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588

NOTE / NOTE

Age- and sex-specific settlement patterns of


chamois (Rupicapra rupicapra) offspring
A. Loison, G. Darmon, S. Cassar, J.-M. Jullien, and D. Maillard

Abstract: The social, spatial, and genetic structures of populations depend on where offspring settle and reproduce
in relation to their parents home range. However, the patterns of settlement in wild populations of large mammals
are often poorly described owing to the difficulty of monitoring motheroffspring pairs over a long period. Here,
we investigated sex-specific settlement patterns in chamois (Rupicapra rupicapra (L., 1758)) based on the study of
31 motheroffspring pairs. We calculated the distance between the center of the mothers range and the center of
her offsprings range, and tested whether this distance differed when the offspring was immature (i.e., a yearling)
and after offspring sexual maturity (>2 years of age). We found no sex effect on the distance between centers of
mother and offspring ranges for yearling offsprings. However, mature sons ranged farther away from their mother
than mature daughters. Daughters appear to settle close to their mothers home range. The distance at which a
daughter settles compared with her mothers range seems to be determined before 2 years of age. On the contrary,
the distance between the center of the locations of yearling males and the center of locations of their mother does
not predict how far away males will eventually settle when mature. We discuss the implications of these patterns
for generating female social structures, as well as population spatial and genetic structures.
Resume : Les structures sociales, spatiales et genetiques des populations dependent du lieu ou` les rejetons vont sinstaller
et se reproduire par rapport au domaine vital de leur parents. Cependant, les patrons dinstallation dans les populations
sauvages de grands mammife`res sont souvent peu decrits car il est difficile de suivre les couples me`res-jeunes sur une
periode longue. Ici, nous avons etudie les patrons dinstallation chez le chamois (Rupicapra rupicapra (L., 1758)) en
fonction de leur sexe chez 31 couples me`res-jeunes. Nous avons calcule la distance entre le centre du domaine vital de
la me`re et le centre de celui de son jeune et teste si la distance differait quand le jeune etait immature (jeune de 1 an)
et avait atteint la maturite sexuelle (>2 ans). Nous navons pas trouve deffet du sexe sur la distance entre les centres
des domaines de la me`re et du jeune pour les jeunes immatures. Cependant, les fils adultes sont partis sinstaller plus
loin de leur me`re que les filles adultes. La distance a` laquelle une fille sinstalle par rapport au domaine vital de sa
me`re semble determinee avant lage de 2 ans. Au contraire, la distance entre les centres des localisations des males de 1
an et le centre des localisations de la me`re ne permet pas de predire la distance dinstallation du fils lorsquil est adulte.
Nous discutons de limplication de ces patrons dans lemergence des structures sociales femelles et des structures spatiale
et genetique des populations.

Introduction
Mammal offspring are obliged to stay close to their
mother until the end of the lactation period, and often do so
longer, as parental care usually extends beyond lactation
(Clutton-Brock 1991). However, when the offspring grow
older, the intensity of parentoffspring conflict increases
because of competition for food and reproduction, or because
of the presence of new siblings (but for cooperative mammals

see Solomon and French 1997). Offspring are then faced


with the choice to remain close to their natal site (philopatry) or to leave (dispersal). The balance between the
costs and the benefits of philopatry and dispersal tactics
may be affected by mating system, sex or social structure,
meta-population context, or local population density, as
well as a wide range of individual characteristics like
body condition or phenotype (Greenwood 1980, Matthysen
2005). In polygynous species where mothers invest heavily

Received 18 September 2007. Accepted 4 March 2008. Published on the NRC Research Press Web site at cjz.nrc.ca on 16 May 2008.
A. Loison,1,2 G. Darmon, and S. Cassar. Universite de Lyon, universite Lyon 1, Centre National de la Recherche Scientifique (CNRS),
Unite Mixte de Recherche (UMR) 5558, Laboratoire de Biometrie et Biologie Evolutive, 43 boulevard du 11 novembre 1918,
Villeurbanne F-69622, France.
J. Jullien and D. Maillard. Office National de la Chasse et de la Faune Sauvage (ONCFS), Centre National dEtude et de Recherche
Appliquee (CNERA) Faune de Montagne, 95 rue Pierre Flourens, BP74267, 34098 Montpellier CEDEX 05, France.
1Corresponding
2Present

author (e-mail: anne.loison@univ-savoie.fr).


address: Universite de Savoie, CNRSUMR 5553 Laboratoire dEcologie Alpine, 73376 Le Bourget du Lac, France.

Can. J. Zool. 86: 588593 (2008)

doi:10.1139/Z08-031

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Loison et al.

in offspring, females are usually more philopatric than


males, a pattern normally accounted for by three nonexclusive hypotheses: (1) the resource competition hypothesis,
by which females should be more philopatric owing to
higher costs for females to disperse compared with males
(Greenwood 1980); (2) the local mate competition hypothesis, according to which males should disperse more than
females owing to the strong malemale competition for access to females (Johnson and Gaines 1990); and (3) the inbreeding avoidance hypothesis, which would lead the sex
with the largest variance in reproductive success to disperse, i.e., males in polygynous species (Moore and Ali
1984; Pusey 1987). Female philopatry can lead to matrilineal structures, where females stay within their mothers social group or subpopulation area (red deer (Cervus elaphus
L., 1758): Albon et al. 1992; Conradt et al. 1999; Soay
sheep (Ovis aries L., 1758): Coulson et al. 1999; bighorn
sheep (Ovis canadensis Shaw, 1804): Festa-Bianchet 1986;
white-tailed deer (Odocoileus virginianus (Zimmermann,
1780)): Nelson and Mech 1987). Investigating the sexspecific patterns of settlement is therefore a major step
toward understanding the social, spatial, and genetic structures of a population.
Indirect methods based on genetics have enabled the
investigation of genetic spatial structure and the consequences of effective dispersal (e.g., Coltman et al. 2003).
However, direct observations of spatial and social behavior
are required to determine the timing of dispersal and the
small-scale consequences of philopatry on social structures
and local densities (Byers 1997; Coulson et al. 1999), as
well as increasing our understanding of the dispersal process
which is often complex and protracted. There are, however,
few observational studies on large mammals in the literature
because of the scarcity of wild populations in which the
natal site of individuals and kinship are known (but see
Festa-Bianchet 1986, 1991; Byers 1997; Coulson et al.
1999; Clutton-Brock and Pemberton 2004). Here, we
investigate whether male and female offspring settle close
to their mother in a chamois (Rupicapra rupicapra (L.,
1758)) population based on the monitoring of individually
marked motheroffspring pairs. We focused on sex-specific
timing of movements and sex-specific distances between
center of activities of mothers and center of activities of
offspring. In a dimorphic and polygynous species such as
the chamois (Loison et al. 1999b), we expected sons to
settle farther away from their mothers than daughters,
according to the inbreeding avoidance or local mate competition hypotheses for males (Johnson and Gaines 1990).
However, the larger distance between mothers and offsprings centers of activities should be only apparent when
males are mature (Dobson 1982). If dispersal is caused by
resource competition, we should expect either no sex difference in settlement patterns or a higher distance between the
mothers and the offspring of the sex most likely to compete
for resources. In the chamois case, this would be females,
which live in labile social groups (Boschi and Nievergelt
2003; Crampe et al. 2007). After testing for age and sex effects on how far offspring settle from their mothers distances, we discuss the small-scale consequences of philopatry
versus dispersal in terms of subpopulation and group spatial
structure.

589
Fig. 1. Map of the study area, with the location of the three marking sites. The border of the Game Reserve is indicated in black.
The darker the shading, the higher the elevation (300 m per shade
of gray).

Material and methods


Study site
The study site was the National Game Reserve (NGR)
Les Bauges in the northern Alps (45840N, 6813E) situated
in France. The NGR Les Bauges encompasses 5 205 ha
and is part of a typical pre-alpine massif covering a total
area of 15 600 ha, separated from other mountain units by
large valleys and the Annecy lake. The NGR Les Bauges
is structured around main summits separated by sharp
valleys (Fig. 1) and constitutes an isolated geographical
unit. Elevations range between 700 and 2200 m with about
2% over 2000 m. One-half of the area is covered by forest.
The treeline is at about 1500 m. Population size was at least
1250 alpine chamois (Houssin et al. 1994; Loison et al.
2006).
Chamois have been captured every year since 1985 at
three marking sites (Fig. 1) using cage-traps and drop nets,
the latter since 1991. Since 1985, more than 1300 chamois
have been captured, of which 650 have been marked with
visual collars, and released, while the remaining were
exported for reintroduction purposes outside the study area.
Each collars combination of colors and signs allows individuals to be identified. Both traps and drop nets were
baited with salt. Kids were fitted with expandable collars,
while adults were fitted with fixed-length collars (RichardHansen 1992; Loison et al. 1999c, 2006; Crampe et al.
2002). Every individual was aged using horn-ring counts
(Koubek and Hrabe 1983; Schroder and Von Elsner-Schack
1985) and weighed. Drop nets allowed the capture of entire
groups, whereby mothers could be caught with their kid.
The motherkid relationship was assessed after capture and
marking from observation of suckling and nursing of mothers and kids. The monitoring of the study area was based on
the survey of seven paths on foot. As often as possible, from
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590

May to September and weather permitting, one or two teams


of observers would walk a path, identifying marked animals
and recording their position on a map using Lambert 2 coordinates with a 100 m  100 m grid. These observations
were recorded on the open areas of the study site (50% of
the study site).
Statistical analyses
From 1985 to 2004, 39 kidfemale pairs were individually
marked. These pairs were captured in marking sites 2 and 3
(Fig. 1), which are separated by a wooded valley. Of the 39
pairs, we only considered those of which offspring were
resighted at least once as yearling and (or) older individuals.
We did not consider offspring <1 year of age because the
kid usually stay with their mother up to their first birthday
(Richard-Hansen and Campan 1992; Ruckstuhl 1999).
Therefore, the location of the kid would not give any information about where the kid chose to settle compared with its
mothers range once the motherkid bond loosens (RichardHansen and Campan 1992). We therefore excluded eight
pairs for which offspring were only seen as a kid. Among
the 31 remaining pairs, 13 were motherdaughter pairs and
18 were motherson pairs. We had a total of 412 locations
for female offspring (mean number of location per offspring
was 31.7, with a median of 7) and 137 for male offspring
(mean number of location per offspring was 7.6, with a
median of 4). Most females in this population gave birth for
the first time at age 2 or 3 (Houssin et al. 1994; Loison
1995). We therefore distinguished locations when the daughter was a yearling from locations when the daughter
was 2 years old. The age at dispersal for males is not well
known. According to the inbreeding avoidance hypothesis
(see Introduction), it should occur before actual participation
to reproduction, which is unlikely to be before 3 or 4 years
of age in chamois (Von Hardenberg et al. 2000; A. Loison,
G. Darmon, and J.-M. Jullien, personal observations). Indirect evidence from capturemarkrecapture estimates of
male survival (Loison et al. 1999a) indicates that dispersal
is most likely to occur at 2 and 3 years of age. Therefore,
we also contrasted the distance between yearling sons to
their mother (before assumed dispersal age) to the distance
of 2-year-old sons to their mother. For the sake of simplicity, we used yearling offspring to designate the yearling
age stage, and mature offspring to designate the
stage 2 years old.
We calculated the mean distance between the center of
the mothers location and the center of her offsprings location when the offspring was a yearling and when the offspring was mature. The center of the mothers locations
was obtained by calculating the centroid of all mothers locations and the center of the offsprings range was likewise
calculated as the centroid of offsprings locations, for each
age class. The number of locations for most offspring was
too low (median of 7 and 4 for daughters and sons, respectively; see above) to allow estimating individual home-range
size, and therefore, to estimate for example the range overlap between the mothers range and her offsprings range or
to explore seasonal differences in distances between the
mothers center of locations and the offsprings center of
locations. To account for the differences in the number of
locations per offspring, we performed the statistical analyses

Can. J. Zool. Vol. 86, 2008

(see below) by weighing the estimated distance between the


mothers center of locations and the offsprings center of
locations by the number of locations per offspring (Burnham
et al. 1987). Indeed, the squared distance calculated from a
reference point (here considered to be the mothers centroid)
to the centroid of a sample of locations is distributed as a
2, and the variance of the estimated distance should be
inversely proportional to the number of locations. We
checked this based on offspring for which we had >10 locations (for a given age class) using bootstrap and found that
this was indeed the case. This allowed us to resume the
analyses accounting for the differences in the number of
locations for each offspring (Burnham et al. 1987).
We tested for the effects of sex and stage on the distance
between the mothers center of locations and the offsprings
center of locations using linear mixed effect models. We
used mixed models to account for the fact that the same offspring could be included in the analysis when yearling and
when mature, and because we expect individual variability
in the tendency to move away from their mother. We logtransformed the distances to account for the skewed distribution of distances, and we weighted the motheroffspring
distance by the number of locations we had for each offspring (see above). We therefore fitted 5 models (sex 
stage, sex + stage, sex only, stage only, and constant) with
offspring identity as a random factor. Parameters for fixed
effects (sex and stage) were estimated using restricted
maximum likelihood (REML). We used Akaikes information criterion (AIC) for selecting the best model according to the parsimony principle (Lindsey 1999; Burnham and
Anderson 2002), using maximum likelihood estimation (not
REML) to obtain comparable values for the AIC. All statistical analyses were performed using the R software (R
Development Core Team 2005).
Afterwards, we compared the distance between the
mothers and offsprings centers of locations with the mean
diameter of a female home range. The home range of a
female chamois is estimated to be around 100 ha in this
area (Loison 1995), which corresponds to a circle with a
diameter of 1130 m.

Results
Mean (SD) weighted distance between the mothers and
offsprings centers of locations was 160.1 216.2 m (range
20.91312.7 m) for yearling daughters, 220.7 255.6 m
(range 63.31715.6 m) for yearling sons, 562.7 788.1 m
(range 73.73340.8 m) for mature daughters, and 1519.1
970.9 (range 44.24581.4 m) for mature sons. The selected
model included the interactive effects of both sex and age
(Table 1), which indicates that sex effects differed for yearling and mature offspring. Motheroffspring distances did
not differ per sex for yearlings (F[1,29] = 0.3, p = 0.586), but
did for mature offspring. Indeed, the center of locations of
mature sons were significantly farther away from the center
of locations of their mothers than was the center of locations
of mature daughters (F[1,16] = 4.5, p = 0.049). In addition,
daughters were significantly farther away from their mothers
when mature than when yearling (F[1,16] = 29.96, p < 0.001).
When calculated at the within-individual level, the weighted
mean differences in distance between the centers of loca#

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Loison et al.

591

Table 1. Models and model selection for explaining chamois


(Rupicapra rupicapra) motheroffspring distance.
Model
Sex + stage + sex  stage + (ID)
Sex + stage + (ID)
Sex + (ID)
Stage + (ID)
Constant + (ID)
Sex + stage + sex  stage

df
6
5
4
4
3
5

AIC
697.48
717.22
992.84
711.17
992.17
1571.14

AIC
0
19.74
295.36
13.69
224.69
873.66

Fig. 2. Histogram of the distance between mothers and sons (above)


and mothers and daughters (below) when chamois (Rupicapra rupicapra) offspring were yearling (shaded bars) and mature (solid
bars). For sons, the distance was available for both yearling and
mature sons in 11 pairs, only for yearling in 3 pairs, and only for
mature sons in 4 pairs. For daughters, the distance was available for
both yearling and mature daughters in 7 pairs, only for yearling in 4
pairs, and only for mature sons in 2 pairs. The thick horizontal line
corresponds to the diameter of a mean adult female home range
(1130 m).

Note: Sex and stage (yearling vs. mature offspring) are factors. ID
refers to the offsprings identity and was accounted for as a random
factor. AIC denotes Akaikes information criterion used for model selection. AIC compared models with the selected models (i.e., the
models with the lowest AIC values).

tions of yearling and mature offspring were 36.7 119.5 m


for daughters (calculated on 7 pairs with both yearling and
mature values) and 818.2 1093.2 m for sons (calculated
on 11 pairs with both yearling and mature values). The distance between mothers and sons centers of location when
sons were immature was not correlated with the distance
calculated when sons were mature (R2 = 0.02, t = 0.47, p =
0.650, N = 11) (Fig. 2). On the contrary, the mother
mature daughter distance was significantly correlated to the
mother immature daughter distance (R2 = 0.96, t = 10.61,
p < 0.001, N = 7) (Fig. 2).
Only 1 daughter out of 13 (8%) monitored was actually
observed farther away than the mean diameter of a female
home range, both when she was a yearling and an adult
(motherdaughter pair no. 7; Fig. 2). In the case of sons, 2
out of 14 yearling sons (14%) and 8 of 15 of the mature
sons (53%) were farther away than the mean diameter of a
female home range.

Discussion
Based on the long-term monitoring of motheroffspring
pairs, we found that offspring settlement patterns are sexand age-dependent in chamois. Chamois is a polygynous
mammal (Loison et al. 1999b) for which dispersal is usually
male-biased (Greenwood 1980). The patterns shown here
confirmed that males tend to move farther away from their
mother than daughters do after the immature stage (Levet et
al. 1995; Loison et al. 1999b). Based on our data, inbreeding
avoidance and local mate competition could be at work;
however, resource competition can be ruled out, as females
are highly philopatric, while in this social species, females
should have been the most sensitive to local food competition (see the Introduction).
For most female offspring, home ranges were likely to
overlap with their mothers home range. These spatial patterns may promote the emergence of matrilineal structures,
i.e., groups of related females. However, the spatial scale at
which these structures may occur is not yet clear. Indeed,
previous studies on this population (Loison 1995) and others
(Richard-Hansen 1992; Richard-Hansen and Campan 1992)
failed to show preferential association of pairs of marked
females, i.e., often seen in the same social group, where a
group is defined as individuals <50 m apart (Richard-Hansen
1992). This also held true even for motheroffspring pairs

following the offsprings independence (Richard-Hansen


1992). Hence, studies of marked individual chamois reveal
a looser social structure than has been often described for
this species (e.g., Locati and Lovari 1991), at least in highdensity populations (Richard-Hansen 1992; Loison 1995).
However, the philopatry of daughters shown here (see also
Loison et al. 1999c) implies that matrilineal structures
could be found at a larger spatial scale than that of the social
group (Loison 1995). Such organization (labile groups associated with stable structures at a higher socio-spatial
scale) has also been seen in other social ungulates where
associations and spatial habits have been studied using
marked individuals (e.g., pronghorn (Antilocapra americana Ord, 1815): Byers 1997; soay sheep: Coulson et al.
1999; Coltman et al. 2003; Clutton-Brock and Pemberton
2004; bighorn sheep: Festa-Bianchet 1986, 1991;
LHeureux et al. 1995). Recent longitudinal studies in ungulates suggest that females may improve their reproductive
success as they age, probably partly owing to increased
experience (Weladji et al. 2006). This experience is certainly partly gained by a better knowledge of the home
range and may be acquired already during the early ontogeny (Richard Hansen 1992). The importance of experience
in improving reproductive success, and thereby fitness, certainly contributes to preventing dispersal in females (Conradt et al. 1999).
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592

Although observations of adult males suggest that they


have a larger home range than females in our study site
(Loison 1995), we do not have a reliable estimate of the
size of their home range. Therefore, the comparison of the
distance between the mothers and her offsprings centers of
locations with the mean diameter of a female home range
cannot reliably be interpreted in terms of the overlap
between the home ranges of mother and son. However, our
data clearly supported a higher trend for males leaving their
natal range, especially after 2 years of age, which support
the inbreeding avoidance hypothesis (Pusey 1987) but could
also be explained by the local mate competition hypothesis.
Social behaviors of mothers and sons have not been studied
before in chamois using marked individuals. Males appear to
choose where to settle later than females. The age at which
they settle should be investigated further in relation to the
actual age of male participation to the rut. In addition,
whether males use different ranges during the mating season
compared with the rest of the year should also be explored
to better understand the connection between settlement
patterns and inbreeding avoidance. Surveys should therefore
span longer than the summer period to access seasonal
variation in range use. Differences between sexes in the
timing of settlement have rarely been investigated in the
wild in ungulates, but sex differences in distances and association patterns between mother and offspring could start as
early as the first weeks of life (Schwede et al. 1994).
Sexual segregation occurs at the population level,
whereby adult males are not often seen within groups of
females (Loison 1995; Bonenfant et al. 2007). Family
groups often consist of females, kids, yearlings, and immature males (Gerard and Richard-Hansen 1992; Bonenfant et
al. 2007). In addition to this social segregation, which does
not require sex-biased dispersal, our data suggest that males
in a given area are less likely to be closely related to the
females than females are to other females (for similar patterns in Soay sheep see Coltman et al. 2003). We still lack
data sets for large mammals to better understand why
individuals choose to settle where they do and what triggers
dispersal (Matthysen 2005). To disentangle these processes,
the long-term monitoring of known motheroffspring pairs
is required and should be complemented with studies of
genetic structures (Rousset 2001).

Acknowledgements
This study has been possible because of the long-lasting
efforts of the National Hunting and Wildlife Office
(ONCFS) to catch motherkid pairs, with the help of many
volunteers. We thank them all, as well as the many students
performing long hours of observation of marked individuals
in the field. We are also grateful to J.-M. Gaillard, N.G.
Yoccoz, and M. Hewison who kindly provided useful inputs
and comments.

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