of our programs is to assess and monitor the status and trends of Little Owl populations.

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Table 11.5 Overview of published Little Owl broadcasting methods

Inter-station Seasonal Complementary
Reference Calls Timing Sequence No. visits Locations distance period methods

Exo and Hennes 8 territorial 30 min after 15 s broadcast/1 min 1 Potential habitats 300–400 m End Feb– –
(1978) calls/30 s sunset to silence/30 s mid-Apr
midnight or broadcast/1 min
2 a.m. to silence/1 min
sunrise broadcast/2–3 min
silence
Bloem et al. Territorial calls Until 2 h after 10 ghuk calls/1 min 3 Potential habitats 250 m Mid-Feb– Visual
(2001) sunset or silence/10 ghuk calls/1 mid-Apr observations
2 h before min silence/10 ghuk in day
sunrise calls/2–3 min silence. If
no response repeat
Verwaerde et al. Alarm/ Sunset to 78 s broadcast/1 min 1 Center of min 8 500 m Mid-Feb– –
(1999) territorial/ midnight or silence/78 s broadcast/ clustered 500 m mid-Mar
kiew calls 2 h before 1 min silence/78 s UTM squares
sunrise broadcast/5 min
silence
Mastrorilli (1997) Territorial/ 1 h after 3 min silence/3–4 min 5 Route selected Depending on Jan–Dec Visual
Mastrorilli alarm calls sunset broadcast/3 min with spot available sites observations
(2001) silence/2–3 min contained in in day
Mastrorilli broadcast/3 min study area and
(2001) silence spot inserted as
a function of
habitat
Bretagnolle et al. Territorial calls Sunset to 1 min silence/1 min 2 Potential habitats Min 500 m; Mar and Jun –
(2001) midnight broadcast/1 min around villages preferably
silence 750–1000 m
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544

Table 11.5 (cont.)

Inter-station Seasonal Complementary
Reference Calls Timing Sequence No. visits Locations distance period methods

van’t Hoff (2001) Territorial calls Unknown Unknown Unknown Potential habitats Depending on Unknown Visual
available sites observations
in day
Lorgé (pers. Territorial calls Sunset to 15 s ghuuk call/30 s 1–3 max Potential habitats 300–400 m Mid-Feb– Yearly control of
comm.) midnight silence/35 s ghuuk (if no mid-Apr nestboxes
call/60 s silence/ 40 s reply)
ghuuk call by pair/
5 min wait for response
Navarro et al. Territorial calls 2 h before Two broadcasting 5b Potential habitatsc 500 m Apr 19–May Surveys at sunset
(2005) (of male owl) sunset; 2 h sequences of: 2 min 17, 2002h would be
after tape and 1 min silence, helpful when
sunseta and then an additional looking for
2 min tape and 1 min nests, as owls
silence could be
observed
responding to
playback
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Survey Protocol Territorial calls Sunset to 1 min silence/2 min 3d or Center of min 8 500 m Feb–Apri Visual
for the Little midnight or broadcast/1 min 4–6e clustered 500 m Mar–Mayk observations
Owl April 2 h before silence or 4f UTM squaresg in day in low-
2005 – sunrise 1 min silence/2 min density areas
proposed broadcast/1 min
standard silence
1 min silence/2 min
broadcast/3 min
silence
a
Results indicated that nearly twice as many owls responded after sunset as before.
b
Calling was done before and after sunset on these five occasions.
c
A total of 14 survey stations were used in this study.
d
For habitat selection.
e
For demographic monitoring (number needs more testing).
f
For density estimates of terriorial pairs and singles.
g
In high-density areas and not trivially unsuitable habitat.
h
Peak vocal activity in southeast Spain.
i
Western Europe.
k
Eastern Europe, Middle East and Asia.
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A key step in achieving our shared goal will be based in standardizing the survey
methods and associated terminology (see Supplementary Glossary).
Field observations during Little Owl surveys have indicated that there appears to be a
population-density-based response involved with response rates (Exo and Hennes
1978, Pirovano and Galeotti 1999, Centili 2001). That is, owls residing in high popula-
tion densities appear to respond more readily to playback. This could be because of the
additional commitment needed by resident owls to continually affirm their territorial
boundaries in the face of increased competition for space and resources. More detailed
field trials are needed to determine response rates of Little Owls occurring at low,
moderate and high population densities.
Computational bioacoustics can be used in a wide range of applications in a cost-
effective manner. Ntalampiras and Potamitis (2021) focused on the problem of
detecting calls associated with individual Little Owls. To this end, variational auto-
encoders, consisting of deep encoding–decoding networks, were employed. A dataset
for new individual bird detection was used from 16 individual Little Owls consisting
of 952 bird acoustic events evenly distributed among individuals, with the audio
signals sampled at 44.1 kHz. The encoder encompassed a series of convolutional
layers leading to a smooth high-level abstraction of log-Mel spectrograms that
characterized bird vocalizations. The decoder operated on this latent representation
to generate each respective original observation. Individual detection was carried
out by monitoring and thresholding the expected reconstruction probability.
Further field testing is needed to validate the use of individual recognition during
fieldwork.

Demographic Data
Standardization is needed for the description and measurement of demographic data
across the breeding range. We consider the work by Van Harxen and Stroeken (2016) to
be an important step in this undertaking. We strongly encourage readers to examine
Forsman et al. (1996), Franklin et al. (2004) and Ganey et al. (2004) for their particularly
valuable contributions to the scientific assessment and analysis of demographic
parameters in the three subspecies of Spotted Owls. In particular, the focus of
demographic analyses, based on mark–re-capture methods, was directed at adult
and juvenile survival, fecundity and population trends. The consistent collection
and reporting of ringing and radio-telemetry data on juveniles and adults, especially
with regard to dispersal, will be of particularly significant value in understanding this
key aspect of population dynamics. For owl dispersal data, we strongly urge readers
to examine the monograph by Eric Forsman et al. (1996), to better understand the
compilation and analysis of substantial ringing and radio-telemetry data. Based on
the work of Van Nieuwenhuyse and Leysen (2001), international standardization is
also suggested for the analysis of habitat data, as this issue is closely associated with
demographic performance.
Nestboxes will likely remain one of the main venues for gaining demographic data
on Little Owls. Following the thoughtful work of Paul Marié in Soignies (Bultot et al.
2001), an experimental design should be applied when placing nestboxes in research

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 11.2 Conservation Strategy for the Little Owl

areas. Marié installed nestboxes at regular intervals, thus minimizing aspects of habitat
quality on the resulting aspects of box use. Further, this method allowed additional
types of analyses in subsequent years. Even with volunteer-based activities, an experi-
mental design of actions should be undertaken. We urge the development of standard-
ized guidelines for the installation of nestboxes to aid in this effort.

Other Aspects
Further research is also needed to better use birds released from care centers as experi-
mental samples to further improve the guidelines for release.
It should be stressed that there is a strong need to make data collection methods
straightforward and clear. Simple but useful methods can help to narrow the “gap”
between scientists and volunteers. Furthermore, simplified methods allow us to let
larger groups of interested people and nonprofessional birders contribute to the conser-
vation of Little Owls. Optimal exploitation and publication of existing volunteer data
should be undertaken, along with statements that describe some of the methodological
shortcomings. This will serve to make volunteer-based data more apparent, and to
validate the important conservation efforts of these caring people.

11.2.2.4 DATA MANAGEMENT

The importance of good data management cannot be overstated. Future research and
international co-ordination should emphasize the use of data management method-
ologies, including storing and exploiting data using data warehousing techniques. To
avoid the inefficient use of funds and data, and subsequent delays in critical conser-
vation efforts, we recommend that all future Little Owl projects have mandatory
standards for metadata, data collection and storage, and a data exploitation steering
committee. To use all information available throughout the complete distribution
range, more literature overviews like Génot (1989), Mastrorilli (2005; Italy), and
J. Shergalin (personal communication; Former USSR) should be made and published
on paper or via the Internet. An International Little Owl Data Warehouse should serve
to support a complete overview of literature, data, conservation insights and easy access
to and distribution of the available knowledge.

Standardized Data Sources

We believe that within-country datasets (especially in developing countries) are
needed, or need to be upgraded. With careful and reasonable design, these upgraded
datasets can support international efforts such as:

• Global Biodiversity Information Facility (GBIF): www.gbif.org/. The mission of the

GBIF is to facilitate digitization and global dissemination of primary biodiversity
data, so that people from all countries can benefit from the use of the information.
See data from Iran after Ilanloo et al. (2020).

• The Species Analyst Project (Peterson et al. 2003): http://speciesanalyst.net/. The

Species Analyst is a research project developing standards and software tools for
access to the world’s natural history collection and observation databases. The
project has given rise to, or provides the technical infrastructure for a number of

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 Chapter 11: Management and Conservation

other projects including: FishNet, HerpNET, Canadian Biodiversity Information

Facility and the Global Biodiversity Information Facility.

• EURING (https://euring.org/) is the co-ordinating organization for European bird

ringing schemes. It aims to promote and encourage scientific and administrative co-
operation between national ringing schemes, development and maintenance of
high standards in bird ringing, scientific studies of birds, in particular those based on
marked individuals, and the use of data from bird ringing for the management and
conservation of birds. These objectives are achieved mainly through co-operative
projects, the organization of meetings and the collection of data in the EURING Data
Bank. All European bird ringing schemes within the Little Owl distribution area
kindly agreed to make the ringing data available for this volume, as reported in the
EURING Data Bank ring recovery database (du Feu et al. 2016).

• Xeno-canto (https://xeno-canto.org/) is a website dedicated to sharing bird sounds

from all over the world. Whether you are a research scientist, a birder or simply
curious about a sound that you heard in the garden, xeno-canto invites you to listen,
download and explore the bird sound recordings in the collection. It is also a
collaborative project for people to share their own bird recordings, help to identify
mystery recordings or share their expertise in the forums. Xeno-canto uses the ever-
evolving possibilities of the internet to popularize bird sound recording worldwide,
improve accessibility of bird sounds and increase knowledge of bird sounds.

• Macaulay Library (https://search.macaulaylibrary.org/) is the world’s premier

scientific archive of natural history audio, video and photographs. Their mission is
to facilitate others to collect and preserve such recordings and to actively promote
the use of these recordings for diverse purposes spanning scientific research,
education, conservation and the arts. Access to audio and video recordings in the
archive is available for research, educational and commercial use.

• Article 12 of the Birds Directive (www.eionet.europa.eu/etcs/etc-bd/activities/

reporting/article-12) requests all Member States of the the European Union (EU28)
to monitor bird species and publish a report every six years. This information
includes population sizes and trends (short- and long-term) for breeding and
wintering populations, as well as pressures and threats for Special Protection Area
trigger species. In addition, population status and trends are assessed at the EU level.
The reporting period 2013–2018 data were used by the European Environment
Agency (2021). The reported data as well as the assessments of status and trends at
the EU level can be visualized using the web tool http://bd.eionet.europa.eu/
article12/.

• EBBA2 (www.ebba2.info/) is the New European Breeding Bird Atlas of the European
Bird Census Council that took up this exciting new challenge for European
ornithology providing vital data for conservation. It is one of the most ambitious
biodiversity mapping project ever attempted. The new Atlas documents changes in
breeding distribution of all European bird species. EBBA2 covers five years of
fieldwork in more than 50 European countries, including the European part of

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 11.2 Conservation Strategy for the Little Owl

Russia, Caucasus and Turkey, between 2013 and 2017. The EBBA2 project
conducted the mapping of more than 500 breeding species in more than
5000 squares of 50  50 km; 120 000 volunteer fieldworkers were involved in the
data collection process.

• ResearchGate (www.researchgate.net/) empowers researchers to address the problems

typical of the way science is created and shared. Their mission is to connect the
world of science and make research open to all. The 20 million researchers in the
community come from diverse sectors in over 190 countries, and use ResearchGate
to connect, collaborate and share their work. Many new publications on Little Owls
were made available through this platform.

Monitoring Data
Thanks to the standardized data collection methods detailed above, inferences about
population trends from all demographic study areas will be comparable, and data on
annual survival, reproduction and owl turnover rate at sites in each area will be
estimated using standardized methods and protocols. These methods reflect the best
available science and provide consistency and rigor to the data collection and analysis
process. For example, consistency will be provided for field surveys, the estimation of
reproductive effort, for capture, ringing, sex and age identification, and for preanalysis
data screening. Estimates of survival rates will be based on Jolly–Seber open-population
models (for example, Program MARK), as described by Franklin et al. (2004). Estimates
of reproduction will be based on empirical counts of numbers of young produced by
each female in the sample. Similar rigor will be applied to estimating adult survival,
fecundity, juvenile survival and estimating rates of population change.
Data about demographic trends in survival, reproduction and annual rate of popula-
tion change will be summarized annually for each demographic study area. A more
comprehensive meta-analysis of all datasets (for example, Franklin et al. 2004) will be
conducted every five years. Specific analyses will identify estimates of adult survival,
fecundity, juvenile survival and rates of population change. Interpretation of results
will continue to address uncertainties about the significance of adult emigration,
possible biases in estimating fecundity, the effects of aging, and differential detectabil-
ity of nesting and non-nesting pairs. Reports will include an annual summary for each
demographic study area and a more detailed report for the meta-analyses.
For additional details on data management and methods for population and meta-
population analyses, please see Franklin et al. (2004), Ganey et al. (2004) and Forsman
et al. (1996). Data sheets and an associated field manual are being developed to reflect
these data components.
The monitoring program outlined here will require a co-ordinated international
effort. The key to successful implementation is a co-ordinated network of personnel
and co-operators who will implement individual elements of the monitoring strategy.
The annual population surveys, periodic habitat assessments, cumulative data analyses
and integrated syntheses of the individual monitoring efforts implement the monitor-
ing strategy as a whole. Steps to accomplish the strategy are assigning specific tasks to
an administrative body, gathering and analyzing the data through standardized

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 Chapter 11: Management and Conservation

methods (some of which await development) and implementing the monitoring

program on schedule.
We recommend that a Little Owl Monitoring Lead be assigned. This individual will
work with international counterparts (such as representatives of the International Little
Owl Working Group (ILOWG), BirdLife International, Worldwide Fund for Nature,
Global Owl Project, etc.) and oversee the organization and implementation of the
Monitoring Plan. Fortuitously, the International Little Owl Working Group reflects
personnel and co-operators enlisted to conduct the demographic studies, assess habitat
conditions and develop predictive models. The Monitoring Lead and ILOWG can co-
ordinate the participation of land managers and key resource management personnel to
assure adequate funding and that survey results are integrated into the annual summary
reports. The ILOWG also should establish contacts with other scientists outside the
European community who are conducting Little Owl monitoring and research.
The Monitoring Lead and associated programmatic staff will require agency or NGO
support, with an estimated equivalent of two to three permanent, full-time positions
among the participating agencies. In addition to agency support of co-ordination
activities, support will be needed for permanent, full-time agency personnel engaged
in implementing the plan. Assuming that demographic studies are continued under the
current co-operative working relations and habitat assessments are done in-house, these
positions would involve research scientists, research wildlife biologists, statisticians and
GIS specialists.

11.2.3 Measuring the Implementation of the Strategy

Habitat condition and trend information will be estimated every 5–10 years after the
baseline habitat map is developed. Monitoring over time will allow the estimation of
changes in amount and distribution of owl habitat and relating such changes
to implementation of conservation measures. Habitat trend reports will tabulate
information about hectares of habitat by land allocation and their percentage change
over time. It also may be possible (and useful) to estimate habitat status and trends at
the home-range scale. Such measurements would be based on the use of extensive
inventory plot data. Here the appropriate habitat indicators would be attributes
associated with home ranges (e.g., nest trees, nest rock piles, road density, area of
pastureland and old orchards) and how their number and distribution are changing
through time.
We need to determine if the overall Little Owl Conservation strategy is effective. Is it
being implemented as we intended? Is it having the effects we desire for Little Owls?
What changes should be made to make the program more effective? It is not our
intention to create additional workloads for already overworked biologists and man-
agers; however, conservation, like any serious profession, needs to highlight its prod-
ucts and demonstrate its accountability.
We propose the establishment of a Little Owl Balanced Scorecard (following Kaplan and
Norton 1996) that contains the key performance indicators needed to obtain a holistic

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 11.2 Conservation Strategy for the Little Owl

view of the status of the strategy. The accomplishments of work on Little Owls should be
provided in annual summary reports. We recommend that a panel of science and manage-
ment personnel be assigned the tasks of: (1) reviewing the annual monitoring summary
reports and (2) preparing the interpretive Scorecard report assessing progress in meeting the
goals, objectives and expected values for the Little Owl under the array of applicable
legislation and national policies. Importantly, the Scorecard report will profile the state of
Little Owl populations and their habitat, evaluate the effectiveness of current conservation
measures in arresting and reversing the decline in owl habitat and populations, and
maintaining the viability of owl populations, point out areas of progress and concern,
and, as necessary, make recommendations for changes in management practices. This
Scorecard report will provide decision-makers with a scientifically credible evaluation of the
state of Little Owl populations and habitat, and would be a reference document for
decision-makers during periodic land-use planning reviews.
To fill key gaps in our collective knowledge we would monitor the number of topics
that have and will become available, the number of people working on them and follow
the number realized versus needed. For breeding data we suggest keeping track of the
number of owl sites (with each site having detailed latitude/longitude co-ordinates),
track the actual associated breeding data, and offer comparisons of demographic and
other results from the different study areas.
More specifically, components of a Little Owl Balanced Scorecard would reflect aspects
of: Knowledge, Limiting Factors, Landscape Conditions, Legislation and Policies, and People.
For example, concurrent with demographic data collection, range-wide Landscape
Conditions would be monitored to track changes in conditions by using owl habitat maps
derived from the national and international vegetation maps and imagery. This would
reflect analysis at several spatial scales (i.e., nest site, home range, landscape). Landscape
components, such as the area of pastureland, would be monitored on the basis of critical
structural and management activities that reflect ecological conditions for the owl.
An evaluation of Policies and Legislation relevant to land-use practices that affect
Little Owls should include: (1) country-specific evaluations and (2) an international
evaluation. Compliance with existing policies and legislation is part of this
review process.
For People, part of the Scorecard evaluation could be a status report on the current and
anticipated infrastructure of people and programs working on Little Owl core studies
and conservation projects. Implementation of the monitoring plan will require a co-
ordinated international effort. As part of this review process, the status of the imple-
menting components of: Monitoring the Owl and Its Habitat, Managing Little Owl
Populations, Standardizing Methodologies, Data Management, and Measuring Success –
Assessing the Strategy will be reported. For example, is the monitoring plan being
adopted across the range of the owl? Is a standard protocol for demographic surveys
and habitat assessments in place? Have within-country and range-wide results been
presented? A meta-analysis of owl population data from the demographic study areas
could be completed every five years. Range-wide habitat maps would be re-compiled
every 5–10 years in synchrony with monitoring plan schedules.

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 Chapter 11: Management and Conservation

11.2.4 Reporting of Results and Linkage to Decision-Making

Assurance of the quality of data collected and the methods used to summarize, analyze
and interpret the data will be applied to both aspects of the monitoring plan: popula-
tion survey and habitat assessment. For population surveys, survey timing in demo-
graphic study areas, determination of the sex and age of individual owls, and capture
and marking methods will be conducted according to the forthcoming Survey Protocols
for the Little Owl. Summarization and analysis of the data from the demographic study
areas will be subject to these protocols. Survival and reproduction data along with
estimates of population trend will be analyzed by using the procedures described in
Franklin et al. (2004) and Forsman et al. (1996). Consistency in the demographic data is
paramount, and participants contributing data to the meta-analysis will agree to adhere
to a rigid and formal protocol for analytical sessions (e.g., Anderson et al. 1999).
For landscape-scale habitat assessment we will rely on the quality assurance protocols
for the production of the range-wide vegetation map. Quality assurance for the derived
Little Owl habitat maps will rely on the knowledge of region-specific experts as they
define the habitat attributes at both the demographic study-area scale and regional scale.
An uncertainty estimate is essential for decision-makers to assess risks associated with
their decisions and for scientists to assess the efficacy of the data and associated models.
The accomplishments of monitoring for Little Owls will be provided in annual
summary reports. A meta-analysis of owl population data from the demographic study
areas would be completed every 5 years.
The status and trends of owl habitat and projected population responses will provide
managers with feedback about existing conditions and allow comparison with future
expected conditions. The results of these comparisons will provide information for
review of the adequacy of management direction. If the trend, or rate of improvement,
in habitat conditions is significantly below expectations, then a change in management
practices may be required. These changes may involve land-use allocations or manage-
ment standards and guidelines.
There will be no sirens or alarms wired to these elements to sound a signal for
change. We will be responsible for interpreting the monitoring results, and if needed,
signaling ourselves for the need for change. In the end, this monitoring plan will
provide data only about owl habitat and populations. Knowing how much habitat
there is, the survival rates of owls and how many young they produce are important
indicators, but we will be required to assess these indicators and decide whether conser-
vation is proceeding as planned and yielding the results we expected.

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 Chapter 12: Research Priorities

12.1 Research Priorities

The framework of this chapter was taken from Rich et al. (2004), who proposed a large-
scale conservation plan for American landbirds. We complement this framework with
some ideas of Noon and Franklin (2002) who have given an excellent overview of the
current knowledge, and what is still needed, of the ecology of Spotted Owls Strix
occidentalis. This chapter applies the given approaches and insights to the context of
the Little Owl.
Until the first edition of this volume, Little Owl research had been mostly descrip-
tive. Research has proceeded in a logical, sequential fashion, beginning with descriptive
studies of life history, food, behavior and habitat ecology. Initial studies focused on
characterizing the geographic distribution of the species and its general habitat associ-
ations. That work was followed by detailed investigations into patterns of distribution
within its geographic range, specific habitat relationships within these areas, estimates
of minimum area requirements and the extent of geographic variation in these pat-
terns. However, the mechanisms underlying the observed patterns of variation were at
that time mostly unknown. Causal models to better understand relationships and to
seek mechanistic explanations for the patterns observed have been gradually developed
since then. New research is getting more and more applied, and moving away from
descriptive, correlative and short-term work in small geographic areas, to large-scale
replicated studies, controlled experiments and long-term studies of demography
(Donovan et al. 2002). Results that lead to concrete recommendations for habitat
management are becoming more and more available.

12.2 Recent Insights

The first edition of this book referred to some research priorities based upon what was
known at that time. The major contributions since then are mainly on taxonomy,
habitat selection, habitat quality and population regulation.
Fundamental additional knowledge on the taxonomic diversity of the species was
obtained through DNA analysis (e.g., mitochondrial and nuclear) by Pertoldi et al.
(2012) and Pellegrino et al. (2014, 2015, 2016). An example of an important taxonomic
clarification was recently made in China, with the confirmation of a new subspecies,

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 Chapter 12: Research Priorities

Athene noctua impasta (Qu et al. 2002), based on DNA evidence. Pellegrino et al. (2020)
studied plumage coloration and morphometrics using over 200 museum skins.
The general knowledge of Little Owl habitat selection and habitat use stems mainly
from research carried out in western European agriculturally dominated landscapes.
The Little Owl has been characterized as having an important plasticity in habitat use
(Van Nieuwenhuyse and Leysen 2001). The species displays an ability to cope with local
circumstances in some situations but not in others. Habitat selection studies at different
spatial scales (Apolloni et al. 2018), fully integrated into one single model (Fattebert
et al. 2018) and across international borders (Tschumi et al. 2020) were increasingly
able to understand how the owl chooses its habitat. Combined with specific projects
that monitored the demographic performance of Little Owls (Bekaert 2006, Jacobsen
et al. 2016), habitat quality assessments were carried out and modeled, taking into
consideration the heterogeneity of the landscapes (Michel et al. 2017) occupied by
Little Owls and providing a significant advance for cost-efficient monitoring and the
possible identification of population source and sink areas (Pulliam 1988).
So far only a few Little Owl populations have been continuously monitored (in the
countries of Belgium, France, the Netherlands, Switzerland and Germany) for more
than 10 years – a time period long enough to begin to yield meaningful results on
population performance (see Knötzsch in Schönn et al. 1991, Ullrich in Schönn et al.
1991, Gassmann and Bäumer 1993, Kämpfer-Lauenstein and Lederer 1995, Furrington
1998, Bloem et al. 2001, Génot 2001, Meisser and Albrecht 2001 and Van
Nieuwenhuyse et al. 2001a). More insights on important demographic parameters
(e.g., immigration and emigration, juvenile and adult survival, age structure of the
breeding population) and population-regulating forces like roosts (Bock et al. 2013)
and predation (Michel et al. 2016) have been published.
For the first time, the entire EURING database of the Little Owl was studied, elevating
studies on samples to populations. Maximal exploitation of existing ringing data was
obtained, similar to the data from Wallonia in Bultot et al. (2001), but now across
Europe and showing that even nonexperimental data can yield important knowledge
about the species. This knowledge is crucial to assess the local context of breeding
performance and health of the population.
Experiments are emerging in recent research. Such projects have the fundamental
advantage that specific factors can be controlled for, leading to dedicated hypothesis
testing with excellent design of experiment (DoE) that optimizes and directs data
collection, saving time and effort compared to empirical data collection in
uncontrolled circumstances.
Experimental progress has been really huge in the last two decades, starting with the
bill coloration of adults (Avilés and Parejo 2012) and nestlings (Avilés and Parejo 2013).
Females with more yellow beaks had significantly larger nestlings while juveniles with
more yellow beaks received more food than dull colored or gray bills.
Predation risk was experimentally studied in a crossed design in which Parejo et al.
(2018) manipulated social information on a threat at the patch scale by broadcasting
calls of Little Owls indicating different levels of danger. Competitors sharing top
predators with the callers and prey of the callers interpreted alarm and nonalarm calls

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 12.3 Research Topics

differently. Competitors chose preferentially alarm and nonalarms patches over control
patches to breed, while prey selected alarm patches, taking into account the stress level
of their predators. Experiments using special predator-proof nestboxes or removal of
predators, and adding fake predators like plastic snakes (Pajero and Avilés 2011) yielded
insight into how owls choose their breeding locations, taking into account
different signals.
Food supplementation experiments were executed by Thorup et al. (2010), Perrig
et al. (2014, 2017) and Grüebler et al. (2018), identifying the importance of food as
limiting factor. Partial cross-fostering of nestlings was studied by Perrig et al. (2014)
supplementing only some nestlings from the same family. Thus, intrinsic factors acting
on survival and phenotypic development could be statistically controlled for. Two
synchronous broods were denoted as partner broods. One of the two was randomly
assigned to experimental treatment (i.e., food supplementation) and compared with
the complementing brood. Similar experiments analyzed the parental sex allocation
and sex-specific survival that drive the offspring sex-ratio bias in Little Owls. Tschumi
et al. (2019) found a clear positive discrimination towards females leading to signifi-
cantly stronger female than male nestlings.

12.3 Research Topics

12.3.1 Identifying Critical Habitat Components
The general habitat associations of Little Owls are well known. However, we still lack
important information on specific structural features, landscape configurations and
amounts of habitat that are required. Such information is critical for guiding the
development of effective management strategies for meeting large-scale population
objectives. For the Little Owl, this is an important challenge, since different landscape
elements might serve similar purposes for the species and some landscape features
might have a different impact on habitat suitability at different concentrations. The
insights that we obtain might be misleading, since current densities might not be
revealing real habitat quality (Van Horne 1983). This has been shown in the
Netherlands, where isolated high-quality habitats are becoming less favorable due to
the degree of isolation, while clustered suboptimal habitats remain occupied (van’t Hoff
2001). In France, Blache (2004) found the majority of occupied territories in exception-
ally homogeneous habitats rather than mosaic-like landscapes, suggesting the occupa-
tion of suboptimal rather than optimal habitat. This led to research that combined
habitat selection with demographic performance (i.e., survival and reproduction) to be
sure that we are modeling for habitat quality and not purely for habitat occupation
(Rosenzweig 1991). A growing number of multiscale habitat assessments have been
undertaken on Little Owls (i.e., Blache 2004, Martinez and Zuberogoitia 2004b,
Cornulier and Bretagnolle 2006, Zabala et al. 2006, Van Nieuwenhuyse et al. 2008,
Šálek et al. 2016, Apolloni et al. 2018). There is a growing need for integrated models in
which the different spatial scales are combined into one model. Fattebert et al. (2018)
developed multilevel habitat suitability models for the Little Owl over a large area,

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 Chapter 12: Research Priorities

aiming at identifying the most promising conservation areas. They evaluated habitat
suitability for the species over a large extent by using publicly available, high-resolution
governmental land-use data and Little Owl occurrence data obtained from citizen-
science species observations in combination with radio-tracking data. Their study
encompassed large geographical areas of two European countries differing in their
agricultural policies. The all-in-one model resolves two main issues in the conservation
of an endangered species: identification of large-scale suitability patterns to develop
conservation strategies and identification of fine-scale clusters of high-quality habitat
patches where conservation measures can be applied.
Despite a major progress in some areas, there is a clear need for replication in other
geographies and in larger areas. Finally we also need more insight into the selection
process of natural habitats. The use of comparable satellite imagery was expected to be
crucial for this but has proven insufficient. There is a need for more detailed infor-
mation on intensity of land use and the availability of resources like roosts, the age of
trees and the grazing regime. Tschumi et al. (2020) showed that the relationship
between geodata used as proxies and ecologically relevant resources may differ
according to history, policies and socio-cultural context, constraining the viability of
habitat selection models across political borders.

12.3.2 Demographics
Measurement of demographic parameters (nest success, productivity, survival, disper-
sal) is needed to identify factors limiting populations and to contribute to understand-
ing of metapopulation dynamics (e.g., gene flow, source versus sink populations). Until
now, minimum viable population and metapopulation studies have only been under-
taken for a small population in the Northern Vosges (France) (Letty et al. 2001a) and
Denmark (Andersen et al. 2015). Measurement of survival, particularly for juveniles as
they move from their natal sites to eventually acquire their own territories and mates, is
needed to both assess when Little Owls are most at risk and to identify sources of
mortality. Some initial meta-research was done by Bekaert (2006). An analysis of
capture–re-capture data from five research areas in Flanders (Zuiderkempen
(1991–2005), Dijleland (1998–2005), Hageland (1992–2005)), Wallonia (1989–2005)
and the Netherlands ((Achterhoek) (1986–2005)) yielded 8550 capture–re-capture
observations collected over 86 man-years of conservation and data collection for
5649 Little Owls within diverse anthropogenic habitats, confirmed that reproductive
success, just like survival, is influenced by a combination of extrinsic (environmental
factors) and intrinsic factors (individual quality) (Bekaert 2006).
In the near term, there is a need for more statistical analysis of existing multiyear
datasets that are available among the many Little Owl volunteer working groups.
Consolidating these data with those of other countries should be able to supersede
the local level up to the regional and even global level.
Age-specific estimates of survival and fecundity can be used to parameterize popula-
tion metrics to project future population trajectories assuming constant vital rates
(similar to Blakesley et al. 2001). Estimates of survival rates could be based on existing

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 12.3 Research Topics

capture–re-capture data, and fecundity rates could be estimated by determining the

reproductive output of birds of known age. There are examples of female owls where the
entire lifetime reproductive success is now becoming available. Since Hardouin (2002)
showed that individual recognition of individuals is feasible through vocal recognition
and as technology becomes available that might classify individuals directly in the field
(Ntalampiras and Potamitis 2021), new opportunities arise to better identify each
individual bird on a large scale with RFID rings (Hameau and Millon 2022) without
having to catch them. This is very promising for high-density regions where crucial
factors that determine the distribution of the species might be further researched, e.g.,
spatial auto-correlation, social interactions, occurrence of floaters and exchange
between populations. There is currently enough historical data available to study the
auto-correlative processes of the disappearance and expansion of population clusters as
is currently happening in Denmark, the Czech Republic, Slovakia, Slovenia, and
Germany and Austria.

12.3.3 Examining Responses of Little Owl Populations

to Land Use
There is a critical need to determine the effects of various types of land use and small-
scale landscape elements on Little Owl populations in order to devise effective measures
for minimizing the negative consequences of such land use. Land uses affecting owl
populations include livestock grazing, silviculture, recreation, fire management, oil and
gas development, mining, water control and development, agriculture, roads, subur-
banization, communication towers and wind-power development. Only by under-
standing the responses and tolerances of owls to land use and management regimes
can effective mitigation actions be developed. Research should involve stakeholders
from the beginning so that solutions will be accepted and used by those who control
the land. First results have been given by Onkelinx and Everaert (2018), Apolloni et al.
(2018) and Tschumi et al. (2020), who modeled the landscape as a function of habitat
quality and management impacts, and predicted future land re-allocation.

12.3.4 Examining the Effects of Abiotic Environmental

Factors
More research is needed on the importance of abiotic factors on owl population regulation,
including climate change, drought and contaminants (acid deposition, pesticides).
The special case for Little Owls is the abundant predation of earthworms and hence a
possible accumulation of heavy metals or PCBs. Previous results are limited to habitats
like the Rhine floodplains (Groen et al. 2000) that are not always representative of larger
areas. Special attention needs to be paid to vermicide contamination in Little Owls.
Initial attempts were undertaken by BirdLife International and Durham University
(2022) to estimate the prospective impacts of climate change on the distribution of
Little Owls. Distribution maps were created for 2025, 2055 and 2085 based on projected
future climate change. Maps were generated by relating the current range of the species

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 Chapter 12: Research Priorities

to current climate and then projecting this relationship onto future climate simula-
tions. This projected future range was based on climate simulations produced by the
Hadley center (HADCM3 model using the B2a scenario of future greenhouse gas
emissions) (Hole et al. 2009). Projections indicate a gradual extension of the distribu-
tion in sub-Saharan regions, extensions towards the south from Somalia towards Kenya
and the highly unlikely establishment of the species at the Angola-Namibia border,
gradually stretching east towards Botswana and Zimbabwe to southern Mozambique.
More in-depth research is needed into the more local impacts of global warming on
the species.

12.3.5 Adoption of the Information Theoretic Approach

Being a plastic species, statistical analysis of Little Owl data might yield unexpected
results through null-hypothesis testing, which could be considered as a “fishing exped-
ition” hoping to hook a significant p-value (Noon and Franklin 2002). An information
theoretic approach avoids the numerous problems with using null-hypothesis testing
and instead focuses on posing a set of multiple a priori research hypotheses for a
particular dataset, and then ranking and weighting those hypotheses to find the most
plausible, given the data, using an objective criterion. Additional features of this
approach include multimodel inference because it is unlikely that a single model will
be uniquely supported by the data; the possibility of averaging effects across competing
models also exists (Burnham and Anderson 1998). The link between statistical methods
and the biology of the organism occurs through a set of a priori hypotheses. These
begin as verbal ideas of what factors may influence the observed system, which are then
translated into statistical models (Franklin et al. 2004). Furthermore, alternative data-
driven artificial intelligence modeling methods could be explored.

12.3.6 Focus on Process Variation

There is a clear need for a mutual dependence between model formulation and prior
biological knowledge (Noon and Franklin 2002). White (2000) emphasized three
sources of variation in the dynamics of wild populations as important, effecting life-
history traits: (1) temporal, (2) spatial and (3) individual sources. Factors can be expli-
citly incorporated as co-variates into a set of a priori statistical models that seek to
explain variation in demographic rates of possible causal factors in order to address the
variation of population processes. This requires partitioning the variance of the depend-
ent variable into its sampling and process components (White 2000). By removing
variation attributable to the sampling process, subsequent statistical modeling focuses
exclusively on the process component of variation. The presence of spatial auto-
correlation has already been shown by Van Nieuwenhuyse and Bekaert (2002) in the
Little Owl. Elimination of this auto-correlation of Little Owl presence in habitat
selection studies should yield statistically more relevant results in terms of population
processes. Spatial auto-correlation in independent variables needs further study to
assess its impact on Little Owl models.

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 12.3 Research Topics

12.3.7 Search for Mechanisms

Morrison (2001) criticized studies on wildlife habitat use and selection for being mostly
descriptive in nature and concentrating solely on identifying patterns of use, rather
than the causes of those patterns. Mechanistic explanations for habitat use in Little
Owls is still missing, making it difficult to understand why certain components must be
managed in a given manner. In addition, habitat is rarely considered in terms of quality
(Van Horne 1983), where variation in demographic traits, such as survival and
reproduction, may be explained by different habitat components.
Little Owl research needs to strive for mechanistic explanations of why certain habitat
components separate high-quality from low-quality habitats by combining habitat selection
studies with demographic studies. Establishing real causes can either be done through
experimentation or through structural equation modeling (Pugesek 2003). Initially, habitat
quality studies rather than habitat selection were carried out in Switzerland and Germany by
Michel et al. (2017), Apolloni et al. (2018) and Tschumi et al. (2020). Replication of this
research in other regions of the global distribution is highly recommended.
The Little Owl is a perfect species for experimental research due to its extraordinary
ease of manipulation, its relative commonness in human-dominated landscapes and the
availability of individuals from bird care centers. Manipulation of occupied habitats by
adding or removing perches (Apolloni et al. 2018), adopting rotation mowing of hay-
lands (Tschumi et al. 2020) and adding additional daytime roosts (Grüebler et al. 2018)
has been shown to influence the hunting process and survival of the owls. Supplemented
feeding experiments have allowed for the first calibrations of the energetic balance for
specific prey items (Thorup et al. 2010, Perrig et al. 2014, Staggenborg et al. 2017).
Density dependence studies could use experiments by adding new nesting cavities
and even by adding some extra birds to the population under study. Hardouin (2002) is
one of the few that undertook some manipulative research using playback of recorded
Little Owls at other locations than normal, to test the recognition of individuals and
territorial aggression.

12.3.8 Long-Term Observational Studies

Observational studies serve three useful purposes: (1) providing clear and explicit
hypotheses that can be further examined with experiments, either large-scale manipu-
lative experiments or smaller-scale experiments that test key predictions posed by a
larger question, and (2) estimating variation in demographic parameters, such as
weather (Sunde et al. 2014) or climate, that may influence that variation and (3) long-
term observational studies are required to capture sufficient environmental variation
and examine hypotheses on the large-scale effects of that variation on Little Owl
populations (e.g., Noon and Franklin 2002).

12.3.9 Cumulative Effects

Because owl populations are affected by multiple factors, understanding the cumulative
effects of these factors is critical to all future management strategies. Hence, Design of

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 Chapter 12: Research Priorities

experiment (DoE) techniques and statistical interaction effects will need to be used to
cancel out or better quantify specific combinations.

12.3.10 Investigating Interactions Among Little Owls and

Other Flora and Fauna
We need to understand the relative importance of disease, predation, nest parasitism
and introduced species. Effects may be magnified by land use and abiotic factors, so
these should not be studied in isolation. Also, there is significant conservation value to
be achieved in co-ordinating Little Owl research efforts with those of other agro-
pastoral species (e.g. Parejo and Avilés 2011).

12.3.11 Combining Research and Management

Owl conservation plans are built upon information about the ecological and environ-
mental factors affecting bird populations, which is inadequate for many species.
Research should be combined with ongoing management to evaluate assumptions
and contribute new information for revision and improvement of those plans (adaptive
management). Combining research and management is also fundamental to testing the
effects of management actions on bird population response. Experimental research
within agricultural landscapes is fairly easy for Little Owls and should be expanded.
The impact of habitat management could be assessed and used for scenario building in
new projects.

12.3.12 Improving Monitoring

There is need for research on monitoring methods, dispersal assessments and analysis
procedures, e.g., via MARK. In particular, there is a pressing need for detailed and
rigorous testing of the response rates of Little Owls surveyed through the use of
playback. This is especially needed for areas of low, moderate and high population
densities. This information, once obtained, should quickly be incorporated into subse-
quent peer-reviewed and published versions of the Survey Protocol for the Little Owl.
We strongly urge additional research on juvenile dispersal – from natal grounds to
ultimate paired territory-holding status and further exploring the EURING database to
its full extent. This will involve radio-telemetry on relatively large numbers of owls, but
will be the primary way that this crucial information on dispersal survival and inter-
population connectivity will be acquired. Data about demographic trends in survival,
reproduction and annual rate of population change will need to be summarized for
each of the Vital Sign demographic study areas. A comprehensive meta-analysis of all
data sets (for example, Franklin et al. 2004) will need to be undertaken, with specific
analyses identifying estimates of adult survival, fecundity, juvenile survival and rates of
population change. This will require a new level of data organization and co-ordination
for researchers and managers working on the Little Owl. The development of the first
range-wide habitat map is technically within reach, but still no easy task. Detailing the

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 12.3 Research Topics

specifics of this satellite-imagery-based mapping process will help to insure that

future updates will be accomplished every 5–10 years in synchrony with monitoring
plan schedules. Finally, formal connections to major policies and legislation need
to be developed, with Little Owl demographic and habitat data providing a key set
of measurement tools (metrics) for evaluating and supporting such agreements
as the Convention on Biological Diversity (www.sdinfo.gc.ca/docs/en/biodiversity/
Default.cfm).

12.3.13 Expansion of Investigated Geographic Range

Few publications deal with Little Owl populations occurring in natural habitats. While
western Europe will probably remain the scientific nucleus for the species, substantial efforts
need to be undertaken in Northern Africa, the Middle East and Asian portions of the owl’s
range. A number of countries still lack population estimates and distribution maps. For some
areas, basic distributional and ecological studies are still needed for the owl. An asset to help
bring information and insights to these areas more rapidly can be offered through enlarge-
ment of the International Little Owl Working Group (ILOWG). Initial steps towards this are
being shown by active participation of members from the former Soviet Union and China in
the ILOWG. There is a need for additional DNA studies to clarify some subspecies-level
classification of taxonomy of the Little Owl; this would involve some relatively low-cost
projects in the Middle East and Asian portions of the owl’s range. Funding and institutional
support are, of course, the foundation for ensuring that the necessary research is under-
taken. Providing adequate resources will require co-operation and collaboration among
management agencies, research facilities, industry and nongovernmental organizations,
all of which have a role to play in support of Little Owl research. The principal role that the
ILOWG plays is bringing different research and conservation teams together. This co-
operation is comparable to the origin of this book, i.e., citizen science and academics, always
a fruitful combination if they find and understand each other.

12.3.14 Increasing Research and Experiment Maturity in

Emerging Countries
Replication is key to extended knowledge at the population level. This calls for inter-
national co-operation between teams that have very well-established conservation pro-
jects running with teams that are experts in experimentation, analysis and publishing.
A special role is foreseen for mature research centers like the Swiss Vogelwarte or the
Danish research group to help emerging countries by offering simple replication of experi-
ments and guidance in data manipulation and data processing. New research is still too
often dedicated to diet studies as the most obvious starting point. This should no longer be
needed when international co-operation is initiated (e.g., Ilanloo et al. 2020 in Iran) using
relevant experiments and data modeling expertise and replicating research that has proved
its value to the gathering of knowledge about the Little Owl. The Vogelwarte could easily
copy and paste its cross-border research (Tschumi et al. 2020) into Flanders versus Zeeuws-
Vlaanderen, which borders to the north (the Netherlands).

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 Chapter 12: Research Priorities

12.4 Conclusion
The update of this volume is a perfect example of how fast international co-operation,
aided by standardized methods for data collection, management and analysis, and
dissemination, within all European countries has led to more fundamental insights
beyond purely descriptive work and optimized by experimental research. Two decades
have led to an almost doubling of Little Owl publications, mostly in English, across the
entirety of Europe and gradually expanding over most of its distribution range.
Technological advances like PDF, GIS, advanced statistics, international standardized
databases of geocoded and time-stamped pictures and sound recordings have led to
online exchange of papers, data, advice, images, sound recordings and maps, facilitat-
ing analysis and editing compared to working on paper using pen, postage stamps and
at best a simple word processor. The ultimate difference in the way of working has
definitely been the digital artwork by Joris De Raedt who worked with photographic
input of owls and landscapes, collected by travelling from Portugal to China, to all the
places where subspecies have been discovered, using Google Maps, in a matter of days
instead of years. The drawings were custom made, as ordered by the authors, adapted,
standardized and optimized in a matter of days, leading to excellent, tailor-made plates
and illustrations that each time were saying exactly what they were meant to say. Our
hope is that this state-of-the-art contribution to the knowledge and understanding of
the species, its study methods, proven conservation and management approaches will
undoubtedly lead to even more international co-operation between conservationists
and well-trained academics and field workers across the entire distribution range of the
Little Owl. No better argument can be given for the fact that Little Owls really matter to
someone. Let’s use this book and this momentum to the utmost in the next two
decades as a yardstick for advanced citizen science and conservation management.

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 Index

abandoned houses, 236, 335 Athene noctua orientalis, 32, 40, 65, 78
activity peaks, 468 Athene noctua plumipes, 31, 39, 41, 69, 78
aerial photograph, 499, 503 Athene noctua saharae, 33, 42, 60, 77
Afghanistan, 30, 48, 78, 88, 227, 240 Athene noctua sarda, 33, 41, 56, 77
Africa, 1, 29, 31, 33, 46, 69, 73, 77, 86, 87, 88, Athene noctua somaliensis, 32, 42, 65, 78
90, 127, 148, 151, 235, 240, 242, 285, Athene noctua spilogastra, 31, 42, 43, 65, 78
286, 304, 488, 561 Athene noctua vidalli, 143
aggression, 404, 559 Austria, xvi, 11, 73, 146, 155, 163, 202, 203,
Albania, xvi, 77, 163, 212 205, 244, 254, 258, 280, 284, 305, 335,
albinism, 107 378, 423, 445, 484, 525, 557
Algeria, xvii, 33, 73, 77, 235, 236, 281, 484 auto-correlation, 464, 558
altitude, 35, 73, 216, 243, 266, 386, 388, 426, average clutch size, 311, 327, 345, 351, 352,
467 355
anatomy, 89, 114 average population densities, 154
anthelmintics, 413 aviary, 115, 126, 275, 358, 531, 533, 534,
anthropogenic habitats, 239, 457, 556 536
apple tree, 260, 329, 336, 337, 422 Azerbaijan, xvi
Arabian Peninsula, xvi, 29, 33, 60, 73, 77, 151,
234, 240 Bahrain, 234
Armenia, xvi, 152, 224 Balkan, 30, 46, 48, 51, 87, 143, 146, 216, 476,
ash, 522 484
Asia, 1, 10, 45, 46, 127, 218, 220, 240, 241, 275, Baltic states, 30
285, 286 Barn Owl, 80, 86, 90, 92, 114, 115, 144, 267,
Athene noctua bactriana, 30, 37, 56, 77, 80, 226, 282, 429, 431, 434
337 bee hives, 335
Athene noctua glaux, 29, 37, 38, 60, 77 beetles, 44, 65, 253, 254, 255, 272, 274, 275,
Athene noctua impasta, 20, 35, 38, 69, 78, 554 279, 280, 281, 282, 283, 284, 286, 303,
Athene noctua indigena, 30, 39, 56, 77, 224, 337 355, 413, 471, 486, 510, 513, 517, 521,
Athene noctua lilith, xiii, 34, 39, 40, 60, 77 528
Athene noctua ludlowi, 34, 39, 40, 69, 78, 80, begging, 97, 101, 139, 144, 399
229, 433 behavior, 327, 329, 345, 386, 389, 392, 396,
Athene noctua noctua, 29, 35, 36, 50, 77, 80, 119, 397, 403, 404, 411
337 Belarus, 30, 223, 335

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Belgium, xiii, xvii, 15, 73, 125, 146, 154, 155, cattle trampling, 254
159, 163, 169, 173, 186, 191, 248, 263, causes of mortality, 408, 410
267, 272, 349, 360, 378, 384, 410, 414, cavity nesters, 261, 421, 430
419, 427, 432, 433, 445, 446, 449, 473, census, 136, 412, 502, 537
482, 484, 488, 493, 554 cere, 90, 107, 117, 129
Bhutan, 80, 229 Chad, 151
Bibliography, 482 cherry, 260, 285, 521
bigamy, 385 chestnut, 198, 242, 518, 520, 521, 523
bill coloration, 100, 120 chimney, 528
bill size, 122, 129 China, 1, 31, 34, 35, 38, 40, 46, 48, 73, 78, 79,
binocularity, 115 80, 151, 229, 230, 231, 282, 285, 484,
biocides, 278 553, 561, 562
biodiversity, xvi, 227, 489, 505, 547 churches, 206, 210, 335
biology, ix, 5, 259, 413, 414, 436, 475, 486, 493, claws, 37, 40, 41, 44, 65, 69, 90, 107, 133, 275,
496, 558 284, 390, 404
bird-care centers, 559 cliffs, 226, 229, 232, 240, 241, 242, 243, 336
birth, 9, 83, 190, 325, 327, 382, 398, 445, 446, clustered pairs, 349, 473, 475
449, 453, 460, 472, 497, 506, 508 clustering, 84, 265, 266, 380, 411
body weight, 373, 425, 456, 530 cockchafers, 274, 278, 279, 282, 286, 292, 299,
Bosnia-Herzegovina, 73 301, 344, 355, 401
breeding cavity, 391, 470, 523, 526 collisions, 194, 211, 251, 405, 408, 410, 411,
breeding ecology, 431 481, 525, 533
breeding performance, 179, 305, 414, 424, 432, Common Vole, 205
554 competition, 15, 22, 27, 120, 158, 275, 282,
brood patch, 356, 396 407, 435, 450, 472, 473, 524, 546
Bulgaria, xvi, 88, 152, 214, 216, 242, 263, 284, connectivity, 423, 436, 440, 494, 496, 497, 503,
484 560
burrows, 31, 90, 226, 240, 241, 275, 333, 336, conservation plan, 493, 525, 560
337, 352, 433, 522 conservation projects, 489, 551, 561
conservation status, 5, 200
cainism, 380 conservation strategy, 1, 480, 481, 488
calling males, 158, 159, 174, 186, 191, 202, contamination, 363, 414, 417, 486, 557
207, 208, 267, 469, 473 copulation, 138, 143, 326, 329, 330
camera traps, 530 correspondence analysis, 484, 485
captive breeding, 6, 476, 477, 535 Corsica, 21, 77, 190
captivity, 15, 83, 87, 97, 124, 275, 349, 356, courtship, 137, 138, 143, 257, 324, 326, 329,
428, 431, 434, 531, 533, 534, 535, 536 386, 467, 470, 524
capture–re-capture, 382, 457, 497, 556 Crete, 21, 22, 77, 482
Caspian Sea, 77, 79, 221, 225, 226 crickets, 278
caterpillars, 271, 283, 292, 297, 299, 303, 304, Crimea, 79, 223, 241, 280
306, 402, 517, 518, 521 Croatia, xvi, 29, 73, 209

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cross-fostering, 425, 530, 555 earwigs, 279, 280

Cyprus, xiii, xvi, 21, 34, 48, 77, 86, 87, 129, ecotone, 262
143, 146, 212, 420, 482 ectoparasites, 431
Czech Republic, xvi, 27, 146, 155, 163, 206, edge effects, 254, 429
207, 208, 252, 253, 267, 405, 408, 421, education, xv, 490, 519, 548
423, 445, 449, 482, 486, 493, 526, 531, egg tooth, 97, 360
535, 557 egg-laying, 342, 344, 345, 349, 374, 393, 397,
Czechoslovakia, 378 424, 432, 473, 529
eggshell, 360, 414
data management, 480, 491, 547, 551 Egypt, xvii, 29, 30, 32, 39, 48, 77, 235, 285
daytime roosting, 259, 283 electrocution, 408, 420
Dead Sea, 232 elevation, 148, 152, 197, 206, 208, 214, 224,
death, 5, 9, 83, 126, 354, 360, 408, 411, 413, 269, 335
419, 421, 472, 497, 508, 527, 535 embryonic development, 89, 90, 119, 432
decision, 84, 495, 535, 551, 552 emigration, 7, 407, 436, 440, 445, 462, 473,
decisions, 256, 264, 470, 494, 552 549, 554
declining population, 163, 382, 461, 463, 494, Eritrea, 1, 20, 31, 48, 65, 78
533 Ermine, 363
decoy, 467 Estonia, 146, 148, 153, 220, 223
demographic study areas, 492, 495, 502, 503, Ethiopia, 31, 48, 78, 151, 227, 235
505, 506, 549, 551, 552, 560 Europe, xv, 1, 5, 10, 14, 22, 29, 30, 39, 45, 46,
demography, 410, 435, 530, 553 48, 51, 73, 86, 87, 127, 146, 148, 152,
Denmark, xvii, 1, 11, 73, 146, 153, 155, 163, 153, 155, 163, 168, 186, 198, 216, 239,
183, 186, 187, 189, 252, 253, 274, 405, 242, 244, 245, 251, 252, 257, 263, 268,
408, 424, 425, 433, 449, 455, 456, 469, 282, 285, 286, 301, 305, 324, 333, 342,
470, 529, 531, 556, 557 345, 361, 374, 378, 392, 406, 408, 410,
density dependence, 559 414, 418, 421, 423, 436, 466, 473, 476,
detection rate, 254, 503 480, 481, 482, 488, 489, 490, 532, 554,
diet, ix, 1, 5, 120, 236, 251, 256, 257, 258, 260, 561, 562
271, 278, 279, 280, 281, 282, 283, 285, extensive agriculture, 218
303, 304, 306, 315, 414, 420, 429, 430, extra-pair copulations, 385
484, 510, 516, 530, 561 eyelids, 89, 107, 390, 431
digestive apparatus, 145 eyes, xi, 3, 10, 27, 30, 89, 97, 107, 114, 115,
discriminant analysis, 267 116, 388, 390, 399, 431
disease, 382, 431, 521, 560
dispersal, 197, 405, 440, 441, 442, 478 farmers, 18, 238, 250, 480, 489, 490, 509, 536
distribution, 236 fecundity, xv, 382, 454, 455, 492, 497, 546,
distribution maps, 557 549, 556, 560
DNA, 20, 83, 86, 88, 385, 455, 536, 553, 561 fidelity, 262, 426, 450
Dormouse, 363, 487 field edges, 248, 253, 255, 269, 306, 422, 489,
drowning, 405, 407, 408, 409, 507, 528 518

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fingerprinting, 83, 385, 534, 536 255, 258, 259, 263, 269, 280, 287, 305,
Flanders, xvi–xvii, 11, 19, 159, 169, 170, 172, 334, 336, 343, 355, 360, 374, 378, 380,
175, 242, 243, 257, 263, 267, 279, 382, 385, 388, 395, 405, 408, 411, 421, 422,
431, 437, 457, 460, 473, 489, 493, 525, 426, 428, 429, 431, 432, 433, 436, 437,
556, 561 445, 446, 449, 450, 453, 474, 482, 484,
fledging period, 252, 408, 529 515, 529, 531, 554, 557, 559
fledging success, 6, 183, 324, 378, 432 grasshoppers, 44, 65, 274, 280, 281, 284, 303
floater, 324, 403, 487 grasslands, 39, 154, 192, 213, 214, 248, 251,
fluctuations, 163, 205, 217, 220, 458, 462, 504 254, 255, 258, 266, 268, 269, 285, 312,
food supplementation, 425, 476, 529, 530, 555 423, 471, 486, 507, 513
food supply, 116, 183, 212, 317, 425, 428, 471, Greece, 10, 11, 30, 39, 77, 146, 155, 163, 212,
509, 528, 529, 531 482, 484
foraging behavior, 258, 306
foraging efficiency, 6 habitat heterogeneity, 423, 492
foraging grounds, 251, 253, 264, 492 habitat loss, 171, 175, 176, 405, 423, 493
foraging habitat, 251, 252, 268, 333, 424, 425, habitat selection, 148, 154, 239, 242, 249, 252,
440, 503, 504, 507, 530 256, 257, 264, 265, 266, 267, 268, 270,
fragmentation, 83, 155, 170, 262, 406, 412, 406, 433, 435, 473, 488, 492, 535, 553,
423, 474, 475, 481, 488, 494, 496, 533 555, 558
France, ix, xvii, 5, 9, 10, 11, 15, 27, 48, 73, 83, habitat use, 253, 433
124, 126, 146, 152, 154, 155, 163, 190, habitat-based models, 494, 497
191, 200, 243, 255, 257, 259, 263, 265, handling prey, 100, 275
275, 279, 284, 305, 333, 336, 337, 343, hatching, 6, 89, 90, 92, 107, 116, 117, 119, 144,
349, 355, 360, 369, 378, 382, 405, 410, 292, 297, 299, 313, 317, 357, 360, 363,
411, 414, 416, 417, 419, 420, 421, 422, 364, 371, 380, 399, 456, 529, 530
428, 431, 432, 433, 436, 445, 449, 450, hatching success, 360, 376
467, 471, 473, 474, 482, 484, 493, 525, haystacks, 333, 335
531, 533, 534, 554, 555, 556 heavy metals, 405, 414, 417, 418, 557
fruit trees, 334, 336, 337, 422, 508, 510, 520, hedges, 3, 191, 194, 242, 248, 249, 255, 257,
521, 522 265, 266, 306, 408, 421, 422, 468, 486,
full moon, 468 510, 513, 517, 518, 519, 525
high-stem orchards, 174, 248, 336, 481, 489,
genetic proximity, 83, 534 508, 515
genetic variability, 80, 83, 455, 534 home range, 256, 268, 270, 469, 471, 494, 495,
genetics, ix, 1, 47, 88, 484, 535 496, 497, 503, 505, 506, 535, 550
geographic range, 176, 553, 561 hoot, xi
geographic variation, 33, 279, 553 hooting, 136, 137, 139, 397
Georgia, xvii, 77, 152, 224 hovering, 144, 388
Germany, xvii, 11, 15, 17, 31, 41, 73, 83, 84, human influences, 407
126, 129, 138, 146, 152, 155, 163, 176, Hungary, xvii, 10, 11, 45, 73, 87, 155, 213, 263,
183, 184, 185, 200, 237, 249, 250, 251, 267, 268, 284, 343, 411, 449, 482, 484

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hunting perches, 239, 242, 257, 468, 487 Korea, 78, 231
hunting success, 119, 254, 255, 258, 272, 433 Kuwait, 151, 163, 227, 235

immigration, 6, 84, 243, 405, 407, 436, 440, land use, 192, 248, 266, 270, 407, 408, 421, 422,
453, 454, 455, 459, 473, 474, 481, 554 481, 493, 508, 509, 510, 551, 557, 560
inbreeding, 533 larders, 271, 275, 277, 287, 310, 312, 314, 317,
incubation and nestling period, 278, 286, 292, 374
301, 304 Latvia, 1, 73, 146, 148, 153, 220, 222, 223
incubation breaks, 356 laying date, 261
incubation period, 292, 356, 357, 363, 364, laying interval, 345
467 Libya, 235
incubation time, 124, 356, 363, 364 limiting factors, 6, 148, 199, 264, 407, 435
India, 29, 30, 34, 228, 229, 488 Lithuania, 220, 223
indicators, 495, 496, 497, 506, 550, 552 low-stem orchards, 242, 248, 257, 434
intensification of agriculture, 155, 205, 211 Luxembourg, xvii, 155, 170, 176, 484, 489
inter-grading, 77, 80
intra-specific, 86, 136 management activities, 486, 495, 551
Iran, 30, 45, 77, 80, 151, 154, 226, 227, 242, management recommendations, 264
281, 285, 423, 481, 503, 547, 561 mating, 137, 138, 143, 325, 326, 329, 330, 385,
Iraq, 34, 48, 77, 80, 151, 227 396, 406, 440
Ireland, 179, 181, 444 measurements, 94, 97, 122, 131, 337, 356, 491,
iris colobome, 432 550
iris yellowness, 47, 117 mechanisms, 259, 425, 427, 429, 455, 489, 497,
isolated pairs, 223, 240, 349, 378, 473, 475 529, 553, 559
isolation, 22, 138, 349, 388, 453, 454, 455, 475, mechanization of agriculture, 3
555, 560 mesoptile, 101
Israel, xvii, 29, 77, 79, 80, 86, 163, 232, 234, metapopulation, 436
281, 285, 286 micromammals, 272, 277, 287, 296, 306, 309,
Italy, xvii, 11, 15, 17, 26, 29, 33, 46, 48, 51, 77, 310, 311, 313, 374, 398, 400, 402
80, 86, 143, 146, 152, 155, 163, 191, 197, Middle Ages, 9, 12, 13
198, 199, 200, 258, 263, 280, 281, 284, Middle East, 10, 86, 88, 127, 148, 218, 304, 561
333, 396, 421, 434, 449, 476, 482, 484, migration, 7, 92, 192, 197, 221, 388, 481
538, 547 minimum convex polygon, 463
mitochondrial cytochrome b gene, 84
Jordan, xvii, 9, 34, 232 mobbing, 15, 396, 434
juvenile dispersal, 440, 441, 444, 560 Moldavia, 284, 305, 336
Moldova, xvii, 77, 217
Kashmir, 78, 80, 127, 151, 228 Mongolia, 9, 11, 31, 46, 73, 78, 229, 241
Kazakhstan, 30, 73, 77, 78, 79, 129, 220, 225, monitoring areas, 492, 505
275, 281, 343, 345, 429, 441, 488 monitoring program, 1, 195, 197, 492, 504,
Kenya, 558 505, 506, 549

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monograph, ix, 546 over-fertilizing, 189

Montenegro, 80, 211 owl market, 17
Morocco, xvii, 33, 45, 77, 88, 235, 236, 240
mortality rate, 324, 380, 411, 436, 440, 446, pair bond, 324, 325
535, 536 pair formation, 136, 324
mosaic-like, 154, 170, 263, 267, 555 Pakistan, xvii, 30, 73, 80, 151, 228, 229
molting, 47, 56, 107, 437 Paleolithic, 9
molting period, 47 parasites, 336, 391, 405, 407, 430, 431, 473,
486, 521, 525
natal territory, 419, 440, 446 PCB, 414
natural cavity, 3, 258, 263, 352, 426, 459, 468, pellets, 271, 281, 282, 283, 284, 286, 304, 337,
487, 525 389, 401, 431
nearest-neighbor distance, 158, 460 perches, 6, 139, 163, 239, 240, 243, 244, 251,
neoptile, 101 252, 254, 255, 257, 258, 260, 268, 274,
Nepal, 34, 229 275, 388, 421, 471, 486, 509, 520, 525,
nest dimensions, 337 527, 559
nestbox, 261, 426, 525 pesticides, ix, 170, 171, 189, 255, 405, 408, 413,
nesting cavities, 241, 261, 263, 407, 472, 486, 414, 421, 471, 481, 521, 557
487, 509, 559 piles of stones, 10, 65, 199, 236, 260
nesting sites, 163, 189, 199, 213, 219, 235, 268, plasticity, 244, 261, 526, 554
333, 336, 407, 421, 457, 460, 509, 524, playback, 158, 172, 174, 203, 208, 267, 397,
526, 533 502, 537, 538, 539, 542, 546, 559,
nestling development, 425, 530 560
nest showing, 138, 143 plumage, xiii, 21, 27, 45, 47, 48, 56, 87, 89, 90,
Netherlands, xiii, xvii, 11, 73, 124, 127, 146, 104, 109, 114, 119, 390, 397, 554
155, 163, 174, 176, 186, 255, 259, 266, Poland, xvii, 73, 146, 155, 163, 218, 219, 221,
272, 277, 278, 287, 290, 306, 314, 317, 282, 305, 405, 408, 445, 482, 484
325, 334, 335, 336, 341, 343, 344, 345, pollard willows, 191, 242, 336, 392, 487, 489,
352, 353, 354, 355, 356, 364, 366, 381, 508, 510, 522, 525
386, 395, 396, 405, 414, 427, 436, 437, pollarded tree, 522
442, 446, 449, 450, 456, 457, 460, 475, population density, 138, 152, 205, 207, 208,
476, 484, 488, 492, 493, 525, 535, 554, 219, 226, 230, 244, 254, 258, 349, 468,
555, 556, 561 472, 473, 546
neuronal cells, 114 population dynamics, 84, 176, 209, 324, 425,
New Zealand, 1, 18, 73, 237, 243 435, 440, 441, 454, 476, 484, 486, 496,
NGO, 550 497, 504, 529, 537, 546, 556
Niger, 73 population estimates, 6, 136, 148, 153, 155,
170, 179, 184, 191, 199, 204, 211, 484,
old orchards, 1, 242, 336, 550 561
olive plantations, 194, 236 population models, 493, 549
Oman, 88, 235, 240 population regulation, 6, 255, 380, 553, 557

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Portugal, xvii, 11, 73, 88, 143, 146, 155, 163, response rate, 537, 538, 539
194, 195, 197, 242, 254, 258, 260, 272, retina, 89, 114, 115
281, 412, 419, 426, 428, 430, 436, 510, Rhodes, 77
562 ringing date, 328, 366, 373, 375
postfledging, 260, 412, 525, 529, 531 ringing study, 124
posthatching, 469 Ripley’s K function, 266
posture, 386, 396 road density, 266, 550
poultry, 192, 221, 278, 531 rocks, 1, 30, 37, 221, 230, 232, 236, 239, 240,
predation, 189, 244, 255, 256, 259, 260, 261, 333, 335, 336, 434
262, 263, 268, 278, 354, 355, 368, 370, rodenticides, 413, 528
382, 395, 396, 410, 423, 427, 428, 429, rodents, 1, 15, 22, 89, 119, 163, 221, 236, 250,
435, 455, 486, 488, 535, 554, 557, 560 255, 277, 279, 280, 281, 282, 284, 287,
predator avoidance, 259, 260, 427, 536 301, 306, 309, 310, 312, 349, 400, 412,
preening, 390, 391 528, 536
prehistoric, 9, 10, 22, 86 Romania, 29, 217, 263, 305, 482, 484
prey accessibility, 244, 251, 486 roosting opportunities, 386, 521
prey availability, 194, 244, 254, 255, 268, 408, roosting sites, 249, 260, 405, 470, 487
411, 486, 508 ruins, 1, 30, 31, 37, 199, 218, 224, 229, 231, 333
prey capture, 116, 441 Russian Federation, 78
prey density, 254, 423, 473, 508
protection, 9, 278, 363, 426, 480, 489, 518, 519, Sahara, 33, 42, 73, 77, 151, 235, 236, 237, 333
526, 527 Sardinia, 21, 33, 48, 77, 86, 197, 198
satellite, 264, 492, 503, 556, 561
Qatar, 235, 282, 285, 286 Saudi Arabia, 77, 235
quarries, 242, 333, 334 scapulars, 56, 101, 113
seeds, 285, 513, 515, 517, 519
radio-telemetry, 406 Serbia, xvii, 80, 211
radio-tracking, 270, 408, 556 sexual, 47, 120, 121, 456
railway, 267, 408, 412 shooting, 15, 370, 420
rain, 44, 65, 331, 355, 391, 393, 523, 524, 539 singing, 138
Red Sea, 1, 31, 33, 78, 235 sleeping, 34, 388, 393, 396, 397
regression analysis, 154 Slovakia, xvii, 27, 29, 77, 146, 155, 163, 206,
regression model, 92, 430 207, 208, 268, 405, 408, 484, 557
re-inforcement, 83, 184, 428, 532 Slovenia, xvii, 24, 27, 29, 77, 146, 155, 163,
reintroduction, 6, 531, 536 209, 210, 211, 408, 434, 522, 557
replication, 505 small-scale landscapes, 175, 457, 459
reproduction, 120, 176, 202, 244, 261, 264, snakes, 262, 282, 284, 407, 555
287, 310, 312, 324, 352, 373, 383, 403, snow cover, 148, 192, 207, 255, 343, 434
406, 430, 436, 441, 455, 457, 458, 462, social contact, 138
472, 484, 493, 494, 502, 503, 506, 530, social contexts, 117
552, 555, 559, 560 social information, 435, 554

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social interactions, 266, 473, 557 tail length, 108, 122, 129
soil, 170, 214, 255, 285, 414, 488, 510, 515, Tajikistan, 152, 163, 227, 343
516, 521, 522 talons, 97, 391, 396
Somalia, 1, 11, 33, 73, 151, 227, 235, 240, 558 tarsus, 48, 89, 107, 120, 122, 126, 129, 130
Somaliland, 32, 48, 73, 78 Tawny Owl, 14, 80, 114, 138, 171, 243, 260, 266,
Spain, xiii, xvii, 11, 30, 44, 45, 73, 88, 114, 126, 395, 412, 427, 429, 434, 441, 536, 542
146, 152, 155, 163, 191, 192, 193, 242, taxonomy, ix, 6, 20, 32, 88, 486, 553, 561
251, 258, 260, 266, 274, 280, 282, 343, telemetry, 249, 268, 388, 410, 429, 437, 445,
408, 412, 417, 419, 430, 435, 449, 476, 470, 484
482, 486, 493, 535 temperature, 97, 192, 257, 260, 337, 344, 433,
spatial scales, 147, 148, 239, 264, 265, 267, 268, 458, 461, 462, 470, 524
270, 494, 496, 497, 503, 504, 505, 551, territorial call, 395, 397, 538
554, 555 territory size, 405, 467, 468, 469
standardizing, 490, 546 threats, 146, 480, 481, 484, 508, 509, 526
starvation, 380, 382, 410, 433, 437 Tibet, 34, 35, 39, 40, 73, 78, 80, 127, 230, 488
statistics, 1, 120, 159, 372, 503, 562 traffic mortality, 171, 258, 411, 412
steppe-like habitat, 163, 195 Tunisia, xvii, 42, 48, 236
stomach, 145 Turkestan, 32, 227
Stone Marten, 171, 205, 324, 354, 363, 427, Turkey, xvii, 11, 30, 34, 77, 86, 88, 143, 152,
428, 488 163, 218, 242, 263, 343, 484, 549
straw bales, 335 Turkmenistan, 107, 111, 145, 226, 280, 343,
Sudan, 31, 73, 78, 151, 227, 235 345
sunbathing, 391
supplementation, 6, 83, 87, 243, 424, 425, Ukraine, xvii, 73, 79, 223, 430
472, 476, 529, 530, 531, 533, 535, 536, United Arab Emirates, 163, 235
555 United Kingdom, xv, xvii, 1, 73, 155, 159, 178,
supplementing, 425, 476, 530, 555 179, 180, 181, 242, 263, 277, 278, 279,
surveillance, 244, 364 305, 335, 343, 369, 410, 411, 414, 421,
survey method, 154, 487 437, 440, 446, 449, 482, 486, 510, 539
surveys, 153, 178, 186, 189, 194, 199, 211, 411, urban development, 194
473, 484, 486, 491, 492, 502, 505, 537, uropygial gland, 145, 391, 397
538, 539, 542, 549, 551 Uzbekistan, 152, 163, 225, 227, 240, 280, 282,
survival rates, 260, 382, 424, 441, 455, 456, 342, 345
492, 506, 530, 549, 552, 556
Switzerland, xvii, 11, 13, 77, 146, 155, 159, 163, vegetation height, 252, 254, 255, 268, 471, 513,
171, 200, 202, 249, 250, 269, 270, 274, 515
277, 278, 280, 299, 305, 336, 337, 343, vegetation maps, 264, 495, 504, 505, 551
360, 378, 382, 417, 422, 423, 431, 433, vermicide, 557
436, 445, 449, 450, 482, 484, 489, 493, viability, 239, 380, 453, 474, 476, 503, 504,
515, 529, 531, 554, 559 533, 535, 536, 551, 556
Syria, 9, 34, 77, 231, 285 video, xv, 92, 301, 386, 394, 526, 548

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visual acuity, 114 warmth pyramid, 97

vital sign, 491 weather conditions, 96, 199, 260, 274, 287,
vocal activity, 467 296, 309, 355, 373, 432, 459, 462,
voice, xi, xv, 10, 89, 397 526
volunteers, xii, xv–xvi, 1, 5, 154, 174, 487, 507, wing length, 29, 48, 92, 94, 95, 97, 100, 108,
508, 547 122, 126
wing surface, 388
Wallonia, xvii, 11, 170, 173, 349, 437, 445, 457, Wood Mice, 272, 277, 278, 286, 287, 306, 309,
460, 473, 554, 556 310, 312, 313, 315, 344, 393
walls, 31, 44, 145, 205, 224, 226, 236, 240, 241,
249, 255, 333, 335, 386, 388, 421, 487, Yellow Sea, 78, 151, 230, 231
514, 523, 526, 528 Yemen, 235, 240

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