Table Of ContentTHE RED FOX
Symposium on Behaviour and Ecology
BIOGEOGRAPHICA
Editor-in-Chief
J. SCHMITHUSEN
Editorial Board
L. BRUNDIN, Stockholm; H. ELLENBERG, Gottingen; J. ILLIES, Schlitz;
H. J. JUSATZ, Heidelberg; C. KOSSWIG, Istanbul; A. W. KUCHLER, Lawrence;
H. LAMPRECHT, Gottingen; A. MIYAWAKI, Yokohama; W. F. REINIG, Hardt;
S. RUFFO, Verona; H. SICK, Rio de Janeiro; H. SIOLI, Pion;
V. VARESCHI, Caracas; E. M. YATES, London
Secretary
P. MULLER, Saarbriicken
VOLUME 18
SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. 1980
THE RED FOX
Symposium on Behaviour and Ecology
edited by
Dr. ERIK ZIMEN
SPRINGER-SCIENCE+BUSINESS MEDIA. B.V. 1980
Library of Congress Cataloging in Publication Data
Main entry under title:
The red fox
(Biogeographica; v.18)
1. Red fox-behavior. 2. Red fox-ecology. 3. Mammals-behavior. 4. Mammals-ecology.
I. Zimen, Erik, 1941- II. Series: Biogeographica (The Hague)=v.18.
QL737.C22R42 599.74'442 80-18261
ISBN 978-90-6103-219-9 ISBN 978-94-017-5592-4 (eBook)
DOI 10.1007/978-94-017-5592-4
Copyright © 1980 Springer Science+ Business Media Dordrecht
Originally published by Dr. W. Junk bv Publishers, The Hague in 1980
Softcover reprint of the hardcover I st edition 1980
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or
transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise,
without the prior written permission of the publishers Springer-Science+Business Media, B. V.
CONTENTS
1. Introduction. A short history of human attitudes towards the Fox
by Erik Zimen . . . . . . . . . . . . . . . . 1
2. Habitat requirements of the Red Fox by H. G. Lloyd 7
3. The Red Fox-Vulpes vulpes (L., 1758)-in Europe by Michael
Stubbe . . . . . . . . . . . . . . . . . . . . . . . . . . 27
4. Comparison of the diet of the Red Fox (Vulpes vulpes L.) in
Gelderland, Holland, Denmark and Finnish Lapland by Darrell
Sequiera . . . . . . . . . . . . . . . . . . . . . . . . . 35
5. Prey consumption of a Red Fox population in southern Sweden
by Torbjorn von Schantz . . . . . . . . . . . . . . . . . . 53
6. The diet of the Red Fox - Questions about method by Hugo
Witt . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
7. Population ecology of the Red Fox-Vulpes vulpes (L., 1758)-in
the G.D.R. by Michael Stubbe . . . . . . . . . . . . . . . 71
8. Ein Modell fiir die Fuchspopulationdynarnik in der Bundesre-
publik Deutschland by A. V. Braunschweig . . . . . . . . . . 97
9. Population dynamics of the Red Fox (Vulpes vulpes) in Sweden by
Jan Englund . . . . . . . . . . . . . . . . . . . . . . . 107
10. Social factors affecting reproduction amongst Red Foxes, Vulpes
vulpes by D. W. Macdonald . . . . . . . . . . . . . . . . . 123
11. The Red Fox in a small game community of the south taiga region
in Sweden by Erik _Lindstrom . . . . . . . . . . . . . . . . 177
12. Aspects of the social structure of Red Fox populations: A
summary by Freek J. J. Niewold . . . . . . . . . . . . . . . 185
13. Yearly variations of recovery and dispersal rates of fox cubs
tagged in Swedish coniferous forests by Jan Englund . . . . . . 195
v
14. Zur Feindvermeidung freilebender Rotfiichse by Felix Labhardt 209
15. Foxes, wolves and conservation in the Abruzzo mountains by D.
W. Macdonald, L. Boitani and P. Barrasso 223
16. Epidemiology of fox rabies by A. I. Wandelar 237
17. Characteristics of the spread of a wildlife rabies epidemic in
Europe by K. Bogel and H. Moegle . . . . . . . . . . . . . 251
18. Short report on materials and methods used in a study of the
effects of rabies on the dynamics of fox populations in France,
with some preliminary results by M. Artois and L. Andral 259
19. Epizootiology and control of rabies in Central Europe by G.
Wachendorfer and J. W. Frost . . . . . . . . . . . . . . . 263
20. Fox social ecology and rabies control. Editor's closing remarks by
Erik Zimen . . . . . . . . . . . . . . . . . . . . . . . 277
VI
INTRODUCfiON
A SHORT HISTORY OF HUMAN ATTITUDES TOWARDS
THE FOX
Erik Zimen*
Save for the wolf, probably no animal has stirred the fantasy of our
forefathers more than the fox. In the old Greek fables of Aesop it was the
cunning jackal who outwitted his colleagues, but in Europe, by the Middle
Ages, time and repetition had gradually transformed him into a fox. Al
though the fox looks weak and slight contrasted with the wolf, bear and lion,
his darting intelligence and impudent daring usually get him out of all the
scrapes his insatiable appetite gets him into. Other fables tell how more
"naive" animals managed to trick him -like the famous Aesop fable, retold
by Hans Sachs, of the stork who took his revenge, after the fox offered him
soup i~, a shallow dish, by inviting the fox to partake of food in a
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narrow-necked jug. Or the "Brer Rabbit" stories supposed to be told by an
old Negro slave, and dating back through the centuries. The simpleton
always prevails, and the whole fascination of these fables lies in the fox's
age-old reputation for shrewdness far in excess of his fellows.
Whatever moral the fables offered, their detailed and apt descriptions of
animal behaviour mirror our forefathers' sound basic knowledge of their
natural environment, the red fox included, a knowledge that is our common
cultural heritage. Our children think of the fox as smart, cunning and
crafty-a well-deserved reputation if we consider his enormous geographical
distribution covering almost the whole holarctic region (Fig. 1), the largest
natural distribution of all mammals, again with the exception of the wolf.
But the fox, not the wolf, has managed to survive over almost all his former
range, in spite of extermination efforts and habitat destruction. His numbers
may even have increased considerably in certain areas, and regions tem
porarily lost seem to be resettled. Today he even lives in urban areas of
several large cities, including London.
Through the ages the adaptive .success of the fox has had a mixed
reception by man, best demonstrated perhaps in the British Isles. Some
thousand years ago it was the wolf who was valued by the nobles for his
sports utility, but detested by farmers and villagers alike for his destructive
potential to sheep and cattle. In those days wood was the main fuel in
Britain, not only for home use but for the early smelting industries, and
demand for these purposes and for shipbuilding destroyed the forests and
deprived the wolf of his last retreats during the Middle Ages. The remaining
* Lehrsti.ihl fUr Biogeographie, Universitiit des Saarlandes, D-6600 Saarbri.icken, G.F.R.
Biogeographica, Vol. 18: The Red Fox, ed. by E. Zimen 1
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wolves were hunted down and killed at the end of this period, save for very
remote areas of Scotland and Ireland, whereas on the continent the wolf
managed to survive over a large part of his range up to the end of the last
century.
The economic motives and their linked emotions in the British farmers
and villagers triumphed with the eradication of the wolf, and the nobles
turned rather reluctantly to foxhunting, considered to be only a poor
substitute for the tougher wolf-hunts. But slowly new cultural ritualizations
emerged, leading to the highly sophisticated modern foxhunt. The wolf had
left a hiatus for the farmers too, who now adopted the next smaller
carnivore-the fox-as the object of their detestation, and developed
equally sophisticated methods of persecution. The nobles bred pack-hunting
foxhounds, and the farmers bred small but tough little earthdogs (Fox
Terriers) which were trained to burrow underground and flush the fox from
its earth.·
The climate of Britain was pretty well perfect for sheep-rearing, and we
can read that "On the pleasant downs within a six-mile radius of Dorchester
more than half a million sheep were feeding" in 1700, for example, and "the
average weight of sheep ... doubled between 1710 and 1795". Britain
produced sheep with "the most valuable fleeces in the world", and peasants
grew rich by selling wool. Most sheep were let alone grassing out on the
range all year around. On the European continent, however, where the wolf
survived so much longer, sheep were usually herded and guarded by
shepherds and their dogs. Probably the fox had no opportunity to develop
techniques of predation on lambs, for which he is condemned in Britain. On
the continent the fox competed with the hunters of small game, and here
again sophisticated methods of fox control were developed, including the
breeding of the Teckel.
As so often happens, this hatred-induced cultural ritualization becomes
divorced from the ultimate cause for its appearance; efforts of farmers and
hunters to control the fox are out of all proportion to the animal's actual
destructive potential. Are our conceptions of his size at fault? A fox is a
small animal, seldom exceeding 10 kg in weight. Nor is he able to kill any
animal considerably larger than himself. A popular German children's song
is "Fox, you've stol'n the goose away ... " We kept a goose in our fox
enclosure for weeks, but the picture we wanted never materialized; in fact
the goose tyrannized over the foxes. Nevertheless, a fox in a chicken pen can
certainly be destructive. And even if the effects of fox predation on his
wild-living prey species is mainly compensatory in the sense of Errington,
the rabbit, hare or pheasant he kills is definitely not available to the hunter,
so that naturally, to any hunter who wants to maximize his bag, the fox is a
competitor for their mutual prey species.
The resulting widespread fox antipathy in small-game hunters is expressed
in the "closed seasons" hunting law/omission. Of the seven Central Euro
pean countries from which I could get information, only two have closed
seasons for the fox. According to our data from the southwestern part of
Germany, a high proportion of female foxes are shot in spring and early
3
