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ASPECTS OF THE POPULATION DYNAMICS

OF THE BADGER (MELESMELES L.)

HansAhnlund

Stockholm 1980
 ASPECTS OF THE POPULATION DYNAMICS OF THE BADGER
(MELES MELES L.)

av
Hans Ahnlund
Fil. kand.

AKADEMISK AVHANDLING

som med tillstånd av Matematisk-Naturvetenskapliga fakulte­

ten vid Stockholms universitet för avläggande av filosofie
doktorsexamen framläggs till offentlig granskning på Zoolo­
giska institutionen, Sandåsgatan 2, tisdagen den 20 maj 1980
kl. 10.00.

Abstract
Reproduction and mortality in Swedish badgers has been investigated.
Material from animals killed mainly in central Sweden was studied in
order to obtain information on pregnancy rate, litter size and morta­
lity in different years. In a later part of the work, a free-living
population was studied by means of capture-recapture technique, for
data on population density, movements and turnover rate.
Both sexes become sexually mature when about one year old. Preg-
nacy rates and litter sizes varied little between years, resulting in
a very constant productivity of the population.Prenatal mortality was
small but there was evidence of a high mortality among cubs in their
first summer. Later in life, mortality appears to be rather constant.
The results are discussed in the light of recent data obtained else­
where, showing the badger to be a social animal, living in groups.

Stockholm 1980
ASPECTS OF THE POPULATION DYNAMICS

OF THE BADGER (MELES MELES L.)

HansAhnlund

Department of Zoology
University of Stockholm, Sweden

Stockholm 1980
CONTENTS

1. Introduction
2. Age determination
3. Sexual maturity and breeding season
4. Pregnancy rate and litter size
5. Juvenile and adult mortality
6. Spatial organization and movements
7. Discussion
8. Acknowledgements
9. References

Abstract

Reproduction and mortality in Swedish badgers has been

investigated. Material from animals killed mainly in
central Sweden was studied in order to obtain information
on pregnancy rate, litter size and mortality in different
years. In a later part of the work, a free-living popula­
tion was studied by means of capture-recapture technique,
for data on population density, movements and turnover
rate.
Both sexes become sexually mature when about one year old.
Pregnancy rates and litter sizes varied little between
years, resulting in a very constant productivity of the
population. Prenatal mortality was small but there was
evidence of a high mortality among cubs in their first
summer. Later in life, mortality appears to be rather
constant. The results are discussed in the light of recent
data obtained elsewhere, showing the badger to be a social
animal, living in groups.

Sundt Offset Stockholm 80

2
 ASPECTS OF THE POPULATION DYNAMICS OF THE BADGER

(MELES MELES L.)

This paper is a summary of the following articles:

I. Ahnlund, H. 1976. Age determination in the European

badger, Meles meles L. - Zeitschrift fiir Säugetier-
kunde 41(2):119-125.

II. Ahnlund, H. 1980. Sexual maturity and breeding season

of the badger, Meles meles in Sweden. - J. Zool., Lond.
190:77-95.

III. Ahnlund, H. 1980. Productivity and mortality of the

badger, Meles meles L. in central Sweden. Manuscript.

IV. Ahnlund. H. & Lindahl, L. 1980. The size, age struc­

ture and dynamics of a free-living badger population
in Sweden. Manuscript.

3
1. INTRODUCTION

This study set out with the aim of investigating how badger
populations are regulated, with emphasis on data concerning
reproduction and mortality. The first part of the investi­
gation (I-III) deals with data collected from killed animals,
while (IV) is a field study.

Objectives of the study originally included:

(a) A description of the reproductive cycle of badgers with
special emphasis on age-specific productivity in diffe­
rent areas and in different years.
(b) An estimation of mortality rates before and after birth,
partly through studies of the age distribution in killed
samples.
(c) Identification of environmental factors affecting repro­
ductive- and mortality rates.

The above, rather naive classification of problems has changed

somewhat during the course of the study. A trivial reason for
this is that deficiency of material has limited the possibili­
ties of making comparisions , e.g. between different areas.
Another, and more fundamental change is related to the deve­
lopment of population ecology in later years, with more in­
terest given to evolutionary aspects and to questions relating
the behaviour of animals to their chances to survive and repro­
duce. Partly for this reason, (IV) was initiated in order to
collect data on population density, home range size and move­
ments on a study area in central Sweden. A third cause for a
change of interest lies in the fact that recently our know­
ledge about several important aspects of the biology of bad­
gers has greatly increased (9,10,21,22,23,24).
Results of the study will be presented and I finish the
paper with a concluding discussion.

4
2. AGE DETERMINATION.

Comparisons with other studies concerning reproduction and

mortality of badgers is difficult due to the fact that ageing
either has not been employed or the methods used (tooth-wear
analysis)(6,29,37) have been shown to be very inaccurate (40).
Accurate age determination is important in population
studies (8). It was found that age of Swedish badgers could
be determined from the closure of epiphyses of the tibia
until about two years of age, and later from the number of
annuli in the tooth dentine (I). When the paper was written,
no known-age animals were available for study, but a few
such have since been obtained and have confirmed the vali­
dity of the method (2).
Interestingly, the formation of distinct zones in the
teeth seems to vary in different areas; in animals from south­
west England the annuli are very poorly developed and often
impossible to read, whereas Scottish badgers show lines al­
most as distinct as Swedish animals (2).

3.SEXUAL MATURITY AND BREEDING SEASON.

Most studies have shown that badgers of both sexes become

sexually mature during their second spring at an age of about
one year (11,3,29). Most males then show presence of sperm
with an accompanying weight increase of testes and epididy­
mides and the majority of females ovulate during spring and
early summer. In Sweden, it was found that a fraction of
both sexes do no breed as yearlings (II).
Different authors have reached somewhat varying conclusions
concerning the reproductive cycle of female badgers, some
perhaps real and related to environmental factors, while
others may be caused by methodological problems, e.g. age
determination (11,6,29).
Generally, the cycle is characterized by the presence of
delayed implantation (12,14) and shows some unusual features.
Adult females, giving birth to young in early spring (early
March in Sweden, February in south England and late January

5
in south-west France (2,6,29)), in most cases seem to ovulate
and become pregnant very soon afterwards. By the end of March
about half of the adult animals in central Sweden are pregnant
(II). A parous female was observed in captivity to copulate
frequently about two weeks after the estimated birth date (30).
In contrast, nulliparous females, whether one or two years
old show a retarded pattern with the first ovulations occurring
in April. Despite these differences, all age classes seem to
reach their final level of pregnancy at about the same time
(II) .
Sexual activity of male badgers shows a maximum in March,
coinciding with the breeding season of parous females. Mini­
mum activity occurs in late autumn when the majority of males
lacked stored sperm (II). However, the annual variation is
rather small and apparently fertile males are found in almost
every month of the year (11,3).

4. PREGNANCY RATE AND LITTER SIZE.

In the years investigated, pregnancy rates among badger fe­

males showed great consistency. 90-95% of females two years
and older were pregnant each year, while among yearlings
approximately 50% possessed blastocysts or placental scars
(III) .
A comparison with other studies reveals that a high in­
cidence of pregnancy seems to be the rule for most badger
populations (6,10,29). Whether the pregnancy rate of one-
year old females varies in different areas is difficult to
judge from information published to date, due to inadequate
ageing techniques (cf. above).
Litter sizes, calculated from blastocysts and placental
scars also varies very little between years (HI). The number
of post-implantation uteri is too small to permit a compa­
rison between years, but the litter size is similar to that
calculated from blastocysts.
Counts of placental scars generally result in smaller litter
sizes than do counts of blastocysts and foetuses. In part,
this is explained by post-implantation losses, but it may
also be an effect of misinterpretation of scars.

6
 It was found that implantation seems to take place in al­
most all females and in most cases to result in live births.
Partial losses from litters seem to be most common among
primiparous and old females (III).
Litter sizes found in the present study are similar to
figures reported in other studies (10,28,29,37).
Combined with data on the age distribution, pregnancy rates
and litter sizes yield productivity figures for the entire
population which vary very little from year to year (III).

5. JUVENILE AND ADULT MORTALITY.

Quantitative data on mortality rates of wild mammals are rare,

due to the very difficult collection of information (7).
In the present study, cub-mortality was estimated from data
on the productivity and from the age distribution in the
sample (II). The resulting value depends on the mortality rate
of adult animals, which tentatively has been estimated at
approximately 2l5% annually (III,IV). Nothing is known about
how adult mortality is partitioned between seasons, but it
can be safely assumed that at least 50-60% of the cubs die
during their first summer. Field data (IV) indicate that most
of the mortality occurs very early and similar results have
recently been obtained elsewhere (10,21).
Cub mortality has been attributed to a variety of factors,
e.g. infectious disease (38) and starvation (28), though few
data exist. I have hypothetized (III) that intraspecific
aggression could contribute to the high cub mortality (cf.
below).
Human-caused mortality in the form of road kills and hun­
ting is probably a major source of death among adult badgers
(IV,16). Among badgers found dead in south-west England, it
was found that road kills was the largest single cause of
death. Of special interest is the fact that bite-wound in­
flicted by conspecifics was the second largest source of
natural death, surpassed in importance only by deaths from

7
bovine tuberculosis which occurs in this area (15,16,25).
There are some indications that juvenile mortality is less
in areas with low population density (III). I have found no
signs of a raised mortality among one- and two-year old
animals as reported from elsewhere (28). Instead, the death
rate seems to be fairly constant for most of the life-span,
possibly in part a result of human depredations (III).

6. SPATIAL ORGANIZATION AND MOVEMENTS.

Badgers were earlier reported to be non-territorial (27).

Recent studies have shown that rather the reverse is true;
in many areas territorial fights or signs of aggressive
encounters have been documented (IV,9,10,16,21,22). It has
been demonstrated that in various parts of England badgers
live in groups within joint territories which are actively
defended against neighboring groups (10,21). Data from cen­
tral Sweden are inconclusive, but indicate that a similar
pattern prevails also in this area (IV).
Group-territory size and size and composition of groups
shows a considerable variation (IV,10,21). There are indi­
cations that variations in population density result pri­
marily from variations in territory size, but too few data
exist to state this conclusively
Concerning dispersal of young animals, capture data and
a few other observations indicate that young females tend
to disperse to a higher degree than males (IV)(cf. below).

7. DISCUSSION.

In the following, I have assembled information from seve­

ral sources in an attempt to discuss badger population dy­
namics and reproductive strategies. Such a venture will
make painfully clear the large gaps that still exist in our
knowledge about this animal, but may hopefully be of some
inspiration for future studies.

8
 The recent discovery that badgers are social animals,
living in groups (9,21) necessarily influences interpretation
of data on reproduction and mortality.
According to one theory (22), food distribution is an im­
portant factor for determining the spatial organization of
badgers. Earthworms, which seem to be the principal food of
badgers in most areas (22,23,26,35,38) are available at the
surface in temporally unstable patches, related to local
weather conditions, but often in large quantities (22) . The
theory predicts that territory size should be related to the
dispersal of suitable feeding places, while the number of
animals in a group should be a function of the richness of
the food patches (22).
Living in groups will, however, involve several consequ­
ences of importance in reproductive contexts. So, for in­
stance, it is to be expected that competition for resour­
ces and reproductive opportunities will occur within groups.
On the other hand, if groups are relatively permanent, the
probability increases that animals will be related, allow­
ing the evolution of altruistic behaviour through kin selec­
tion (see 4).fThe selective forces that originally caused
the evolution of sociality will be modified by such con­
ditions, and I think that it is in this framework we must
discuss questions relating to reproduction and mortality of
badgers.
Delayed implantation, a characteristic of many Mustelids,
has often been regarded as a mechanism that enables optimal
timing of birth, but it can also profitably be looked upon
as a means of advancing the mating (33,41)? the question
reverts to what determines the mating time.
In badgers, we have seen that females who have given birth
breed earlier than nulliparous animals (II). Since this
condition prevails also among two-year old animals breeding
for the first time, it is likely that the two categories
have different optima as regards mating time. It is some­
what difficult to percieve in what way a post-partum estrus
might benefit a female, but some suggestions have been made

9
(II), relating the phenomenon to the influence of estrus
females on males; perhaps a more plausible explanation is
that it is advantageous for a parous female to "get it over
with" as early as possible, before the young start to make
too heavy demands.
On the other hand, a young female breeding for the first
time might benefit from delaying the estrus period. It
appears that females disperse from their natal area when
one or two years old (IV), and it it possible that breeding
occurs in conjunction with this event. It is likely that
the most suitable time for dispersal occurs later than the
time when parous females breed, which could explain the
difference in breeding time.
Whatever factors that have caused the evolution of this
pattern, it is interesting to note that most reproductive
activities of badgers are performed "on an empty stomach".
Implantation and the entire period of active pregnancy,
birth of the young and start of lactation, as well as most
mating activity takes place during a time when food, at least
in areas further north is severely limited or nonexistent.
(In central Sweden, badgers of both sexes are loosing
weight at least until early May.(2)) This touches on the
question of how badgers hibernate, an area where our know­
ledge is entirely lacking.
Ovulations during delay (11,17,29), occuring most frequ­
ently among young females, might be an adaptation under
similar selection pressures that have determined the mating
time (II). Data from mink (11) indicate that an infertile
remating during a secondary estrus period will not affect
conceptuses already present, whereas a fertile remating
increases the litter size through superfetation. If a simi­
lar mechanism is present in the badger, it could be seen
as a mechanism of insuring pregnancy.
The fact that only a fraction of individuals breed when
one year old (II) raises some questions. The delay in ma­
turity in some animals may be caused by nutritional factors
(33) or by some kind of social suppression. Socially in­

10
duced delay in puberty or breeding suppression has been do­
cumented in many mammals (e.g 1,32,36,39,43).
Badger females that breed when one year old may be in
better physical condition than those which do not, though
body weights in the two groups do not differ (2). As men­
tioned, breeding might be related to dispersal; reproduc­
tive suppression in the presence of parent animals has been
described in other mammals (36,39). In this context it can
be mentioned that in captivity young badger females tend
not to mate with familiar males with whom they have grown
up (30).
Additional factors which probably affect the age at first
breeding include the relative reproductive success of year­
lings and the risks involved by breeding as a yearling (42).
Concerning the first point, it might be thought that females
breeding for the first time when two years old would then
succeed better than those who have bred as yearlings. How­
ever, the opposite seems to be true; the litter size of the
former group is smaller and similar to that of yearlings (2).
Now, litter size is probably not a very good measurement of
reproductive success since survival of the cubs seems to be
crucial. Despite the fact that about half of the females
produce cubs when two years old (III), no animal in this age
class was found to be lactating on a study area (IV), pro­
bably indicating a high cub mortality. As to the trade-off
between mortality and early reproduction, it is extremely
difficult to measure and no data exist.
Since the social system of badgers seems to be of a kind
of resource defence type (5,22), it is likely that food at
times is a limiting factor. Moreover, due to the rather
slow development of the cubs, with dispersal appearently
not occurring until the age of one year (IV,28), any young
born into a group will consume resources for a long time.
For this reason, it is to be expected that reproductive
competition should occur between females in a group.
As argued earlier (III) there are some reasons for be­
lieving that reproductive competition between females could
take the form of infanticide (19,34). One is that inhibited

11
reproduction, as described in other group-living carnivores
(13,32,43) and probably a manifestation of reproductive com­
petition, seems to be absent among adult badger females and
is unlikely for theorethical reasons (III). Another is the
possibility that, as a result of the dispersal pattern (IV),
females in a group will be unrelated, reducing the likeli­
hood of altruistic relations between them (cf. 19). Empirical
data relating to this are lacking, but an impression gained
from field studies is that reproduction seems to be restric­
ted to certain individuals (IV) or in some way restricted
within groups (21). In addition, wounds from intraspecific
(and probably intrasexual (2,22)) aggression occur with
about equal frequency and are as severe on females as on
males (IV,16).
The relations between males can as yet only be guessed at.
There are observations from the wild of two males mating in
turn with a female without much aggression (18,31) and the
same has been observed in captivity (30), in which case the
males were brothers who had grown up together. Otherwise,
the general impression of captive animals is that males
react very aggressively towards new, sexually mature males,
also after a long acquaintance (2,30). With similar arguments
as put forward for females, there is a possibility that males
occupying and defending an area are genetically related.
Reduced sexual competition could then be expected to evolve
through kin selection.
I stress the very speculative nature of these arguments,
but I feel that increased knowledge about the kinships in
natural badger groups is essential for a better understan­
ding of badger population dynamics and social organization.

8. ACKNOWLEDGEMENTS.

I wish to thank Prof. Lars Silén and Dr. Jan Englund for
much support and help. Dr. Christer Wiklund and many other
colleagues at the Department of Zoology helped with dis­
cussions and criticism for which I am very grateful. I will

12
also express my gratitude to many people at Tovetorp Rese­
arch Station for valuable help in different ways and to all
hunters and other persons who provided material from killed
badgers.
The work was supported financially by grants from
"Anslaget för främjande av ograduerade forskares verksamhet
vid Stockholms Universitet", Hierta-Retzius"Stipendiefond,
Helge Ax:son Johnsons Stiftelse and the Swedish National
Environment Protection Board, this is gratefully acknow­
ledged.

13
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SUNDT OFFSET STHLM 80