Ecography 30: 51  65, 2007

doi: 10.1111/j.2006.0906-7590.04646.x
Copyright # Ecography 2007, ISSN 0906-7590
Subject Editor: Douglas Kelt. Accepted 16 October 2006

Effects of spatio-temporal variation in food supply on red

squirrel Sciurus vulgaris body size and body mass and its
consequences for some fitness components

Lucas A. Wauters, Marleen Vermeulen, Stefan Van Dongen, Sandro Bertolino,

Ambrogio Molinari, Guido Tosi and Erik Matthysen
L. A. Wauters (l.wauters@uninsubria.it), A. Molinari and G. Tosi, Dept Environment-Health-Safety, Univ. of Insubria Varese, Via
J.H. Dunant 3, IT-21100 Varese, Italy. * M. Vermeulen and E. Matthysen, Dept of Biology, Laboratory of Animal Ecology, Univ.
of Antwerp, Universitsplein 1, BE-2610 Antwerp, Belgium. * S. Van Dongen, Dept of Biology, Group of Evolutionary Biology,
Univ. of Antwerp, Groenenborgerlaan 171, BE-2020 Antwerp, Belgium. * S. Bertolino, DIVAPRA Entomology and Zoology, Univ.
of Turin, Via L. da Vinci 44, IT-10095 Grugliasco (TO), Italy.

Food availability is likely to influence body condition and, in turn, fitness. The intensity of this response may
vary between populations of the same species on a small spatial and temporal scale. We used 5 yr of data from 6
Eurasian red squirrel Sciurus vulgaris populations from the southern Alps to explore differences in body size and
body mass among neighbouring populations, in relation to habitat type and variation in food supply. We also
investigated sexual dimorphism in these traits and whether phenotypic variation affects local survival and female
reproductive success. Mean hind foot length, a measure of body size, did not differ between sexes but differed
between areas. Seasonal variation in body mass was small with no evidence for fattening in autumn. Females
were slightly heavier than males, but this difference was largely explained by mass gain of females during
reproduction. The size of conifer seed crops, the major food supply, varied strongly over years and between
habitats, but this variation corresponded only weakly with autumn body mass. Differences in size and mass
between populations were partially explained by habitat-related differences in body size and variability of seed-
crops, suggesting differential selection for smaller squirrels in spruce-larch forests against selection for larger and
heavier animals in mixed broadleaves and conifer forests and in Scots pine forests with more stable seed
production. The probability of reproduction by females increased with body mass, but varied strongly between
habitats and years, with more females reproducing in years with rich seed-crops. In both sexes, body mass
positively affected probability of settlement and length of residency. Our results suggest that in temporally
variable environments that differ in overall amount of food resources, individual variation in body mass is
related to habitat type, and that having a relatively high body mass, within each population, positively affects
male and female settlement success and local survival, and female reproductive success.

The amount of food animals consume will affect their
fitness in various ways. Food assimilation produces
energy necessary for body growth and/or maintenance
of body condition and for reproductive investment, as
well as essential nutrients such as proteins, carbohy-
drates, minerals, and vitamins (Robbins 1993, Cuthill
and Houston 1997). A diet rich in lipids allows a high
rate of energy intake and, in a large variety of species,
the accumulation of fat reserves that cause a (seasonal)
increase in body mass. In many vertebrates, a relatively
high body mass in relation to body size is an indication
of good condition: such individuals are more likely to
survive critical periods of food shortage or extreme
weather conditions (Blem 1990, Millar and Hickling
1990, Schulte-Hostedde et al. 2001). Moreover, heavier
animals often are more likely to reproduce successfully,
particularly in female mammals, and tend to be better
competitors (social dominance) than conspecifics of
smaller size or lesser body mass (Wauters and Dhondt
1989a, 1995, Robbins 1993, Law 1995, Cuthill and

51
Houston 1997, Lin and Batzli 2001, Diaz and Alonso
2003). Finally, in some species, animals in good
condition are better at escaping predators and are less
vulnerable to parasites or diseases (Ostfeld et al. 1996,
Van der Veen 1999, Wirsing et al. 2002). Inevitably,
there are also costs associated with being heavier: larger
or heavier animals have higher absolute energy require-
ments than smaller conspecifics and therefore are forced
to forage more intensively to meet their energy
demands. These increased foraging costs may augment
the possibility of predation, aggressive encounters, and
contact with parasites (McNamara and Houston 1987).
Hence, body condition will affect an animal’s behaviour
and fitness, and understanding the relationships be-
tween phenotypic characteristics and environmental
factors that influence body condition is of general
interest to ecologists.
Many ecologists have explored variation in body
condition in relation to predation rate, age-related
survival, and reproductive success in small and med-
ium-sized mammals (Wauters and Dhondt 1989a,
1995, Becker 1993, Dobson and Michener 1995,
Humphries and Boutin 1996, Green 2001, Wirsing
et al. 2002). Most of these studies were relatively short-
term and in a single population or habitat, and there are
few papers that explore effects of habitat-type and/or
seasonal and annual variation in food resources on
within and between population patterns of variation in
body condition (Andersson 1994, Moss and Croft
1999, Naeff-Daenzer et al. 2001).
Differences in size or body mass between neighbour-
ing populations may be the result of genetic variation
with individuals responding to differential selection for
body size and/or body mass on a small spatial and even
temporal scale (Hoffmann and Merilä 1999, McAdam
and Boutin 2003a, b, Garant et al. 2005). In North
American red squirrels Tamiasciurus hudsonicus , juve-
nile growth in both body mass and size had significant
amounts of genetic variation, but also experienced large,
heritable maternal effects (McAdam et al. 2002). A
long-term study on great tits Parus major showed that
differences in body mass at a small spatial-scale,
between (sub)populations occupying different habitats
within a large, contiguous forest, was an expression of
divergent evolution (Garant et al. 2005). In fact,
evolutionary theory predicts that local, adaptive, popu-
lation divergence will depend on the balance between
diversifying effects of natural selection and the (oppo-
site) homogenizing effect of dispersal, hence gene-flow
(Endler 1986, Slatkin 1987). Therefore, it is important
to understand how body size and body mass vary
among populations of a species occurring in different
habitats in a limited geographical range, and how these
phenotypic traits change with fluctuating environmen-
tal conditions (in particular fluctuations in food
availability) within populations.
52
Here we use a 5-yr data set from 6 Eurasian red
squirrels Sciurus vulgaris populations from the southern
Alps to explore effects of habitat type, characterised by
variation in type and overall availability of the major
food supply (tree seeds), and effects of season and year,
representing temporal variation in food supplies, on
patterns of variation in two phenotypic traits: body size
and body mass. We also investigate sexual dimorphism
in these traits. Finally, we explore whether phenotypic
variation affects length of residency (local survival) of
males and females, and reproductive success of female
red squirrels.
Materials and methods
We selected 6 study areas within mature, secondary
montane and subalpine mixed conifer forests of the
Italian Alps, with elevations ranging from 1100 to 2100
m a.s.l. (the upper tree-line). These areas are distributed
over 2 geographic regions: COG and RHE are located
in the Cogne and Rhemes Valleys of the Gran Paradiso
National Park, in the western Alps, while Cedrasco
(CED), OGA, Valfurva (VAL), and Bormio (BOR) are
in the Valtellina Valley in the central Alps. Specific
location and distances between study areas are presented
elsewhere (Fig. 1 in Trizio et al. 2005). RHE and VAL
are dominated by Norway spruce Picea abies , OGA by
Scots pine Pinus sylvestris , and BOR by Arolla pine
Pinus cembra . COG is spruce-larch Larix decidua forest
and at CED the forest is mainly composed by silver fir
Abies alba and spruce with small proportions of larch,
Scots pine, and some beech Fagus sylvatica at lower
elevations (Wauters et al. 2004a).
The proportion of spruce, silver fir, larch, Scots pine,
and Arolla pine differed among study areas, but also
varied within study areas at a fine-grained level. There-
fore, to estimate seed availability as our measure of food
abundance, we first determined woodland composition
by establishing a 20 20 m (400 m2) vegetation plot
around each trapping station across the trapping grid
(n 20 30). In each vegetation plot, we counted the
number of trees of each species, and measured the
diameter at breast height (DBH in cm) of 2 trees of
typical size for each species on the plot (forests consisted
of large areas of even-age stands with trees of similar
size), hereafter called ‘‘sample trees’’ (Wauters et al.
2005). Dead trees were recorded separately since their
presence indicated a naturally structured forest. Each
year in August, we counted green cones in the canopy of
all coniferous sample trees from a fixed position using
10 40 binoculars and scaled to estimate the total
number of cones per tree (Wauters et al. 2005).
Counted cones were multiplied by species-specific
average number of seeds per cone (unpubl.) to obtain
an estimate of annual seed production per tree in each
vegetation plot. To combine data of different seed
species, we converted the number of seeds tree 1 for
each species to energy values (kJ tree 1), using our own
values of seed-specific energy-content. We then multi-
plied the number of trees of each species with the
species-specific seed energy tree 1 to calculate seed
density in each vegetation plot (multiplied by 25 and
thus expressed in kJ ha1). The mean value of all
vegetation plots was calculated as the measure of average
annual tree seed abundance in each study area. Seeds
produced in the summer-autumn of year t were
considered available for the squirrels during the follow-
ing ‘‘squirrel year’’ (Wauters and Lens 1995), i.e. from
August of year t to July of year t1.
In each study area, squirrels were live-trapped
bimonthly from April to October 2000 2004, using
20 to 30 ground-placed Tomahawk traps (models 201
and 202, Tomahawk Live Trap, WI, USA). Traps were
placed on a grid, with distances of 100150 m between
traps and average trap density of 0.60.7 traps ha1.
Traps were pre-baited with sunflower seeds and hazel-
nuts 4 to 6 times over a 30-d period, and then baited
and set for 812 d, until no new, unmarked squirrels
were trapped for at least 2 consecutive days. Traps,
partly covered by dark plastic to give shelter from rain
or cold, were checked 23 times a day. Each trapped
squirrel was flushed into a light cotton handling bag
with velcro-type fasteners or with a zipper (Koprowski
2002), or a wire-mesh ‘‘handling cone’’ to minimize
stress during handling, and individually marked using
numbered metal ear-tags (type 1003 S, 10 by 2 mm,
National Band and Tag, Newport, KY, USA). It was
weighed to the nearest 5 g using a Pesola spring-balance
(Pesola, Baar, Switzerland), and the length of the right
hind foot (without nail) was measured (0.5 mm) with a
thin ruler (Wauters and Dhondt 1989a, b, 1995). Sex,
age, and reproductive condition were recorded follow-
ing Wauters and Dhondt (1995). Thus, we scored a
female’s reproductive status as: 1) anoestrus (vulva
small, no longitudinal opening, not lactating), 2)
oestrus (or post-oestrus, vulva partly or strongly swollen
with longitudinal opening, enlarged belly during late
pregnancy), or 3) lactating (nipples large, milk excre-
tion can be stimulated, Wauters and Dhondt 1995,
Wauters and Lens 1995).
Indices of body condition
In mammals, fat reserves are considered the best index
of condition but are extremely difficult to measure in
the field for live animals (Green 2001, Blackwell 2002).
There are 2 common methods to derive an index of
body condition in studies on vertebrates: a ratio of body
mass on a measure of structural body size, using hind-
foot length (Krebs and Singleton 1993, Jakob et al.
1996, Becker et al. 1998, Wirsing et al. 2002), or the
residuals of a regression of body mass (or log-trans-
formed body mass) on size (Dobson and Michener
1995, Wauters and Dhondt 1995, Guinet et al. 1998,
Blackwell 2002, Schulte-Hostedde et al. 2005). In the
first case, animals with a higher ratio are considered to
be in better condition than those with a lower ratio, and
in the second case individuals with a positive residual
value have a better condition than those with a negative
residual value. However, mass/size ratios often are
highly correlated with body mass in small mammals
(Lidicker and Ostfeld 1991, Schulte-Hostedde et al.
2001), potentially biasing interpretations of body size
variation among populations or sexes, or the effects of
body size on fitness components (Wauters and Dhondt
1989b, 1995). Moreover, many recent studies have
criticized the use of residuals as a condition measure
based on statistical assumptions inherent to deriving
residuals from regression equations, or the lack of a
relationship between residuals and true variation in fat
reserves, or because residuals are an inappropriate
measure of condition when comparing several popula-
tions, sexes, and/or age-classes that differ in body mass
(Garcia-Berthou 2001, Green 2001, Blackwell 2002,
Freckleton 2002). Thus for these biological and
statistical reasons, we chose to include foot length,
our measure of structural body size, as a covariate in
multivariate models that tested for spatio-temporal
variation and possible gender differences in body
mass, and in multivariate models that explored effects
of body mass on settlement, residency and reproduction
(Garcia-Berthou 2001).
Statistical analyses
Variation in foot length and body mass were analysed
with linear mixed models (Anon. 1999, Verbeke and
Molenberghs 2000), with area, sex, age, season, year
(and, where relevant reproductive condition) as class
variables. Because this study was designed to monitor
individual body mass over longer periods of time, these
models were adjusted for repeated measures by expli-
citly modeling residual correlation structure. In each
analysis we started from a saturated mean model (i.e.
including all fixed effects and interactions) and com-
pared models with different correlation structures.
Model fit was compared using Schwarz’s Bayesian
Information Criterion (BIC), where smaller values
indicate better fit (Verbeke and Molenberghs 2000).
After selecting correlation structure of the residual
correlation matrix, we continued testing fixed effects
using Type-III SS and performed model selection using
a backward procedure. Degrees of freedom and stan-
dard errors of F- and t-tests were obtained using the
Kenward-Rogers method. Interpretation of final models
53
was based on least square means (Verbeke and
Molenberghs 2000). Distribution of residual values of
each finally-selected model was explored using the
Shapiro-Wilk statistic (Verbeke and Molenberghs
2000). When effects on body mass were explored,
foot length was always included as a covariate to correct
for variation in body mass produced by variation in
skeletal size as measured by foot length. Foot length of
adults does not change over time, and variance from
repeated measures was small (0.313) compared to
variance caused by individual differences in size
(2.125), indicating that measurement error in foot
length was much smaller than individual variation
(87% of total variance). Therefore, we used only one
value for individual foot length based on the mean of all
measurements.
Since juveniles differed from adults and subadults in
both mass and foot length (see Results), they were
analysed separately. Only at BOR, juveniles were
trapped in late spring, hence these were pooled with
summer captures to avoid confounding effects of season
with those of study area. Since most squirrels were
caught only once as juveniles, we tested variation in
juvenile body mass with a generalised linear model,
using each individual only once.
We further explored to what extent variation in food
abundance could explain body-mass variation. In a first
step we calculated mean body mass for each combina-
tion of year, season, and area, using only data of spring
and autumn captures. We predict that average body
mass in spring of year t will be correlated with seed crop
size of the previous year (t1) and that average body
mass in autumn of year t will be correlated with seed
crop size of the same year (t) (Wauters and Lens 1995,
Wauters et al. 2004b). We also tested for a possible
effect of the previous year’s seed crop on autumn body
mass, since conifer seeds produced in mast years may
still be harvested throughout the following year’s
summer and autumn (Wauters et al. 2005). Since
measurements of food abundance are year- and area-
dependent, year and area are included in a linear mixed
model as random variables.
To investigate differences in body mass between
anoestrus, oestrus, and lactating females (Wauters and
Dhondt 1989a), we used only the spring trapping data
(April early May) of adult females since in this season
we trapped anoestrus females, females in early oestrus
whose body mass was not or only slightly affected by
pregnancy (intermediate litters with expected parturi-
tion mid May mid June), and lactating females (spring
litters, observed or expected parturition dates mid
March late April). We used a mixed linear model
(122 observations of 96 adult females) to explore effects
of study area, reproductive status, foot length, and a
foot length by study area interaction on variation in
body mass. In a second step, logistic regression was used
54
to examine the relationships between body mass, foot
length, study area, year, and season (data over all
seasons) as independent variables and the probability
for adult female squirrels to reproduce as dependent
variable (Hosmer and Lemeshow 1989). The binary
dependent variable (Yi) was scored 0 when reproductive
condition of a female was classified as anoestrus, it was
scored 1 when a female was classified as pregnant or
lactating. Models were calculated with PROC GEN-
MOD (Anon. 1999) by stepwise forward selection of
significant independent variables (a 0.05) (Hosmer
and Lemeshow 1989). The probability for immigrant
subadult or adult squirrels to settle successfully (defined
as presence in at least 2 trapping sessions) was also
examined by logistic regression by testing the same
variables and factors as for reproduction. In this case,
the binary dependent variable was scored 0 when a
squirrel was trapped only in a single trapping session
(transient dispersing animals, Wauters and Dhondt
1993), it was scored 1 when a squirrel settled. For each
individual, we used mean body mass over all captures.
Squirrels that immigrated in 2004 were excluded from
the analyses since trapping did not last long enough to
classify them as settlers or transients. In a next step,
length of residency, or local survival (number of months
between first and last capture), was estimated for those
squirrels that were captured at least twice in the study
areas (n 195). If data from animals first captured in
2003 or 2004 and still present in autumn 2004 were
excluded, sample size was reduced to 171 squirrels.
However, general linear models based on the entire and
on the reduced dataset gave similar results: therefore we
present only analysis for the entire dataset. Length
of residency was ln-transformed to meet assumptions of
normality (Shapiro-Wilk’s W 0.94). All tests of
significance are 2-tailed and the significance level was
set at 0.05. Unless otherwise indicated values are
presented in the text as mean9SD.
Results
Food availability
Tree seed production varied between years and study
areas (Fig. 1, 2-way ANOVA on Log-transformed food
abundance: year F5,21 7.07, p 0.0005, area F5,21 
5.35, p 0.0025). The largest seed-crops were pro-
duced at CED and VAL, the smallest at BOR and
OGA. The years 1999 and 2004 had the largest crops,
while 2003 and especially 2000 had poor crops. The
year-effect explained 41% of variation in food abun-
dance and the area-effect 31%. Although the year by
area interaction could not be tested, fluctuations in
seed-crop size differed among study areas; in particular
the seed-crops of pines at OGA and BOR were less
 10000 61
CED Males
9000 OGA Females
8000 BOR (a)
60
Seed-crop size (103 kJ/ha)

VAL
7000 RHE

Foot length (mm)

6000 COG 59
5000
4000 58
3000
2000 57
1000
0 56
1999 2000 2001 2002 2003 2004 CED OGA BOR VAL RHE COG
Year
340
Fig. 1. Tree seed production per year (mean1 SE expressed
in 103 kJ/ha 1) as a function of study area.
330 (b)
variable than spruce and larch crops at the other sites
(Fig. 1).

Body mass (g)

320

Foot length and body mass

310
Between July 2000 and October 2004, 1360 measure-
ments of foot length and 2189 of body mass were taken
for 520 and 544 red squirrels respectively. Mean foot 300
length was 57.592.0 mm (range 46.062.0 mm, 90%
between 54.0 and 60.0 mm) and mean body mass was
290
304937 g (range 90400 g) with 90% of measure- CED OGA BOR VAL RHE COG
ments between 245 and 360 g. Both foot length and
320
body mass differed between the 3 age-classes (Mixed
ANOVA models with first order autocorrelation struc-
ture, foot length, F2,1210 190.8, pB0.0001, body (c)
mass F2,1739 746.2, pB0.0001). 315
Body mass (g)

Juveniles
310
Juveniles (n 110 measures, 54.092.5 mm, 46
61 mm) were smaller than subadults (n 182, 56.99
1.6 mm, 53 61 mm) and adults (n 1068, 58.091.5
305
mm, 5362 mm). Juveniles also weighed less (n 136,
209940 g, 90 265 g) than subadults (n 278919 g,
230 340 g) and adults (n 1789, 315925 g, 240
400 g). As expected, juvenile body mass increased with 300
foot length (slope 3.8891.39). There was no difference Spring Summer Autumn
in juvenile body mass between the sexes or between years Fig. 2. Plots of body size measurements of male and female
(Table 1) but juveniles trapped in autumn were heavier red squirrels (mean1 SE) as a function of study area or
than those trapped in summer (Table 1, summer n 51, season. (a) mean foot length per study area, (b) mean body
197942 g, autumn n 57, 221933 g). Moreover, mass per study area, and (c) mean body mass of adults and
juveniles at RHE and BOR were heavier than at CED, subadults per season (years pooled).

55
Table 1. Average (9SD) foot length and body mass of juvenile red squirrels (n108) per study area. Generalised linear model
exploring effects of foot length (as co-variate), study area, sex, year and season on variation in body mass.
Study area (n) CED (5) OGA (11)
Foot length (mm) 55.392.5 54.492.7
Body mass (g) 164931 198949
Generalised linear model
Significant parameters
Foot length F1,100 7.80
Season F1,100 9.96
Area F5,100 2.72
and those from RHE were also heavier than juveniles
from COG (Table 1, and Tukey-test for differences
between means, all p B0.05). Juveniles were of inter-
mediate mass at OGA and VAL.
Foot length of subadult and adult squirrels
Variation in foot length among subadult and adult
squirrels was analysed with a stepwise backward linear
mixed model (compound symmetry correlation struc-
ture) exploring the effects of study area, sex, and year
and all interactions. The model was corrected for
repeated measures with foot length measured once in
each trapping period (113 observations per individual,
1045 observations from 433 different squirrels). None
of the interaction terms were significant (all p0.10),
and foot length did not differ between male and female
squirrels (males, n 627, 57.891.5 mm, females, n 
418, 57.791.5 mm, F1,426 0.34, p0.56), or
among years (F4,612 0.70, p 0.59). However, size
of squirrels differed between study areas (Fig. 2a,
F5,427 12.1, p B0.0001). Squirrels were larger at
CED and smaller at RHE than at all other sites
(differences of least square means, DLSM, all p B
0.01). There were no differences between the other 4
study areas (Fig. 2a, DLSM, all p0.05).
Variation in body mass
In a preliminary test, we compared within-season
variation in body mass at the individual level of
subadults and adults (variation among repeated mea-
sures of an individual’s mass within one trapping
session) with variation in body mass between seasons.
To achieve this we constructed a linear mixed model
with repeated measures at the individual level and each
season (spring, summer, or autumn) treated as an
indicator variable and added to the model as random
factor (nested within individual), next to a random
factor individual which models overall between-indivi-
dual variation. Within-season variation in body mass
was largest in spring (72.10), intermediate in summer
56
BOR (20) VAL (12) RHE (34) COG (26)
56.192.7 54.391.0 54.391.0 54.391.0
219929 199939 221938 205939
Non-significant parameters
p0.006 Sex F1,95 0.69 p 0.41
p0.002 Year F4,96 2.09 p 0.09
p0.024
(63.57), and smallest in autumn (49.04). However,
within-season variation at the individual level  reflect-
ing day to day variability in body mass  was an order
of magnitude larger than between-season variation at
the individual level (variance component 174.4).
Average individual body mass across seasons and
trapping sessions varied even more strongly (variance
component 348.71).
We examined effects of age, sex, study area, season,
and year as factors, and the season by sex, season by
year, season by area, and area by year interactions on
variation in body mass of subadult and adult squirrels,
using foot length as a covariate (Table 2). Interactions
were chosen because of predicted variation in seasonal
patterns of body mass changes between the sexes
(season sex, seasonal changes in body mass of females
related to reproductive condition of pregnancy or
lactation, Wauters and Dhondt 1989a, b, 1995),
between years (season year, spring, and summer
mass was expected to be lower after poor than good
seed-crops), and between areas (season area, autumn
mass-gain might be habitat-dependent, Wauters and
Dhondt 1989b, Lurz and Lloyd 2000). Finally, an area
by year interaction examined if mass changes across
years differed between the study areas in relation to
variation in food abundance between areas (Fig. 1).
Body mass increased 3.91 (90.70) g with each mm
increase of foot length (Table 2a). Corrected for
variation in foot length, adult squirrels were heavier
than subadults. Variation between years, seasons, and
areas was complex, including interactions among these
factors as well as interactions with sex (Table 2a, Fig.
2b). Overall, females were heavier than males (females,
n 425, 316931 g; males, n 635, 305925 g), but
sex-differences changed across seasons (Table 2a). Males
were slightly heavier in spring and autumn (DLSM
spring-summer p0.003, spring-autumn p 0.25,
summer-autumn p 0.064), while females were heavier
in spring and summer (Fig. 2c, DLSM spring-summer
p0.60, spring-autumn p 0.03, summer-autumn
p0.01). Therefore, effects of study area, age, year,
and season on body mass were subsequently examined
for males and females separately.
Table 2. Mixed linear models investigating effects of study area, age, sex, year and season, interactions (see Results), and foot length
as co-variate on variation in body mass of subadult and adult red squirrels. (a) Both sexes, using 2037 observations of 434
individuals, (b) males, using 1253 observations of 253 individuals, and (c) females, using 784 observations of 181 individuals.
a
Excluded from the model.

Fixed effects Statistics Interactions Statistics

(a) Sexes pooled

Foot length F1,450 45.9, pB0.0001
Study area F5,418 10.58, pB0.0001 Areayear F16,1923 4.38,pB0.0001
Age F1,1817 284.1, p B0.0001 Seasonarea F10,1854 3.95, pB0.0001
Sex F1,381 40.8, pB0.0001 Seasonsex F2,1843 10.95, p B0.0001
Year F4,1922 9.49, pB0.0001 Seasonyear F8,1780 2.41, p0.014
Season F2,1858 1.45, p0.23
(b) Males
Foot length F1,266 28.9, pB0.0001
Study area F5,230 10.26, pB0.0001 Areayear F16,1130 3.39, pB0.0001
Age F1,1124 111.5, p B0.0001 Seasonarea F10,1117 4.70, pB0.0001
Year F4,1077 7.54, pB0.0001 Seasonyear F8,1075 5.75, pB0.0001
Season F2,1123 11.7, pB0.0001
(c) Females
Foot length F1,176 17.4, pB0.0001 Areayear F16,721 2.40, p0.0016
Study area F5,171 1.43, p0.22 Seasonareaa F10,696 1.74, p0.069
Age F1,657 171.9, pB0.0001 Seasonyear F7,671 4.40, pB0.0001
Year F4,701 9.81, pB0.0001
Season F2,668 7.05, p0.0009

Males whereas at COG, mean autumn body mass was highest

On average, body mass of male squirrels increased by
3.81 (90.71) g with each mm increase of foot length
(Fig. 3). Variation among study areas changed accord-
ing to season and year (Table 2b, Fig. 4a, b). In the
following paragraphs we summarize the patterns based
on the significance of pairwise comparisons (details not
shown).
In spring, body mass showed hardly any differences
between VAL, RHE, and COG, and in these popula-
tions annual variation was absent or weak (all compar-
isons p0.05, except COG-RHE 2002, p 0.044;
RHE 2003 2004 p0.034; Fig. 4a). In the other 3
study areas, annual fluctuations were more pronounced
in 2004 and 2000, intermediate in 2003 and 2001, and
lowest in 2002 (Fig. 4b). At CED, males weighed less in
2001 than in all other years (DLSM, pB0.001), and
from 2002 to 2004 they weighed more than males in
the other populations (DLSM, pB0.001, Fig. 4b). At
OGA, males weighed less in autumn 2000 and 2002
420
370
f
Body mass (g)

(Fig. 4a). At BOR, males were lighter in 2002 and 2004

than in 2003, consequently, male squirrels at BOR 320 m
weighed less in spring 2002 and spring 2004 than
conspecifics in all other populations (Fig. 4a). Male
squirrels tended to be heavier at CED than at other sites
270
in most years, but there was no significant difference
between mean body mass of animals from CED and Males
OGA in 2002 and 2003. Mass fluctuations over years Females
differed between these two study areas. 220
In autumn, male body mass did not differ between 52 55 58 61 64
BOR, VAL, RHE, and COG, except in 2002 when at Foot length (mm)
COG males weighed less than at VAL and RHE
Fig. 3. Relationship between body mass and foot length for
(DLSM, pB0.001, Fig. 4b). At VAL, males reached male and female red squirrels (subadults and adults) from six
higher autumn mass in 2002 than in 2003 and 2004 study areas in the Italian Alps. Regression lines: (m) male
(DLSM, pB0.05, Fig. 4b). There was no significant body mass (slope t266 5.38, pB0.0001), (f) female body
between-year variation in autumn mass at RHE, mass (slope t176 4.17, p B0.0001).

57
 AREA
BOR CED COG OGA RHE VAL
350

Body mass (g)

300
(a)
250

200
350

Body mass (g)

300
(b)
Season * sex

250

200
350

Body mass (g)

300
(c)
250

200
350

Body mass (g)

300
(d)
250

200
00 01 02 03 04 00 01 02 03 04 00 01 02 03 04 00 01 02 03 04 00 01 02 03 04 00 01 02 03 04
20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20 20
Year Year Year Year Year Year

Fig. 4. Body mass as a function of study area, year, season and sex: (a) spring mass of males, (b) autumn mass of males, (c) spring
mass of females, (d) autumn mass of females.

than in autumn 2003 and 2004 (DLSM, all p B0.05,
Fig. 4b).
Females
Body mass of female red squirrels increased on average
by 4.45 (91.07) g with each mm increase of foot
length (Fig. 3). For females, body mass, corrected for
variation in foot length, did not differ on average
among study areas, but there was a significant study
area by year interaction, suggesting that in some years
females were heavier in some sites than in others (Table
2c, Fig. 4c, d). Overall, females weighed less in 2000
than in other years. The season by area interaction was
not significant, indicating that trends of seasonal mass
variation were similar for all study areas (Table 2c).
There were no differences between years in mean
spring body mass of females in 3 populations (CED,
BOR, and COG), while in the other populations trends
differed. In OGA females weighed less in 2001 than in
2003 and were heavier in 2002 and 2004. Females of
the VAL population also were heavier in 2004 than in
2003, but those from RHE weighed less in 2002 and
2001 and mean body mass was highest in 2003 (Fig.
4c). Only in 2002, females at RHE weighed less than at
the nearby COG study area (DLSM, p0.01, Fig. 4c).
In 2002, the year with the highest average spring body
mass, females from OGA population were heavier than
those from BOR and RHE populations (Fig. 4c).
At CED, mean autumn body mass of females was
lowest in 2002 and highest in 2004 when they also were
heavier than females from the BOR population, that
weighed less than all other populations (Fig. 4d). At
OGA mass variation between years was more pro-
nounced and females were heavier from 2002 to 2004
than in 2000 and 2001 (DLSM, all pB0.05, Fig. 4d).
Females from OGA were also heavier than those from
COG in 2000, than females from COG and RHE in
2001 and 2002 (DLSM, all p B0.05, Fig. 4d). At VAL,
females weighed more in autumn 2004 than in other
years. At RHE and COG patterns were similar: female
red squirrels reached the highest autumn body mass in
2004, followed by 2003 and 2002, they weighed less in
2000 and 2001 (DLSM, all pB0.05, Fig. 4d).
Body mass patterns and food availability
We examined whether annual and site-related variation
in body mass could be explained by variation in the size
of tree-seed crops for each sex separately. Since we
predicted that body mass in spring was affected by the
seed crop of the previous year, but autumn body mass
by the seed crop of the same year, body mass-food
relationships were analysed per season. There was no
increase of spring body mass with larger seed-crops the
previous autumn in either sex (Fig. 5). However,
autumn body mass of both males (p 0.058) and
females (p 0.028) was positively correlated with the
size of the autumn seed-crop, although only 15%

58
 350 1.0
f (a)
0.9

Probability produce offspring

CED 2004
0.8 OGA 2004
330
m 0.7 BOR 2004
VAL 2004
0.6
Mean body mass (g)

RHE 2004
310 0.5 COG 2004
CED 2000
0.4
OGA 2000
0.3 RHE 2000
290 COG 2000
0.2
Males, spring
0.1
Males, autumn
270 0.0
Females, spring
Females, autumn 1.0
(b)
0.9

Probability produce offspring

250
0 3000 6000 9000 0.8
Food abundance (103 kJ/ha) 0.7 CED 2001
RHE 2001
Fig. 5. Relationship between mean body mass in spring (year 0.6 CED 2002
t) and mean seed-crop size of the previous year (year t 1) and 0.5 VAL 2002
mean body mass in autumn (year t) and mean seed-crop size RHE 2002
0.4
of the same year (year t) over all study areas for male and CED 2003
female red squirrels. Regression lines are provided for 0.3 VAL 2003
RHE 2003
relationships of food with autumn body mass of males (m) 0.2
and with autumn body mass of females (f). 0.1
0.0
260 280 300 320 340 360
(males) to 19% (females) of variation in mean autumn Body mass(g)
body mass was explained by variation in food abun-
dance (Fig. 5). Fig. 6. Logistic curves representing the increased probability
of producing offspring as a function of body mass for adult
female red squirrels. Different curves show changes in
probability of offspring production as functions of different
Reproduction study area year combinations.
Body mass among adult females increased with foot
length (F1,118 8.67, p0.0039, slope 4.8891.66) heavier females of 340 g these probabilities increased to,
and differed with reproductive status (F2,118 8.91, respectively, 0.84, 0.91, 0.95 and 0.97 (Fig. 6a). But,
p0.0002), but did not differ between study areas. within each study area, females with a certain body mass
Anoestrus females (n 27, mean9SD 308916 g) had a much lower probability of producing offspring in
weighed less than oestrus (n 70, 324929 g) or 2000 than in 2004 (Fig. 6a). Another example showed
lactating females (n 25, 337923 g) (DLSM: anoes- that probabilities of producing a litter increased from
trus-oestrus t118 3.19, p0.0018, anoestrus-lactating 2001 to 2002 and further in 2003, but that they were
always lower at CED than at RHE (Fig. 6b). Adding
t118 4.10, p B0.0001). Lactating females tended to
season (Wald x2 1.74, DF 2, p0.42), or foot
be heavier than oestrus females, but the difference was
length (Wald x2 0.51, DF 1, p 0.48) did not
not significant (DLSM t118 1.74, p0.085).
improve the fit of the model.
Examining the probability of reproduction over the
entire study period with logistic regression, heavier
female squirrels were more likely to reproduce than Settling success and length of residency
females of lower body mass (Wald x2 31.3, DF 1,
pB0.0001), and probability of producing offspring Of 341 dispersing subadult and adult red squirrels, 167
differed between areas and years (area-effect Wald x2  (49%) settled and 174 (51%) were transients. Heavier
17.0, DF 5, p0.0045, year-effect Wald x2 10.9, squirrels were more likely to settle within the study
DF 4, p0.028, Fig. 6a, b). For example, in 2004 a areas than animals of lower body mass (Wald x2 14.4,
female that weighed 300 g had a probability to produce DF 1, p B0.0001), and probability of settling was
offspring of only 0.53 at CED against 0.68 at BOR, but higher for spring and summer immigrants than for
females of the same mass were more likely to have a animals that immigrated in autumn (season-effect Wald
litter at COG (0.82) and RHE (0.89, Fig. 6a). For x2 12.7, DF 2, p0.0018, Fig. 7a). Adding a

59
 0.8 Table 3. General linear model investigating effects of body
(a) mass, study area, age, sex, and their interactions with body
mass, and foot length as co-variate, on variation in length of
0.7 residency (ln-transformed) of red squirrels.

Parameter Statistics
0.6
Probability to settle

Body mass by study area F5,176 0.68, p 0.64

0.5 Body mass by sex F1,181 1.67, p 0.20
Body mass by age F2,182 1.53, p 0.22
Foot length F1,184 0.32, p 0.57
0.4 Sex F1,185 0.50, p 0.48
Age F2,186 1.64, p 0.20
Study area F5,188 3.30, p 0.007
0.3 Body mass F1,188 12.95, p 0.0004
Spring
R2 selected model0.15 F6,188 5.65, p B0.0001
Summer
0.2
Autumn

0.1 among individual red squirrels (Table 3). Local survival

250 270 290 310 330 350 was higher at OGA than at RHE, COG, and BOR
Body mass(g) (Tukey test). There was no relationship between foot
60 length and residency (Table 3). Effects of sex, age, or
(b) their interactions with body mass were not significant
(Table 3).
50
Residency time (months)

40
30
20
10
0 availability, but variation in seed crop-size corre-
250 300 350
Average individual body mass(g)
Fig. 7. (a) Logistic curves representing the increased prob-
ability of settling as a function of body mass for subadult and
adult red squirrels. Different curves show changes in prob-
ability of settling in relation with immigration period
(season). (b) Relationship between mean body mass of
individual red squirrels and length of residency: different
symbols present the different study areas.
body mass by season interaction (Wald x2 4.52,
DF 2, p 0.10), or foot length (Wald x2 1.24,
DF 1, p0.27) did not improve the fit of the model.
The other factors tested, sex, age, area and year, or their
interactions with body mass, were not related with
probability of settling (all p 0.10). Average residency
was 16911 months (skewness 1.28, kurtosis 2.29,
SE 0.82), with a minimum of 3 and a maximum of
51 months. Length of residency increased with body
mass (Fig. 7b), and body mass together with study area
explained 15% of the variation in length of residency
Discussion
We found differences in body size and body mass of
CED Eurasian red squirrels among nearby populations and
OGA average body mass in the different populations
BOR fluctuated between years. Significant area by year
VAL interactions showed that patterns of changes in body
RHE mass were complex. Differences in size and mass
between populations were partially explained by
COG
habitat-related differences in body size and food
400 sponded only weakly with variation in autumn body
mass. There was no sexual dimorphism in body size,
but patterns of seasonal variation in body mass
differed between the sexes. In spring and summer
females tended to be slightly heavier than males, due
to pregnancy and/or a mass gain during early lacta-
tion. There was no evidence of autumn fattening
(Wauters and Dhondt 1989b, Lurz and Lloyd 2000,
Wauters et al. 2001a) in these alpine populations.
Finally, we found several positive effects of having a
relatively high body mass on fitness components:
heavier females were more likely to reproduce; and,
within each population, having a relatively high body
mass positively affected male and female settlement
success and length of residency. Hence, our results
suggest that in temporally variable environments that
differ in overall amount of food resources, individual
variation in body mass is related to habitat type, and
that having a relatively high body mass is advanta-
geous, increasing both reproductive success and local
survival.

60
Sexual size dimorphism
Patterns of sexual size dimorphism differ between
species of sciurids. In some species of chipmunks (e.g.
yellow-pine chipmunk Tamias amoenus ) females are ca
4% larger (body size) and 1020% heavier than males
(Levenson 1990, Schulte-Hostedde et al. 2001), while
adult males of territorial North American red squirrels
Tamiasciurus hudsonicus are on average ca 10 30 g
heavier than adult females (Boutin and Larsen 1993). In
this species, post-weaning growth and survival were not
correlated and sexual dimorphism was less pronounced
in years of abundant food (Boutin and Larsen 1993).
Although bigger mothers might provide more and/or
better maternal care (Wauters et al. 1993, Dobson and
Michener 1995, Wauters and Dhondt 1995), it is still
uncertain how natural selection has favoured females
growing larger than males in some species, since they all
have promiscuous mating-systems where larger or
heavier males tend to be dominant over lighter (or
smaller) males during mating chases, and, consequently,
obtain a higher mating success (Wauters et al. 1990,
Koprowski 1993a, b, 1998, but see Farentinos 1980).
In contrast, lack of sexual size or mass dimorphism
seems to be the rule for most holarctic tree squirrels of
the genus Sciurus (Wiltafsky 1973, Nash and Seaman
1977, Koprowski 1994a, b). In Eurasian red squirrels
there are no differences in body mass between the sexes
throughout most of its range (Wauters and Dhondt
1989a, Lurz 1995, Münch 2000). In Belgium, males
and females did not differ in body size measurements
and had similar body mass in late autumn and winter,
when females did not invest extra energy in lactation
(Wauters and Dhondt 1989a, b). During months when
many females were pregnant or in early lactation they
weighed more than males (Wauters and Dhondt
1989b). In this study on alpine populations, using
average body mass over all years and seasons, females
were 3.6% heavier than males, a small but statistically
significant difference. In our data set, however, some
females were in reproductive condition (pregnant or
lactating) in all seasons, which might cause an upward
mass bias in adult females (Wauters and Dhondt
1989b, 1995, Humphries and Boutin 1996, 2000).
The significant sex by season interaction, indicating
different seasonal patterns of body mass changes for
males and females, and small differences between male
and female body mass in autumn (males, 307926 g,
females, 312929 g), when only a small proportion of
females is still lactating, agree with this hypothesis.
Finally, females averaged 10 g heavier than males in
only one population, VAL. This population has only
been monitored for ca 2 yr and further data will be
gathered to examine possible sexual dimorphism in
body mass.
Habitat type and annual and seasonal variation in
body mass
The size of conifer seed crops varied strongly over years
in some study areas (e.g. CED and RHE) but remained
rather constant in others (e.g. OGA). This is the
consequence of patterns of seed production of the
dominant tree species. Norway spruce and silver fir
show masting (high and synchronized, episodic seed
production, Smith 1970, Silvertown 1980), thus in
spruce and spruce-fir forests, years of rich seed-crops
alternate with poor to medium seed production, with
seed-crop failure occurring about every 5 yr (Mencuc-
cini et al. 1995, Wauters et al. 2005). Scots pine, in
contrast, produces rather stable seed-crops in most years
(Wauters and Lens 1995, Castro et al. 1999) and total
seed-energy production varied little throughout the
study period. Our results showed that this variation in
seed-production affected between-population differ-
ences and fluctuations in average body mass of red
squirrels over years. Autumn condition was weakly but
positively correlated with autumn seed-crop size. This is
probably directly related with higher feeding rates
causing a more positive energy-balance in autumns
with high seed abundance and lower foraging costs.
Also in squirrel populations in Belgium, good tree seed-
crops positively affected autumn-winter body mass of
red squirrels, thereby increasing immigration success of
subadults and winter survival of subadults and adults
(Wauters and Dhondt 1989a, b, 1993, Wauters and
Lens 1995, Wauters et al. 2004b). Why was the food 
body mass relationship in different alpine populations
rather weak? We suggest this is related to between-site
differences in body size and in temporal variation in the
availability of the major seed species. At OGA, red
squirrels were heavier than expected on the basis of
absolute seed-energy production, but the dominant tree
species, Scots pine, had relatively stable seed-crops over
years. This might help squirrels to use relatively small
home ranges linked with low costs of moving (Wauters
and Dhondt 1992), and have activity patterns that
allow them to maintain higher body mass than in more
variable spruce-larch forests. At CED squirrels were
heavier than in other populations in most years, but
they were also larger in structural size. Finally, in
spruce-larch forests, body mass might be only poorly
related to available food supplies because squirrels
showed a time-lag in responding to changes in seed
abundance, still using large home ranges, and therefore
having high energy-costs of locomotion, in the autumn
with a medium to good seed-crop when it followed a
seed-crop failure (Wauters et al. 2005)
Contrary to expectation, there were no relationships
between spring condition of squirrels and tree-seed
crops produced the previous autumn. There might be
several explanations for this. In spring, seed-crops of the
61
previous year become gradually depleted, because cones
have dispersed most of their seeds (Norway spruce,
Scots pine, larch), or because all cones have been
harvested from the trees and their seeds cached (Arolla
pine). Therefore, a squirrel’s condition might depend of
the amount of stored (cached) food within its home
range, and thus within-home-range food availability,
which was not measured here, might affect body mass at
a fine-grained level (Wauters et al. 2001b). An
alternative, but not mutually exclusive hypothesis is
that secondary food resources, such as fungi (Moller
1983, Wauters and Dhondt 1987, Bertolino et al.
2004), buds, shoots, and male flowers of conifers
constitute an important part of the diet and affect
body-mass variation. Finally, each sex might be
optimizing its daily food-intake to ‘‘face’’ the current
or future costs of reproduction. Breeding females tend
to increase daily energy-intake and thus become heavier
during pregnancy and early lactation (see below),
whereas males might try to maximize body mass to
improve their competitive ability in intrasexual en-
counters during mating chases (Wauters et al. 1990,
Koprowski 1998).
Seasonal body mass variation in alpine populations
(on average B3% of summer mass) was much smaller
than in populations in west and central Europe
(Wauters and Dhondt 1989a, b, Münch 2000), and
daily changes (difference between subsequent measure-
ments within a single trapping session) were of the same
magnitude (see our preliminary analyses). In contrast
with other populations, there was no evidence for alpine
squirrels fattening in autumn (September October). A
similar lack in autumn mass increase was also found for
native red and introduced grey squirrels in conifer
plantations (with several exotic conifer species) in
northern England (Lurz and Lloyd 2000). These
authors suggested that fat accumulation was less
pronounced or even absent in conifer habitats where
autumn and winter food supplies are more predictable,
and manoeuvrability to feed on cones in the canopy is
important.
Size differences between populations
The average body mass of adult red squirrels from
alpine populations was 315925 g (range 240 400 g).
This was less than mean adult body mass from
populations living in high-quality woodlands with
stable food supplies, and high squirrel densities in
Belgium (mean9SE 33097.1 g, t4.14, p B
0.0001, Wauters and Dhondt 1989a, b). Variance in
body mass was similar in both countries (from 270 to
380 g in Belgium). Adult squirrels from alpine forests
were also smaller than conspecifics in Belgium (average
hind foot length: Italy 58.091.5 mm, Belgium 59.99
62
0.25 mm, t2.13, pB0.0001). Subadults were smal-
ler and weighed less than adults in all populations and
years, as found in other habitats throughout the
squirrel’s range (Wauters and Dhondt 1989a, Lurz
1995, Münch 2000).
Among populations from the southern Alps, mean
hind foot length did not differ between male (57.89
1.5 mm) and female squirrels (57.791.5 mm), but
squirrels were larger at CED and smaller at RHE than
at the other 4 sites. This could be the result of
differential selection in populations at CED and RHE
(Garant et al. 2005). CED is part of large contiguous
forests of the Orobic Alps in Valtellina that change
from deciduous woods dominated by chestnuts Casta-
nea sativa at submontane elevations, over mixed
montane forests of beech Fagus sylvatica, silver fir and
Norway spruce (in the study area), to subalpine spruce
 larch forests at elevations above 1800 m a.s.l. Hence,
many animals in this population have access to large
nuts (sweet chestnuts, hazelnuts, and beechnuts) that
allow higher rates of energy intake than foraging on
small-seeded conifers (Wauters et al. 2001a, Steele et al.
2005). Larger squirrels might be advantaged over
smaller ones in handling these large nuts (Weigl et al.
1998). RHE, in contrast, is part of an extensive
subalpine conifer forest dominated by Norway spruce
with patches of larch that increase at higher elevations
(Wauters et al. 2004a, 2005). Here, squirrels only have
access to small conifer seeds that are difficult to extract
from closed cones. Therefore, squirrels are likely to
need more time to fulfil their daily energy demands and
natural selection might favour smaller individuals with
lower absolute energy requirements. Although this
suggestion is speculative, recent studies on birds and
closely related tree squirrels have shown that there is
strong selection on traits of body size and that non-
random dispersal might reinforce population differen-
tiation and adaptation at small spatial-scale (McAdam
and Boutin 2003a, b, Haughland and Larsen 2004,
Garant et al. 2005).
Body mass and reproduction
Body mass among adult females differed with repro-
ductive status, independent of foot length. Anoestrus
females weighed ca 5% less than females in oestrus and
ca 9% less than lactating females. We found that
heavier female squirrels were more likely to reproduce
than females of lower body mass, but probability of
producing offspring also differed between study areas
and years. These results agree with earlier studies on
Eurasian red squirrels in Belgium (Wauters and
Dhondt 1989a, 1995) and eastern chipmunks Tamias
striatus . However, as Humphries and Boutin (1999)
pointed out, the differences found in these cases may be
due to reproductive females increasing their daily rate of
food-energy intake gaining more mass to meet the
growing energy-demands of lactation (Cuthill and
Houston 1997). In Tamiasciurus hudsonicus , mass
gain during early lactation (indicating changes in
body fat levels) was positively related to litter size and
females that gained most mass had the highest levels of
juvenile survival to emergence (Humphries and Boutin
1996). Hence, in female North American red squirrels
early lactation mass gain is an important component of
parental investment under favourable conditions
(Humphries and Boutin 1996). Because of highly
asynchronous breeding in alpine red squirrel popula-
tions, our spring data were taken during the prepartum
period for part of the animals and during early
postpartum for others. Since lactating females tended
to be heavier than oestrus females and both were heavier
than those that did not reproduce, our data suggest that
both mechanisms occur. On the one hand, reproduc-
tion is mass-dependent, with a minimum threshold of
body condition (mass corrected for size) that must be
reached before a female enters oestrus (Wauters and
Dhondt 1989b, Lurz 1995, Wauters and Lens 1995).
On the other hand, successfully reproducing females
increase body mass, independent of size, during early
lactation, suggesting that they supplement their energy
reserves in preparation for pronounced increases in
energy requirements during late lactation (Gittleman
and Thompson 1988, Wauters and Dhondt 1989b,
Humphries and Boutin 1999).
Interestingly, the probability of reproducing among
alpine female red squirrels of a given body mass varied
strongly between habitats and, to a lesser extent,
between years. The between-year differences were
related to variation in tree seed production, and
probability of reproducing increased when seed abun-
dance was high. However, in a given year (e.g. 2004)
females of 280290 g at RHE and COG had a similar
probability (975%) to reproduce than conspecifics
weighing 310315 g at BOR and VAL, and females of
320 325 g at OGA and CED. Since in 2004 good
seed-crops were produced at all sites, these between-
habitat differences suggest variation in life-history
strategies, linked with variation in body size/mass
among red squirrel populations occurring in different
forest types at a small spatial scale. In North American
red squirrels, probability of (first) reproduction was
correlated with cone crop size and further constrained
by foraging and cone-handling efficiency that might
differ between primiparous and multiparous females,
but also between forest types (Becker et al. 1998).
Variation in feeding efficiency (rate of energy-intake per
unit time) while processing cones from different tree
species are likely to exist also in Eurasian red squirrels,
where cone size, ratio of protective cone mass on total
seed mass (Molinari et al. 2006) and, probably,
concentrations of secondary compounds (resins, tan-
nins) differ between conifer tree species.
Finally, body mass in alpine red squirrel populations
also positively affected probability of settlement and
length of residency, a measure of local survival. This was
also the case in high-density populations in mixed
conifer and broadleaf woods in Belgium, where body
mass was positively related with immigration success and
with longevity (Wauters and Dhondt 1989a, 1993).
We conclude that in temporally variable environ-
ments that differ in overall quantity of food resources,
individual variation in body mass of Eurasian red
squirrels is high and body mass, in particular its
condition component, positively affects male and
female settlement success and local survival, and female
reproductive success. Overall, this confirms results from
other populations that live at higher densities in habitats
with larger and more stable food supplies (Wauters and
Dhondt 1989b, 1993, 1995). However, significant
differences existed in both size (hind foot length) and
mass between alpine populations, that could be
explained, at least partly, by habitat-related differences
in total size and degree of (annual) variation of seed-
crops of the dominant conifer species. This suggests
differential selection for smaller red squirrels in spruce-
larch forests (RHE, COG, VAL), against larger and
heavier animals in mixed conifers with some nut-
bearing broadleaves (CED) or in Scots pine dominated
forests (OGA). In North American red squirrels,
direction and strength of selection on growth rates
varied with age, but also with cohort (young born in
rich food-years against young born in poor food-years,
McAdam and Boutin 2003a, b). In populations of
Eurasian red squirrels in high-quality habitats in
Belgium, there was no cohort effect on any of the
components affecting lifetime reproductive success of
females (Wauters and Dhondt 1995). However, in
alpine squirrel populations occurring in habitats with a
high degree of temporal variation in resource abun-
dance (availability), cohort effects may exist. The data
presented in this paper are still on too short a time-scale
to explore this hypothesis in more depth.
Acknowledgements  We wish to thank G. Airoldi, P. Lurz, and
M. Zaninetti for assistance with fieldwork, and D. Preatoni for
preparing figures. Comments by Petit D. Kelt helped to
improve the ms. The study was part of the ASPER (Alpine
Squirrel Population Ecology Research) project, carried out by
Univ. of Insubria, Varese and Istituto Oikos NGO, Milan.
Financial support was provided by the Wildlife Service of the
Province of Sondrio, the Parco Regionale Orobie Valtellinese,
and grants awarded to L.A.W. and S.B. from the Gran Paradiso
National Park, Italy (DGE 25 2000), and the Committee for
Research and Exploration of the National Geographic Society,
Washington DC, USA (grant no. 6997-01). Additional
financial support was obtained from MIUR (Ministero
dell’Istruzione, dell’Università della Ricerca, project COFIN
63
2003, number 2003053710-006) to Insubria Univ., and
logistic support was provided by the Stelvio National Park.
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