Content uploaded by Ludek Bartos
Author content
All content in this area was uploaded by Ludek Bartos on May 04, 2016
Content may be subject to copyright.
4
Hunting seasons in relation to biological breeding
seasons and the implications for the control or
regulation of ungulate populations
marco apollonio, rory putman,
stefano grignolio and lude
ˇkbartos
ˇ
4.1 Introduction
Almost all European countries operate with a principle of restricted hunting
periods for some or all species (with the exception of Portugal, where technic-
ally the season lasts from 1 June in any year to 31 May of the following year,
although most hunting activity is carried out between September and Febru-
ary). There is, however, an enormous diversity in length (and actual time of
year) of the permitted season in different countries (even in adjacent countries:
e.g. seasons in the three adjacent countries of the Baltics: Latvia, Lithuania and
Estonia) and seasons also vary – often quite markedly – between different
regions or provinces of one country (e.g. Italy, Austria, Germany).
It is further apparent that such seasons may also show little relation to
actual biological breeding seasons (rut, parturition, period of dependency of
young) and such mismatch between hunting and biological seasons may have
serious consequences. This chapter explores the wide variation in hunting
season in different European countries and the implications of the mismatch
with biological seasons for welfare, social dynamics – and the ability (or
failure) of hunters to regulate ungulate populations.
There are at least three critical times of year in relation to breeding seasons of
ungulates:
the period of the rut (i.e. period between the first and the last copulation in
the observed population)
pre-parturition (i.e. period between late development of embryos and
parturition; we take this period as that period between the time when the
foetus may reach half of the birth weight and actual birth)
Ungulate Management in Europe: Problems and Practices, eds. Rory Putman, Marco Apollonio and Reidar
Andersen. Published by Cambridge University Press. #Cambridge University Press 2011.
80
the period following parturition when young animals may be nutritionally
or socially dependent on the mother.
In this chapter we will first examine the potential social, ecological and
evolutionary consequences of hunting within these biologically critical
seasons before considering the actual timing of permitted hunting seasons
in different European countries and how far these may ‘respect’ these poten-
tial problems. In addition we will consider the potential consequences of a
‘mismatch’ between hunting seasons and biological seasons (as well as the
problem of hunting seasons which are over-conservative) on our ability
actually to manage populations of overabundant species, before exploring
possible solutions to these various problems.
4.2 Potential problems of hunting during different biological seasons
4.2.1 Potential problems of hunting during the rut – the disruption
of reproductive aggregations
While the reproductive systems of European ungulates vary between species
and may vary between populations even of the same species, all inevitably
involve some degree of aggregation and some mechanism to facilitate male–
male competition, female–male meeting and/or female choice.
There is clearly considerable flexibility in the form of reproductive system
which may be adopted, between and even within a single species, with precise
‘choice’ of reproductive strategy dependent on a number of factors, both social
(Clutton-Brock et al., 1988a, 1993; Bartos
ˇet al., 1998; Willisch and Neuhaus,
2009) and ecological (Langbein and Thirgood, 1989; Carranza et al., 1995;
Thirgood et al., 1999). Thus for example, populations of any given species may
be truly territorial (as roe deer; Liberg et al., 1998) or may establish reproduct-
ive territories (where males defend a reproductive arena and call to attract
females to it, as for example in most populations of fallow deer: Chapman
and Chapman, 1975; Langbein et al., 2008), or alternatively simply defend
an area of rich resources often visited by females for foraging (red deer:
Carranza, 1995; fallow deer: Clutton-Brock et al., 1988b; alpine chamois:
von Hardenberg et al., 2000). Males in some species or some populations
may defend groups of females, rather than a fixed spatial territory, establishing
and defending harems of females (Clutton-Brock et al., 1988a, 1993) or alter-
natively may simply wander freely in search of oestrous females with which to
mate, as in the case of ibex (Willisch and Neuhaus, 2009), mouflon (Tu
¨rcke and
Schmincke, 1965; Briedermann, 1992) and wild boar (Heck and Raschke,
1980; Briedermann, 1986) and as is also reported for some populations of
Hunting seasons in relation to breeding seasons 81
fallow or sika deer (Putman, 1993; Thirgood et al., 1999). In the extreme males
may congregate together on a communal display ground or lek to compete
directly for access to females (Schaal and Bradbury, 1987; Langbein and
Thirgood, 1989; Apollonio et al., 1992; Bartos
ˇet al., 1992, 1998, 2003;
reviewed by Thirgood et al., 1999). But whatever the strategy adopted, that
strategy delivers the potential for intra-male competition and offers
opportunities for females to select mating partners and optimise fitness.
Shooting an individual at such a time may directly cause a temporary, or
perhaps more permanent, disruption of the breeding group formed, dispersing
animals (which may in consequence join other groups), or in the extreme
removing completely the focal male. A female may even abandon mating
altogether if her preferred mate is not accessible (Morrison, 1960). A further
problem which may arise if the disruptions are frequent is that the area used for
the mating aggregation is abandoned by the population and new one/s estab-
lished. Breeding grounds used by most ungulate species tend to be traditional,
with both males and females undertaking deliberate movements to reach them
at the beginning of the rut, or sometimes well beforehand. The abandonment
of a traditional site may be a gradual and incomplete process causing substan-
tial reduction in the amount of breeding opportunities of single individuals.
Not only spatial but also temporal shifts in the rutting activity can be induced
by diurnal hunting: red deer, fallow deer and sika deer males (and therefore
females) are active in reproduction all day and night long in protected, undis-
turbed areas, where their rutting callsare commonly heard in daytime as well as
through the night; where they are heavily hunted, reproductive activity is
virtually entirely restricted to the night (or at dawn and dusk).
A number of immediate and ultimate consequences for reproduction might
arise as the result of various types and degrees of disruption. In species which
display a high degree of polygyny and where, therefore, sexual selection can
be intense, the amount of time at disposal for mate choice and actual mating
can be critical, particularly when females usually have an oestrous period
extending only to some 36–48 hours. If hunting disturbance causes mating
groups to disband or in some other way reduces this time (for example
limiting activity almost exclusively to the night-time) a female can lose the
precise fertile time window of her cycle without being mated. This may have
different consequences in relation to the species affected and the amount of
disturbance.
Failure to conceive
The most serious consequence could even be a failure to conceive during that
season. In species which are monoestrous, such as the roe, a proportion of
82 Marco Apollonio, et al.
females might theoretically fail to breed at all in any given year simply
because of disturbance. However, even amongst polyoestrous species of
European ungulates, all have a comparatively short period of repeated oes-
trus (only wild boar sows show a wide fertile period, being able to conceive
throughout the year). If intense hunting reduces the chances of a mating
during this time, a proportion of females, even in polyoestrous species, may
fail to breed at all.
Disruption of breeding may cause reduction in synchronisation of births
In polyoestrous species, even if all females do eventually breed successfully,
continued disturbance may mean the rut is extended/prolonged. This will
result in lack of close synchrony of parturition, with calves born over a
protracted period. This may increase losses due to increased availability of
vulnerable fawns to predators, since these will be available over a longer time
period (Linnell et al., 1995; Aanes and Andersen, 1996; Kjellander and
Nordstrom, 2003; Jarnemo, 2004; Panzacchi et al., 2009) and/or because late
born calves, entering the winter in poor condition because they have not had
sufficient time to build up necessary body condition, are thus more susceptible
to overwinter mortality (Festa-Bianchet, 1988a; Festa-Bianchet et al., 2000;
Coˆ te
`and Festa-Bianchet, 2001; Gendreau et al., 2005; Pettorelli et al., 2007).
Disturbance of breeding groups may increase
the chance of mating by inferior males
The competition among males that usually takes place at breeding grounds
can be important in enabling females to select males with the best genes and
to avoid inferior competitors, so any disruption of breeding aggregations is a
potential treat to the fitness of the population as a whole.
A possible consequence of any disruption may be an increase of access to
females by inferior males, if as a result of disturbance, females are for
increased periods out of the control of dominant male/s. During the time
that females spend away from dominant male/s they may be mated by inferior
males that may transfer genes with lower value and therefore contribute to
the production of newborns with a lower fitness. In the extreme, if the
dominant male is actually killed, then clearly the probability is much
increased that females may be forced to breed with inferior males (in order
to breed at all while they remain in oestrus).
In a number of ungulate mating systems one of the major determinants of
male mating success is the length of tenure which may be maintained by an
individual male of harem or territory or any single defended female. It is
therefore not surprising that these highly successful males have the highest
Hunting seasons in relation to breeding seasons 83
chance of being shot during the rut, as they spend more time in rutting
activities that not only make them more predictable in terms of use of space,
but also more conspicuous (by, for example, roaring, fighting or courting).
The consequences of shooting prime males in ungulate populations – with the
resultant increase in access to females of less fit males – are widely known
(Coltman et al., 2003) and range from an overall reduction of population
viability (Mysterud et al., 2005) to a decrease in weight and/or horn or antler
size in males (Singer and Zeigenfuss, 2002; Coltman et al., 2003).
It is important to emphasise the strong counter-evolutionary effect of
this kind of ‘selective’ hunting, in that the culling of the fittest is basically
the opposite of what might be desirable by sound management. Fever for
large trophies often leads to reduction of the proportion of the fully mature
males in the population (e.g. in various parts of Central Europe). Absence
of prime males may result in lack of fully developed physical traits such as
body and antler size, etc., utilised in mate selection. As shown previously,
male phenotypic quality affects mate selection (for example, red deer stags
with large antlers are preferred for mating; Bartos
ˇand Bahbouh, 2006) and
males with large antlers had increased lifetime breeding success in an
unhunted population on the Isle of Rum (Kruuk et al., 2002) and also
other breeding characteristics, including offspring sex ratio (body size:
Røed et al., 2007).
Inappropriate harvesting might thus induce an undesirable evolutionary
response when the target characteristic is heritable, while in addition there
might well be unexpected effects on genetically correlated traits. Wildlife
managers must pay attention in order to plan hunting seasons and establish
appropriate hunting practice in order to reduce the genetic effects and the
evolutionary implications (Harris et al., 2002; Festa-Bianchet, 2003).
4.2.2 Culling during the period of late pregnancy
We believe that the killing of females in the last stages of pregnancy (i.e.
females that have already successfully borne almost all costs implied in the
successful development of an embryo) is acceptable in management only if
the declared intent of that management is to stabilize populations with a high
growth rate or effect a decrease in population size. In other contexts we
believe that this practice is undesirable.
Possible damaging effects of harassment
Some types of hunting practices, especially those which employ the use of
hunting dogs, but also noisy drive hunts or the use of vehicles, can cause
84 Marco Apollonio, et al.
considerable distress in the ungulates that are pursued. Such hunting methods
may cause distress not only to the individuals that are actively being hunted
but also for other individuals that may be disturbed by the drives or inciden-
tally chased by dogs.
In the extreme such pursuit may lead to abortion of the foetus, but even
without that, this stress can induce a number of physiological and behav-
ioural modifications. With respect to pregnant females, high levels of stress
(for example long and repeated flights) can cause welfare implications, such
as abortion with possible mother’s death. While it is very difficult to collect
data about effects of stress on reproduction and welfare in free-ranging
wildlife, several laboratory studies have produced data linking prenatal stress
with disturbances in offspring development and behaviour (see, for example,
Paarlberg et al., 1995). Moreover, stress-induced variations, for example
in maternal care, can serve as the basis for a non-genomic behavioural
transmission of individual differences in stress reactivity across generations
(Francis et al., 1999). There are unfortunately no quantitative studies about
the consequential impact on overall population reproductive success, but,
although it is difficult to quantify the effects, if hunting pressure is intense
both in terms of hunting days and in terms of numbers of dogs and beaters
this problem may be not irrelevant.
Even if we are unable to quantify such effects, it would seem probable that
such effects may well have some impact on overall population dynamics of
hunted species. And, as noted, several hunting practices may cause distress
not only to the individuals that are actively being hunted but also for other
individuals, or even other species that may be disturbed by the drives or
incidentally followed by dogs.
Ethical implications
Whatever may be the impact of hunting during late pregnancy on overall
population dynamics, there are also arguments against such practice simply
from an ethical standpoint. Even in situations where, from a strictly technical
point of view, hunting during the period of late pregnancy might be justified
(in situations, for example, where there is seen to be a need to effect a
significant reduction in a given ungulate population), pursuit or shooting of
heavily pregnant females does raise a number of issues of ethics and most
importantly is often considered unacceptable by the more general public.
Periodic outcry in the newspapers on the ‘infanticide’ during wild boar
hunting in January and on similar occasions bears testimony to a rather
general attitude of the public against what is perceived to be ‘cruelty’ in
hunting. Such negative perceptions often become generalised to hunting
Hunting seasons in relation to breeding seasons 85
in general and may help to develop negative views of hunting and
management activities overall.
4.2.3 Culling of mature females when young are still dependent on their mothers
Where the hunting seasons which apply in some countries are such that
culling of adult females may be permitted during the period of dependency
of offspring, there are once again clear implications both from a population
dynamics point of view and from purely welfare considerations.
In this context we should recognise a distinction between the period for
which the young may be nutritionally dependent on the mother and the
period during which the young are socially dependent on the mother (which
in social species may be much longer than the period for which they are
nutritionally dependent).
Neither of these has, to our knowledge, been adequately defined for any
species. Even in terms of nutritional dependency, the recorded period of
lactation is not necessarily a particularly good indicator of dependency, in
that although females may continue to lactate, and juveniles may continue to
take opportunistic advantage of such lactation, for considerable periods, this
does not necessarily imply a requirement for that nutritional subsidy. Fallow
does, for example, may still be lactating some 7 months after parturition (e.g.
Langbein, 1991), but this does not imply that fawns are actually dependent on
that milk, or would suffer loss of condition were it not available.
To generalize, we would suggest that nutritional dependence ends when
physical growth of the offspring is no longer dependent on mother’s energy
budget. While it is difficult to identify an actual time period for that nutri-
tional dependency, the effects of enforced early weaning have been widely
studied in laboratory and domestic animals. Disruption of mother–infant
bonding can induce physical and behavioural problems, including increased
neuroendocrine stress responses, augmentation of fear and aggression, and
reduced maternal behaviour (e.g. Kikusui et al., 2007, 2008), as well as
morphological changes such as myelin formation, dendrite length and spine
density in the brain (Ferdman et al., 2007; Kikusui et al., 2007; Nakamura
et al., 2008).
It is clear that the significance of lactation in ungulates changes as the
young grow and in the later stages of lactation may become more significant
in social bonding than in actual nutritional terms. However, even within this
context, it is again extremely difficult to determine what may be the actual
length of this period of social dependency between offspring and their
mother. The time that mother and young stay together cannot be used as
86 Marco Apollonio, et al.
any clear measure of such reliance since this may change with environmental
conditions and habitat use – and differ from year to year. For example, after
hotter summers the mother–young bond of alpine ibex endures significantly
longer (Grignolio et al., 2003). Further, the period for which mother and young
typically remain together does not necessarily mean that the young are depend-
ent on the mother for that entire period, or that juvenile survival or social
integration is actually compromised if the mother is killed before that time.
That said, the implication of killing the mother before the young are fully
independent (whether socially or nutritionally) has clear implications from
pure welfare considerations, and also has implications for population
dynamics.
Culling of the mother could result in death,
or loss of fitness of dependent young
Even if there are no objective data clearly establishing the time at which
juveniles are no longer nutritionally dependent on the dam’s milk (above) we
may safely assume that at least three months are necessary for most species of
European ungulates if the young are to survive at all. On that basis, any
shooting of lactating females before this time carries with it a considerable
risk of the death of the dependent young by starvation, unless the juvenile is
already accompanying the mother and is shot with it. Best practice would
thus dictate that if a hunter is to shoot a female during the period of lactation,
he/she must ensure that they also kill any accompanying calf.
Problems arise, however, where the hunter may not be aware that there is a
dependent juvenile, because it is not actually accompanying the mother.
Immediately after birth, neonates of almost any species may not be accom-
panying the mother; in addition, in those species (e.g. roe deer, fallow deer)
whose anti-predator strategy makes the offspring ‘hiders’ not ‘followers’
(Lent, 1974), this period where dependent young do not accompany the
mother may be considerably extended. In such situations culling of adult
females will commonly lead to orphaning of dependent young, because the
hunter is unaware of the existence of those offspring.
Such problems are of course most likely to be most acute early in the
calving season simply because the hunter may not even be aware that there is
a calf at all. But even later in the season, where offspring are accompanying
the mother, a strategy of shooting both mother and calf, however appropriate
in theory, may prove hard to achieve in practice. Problems arise with such a
strategy simply for technical reasons (because of the need to shoot two
animals in quick succession). Because calves usually linger for a few moments
in confusion after the death of the mother, many hunters advocate shooting
Hunting seasons in relation to breeding seasons 87
the mother first and then shooting the calf while it is still disoriented. But
from a purely welfare point of view (to avoid any risk of leaving an orphaned
calf ) it is actually more appropriate to shoot the calf first – and risk not being
able to shoot the mother too. As an additional problem, in many social
species (for example chamois or roe deer) female groups often consist of a
number of different females and their young. In this case it is not always
possible to determine accurately the mother–young pair. (Winter groups of
roe deer, for example, are usually based on a family unit, i.e. a doe and her
offspring. Nevertheless, winter groups may fuse or merge and the content of
such a group is typically rather unstable. In Kalø, Denmark, Strandgaard
(1972) monitored content of winter grouping in a population with a high
proportion of marked deer. Group size remained more or less stable with
eight members present on average. Nonetheless, 21 different individuals were
alternating in these groupings. For example, one doe was seen over the period
of a few days two times with her three fawns, five times with only two of
them, and three times without them.)
In either case, whether due to lack of awareness that there are dependent
offspring, or lack of ability to shoot both mother and accompanying calf,
hunting during the period of dependency may lead to premature orphaning of
juveniles. If nutritionally deprived, the young animal may die as a conse-
quence, or may be half-starved so that it takes longer to reach good breeding
condition (or fails altogether to reach breeding condition). In ungulates,
juvenile body mass can be related to maternal care, and body weight is an
important factor affecting offspring survival (Clutton-Brock et al., 1985,
1987a, 1987b; Bender et al., 2007, 2008; Carrio
´net al., 2008; Feder et al.,
2008; Stopher et al., 2008). For example, in mountain goats survival to one
year of age is greater for heavier female kids than for light ones (Coˆ te
`and
Festa-Bianchet, 2001). Even if the animal survives, slower growth rate may
imply a significant delay in reaching the mature body weight and perhaps the
threshold for reaching puberty (see, for example, Hamilton and Blaxter, 1980;
Albon et al., 1986); we should note, however, that in bighorn sheep, female
orphans and non-orphans had the same weight as yearlings and the same
probability of producing their first lamb at two years of age (Festa-Bianchet
et al., 1994).
Finally, in polygynous species with highly skewed probability of breeding
among males, a reduced adult body weight may result in orphaned males
failing to grow to a mature body mass where they are able to secure any
mating opportunities at all. Even though Festa-Bianchet et al. showed that
female orphans among bighorn sheep reached the same weight as yearlings as
did non-orphans and the same probability of producing their first lamb at
88 Marco Apollonio, et al.
two years of age, male orphans were smaller as yearlings compared with non-
orphans and they were unable to compensate for this early weight difference
in later life (Festa-Bianchet et al., 1994). In all cases, early deprivation leads in
effect to a reduction of lifetime reproductive success.
Even when the period of nutritional dependence is over, the bond
between mother and young in ungulates still has an important social
meaning. Often these species are social and females can play a significant
role in obtaining access to food (e.g. socially dominant mothers can
favour optimal feeding of their young; Veiberg et al., 2004), in teaching
population traditions (e.g. migratory route from winter to summer areas:
Festa-Bianchet, 1988b; Nicholson et al., 1997; Lamberti et al., 2004) or in
proper development of anti-predator behaviour (Childress and Lung,
2003; Li et al., 2009; Pipia et al., 2009). All these various social aspects
related to social competence and proper exploitation of environmental
resources, as well as the avoidance of potential threats, can be lost with
the premature loss of the mother, producing individuals with limited
chances of survival and reproductive success with obvious limitations to
population recruitment.
In addition, since many animals ‘acquire’ some of their social status within
the group as a consequence of mother’s status, a young animal whose mother
has been killed before it is socially independent may also suffer from being
rather low in the dominance ranking. In red deer calves social rank was
related to both body weight and mother rank (Veiberg et al., 2004). Loss of
the mother might thus result in an important decrease of social rank. Con-
sidering that social dominance is a fundamental aspect of male evolutionary
ecology in many polygynous mammals, with lifetime reproductive success
strongly related to dominance rank, an artificial modification of social rank is
likely to result, at least for males, in a significant alteration in individual life
history and breeding success.
4.3 European hunting seasons in relation to biological seasons
These various considerations suggest that there may be significant biological
(as well as ethical) issues associated with culling animals during the rut,
hunting during late pregnancy, or killing of adult females during the post-
parturition period. But in many European countries prescribed hunting
seasons do permit hunting at these times. To illustrate we have chosen
here to review the open seasons in different countries for red deer, roe deer,
moose, chamois and wild boar and explore the possible implications. Tables
4.1–4.5 offer a summary of current seasons operative in different European
Hunting seasons in relation to breeding seasons 89
countries/states; these are illustrative only and should not be taken as defini-
tive, since in some cases seasons vary between provinces within a given
country (e.g. Italy, Austria, Germany), or may vary with age-class of animal
(for example, distinct seasons in Wallonia, Hungary or Romania for prime
age stags and ‘poor’/cull stags). Further, seasons are shown here only for
adult males and females and calves of the year; in some countries (e.g.
Germany, Poland, Slovenia, Estonia), there are distinct (and different)
seasons specifically for juveniles/yearlings of both sexes.
Table 4.1 Open seasons for red deer
Country Males Females
Austria 1.05–31.01 1.05–31.01
Belgium 21.09–31.12
(21.09–30.09 large males
only)
1.10–31.12
Croatia 16.08–14.01 1.09–14.01
Czech Republic 1.08–15.01 1.08–15.01
Denmark 1.09–31.01 1.10–31.01
England-Wales-NI 1.08–30.4 1.11–31.03
Estonia 1.09–31.01 1.10–30.11
France 23.08–28.02 all practices;
1.03–31.03 coursing
23.08–28.02 all practices;
1.03–31.03 coursing
Germany
(most states)
Adults 1.08–31.01 (or 15.01);
subadults 1.06–31.01
Adults 1.08–31.01
(earliest 16.06);
subadults 1.06–31.01
Hungary 1.09–31.10 (prime age) or
31.01
1.09–31.01 (old) or
28.02 (young)
Ireland (Republic) 1.09–28.02 1.11–31.01
(28.02 in some counties)
Italy Alps 15.09–31.12 (stopped during
rutting period)
15.09–31.12
Italy Apennines 1.08–30.09 1.08–30.09 and
1.02–15.03 or 25.03
Latvia 1.09–31.01 15.08–31.12
Lithuania 15.08–15.10 1.10–31.12
Netherlands 1.08–15.02 1.08–15.02
Norway 10.10–11.11 10.10–11.11
Poland 21.08–28.02 1.10–15.01
Portugal 1.06–31.05
Romania 10.09–15.12 (prime) or
1.09–15.12
1.09–15.02
Scotland 1.07–20.10 21.10–15.02
Slovakia 1.08–31.12 1.08–31.12
Slovenia 16.08–31.12 1.09–31.12
Spain September to mid February
Switzerland 1.08–31.12 1.08–31.12
90 Marco Apollonio, et al.
4.3.1 Culling during the rut
Considering these five more numerous ungulate species, it is clear that
legislation does not in general take any real account of possible problems
which might arise from hunting during the period of the rut. In fact, the
hunting season overlaps the rutting period of moose, chamois and wild
boar in all countries. For roe deer and red deer, hunting during the rut is
permitted in more than 80% of the countries. In practice, the period of the
rut is often actively exploited by hunters in order to increase the ease of
gaining access to ungulates, especially males, which will at this time be
showing reduced vigilance. In some countries there are even specific
Table 4.2 Open seasons for roe deer
Country Males Females
Austria 1.05–31.12 1.05–31.12
Belgium Wallonia 1.05–15.05 and 1.08–30.11 1.10–30.11
Belgium Flanders 15.05–15.09 15.01–15.03
Croatia 1.05–30.09 1.09–31.01
Czech Republic 16.05–30.09 1.09–31.12
Denmark 16.05–15.07 and
1.10–15.01
1.10–15.01
England-Wales-NI 1.04–31.10 1.11–31.03
Estonia 1.06–30.09 1.09–30.11
Finland 1.09–31.01 and 16.05–5.06 1.09–31.01
France 15.05–31.08 stalking;
1.09–28.02 driving etc.;
1.03–31.03 coursing
1.09–28.02 driving etc.;
1.03–31.03 coursing
Germany (most states) 1.05–15.10 adults 1.09–31.01
subadults 1.05–31.01
Hungary 15.04–30.09 1.10–28.02
Italy Alps 1.09–7.12 1.09–7.12 and 1.02–15.03
Italy Apennines 1.08 or 15.08–30.09 1.08 or 15.08–30.09;
1.01–15.03
Latvia 1.06–30.11 15.08–30.11
Lithuania 1.06–1.11 1.10–31.12
Netherlands 1.05–15.03 1.01–15.03
Norway 10.08–23.12 25.09–23.12
Poland 11.05–30.09 1.10–15.01
Portugal 01.06–31.05
Romania 15.05–15.10 1.09–15.02
Scotland 1.04–20.10 21.10–31.03
Slovakia 16.05–30.09 1.09–30.11
Slovenia 1.05–31.10 1.09–31.12
Spain Mid April–31.07 Mid April–31.07
Switzerland 1.05–31.01 1.05–31.01
Hunting seasons in relation to breeding seasons 91
Table 4.3 Open seasons for wild boar
Country Boars Sows Subadults
Austria All year All year (except
if with piglets)
All year
Belgium 1.10–31.12 1.10–31.12 1.10–31.12
Flanders
Belgium
Wallonia
1.01–31.12/
01.10–31.12
1.01–31.12/
01.10–31.12
1.01–31.12/
01.10–31.12
Croatia All year 1.07–31.01 All year
Czech
Republic
1.08–31.12 1.08–31.12 All year
Denmark 1.10–31.01 1.10–31.01 1.10–31.01
Estonia All year All year All year
Finland 1.06–29.02 1.06–29.02 (except
if with piglets)
France 15.04–14.08 stalking;
15.08–28.02
driving etc.;
1.03–31.03
coursing
15.04–14.08 stalking;
15.08–28.02
driving etc.;
1.03–31.03
coursing
15.04–14.08 stalking;
15.08–28.02
driving etc.;
1.03–31.03
coursing
Germany 15.06–31.01 15.06–31.01 15.06–31.01
Great Britain Not hunted Not hunted Not hunted
Greece 15.09–20.01 15.09–20.01 15.09–20.01
Hungary All year 1.05–31.12 All year
Italy Third Sunday
Sept.–31.01
Third Sunday
Sept.–31.01
Third Sunday
Sept.–31.01
Latvia 1.05–31.01 1.05–31.01 1.05–31.01
Lituania 1.05–1.03 1.10–01.02 1.05–1.03
Netherlands 1.07–31.01 1.07–31.01 1.07–31.01
Poland 1.04–28.02 15.08–15.01 1.04–28.02
Portugal All year All year All year
Romania 1.08–15.02 1.08–15.02 1.08–15.02
Slovakia 16.07–31.12 16.07–31.12 16.07–31.01
Slovenia 1.04–31.01 1.08–31.01 All year
Spain 1.10–28.02 1.10–28.02 1.10–28.02
Switzerland 1.07–31.01 1.07–31.01 1.07–31.01
Table 4.4 Open seasons for moose
Country Males Females
Estonia 15.09–30.11 15.09–30.11
Finland 25.09–31.12 25.09–31.12
Latvia 1.09–15.12 1.09–15.12
Lithuania 1.09–15.11 1.10–15.11
Norway 25.09–31.10 25.09–31.10
Poland* 1.09–30.11 1.10–31.12
*
Hunting stopped from 2001.
92 Marco Apollonio, et al.
traditions of hunting roaring red deer stags (specific calls are used in this
context) or rutting alpine chamois.
With very few exceptions (including Norway and the Italian Alps) the legal
seasons which apply in almost all European countries allow the shooting of red
deer stags during the rut. Scotland has one of the earliest openings of the
permitted season for shooting red stags – 1 July; more generally the commence-
ment of the season is in August or September. In some regions of north Italy, in a
change of practice in recent years, hunting is stopped for two to three weeks
during the actual roaring period. The season for roe bucks also extends through
the rut in the majority of countries. In a few instances (e.g. Norway) hunting does
not commence until the rut is completed and in Italy too the hunting season for
males generally begins from the end of the rut (15 August), but there are regions
when it starts earlier. In Denmark, while the buck season starts well before the rut
as in many other countries (16 May), the season appears deliberately ‘broken’ to
accommodate an undisturbed rut (between 16 July and 30 September).
Hunting of male moose is allowed during the rut in all European countries
where the species is managed. Alpine chamois males (and females) are hunted
throughout their range during the rut (November) with no exception. And
finally, both male and female wild boar can be hunted during the breeding
season (late autumn to early winter, October to December/January) in all
European countries; more specifically it is interesting to observe this is the
only part of the hunting season that is the same in all countries, which
otherwise show very different patterns in their seasons.
4.3.2 Culling during the last weeks of pregnancy
Hunting in the period before parturition is not a problem per se, at least if we
do not worry about some subjective ethical reason (see above), but it may
Table 4.5 Open seasons for alpine chamois
Country Males Females
Austria 1.06–31.12 16.07–31.12
Croatia 1.09–31.12 1.09–31.12
Czech Republic 1.10–30.11 1.10–30.11
France 23.08–28.02 23.08–28.02
Germany 1.08–15.12 1.08–15.12
Italy 1.08/15.08/15.09–15.11/31.12/
31.01
1.08/15.08/15.09–15.11/31.12/
31.01
Romania 15.10–15.12 15.10–15.12
Slovenia 1.08–31.12 1.08–31.12
Switzerland 1.08–31.12 1.08–31.12
Hunting seasons in relation to breeding seasons 93
become a problem if the method of hunting results in high levels of stress
either in the animal culled or others in the area, as in the case of hunting with
dogs. In such context we might note that while the season for roe does in
France finishes at the end of March, for the final month (1–31 March) the
animals may only be hunted by coursing!
Seasons for roe deer more generally are restricted to a period well before
parturition (e.g. 1 January to 15 March, or 15 January to 15 March in the
Netherlands or Flanders) or do not commence until September or the begin-
ning of October (the majority of countries). Only in Spain, Austria and
Switzerland is the season for mature females open from April (Spain) or
May. It should be noted that females culled at this time may well be near-
term or actually have given birth.
In the case of red deer, the closing date for the hind season is in no case
later than the end of March and the opening date is late enough to guarantee
that pregnant hinds would not be culled in late pregnancy or the period
immediately after parturition. In this case, the only exception is Austria with
an open season for females starting from 1 May. Note, however, that we have
suggested above that any disturbance may have serious consequences for
offspring and mother’s welfare. In this context pursuit even of male deer
during this period may thus involve disturbance, movement and stress to
heavily pregnant females. One of the concerns about hunting red stags with
dogs in England (now banned but which used to continue until the end of
April) was that it might cause disturbance to heavily pregnant hinds.
Female moose and alpine chamois are hunted well outside the period of
late pregnancy/parturition time. For moose the close season extends from the
end of December to the beginning of September in the more permissive cases;
for chamois females are not hunted from end December/January (with the
exception of France, 28 February) to beginning of August (16 July in
Austria).
By contrast, in the case of wild boar, sow hunting is allowed, in the
majority of cases, at a time that virtually guarantees that at least a proportion
of mature females are in an advanced stage of pregnancy (31 January or as
later as 31 March). In Austria, Estonia and Portugal females can be hunted
year-round. In respect of this, however, we do recognise that hunting during
the last part of pregnancy may be considered necessary when the aim is to
reduce or control ungulate density. For wild boar, culling during this period
may be simply inevitable in order to enable hunters to exercise some degree of
control over a species which has a very rapid reproductive potential. Despite
this we note that there may be implications of extended hunting seasons on
other, non-target species. In the same way that hunting red deer stags with
94 Marco Apollonio, et al.
hounds in England as late as April potentially caused stress to pregnant
females (above), extended hunting of wild boar – particularly by the most
usual methods of driving and coursing – might actually result in serious
disturbance for other species.
4.3.3 Culling during the period of juvenile dependency
Finally, we note that under the current hunting seasons in many countries,
there remains a potential for killing females while young may still be nutri-
tionally or socially dependent on them in at least the three more abundant
species: wild boar, roe deer and red deer.
We may broadly assume (over the latitudinal range) that the period of
parturition for roe deer is from late April or early May to end of June and
that for red deer is from mid May to the end of June. For roe deer it is
apparent that there is some considerable variation among geographical areas.
Although fawning season is quite ‘tight’, with 80% of fawns born within
20–30 days of median date of parturition (Irvine, 2004), that median date
itself may vary from 11 May to 13 June in different locations (Linnell and
Andersen, 1998). There is, however, no simple relationship with latitude,
although birth dates do follow some pattern, with southern and Atlantic coast
populations giving birth in general before inland, continental ones (Linnell
et al., 1998). For red deer, there appears to be far less variation and populations
in most areasgive birth over the same range of dates (26 May–15 June; Fletcher,
1974). Given these general dates, we may speculate that neonates not accom-
panying the mother may be orphaned if mothers are shot before say mid June
(roe) or mid July (red), while unless culled with the mother, juveniles of either
species nutritionally dependent on lactating dams will die if mothers are shot
before the end of August (see also Putman, 2008; Apollonio et al., 2010).
On such a basis it is clear that, with seasons for mature red deer females in
most countries not opening until September (Italy, Croatia, Slovenia, Hungary,
Romania – all central European, with earlier breeding seasons anyway) or
October (Denmark, Norway, Sweden, Wallonia (Belgium), Poland, Estonia,
Lithuania), cull seasons may be considered outside the period of maximum
welfare risk. Some countries delaythe commencement of the season even further
(November in England and Republic of Ireland). However, we may note
that seasons in the Netherlands, Latvia, the Czech Republic, Slovakia and
Switzerland open as early as the beginning of August.
In the majority of countries in Europe hunting of adult roe deer females is
not permitted before the beginning of September, but in some cases an early
open season for mature females is allowed: this is the case in Austria, Latvia,
Hunting seasons in relation to breeding seasons 95
Switzerland, Spain and the Italian Apennines. In some cases the extension of
the season is not great (e.g. from August in the Italian Apennines), but this
may nonetheless be in an ecological context that may also lead to early birth
dates. In other cases the open season may start from May so there can be little
doubt about the potential and actual possibility that female culls may indeed
leave orphaned dependent fawns.
Moose as a species does not seem to suffer from any real problems with
permitted seasons: open season for adult females ranges from 1 September
(Latvia) to 1 October (Lithuania) so we can rule out the possibility that a
suckling young could be orphaned as birth dates happen in the last ten days
of May to early June (Bowyer et al., 1998). Variation in the start-date for
hunting of female alpine chamois is from 16 July (the earliest, in Austria) to
15 October (Romania), with most seasons in the actual alpine range starting
in August. In Italy culling opens in September. Most births in the alpine
countries are concentrated in the month of May. As a consequence there
seems to be some room for problems connected with early shooting of
lactating females.
In ten countries (Belgium, Spain, Italy, Lithuania, Poland, Denmark,
Czech Republic, Slovenia, Romania, Greece) wild boar sows are not culled
before August but in 12 countries sows may be legally taken well before
this time: it is surely no coincidence that some of the European countries
with the highest number of wild boar (and the consequent highest toll in
damage to agriculture) are within the list, as France, Germany and Austria.
Finally it is interesting to note that in some countries (e.g. Austria,
Finland) killing a sow with dependent piglets is explicitly prohibited. Wild
boar have the largest litter size of any European ungulate species with high
social interactions. When the mother–young bond is still present, a good
practice might be to cull only the piglets. Once again, the problem is even
more exaggerated in relation to wild boar, since even the peak period of
parturition in this species (when most births will occur) may extend over a
period of three months or more (for example in Spain, the birth period
occurs from February to April: Fernandez-Llario and Mateos-Quesada,
1998) and there are occasional females giving birth throughout the
entire year.
4.4 Conclusions
Not all species are equally susceptible to all the problems explored in our
introduction and we must recognise that some of the problems explored are
potential rather than necessarily actual.
96 Marco Apollonio, et al.
The particular breeding biology of individual species (both mating systems
and timing of reproduction) plays a decisive role in making them more or less
vulnerable to the various potential issues we have suggested. It is also clear
that breeding systems vary within individual species (so that there is not even
a single species-specific system which applies universally) and that the timing
of both rutting period and parturition may vary with latitude and geography.
It is important to make it clear therefore that there can be no ‘hard and fast’
general rules.
In addition not all hunting practices have the same kind of impact: single-
hunter selective hunting with rifle from fixed blinds or high seats is at the
lowest level of impact, whereas large drives with dogs and beaters are the
maximum (Swenson, 1982; Kufeld et al., 1988; Kilgo et al., 1998).
Monterı´ais a typical Spanish hunting practice for big game where several
packs of dogs are released within at least 500 hectares of forest or scrubland
(Carranza, 2010). There are some animal welfare concerns surrounding the
use of hounds in hunting, but a positive side of the monterı´asystem is that
each portion of land is hunted only once per year. Moreover, species not
forming large breeding associations (such as roe deer) are presumably less
subject to hunting disturbance during the rut as any given disturbance poten-
tially affects a few deer per time at maximum. However, the potential for
disturbance of non-target species, while rarely taken into account, may
represent a relevant source of distress. Thus, for example the potential impact
of wild boar hunting with hounds on roe deer during the rut could be
important, particularly if it occurs repeatedly in the same areas. These situ-
ations may generate further constraints in setting hunting seasons.
Environmental constraints can also be important in the decision of
hunting times of some species: in the Alps for instance it is hard to hunt
higher than 1500 metres a.s.l. during winter in years of average snow and
environmental difficulties suggest that it is inappropriate to stress ungu-
lates further by hunting them in a period of food shortage and harsh
climate.
Legislators and wildlife managers must thus consider a number of different
factors when setting seasons, which tend to change among different areas.
Hunting season, and hunting practice, must reflect management needs and
objectives: whether directed for conservation of threatened populations or
rare taxa, to support and sustain recreational hunting, or to ensure control of
overabundant species, or impacts of ungulates on agriculture and forestry.
However, to deliver those without risk of welfare or other problems outlined
above, they must also take into account accurate information about (local)
seasonality of the rut and of parturition of different species (as well as the risk
Hunting seasons in relation to breeding seasons 97
of hunting seasons of one species causing problems in other non-target
species sharing the same habitat), length of harassment and, last but by no
means least, social traditions.
Local traditions may play an important, and not always positive, role in
the attempt to find more scientifically sound hunting practices for ungulates.
For example, hunting red deer stags by imitating the call during the rut is so
deeply rooted in the cultural hunting tradition of Central European countries
that specific competitions of calling ability are held among hunters; the
hunting of alpine chamois during the rut allows hunters who adhere to a
more Germanic tradition to obtain with the kill of a mature male his
‘Gamsbart’ – the long hairs that are erected along the backbone by the rutting
males are exhibited in a dense brush on the traditional mountain hunter cap.
Within such constraints, however, we examine, in conclusion, some pos-
sible changes which might be considered in the timing of seasons to try and
overcome some of the shortcomings identified earlier in the chapter. A caveat
is necessary here, however: as we have had occasion to note earlier, not all
species in all countries are managed to maintain intact their potential repro-
ductive output or to manage populations for stability or to encourage actual
expansion in population size. In many instances ungulate populations are
managed instead to try to exercise some control over populations and their
impacts and actually to deliver some reduction in population size or distribu-
tion. In such contexts it is obvious that the primary focus is not concerned
with the conservative management of the population; however, the mainten-
ance of a healthy, balanced population and the avoidance of inhumane
hunting practices remains a goal that has a general validity.
4.4.1 Culling during the rut
Because of the various negative implications we have rehearsed at the begin-
ning of this chapter, as a general rule it would be wise not to hunt ungulates at
all during their rut. In practice, however, for many reasons this would
probably be difficult to establish across the entire range. A partial solution
could be the protection of at least some traditional and important breeding
ground for any given species, as red deer on the Alps. This kind of approach
is obviously limited to situations where there are specific and localised areas
where most individuals of a population do reproduce; it would not be
applicable in situations where breeding grounds are widely distributed across
the whole landscape, as in the case of the red deer in the Scottish Highlands.
An alternative may be to limit hunting just to the second half of the rut,
leaving at least some portion of this important biological activity free from
98 Marco Apollonio, et al.
disturbance. The choice of the first instead of the last half is advisable because
early conception and consequently early birth are linked to better chance of
survival of young; in addition it is generally the most competitive, fitter males
which are active early in the season. In addition, we believe there should be
stringent regulation that males culled should be the poorer individuals (indi-
viduals showing signs of age or ill health, or those with poor antlers for their
age, poor body conformation) since retention within the population of those
males of better quality is crucial to maintaining overall quality and fitness of
the population.
More generally, in those countries where hunting does occur during the
mating season and where there is a long social tradition associated with this
practice, we advise that numbers culled within this period should be at least
restricted. This would help to reduce harassment and also to encourage
hunters to use other hunting practices and periods. Hunters will be more
favourable to suspend culling during the rut if they have successfully hunted
during other periods.
4.4.2 Culling during advanced pregnancy
One possible solution to the problems associated with hunting close to the
period of parturition is to ban hunting when females are close to giving birth.
However, as noted above, actual culling of pregnant females is not a problem
per se; rather it is the use of hunting practices which involve pursuit or
disturbance of a population of females at this time, whether during culling of
females themselves, or during the hunting of males, or even of other ungulate
species. Thus consideration should rather be given to banning of particular
hunting methods rather than necessarily a complete ban on hunting altogether.
We recognise that there are species like wild boar in which a female is
theoretically able to give birth at any time of the year and also at any age (a
female of only 8–9 months old is able to reproduce under favourable circum-
stances). In such cases it is clearly impracticable to ban hunting during
periods where at least some females are bound to be heavily pregnant. In
the few regions without wild boar damage, managers could minimize the risk
of culling sows during late pregnancy by recognising the peak birth months in
their particular region and to stop hunting at that time. But such a solution is
not widely applicable. In almost every European country where they occur,
wild boar are a problem, and attempts to control wild boar populations are
failing dismally. For this reason managers and law makers must encourage
and facilitate hunting activities aimed at reducing wild boar density and
rebuilding a more natural population structure.
Hunting seasons in relation to breeding seasons 99
Even so, in cases like this when the aim of the management is to effect a
major reduction of the population size, welfare and ethical constraints might
suggest that culling of mature females should be concentrated where possible
in the early stages of pregnancy and only, in extreme cases, during later
pregnancy. In addition where culling is required, or permitted later in the
period of pregnancy, consideration should again be given to hunting method,
and during this time culling should be restricted to only those methods which
cause minimum collateral disturbance. In this respect even if a ‘surgical cull’
(for example, stalking with rifles from high seats) of females during these
times could still be considered acceptable, where minimum disturbance is
occasioned to other females in the population, any hunting practice causing
wider stress (like hunting with hounds) should be avoided.
4.4.3 Culling during the period of juvenile dependency
Logically a priority in this context would be to ensure that no culling of
females is allowed during the period when neonatal young may still be
concealed and not accompanying the mother. It is difficult to define the
actual end of hiding behaviour so a prudent suggestion would be a hunting
ban period from the first likely date of parturition in a given area to some
three weeks after the latest possible date of parturition. After that period we
suggest that any shooting of mature females must be avoided in the time in
which juvenile nutrition is primarily dependent on lactation, or, if females
must be shot, that the young should always be shot before the mother.
After this first period of strict nutritional dependence, a more ambiguous
period begins over which the young are still socially dependent upon the
mother. This period is poorly determined and clearly differs between different
species. Typically, the social bond between mother and offspring generally
continues until at least one year of age in most European ungulate species.
We suggest therefore that in a strict conservation perspective, the non-hunting
season for adult females should extend at least until the end of strict social
dependence. However, this is probably impracticable in most situations,
especially for species or populations where a primary aim of management is
to control expanding populations and limit damaging impacts. At the very least
therefore, we would suggest, as above, if females must be shot, that the young
should always be shot before the mother. Moreover it is strongly suggested that
some monitoring programme be put in place (as, for example, monitoring of
average weight of males and females as yearlings) to assess if any management
option that includes the cull of mature females with dependent young has an
impact on population quality. Finally we think that during breeding seasons
100 Marco Apollonio, et al.
hunting is undesirable. If managers plan to cull during these periods, they must
be aware of the potential consequences and they must foresee the best
approaches in order to reduce or to avoid negative biological outcomes.
References
Aanes, R. and Andersen, R. (1996) The effects of sex, time of birth, and habitat on
the vulnerability of roe deer fawns to red fox predation. Canadian Journal of
Zoology 74, 1857–1865.
Albon, S.D., Mitchell, B., Huby, B.J. and Brown, D. (1986) Fertility in female red
deer (Cervus elaphus): the effects of body composition, age and reproductive
status. Journal of Zoology (London),209, 447–460.
Apollonio, M., Festa-Bianchet, M., Mari, F., Mattioli, S. and Sarno, B. (1992)
To lek or not lek: mating strategies of male fallow deer. Behavioural Ecology
3, 25–31.
Apollonio, M., Andersen, R. and Putman, R. (2010) Present status and future
challenges for European ungulate management. In M. Apollonio, R. Andersen
and R. Putman (eds.) European Ungulates and their Management in the
21st Century. Cambridge, UK: Cambridge University Press, pp.578–604.
Bartos
ˇ, L. and Bahbouh, R. (2006) Antler size and fluctuating asymmetry in red deer
(Cervus elaphus) stags and the probability of becoming a harem holder in the rut.
Biological Journal of the Linnean Society 87, 59–68.
Bartos
ˇ, L., Zeeb, U. and Mikes
ˇ, J. (1992) Lekking behaviour in sika deer. In
N. Maruyama, B. Bobek, Y. Ono, et al. (eds.) International Symposium on
Wildlife Conservation: Present Trends and Perspectives for the 21st Century.
Tokyo: Japan Wildlife Research Center, pp.205–208.
Bartos
ˇ, L., Herrmann, H., S
ˇiler, J., Losos, S. and Mikes
ˇ, J. (1998) Variation of
mating systems of introduced sika deer. Revue d’Ecologie – La Terre et la Vie
53, 337–345.
Bartos
ˇ,L.,S
ˇustr, P., Janovsky` , P. and Bertagnoli, J. (2003) Sika deer (Cervus nippon)
lekking in a free-ranging population in Northern Austria. Folia Zoologica 52,
1–10.
Bender, L.C., Lomas, L.A. and Browning, J. (2007) Condition, survival, and cause-
specific mortality of adult female mule deer in north-central New Mexico.
Journal of Wildlife Management 71, 1118–1124.
Bender, L.C., Cook, J.G., Cook, R.C. and Hall, P.B. (2008) Relations between
nutritional condition and survival of North American elk, Cervus elaphus.
Wildlife Biology 14, 70–80.
Bowyer, R.T., van Ballenberghe, V. and Kie, J.G. (1998) Timing and synchrony of
parturition in Alaskan moose: long-term versus proximal effects of climate.
Journal of Mammalogy 79, 1332–1344.
Briedermann, L. (1986) Schwarzwild. Berlin: Neumann-Neudamm.
Briedermann, L. (1992) Reproduction in European mouflon. In F. Spitz, G. Jeneau,
G. Gonzalez and S. Aulagnier (eds.) Ongule
´es/Ungulates 91. Toulouse, France:
IRGM, pp.557–560.
Carranza, J. (1995) Female attraction by males versus sites in territorial rutting red
deer. Animal Behaviour 50, 445–453.
Carranza, J. (2010) Ungulates and their management in Spain. In M. Apollonio,
R. Andersen, and R. Putman (eds.) European Ungulates and their Management
in the 21st Century. Cambridge, UK: Cambridge University Press, pp.419–440.
Hunting seasons in relation to breeding seasons 101
Carranza, J., Garcia-Mun
˜oz and de Dios Vargas, J. (1995) Experimental shifting
from harem defence to territoriality in rutting red deer. Animal Behaviour 49,
551–554.
Carrio
´n, D., Garcı
´a, A.J., Gaspar-Lo
´pez, E., Landete-Castillejos, T. and Gallego, L.
(2008) Development of body condition in hinds of Iberian red deer during
gestation and its effects on calf birth weight and milk production. Journal of
Experimental Zoology A–Ecological Genetics and Physiology 309, 1–10.
Chapman, D.I. and Chapman, N.G. (1975) Fallow Deer: Their History, Distribution
and Biology. Lavenham, UK: Terence Dalton, 271pp.
Childress, M.J. and Lung, M.A. (2003) Predation risk, gender and the group size
effect: does elk vigilance depend upon the behaviour of conspecifics? Animal
Behaviour 66, 389–398.
Clutton-Brock, T.H., Albon, S.D. and Guinness, F.E. (1985) Parental investment and
sex differences in juvenile mortality in birds and mammals. Nature 313, 131–133.
Clutton-Brock, T.H., Albon, S.D. and Guinness, F.E. (1987a) Interactions between
population density and maternal characteristics affecting fecundity and juvenile
survival in red deer. Journal of Animal Ecology 56, 857–871.
Clutton-Brock, T.H., Major, M., Albon, S.D. and Guinness, F.E. (1987b) Early
development and population dynamics in red deer. I. Density-dependent effects
on juvenile survival. Journal of Animal Ecology 56, 53–67.
Clutton-Brock, T.H., Albon, S.D. and Guinness, F.E. (1988a) Reproductive success
in male and female red deer. In T.H. Clutton-Brock (ed.) Reproductive Success:
Studies of Individual Variation in Contrasting Breeding Systems. Chicago, IL:
University of Chicago Press, pp.325–343.
Clutton-Brock, T.H., Green, D., Hiraiwa-Hasegawa, M. and Albon, S.D. (1988b)
Passing the buck: resource defence, lek breeding and mate choice in fallow deer.
Behavioral Ecology and Sociobiology 23, 281–296.
Clutton-Brock, T.H., Deutsch, C.J. and Nefdt, R.J.C. (1993) The evolution of
ungulate leks. Animal Behaviour 46, 1121–1138.
Coltman, D.W., O’Donoghue, P., Jorgenson, J.T., et al. (2003) Undesirable evolu-
tionary consequence of trophy hunting. Nature 426, 655–658.
Coˆ te
`, S.D. and Festa-Bianchet, M. (2001) Birthdate, mass and survival in mountain
goat kids: effects of maternal characteristics and forage quality. Oecologia 127,
230–238.
Feder, C., Martin, J.G.A., Festa-Bianchet, M., Be
´rube
´, C. and Jorgenson, J. (2008)
Never too late? Consequences of late birthdate for mass and survival of bighorn
lambs. Oecologia 156, 773–781.
Ferdman, N., Murmu, R.P., Bock, J., Braun, K. and Leshem, M. (2007) Weaning
age, social isolation, and gender, interact to determine adult explorative and
social behavior, and dendritic and spine morphology in prefrontal cortex of rats.
Behavioural Brain Research 180, 174–182.
Fernandez-Llario, P. and Mateos-Quesada, P. (1998) Body size and reproductive
parameters in wild boar, Sus scrofa. Acta Theriologica 43, 439–444.
Festa-Bianchet, M. (1988a) Nursing behaviour of bighorn sheep: correlates of ewe
age, parasitism, lamb age, birthdate and sex. Animal Behaviour 36, 1445–1454.
Festa-Bianchet, M. (1988b) Seasonal range selection in bighorn sheep: conflicts
between forage quality, forage quantity, and predator avoidance. Oecologia
75, 580–586.
Festa-Bianchet, M. (2003) Exploitative wildlife management as a selective pressure
for the life-history evolution of large mammals. In M. Festa-Bianchet and
102 Marco Apollonio, et al.
M. Apollonio (eds.) Animal Behavior and Wildlife Conservation. Washington,
DC: Island Press, pp.191–207.
Festa-Bianchet, M., Urquhart, M. and Smith, K.G. (1994) Mountain goat recruit-
ment: kid production and survival to breeding age. Canadian Journal of Zoology
72, 22–27.
Festa-Bianchet, M., Jorgenson, J.T. and Re
´ale, D. (2000) Early development,
adult mass, and reproductive success in bighorn sheep. Behavioral Ecology 11,
633–639.
Fletcher, T.J. (1974) The timing of reproduction in red deer (Cervus elaphus)in
relation to latitude. Journal of Zoology 172, 363–367.
Francis, D., Diorio, J., Liu, D. and Meaney, M.J. (1999) Nongenomic transmission
across generations of maternal behavior and stress responses in the rat. Science
286, 1155–1158.
Gendreau, Y., Coˆ te
`, S.D. and Festa-Bianchet, M. (2005) Maternal effects on post-
weaning physical and social development in juvenile mountain goats. Behavioral
Ecology and Sociobiology 58, 237–246.
Grignolio, S., Parrini, F., Bassano, B., Luccarini, S. and Apollonio, M. (2003)
Habitat selection in adult males of Alpine ibex, Capra ibex ibex.Folia Zoologica
52, 113–120.
Hamilton, W.J. and Blaxter, K.L. (1980) Reproduction in farmed deer. I. Hind and
stag fertility. Journal of Agricultural Science, Cambridge,95, 261–273.
Harris, R.B., Wall, W.A. and Allendorf, F.W. (2002) Genetic consequences of
hunting: what do we know and what should we do? Wildlife Society Bulletin
30, 634–643.
Heck, L. and Raschke, G. (1980) Die Wildsauen. Hamburg, Germany: Paul Parey.
Irvine, J. (2004) Calving Dates: a Literature Review. Report to DCS on contract RP28.
Summary in Deer Commission for Scotland Annual Report 2003–04, pp.52–53.
Jarnemo, A. (2004) Predation processes: behavioural interaction between red fox and
roe deer during the fawning season. Journal of Ethology 22, 167–173.
Kikusui, T., Kiyokawa, Y. and Mori, Y. (2007) Deprivation of mother-pup inter-
action by early weaning alters myelin formation in male, but not female, ICR
mice. Brain Research 1133, 115–122.
Kikusui, T., Nakamura, K. and Mori, Y. (2008) A review of the behavioural and
neurochemical consequences of early weaning in rodents. Applied Animal Behav-
iour Science 110, 73–83.
Kilgo, J.C., Labisky, R.F. and Fritzen, D.E. (1998) Influences of hunting on the
behaviour of white-tailed deer: implications for conservation of the Florida
panther. Conservation Biology 12, 1359–1364.
Kjellander, P. and Nordstrom, J. (2003) Cycles voles, prey switching in red fox, and
roe deer dynamics: a test of the alternative prey hypothesis. Oikos 101, 338–344.
Kruuk, L.E.B., Slate, J., Pemberton, J.M., et al. (2002) Antler size in red deer:
heritability and selection but no evolution. Evolution 56, 1683–1695.
Kufeld, R.C., Bowden, D.C. and Schrupp, D.L. (1988) Influence of hunting on
movements of female mule deer. Journal of Range Management 41, 70–72.
Lamberti, P., Mauri, L. and Apollonio, M. (2004) Two distinct patterns of spatial
behaviour of female roe deer (Capreolus capreolus) in a mountain habitat.
Ethology, Ecology, and Evolution 16, 41–53.
Langbein, J. (1991) Effects of density and age on body condition, reproductive
performance, behaviour and survival of fallow deer. PhD thesis, University of
Southampton, UK.
Hunting seasons in relation to breeding seasons 103
Langbein, J. and Thirgood, S.J. (1989) Variation in mating systems of fallow deer
(Dama dama) in relation to ecology. Ethology 83, 195–214.
Langbein, J., Chapman, N.G. and Putman, R.J. (2008) Fallow deer, Dama dama.In
S. Harris and D.W. Yalden (eds.) Mammals of the British Isles: Handbook, 4th
edn. London: The Mammal Society, pp.595–604.
Lent, P.C. (1974) Mother-infant relations in ungulates. In V. Geist and F. Walther
(eds.) The Behaviour of Ungulates and its Relation to Management. Morges,
Switzerland: IUCN, pp.14–55.
Li, Z., Jiang, Z. and Beauchamp, G. (2009) Vigilance in Przewalski’s gazelle: effects
of sex, predation risk and group size. Journal of Zoology 277, 302–308.
Liberg, O., Johansson, A., Andersen, R. and Linnell, J.D.C. (1998) Mating system,
mating tactics and function of male territoriality in roe deer. In R. Andersen,
P. Duncan and J.D.C. Linnell (eds.) The European Roe Deer: The Biology of
Success. Oslo: Scandinavian University Press, pp.221–256.
Linnell, J.D.C. and Andersen, R. (1998) Timing and synchrony of birth in a hider
species, the roe deer Capreolus capreolus.Journal of Zoology (London)244,
497–504.
Linnell, J.D.C., Aanes, R. and Andersen, R. (1995) Who killed Bambi? The role of
predation in the neonatal mortality of temperate ungulates. Wildlife Biology 1,
209–223.
Linnell, J.D.C., Wahlstrom, K. and Gaillard, J.-M. (1998) From birth to independ-
ence: birth, growth, neonatal mortality, hiding behaviour and dispersal. In
R. Andersen, P. Duncan and J.D.C. Linnell (eds.) The European Roe Deer:
The Biology of Success. Oslo: Scandinavian University Press, pp.257–284.
Morrison, J.A. (1960) Characteristics of estrus in captive elk. Behaviour 16, 84–92.
Mysterud, A., Solberg, E.J. and Yoccoz, N.G. (2005) Ageing and reproductive effort
in male moose under variable levels of intrasexual competition. Journal of
Animal Ecology 74, 742–754.
Nakamura, K., Kikusui, T., Takeuchi, Y. and Mori, Y. (2008) Changes in social
instigation- and food restriction-induced aggressive behaviors and hippocampal
5HT1B mRNA receptor expression in male mice from early weaning. Behav-
ioural Brain Research 187, 442–448.
Nicholson, M.C., Terry Bowyer, R. and Kie, J.G. (1997) Habitat selection and
survival of mule deer: tradeoffs associated with migration. Journal of Mammal-
ogy 78, 483–504.
Paarlberg, K.M., Vingerhoets, A.D.J.J.M., Passchier, J., Dekker, G.A. and
vanGeijn, H.P. (1995) Psychosocial factors and pregnancy outcome: a review
with emphasis on methodological issues. Journal of Psychosomatic Research 39,
563–595.
Panzacchi, M., Linnell, J., Odden, M., Odden, J. and Andersen, R. (2009) Habitat
and roe deer fawn vulnerability to red fox predation. Journal of Animal Ecology
78, 1124–1133.
Pettorelli, N., Pelletier, F., von Hardenberg, A., Festa-Bianchet, M. and Coˆ te
´, S.D.
(2007) Early onset of vegetation growth vs. rapid green-up: impacts on juvenile
mountain ungulates. Ecology 88, 381–390.
Pipia, A., Ciuti, S., Grignolio, S., Luchetti, S., Madau, R. and Apollonio, M. (2009)
Effect of predation risk on grouping pattern and whistling behaviour in a wild
mouflon Ovis aries population. Acta Theriologica 54, 77–86.
Putman, R.J. (1993) Flexibility of social organisation and reproductive strategy in
deer. Deer 9, 23–28.
104 Marco Apollonio, et al.
Putman, R.J. (2008) A Report on Current Perception, Legal Status and Expectation
with Respect to Deer Welfare in Other Countries. Contract report for the Deer
Commission for Scotland.
Røed, K.H., Holand, ., Mysterud, A., et al. (2007) Male phenotypic quality influ-
ences offspring sex ratio in a polygynous ungulate. Proceedings of the Royal
Society B–Biological Sciences 274, 727–733.
Schaal, A. and Bradbury, J.W. (1987) Lek breeding in a deer species. Biology of
Behaviour 12, 28–32.
Singer, F.J. and Zeigenfuss, L.C. (2002) Influence of trophy hunting and horn size on
mating behavior and survivorship on mountain sheep. Journal of Mammalogy
83, 682–698.
Stopher, K.V., Pemberton, J.M., Clutton-Brock, T.H. and Coulson, T. (2008) Indi-
vidual differences, density dependence and offspring birth traits in a population
of red deer. Proceedings of the Royal Society B–Biological Sciences 275, 2137–
2145.
Strandgaard, H. (1972) The roe deer (Capreolus capreolus) population at Kalø and
the factors regulating its size. Danish Review of Game Biology 7, 1–205.
Swenson, J.E. (1982) Effects of hunting on habitat use by mule deer on mixed-grass
prairie in Montana. Wildlife Society Bulletin 10, 115–120.
Thirgood, S.J., Langbein, J. and Putman, R.J. (1999) Intraspecific variation in
ungulate mating strategies: the case of the flexible fallow deer. Advances in the
Study of Behavior 28, 333–361.
Tu
¨rcke, F. and Schmincke, S. (1965) Das Muffelwild. Hamburg, Germany: Paul
Parey.
Veiberg, V., Loe, L.E., Mysterud, A., Langvatn, R. and Stenseth, N.C. (2004) Social
rank, feeding and winter weight loss in red deer: any evidence of interference
competition? Oecologia 138, 135–142.
von Hardenberg, A., Bassano, B., Peracino, A. and Lovari, S. (2000) Male Alpine
chamois occupy territory at hotspots before the mating season. Ethology 106,
617–630.
Willisch, C.S. and Neuhaus, P. (2009) Alternative mating tactics and their impact on
survival in adult male Alpine ibex (Capra ibex ibex). Journal of Mammalogy 90,
1421–1430.
Hunting seasons in relation to breeding seasons 105
