465
Site fidelity and effects of body mass on. home-range size of EgypUan mongooses
F. PALOMARES
EsraciOn Sioldgica Doflana, Consejo Superior de lnvestigaciones Gienr(flcas, Aportado 1056, 41080 SeW/ia, Spain
PALOMARES, F. 1994. Site fidelity
and effects of body mass on home-range size of Egyptian mongooses.
Home-range size has been found to be related to body mass of some animals both across species and within species when
the spatial strategies of the sexes differ. I studied home-range size in a polygynous carnivore, the Egyptian mongoose
(Herpesres ichneumon), and compared observed home-range size with predictions based on body mass. First, I tested whether
mongooses actually exhibited site fidelity (for daily and multiday periods). Mongooses always showed site fidelity for a
multiday home range, but in only 59% of the cases for daily home range. Adult males exhibited less daily site fidelity than
did adult females or young. Multiday home-range size was similar among age—sex classes, but males had significantly more
core areas than females or young. Moltiday home-range size was positively correlated with body mass for adult males
(r2
0.98, P 0.0122) and negatively correlated with body mass of adult females (r2 0.40, P 0.0374). Differences
in these relationships and daily site fidelity between adult males and females suggest that the spatial strategies of male and
female Egyptian mongooses are diffesent, with the \arger femaes defending the areas richer in resouTces and the larger males
having more access to females.
=
PALOMARES,
=
F. 1994. Site fidelity and effects of body mass on home-range size of Egyptian mongooses.
L’iniportance du domaine vital est reliée A Ia masse du corps chea certains animaux et ce principe prévaut aussi bien ehez
tome l’espAce que chez les individos d’une espAce lorsque les strategies spatiales diff’erent chez les mAles et les femelles. J’ai
étudie Ia taille du doniaine vital ehez un carnivore polygyne, Ia Mangouste ichneumon (Herpesres ichneuinon), et j’ai compare
las résultats aux valeurs théoriques dCfinies en fonction de Ia masse corporelle. J’ai d’abord tenté de determiner si les
snangoustes soul vraiment fidAles h un she particulier (pour des périodes de 1 jour ou Ae plusieurs jours). Les mangoustes
se sont toujours avCrées tidCles A un domaine vital de plusieurs jours, mais sont restées fidCles au domaine utilisd one journée
dans settlement 59% des cas. Les mAles adultes sont restés moms fidéles A un domaine d’un jour que le femelles ou les
jeunes. La taille des domaines de plusieurs jours était Ia mAme chez toutes les classes d’Age et chez les detix sexes, mais,
chez les males, ces domaines comptaient un plus grand nombia de points dutilisation plus fréquente. La taille de ces domaines
de plusieurs jours Ctait en correlation positive (r2 0,98, P 0,0122) avee Ia masse corporelle chez les males adultes. Chez
les femelles, la taille de ces domaines était en correlation negative (r2
0,40, P = 0,0374) avec Ia masse corporelle. Les
differences entre ces relations et Ia fidélité A un domaine d’un jour entre mAles et femelles adultes semblent indiquer qu’ils
ont des strategies spatiales diffCrentes : les femelles les plus grosses défendent les zones les plus riches en ressources, alors
que les plus gros males s’assurent an accés plus facite aux fbmelies.
[Traduit par Ia Rédaetionj
=
=
Introduction
Home-range size is a frequently investigated phenomenon
in ecology because of its importance to our understanding the
patterns of space use and spatial behaviour of animals.
Accordingly, there is a rich empirical and theoretical history
regarding home ranges (e.g., McNab 1963; Hatesta4 and
Bunnell 1979; Cameron and Spencer 1985; Lindstedt et al.
1986; Clutton-Brock 1989; Sandell and Liberg 1992). These
studies address the adaptive significance of (i) interspecific
variation in home-range size from the perspectives of
metabolic needs, body mass, feeding habits, and mating
system, and (//) intraspecific relationships between spatial
strategy, i.e., the strategy of each sex, and the type and
abundance of resources,
Large species usually have larger home ranges than do
small species (McNab 1963; Harestad and Bunnell 1979;
Gittleman and Harvey 1982; Lindstedt et al. 1986). This is
presumed to be related to greater total.metabolic needs in
the former, and consequently larger borne ranges are
required to accommodate these needs. Assuming a similar
allometric relationship at the intraspecific level, heavier
individuals should have larger home ranges than small
individuals. A fe.w studies suggest that these trends occur
intraspecifically in the Carnivora (e.g., Knick
1990;
Gompper and Gittleman 1991).
relation to the spatial distribution of patches cii these foraging
habitats and occupy exclusive core areas, which suggests that
they defend food resources. In contrast, males are distributed
in relation to females and are territorial (Palomares and
Delibes 1993a). This situation permits a within-species test
of whether the existence of a different spatial strategy for each
sex determines contrasting relationships between hotne-range
size and body mass as predicted above.
In non-Carnivora species with promiscuous or polygynous
mating systems, space use often differs between the sexes,
because males and females use resources differently and have
different behavioural strate’gies to increase their reproductive
success (Schoener and Schoener 1982; Cameron and Spencer
1985; Clutton-Brock 1989; Waiters and Dhondt 1992).
Hence, in the Carnivora we expect spacing patterns to be influenced by sex, beyond differences in body mass (Sandell
1989). Females can be distributed according to food resources,
while males can be distributed according to the distribution
of females. If so, and assuming a positive relationship between
competitive ability and body mass, in the case of a promiscuous or polygynous species we expect a negative correlation
between home-range size and body mass in females because
they would compete for areas with the best food or habitat
patches of higher quality. For males, we expect a positive
relationship because they would try to maximize their mating
success.
Egyptian mongooses, Herpestes ichneumon, behave as
polygynous carnivores in southwestern Spain (Palomares and
Delibes 1993a). Individuals of both sexes forage in rather
specific habitat patches, where their staple prey (rabbits,
Oryctolagus cunicukus; Palomains and Delibes 1991) axe more
abundant and the undergrowth vegetation is denser
(Palomares and Delibes 1993!,). Females are distributed in
The primary goals were to study home-range size in
Egyptian mongooses and to examine general trends and
sex-related differences in home-range size in relation to body
mass. However, before measuring home-range size and
inferring ecological or behavioural factors from its pattern, it
was necessary to demonstrate that animals exhibit site fidelity
(i.e., they do not wander or disperse through the study area;
Munger 1984; Spencer etal. 1990) and to investigate the effect
of sample size and method of measurement on its estimation
(e.g., Bowets 1982; Boulanger and White 1990). Therefore,
possession of a home range by each individual was tested for,
and the way in which sample size affected its size was examined.
Mongooses were considered to possess a home range if they
exhibited site fidelity (i.e., the observed area used by an
individual was significantly smaller than the area that would be
used if its movements were random; Munger 1984).
Study area and methods
The study took place at Coto del key (Doiiana National Park,
southwestern Spain; 37°9’N, 6°26’W). The vegetation of Coto dcl
Rey is characterized by pines (Pious pinea) and eucalyptus
(Eucalyptus sp.). Along small stre.anis, Froxinsis sp., Popular alba,
Pistacia lernircus, and Rubus sp. predominate. Associations of
P. ientiscus occur on places with a higher groundwater table. More
information on the study area can be found in Palomares and Delibes
(1993a).
Twenty-four mongooses were caught and equipped with radio
collars containing tip switches (Wildlife Materials Inc., Illinois,
U.S.A.) (Palomares and Delibes W92a). Three individuals removed
or broke their radio collars. Another was killed by poachers a few
days after its release. Therefore information was obtained from
20 individuals (14 adults (10 F: 4 M), I immatwre (F), arid 5 young
(3 F : 2 M)). The immature female attained maturity while being
monitored and was considered adult for the analyses. Age was
inferred from body mass and dentition (Palomars and Delibes I 992b).
Two radio-tracking sampling schemes were used: (i) 54 days of
intensive tracking of 15 mongooses (3 adult males, 9 adolt females,
and 3 young) located every 30 mm for Il—IS h during daylight, and
(ii) one or two locations daily of each of the 20 individuals. The
homing method (Mech 1983) was used for all locations without
disturbing the animals (Palomares 1990). This method avoided the
error associated with triangulation (Hearer and Tester 1967; Mills
and Knowlton 1989).
To determine whether mongooses had significant levels of site
fidelity (i.e., used a home range and were not wandering or dispersing), the procedure developed by Munger (1984) and Danielson
and Swihart(l 987) for liz.ards and rodents, respectively, was modified
slightly. This frocedure consisted of comparing the size of the
minimum area (estimated by the minimum convex polygon method
(MCP); Hayne 1949), which included the path traversed by an animal,
with the size of an area obtained from random paths generated within
sttitable habitats. If the MCP value based on actua.l movement was
significantly less (using 95% confidence intervals) than the mean size
based on 100 random paths, then the individual was judged to exhibit
site fidelity. MCP was chosen for the analysis because it is easy to
calculate and less sensitive to the use of autocorretated data (which
is the case for daily periods) than probabilistic models (Swihart and
Slade 1985a).
Because site fidelity can vary according to the temporal scale used
(Spencer er a!. 1990). site fidelity for both daily and multiday periods
for every individual was tested for. Data for daily and muleiday
periods came from full days of intensive tracking and the total number
of routine locations, respectively. Mongooses are diumal in the study
area (Palornarecs and Delibes 1992c), so between 21 arid 30 autocorrelated locations (depending on date) were available per day
during intensive tracking. For reasons of independence, only diurnal
locations separated by at least 4 h were used, and only one per night
was used for multiday home-range estimates (Swihart and Slade
t985&; Reynolds and Laundré 1990; Gese et al. 1990). To test
possible changes in site fidelity throughout the period of sampling
individuals (i.e., wandering or dispersing movements), multiday
periods were also subdivided into blocks of 10 fixes, and site fidelity
was independently tested for these blocks. Further details of the
procedure and its applicability to testing she fidelity and wandering
movements will be published elsewhere.
Multiday home-range size was estimated by two methods using the
RANGES software package (Kenward 1990): the above-mentioned
MCI’ and 95% harmonic mean isopleth (HM95) centering of fixes in
40 )< 40 grid cells (Dixon and Chapman 1980; Spencer and Barest
1984). The extent and number of core areas (Adams and Davis 1967)
inside the home-range of each individual were estimated with the
65% harmonic mean isopleth (HM65). This value was obtained using
the procedure proposed by Harris at a!. (1990) and Kenward (1990),
in which area is plotted against harmonic mean isopteth value..
Multiday home-ranges were estimated for age—sex classes (adult
males, adult females, and young).
Observed muttiday home-range sizes were compared with those
predicted by the allometric equation of Lindstedt et al. (1986) for
carnivores living at temperate latitudes.
Relationships between multiday home-range size and each separate
factor (number of fixes and body mass) were examined by simple
linear regression analysis (Kleinbaum et al. 1988). The relationship
between body mass and home-range size was only examined for
adults. Body mass was measured at first capture.
Sample size influenced home-range size estimations (see Results).
Therefore, to minimize the influence of sample size when examining
the relationship between home-range size and body mass, a method
proposed by Harris et aJ. (1990) was used: home-range sizes were
estimated on the basis of a standani number of fixes, which was close
to the figure at which most home-range size estimates reach an
asymptote. Between 30 and 40 fixes were close to this figure for the
mongooses studied (Palomares 1990). For 15 individuals, 32—40
fixes were available, and home-range sizes of the remainder were
estimated using 27—29 fixes for three individuals and 20 and 21 fixes
for the other two individuals. I selected the first 40 fixes for
individuals for which mere than 40 fixes were available.
Home-range size estimations using the MCP methed and a
standardize number of fixes (i.e., as close as possible to 40 fixes,
hereafter referred to as MCP4O) were used to test whether body mass
and home-range size were related in adults. The latter method
(MCP4O) was selected because it was less sensitive to the skewed
distribution of fixes obtained from two individuals that concentrated
their activity at the edge of Dohana National Park, where crops
constituted a barrier to their movements. The HM95 method using
30—40 fixes tended to encompass large unused areas (see Spencer
and Barrett 1984), and thus to overestimate home-range size.
correct for deviations from normality, logarithmic
To
transformations were performed on home-range size and ‘body mass.
Throughout this paper, I avoid pseudoreplication by using the Sean
when more than one value was available for each individual (Hurlbert
1984).
Results
Site fidelity
For all individuals (it = 20), multiday space use was more
concentrated than random movements. However, an immature
female had an MCP area equal to or larger than the simulated
4€?
PALOMARES
TABLE I. Mean (and standard deviation) daily home-range size
TABLE 2. Means, standard deviations (SD), and ranges (km2) of
(estimated by the minimum convex polygon (MCP) method),
multiday home-ranges calculated using different methods, and the
number of core areas (NCA) and its size (F1M65) (from the 65% isopleth determined by the mean harmonic method) of adult male, adult
female, and young Egyptian mongooses
percentage of multiday MCP covered daily, percentage of days on
which site fidelity was exhibited (i.e., the actual daily MCP was smaller
than the simulated one), and percentage of days on which actually daily
MCP was higher than the simulated one for adult male, adult female, and
young Egyptian mongàoses
MCP
(1cm2)
Adult males
(n 3, 16 days)
Mean
SD
(n = 9, 26 days)
Mean
SD
Young
Mean
SD
% of multiday MCP
Site
fidelity
MCP higher
than
simulated
MCP
Adult males (n
Mean
SD
Mm
Max.
0.96
26.1
0.36
16.6
0.42
21.6
0.20
22.4
0.51
0.07
25.7
11.1
38.1
16.0
58,!
19.1
81.4
24.3
11.1
16.7
56.7
20.8
36.7
32.1
as the sampling unit.
MCP area during four of the first five 10-fix blocks of
observed paths, suggesting that this female was dispersing
during this period. Fixes from this period were not used in
following analyses. No other individual showed such a
pattern.
Daily movements were more concentrated than random
movements on 59% of days (n = 54). Interestingly, adult
males exhibited proportionally less site fidelity and higher
actual than simulated daily MCP values than did adult females
and young (Table 1).
Multiday home-range size
Mean multiday home-range sizes did not differ significantly
among age—sex classes (Table 2). Observed multiday homerange values (mean = 3.10 km2, SD = 2.12 km2, n
20;
MCI’ method) were similar to those predicted by Lindstedt
et aL’s (1986) equation (mean
2.85 km2, SD = 0.44 km2.
n = 20). Observed and predicted home-range sites were not
significantly correlated for the whole data set (Spearman rank
correlation coefficient, r, = —0.01, P = 0.9843), but
significant correlations were found when each sex—age class
was analysed separately. The correlation was negative in adult
females (r, = —0.75, n = 11, P
0.0249) but positive in
adult males (r5 = I, n = 4, P < 0.0001).
Individual multiday home-range size and area covered
during daily movements (both obtained by MCP method)
were not significantly correlated (r, = 0.43, P = 0.1041,
n = 15). The percentage of the multiday home-range size
covered daily by males was similar to that for females and
young (Table 1). However, the mean daily MCP for males
was larger than that for both females (Mann—Whitney U test;
Z = 2.03, P = 0.0422) and young (Z = 1.74, P = 0.0809;
Table 1). On average, daily home-range size for males was
2.29 times (SD = 0.86, n = 3) and 1.89 times (SD = 0.71,
n = 3) larger than that for females and young, respectively.
On average, core areas never represented more than 28%
of the multiday HM95 range (Table 2). The sizes of core areas
were not statistically different among age—sex classes; on
=
HM95
RM6S
NCA MCP4O
4)
3.52
2.58
3.07
1.94
0.85
0.84
4.0
1.8
2.56
0.55
6.82
0.59
5.05
0.17
2.07
2
6
0.30
0.03
0.92
(1.4
1.12
I
0.57
2
3.42
0.28
0.36
1.08
0.5
1
2
0.65
l1.l5’
0.78
1.56
0.55
3.98
Adult females
(is = 11)
SD
2.43
Mm.
0.57
Max.
7.73
2.67
0.29
8.77
1.08
1.45
4.28
1.04
1.38
4.21
ye
0.08
0.90
1.94
pC
0.960
0.636
0.380
Young (is = 5)
SD
Mm.
Mat
1.45
3.01
0.004 0.677
Nora: MCP, minimum convex polygon: HM95, 95% mean harmonic isopleth;
MCP4O. minimum convex polygon using as close as possible to 40 fixes.
‘4Values obtained from Kroskal-wallia Lest.
tThere were significant differences between males and females and males and
young (Z = 3.03 and 2.06, P = 0.002 and 0.040, respectively; Mann—Whitney
U test), but not between females and young (2 = 1.55, P = 0.121).
CPrgability of differences among sex—age group of individuals.
average, however, males had 4 core areas, significantly more
than females (mean = 1.2) or young (mean = 1.6) (Table 2).
Multiday home-range size and number of fixes
MCP and HM95 home-range values were affected by
the number of fixes used for the calculations (Y = 1.149 +
0.682x and 1.099 ± 0.678x, r2 = 0.47 and 0.44, respectively;
P < 0.0014 in both cases; number of fixes ranged between
20 and 259). Using MCP4O, a reduction in home-range size
of between 15 and 35% was observed (Table 2). Differences
in MCP4O home-range values among age—sex classes
remained nonsignificant (Table 2).
Muifiday home-range size and body mass
Body mass accounted for variation in home-range size for
adult mongooses (Fig. 1). A positive correlation was observed
for males (r2 = 97.6%, P = 0.0122). In females there was a
negative correlation (P = 0.0374), and body mass explained
39.8% (r2) of the variation in home-range size.
Discussion
The results of testing site fidelity, estimating the size and
configuration of the range, and examining the relation
between range size and body mass show differences in space
use and social behaviour of Egyptian mongooses of different
sexes and ages.
Daily site fidelity was significantly greater for females and
young than for males. Males exhibited territorial behaviour
(Palomares and Delibes 1993a) and had more core areas inside
their ranges than did either females or young (Table 2); daily
patrolling or movements between core areas could therefore
468
(5
4
C
0
C.)
0
-J
Log Body Mass (kg)
FEMALES
I..
4
0
0..
C,
0
O4
0.42
0.44
0.46
0.48
0.5
Log Body Mass (kg)
1. Relation of home-range size to body mass of adult male and
female Egyptian mongooses, measured by the minimum convex
polygon method and usingas close to4O locations as possible (MCP4O).
FiG.
include more area than expected from random. If this
behaviour actually influenced male movements, daily MC?
home-ranges of males should be both larger than those of
females and young and larger than expected from only body
mass and energy requirements (because males are heavier than
females; Palomares and Delibes 1992b). The mean daily MC?
for males was significantly larger than for either females or
young (Table 1), and daily home-range sizes for individual
males were 2.29 and 1.89 times larger than those for females
and young, respectively. However, the ratios of expected
(based on energetic considerations) males’ mean daily MC?
home-range size (i.e., males’ home-range size. = females’
home—range size >z males’ mass°75 /fernales’ mass°’5 Sandell
1989) to observed mean daily MCP for females and young
were 1.16 (SD = 0.06, it 3) and 1.72 (SD 0.09, a 3),
respectively. Males’ daily ranges were thus larger than those
expected from energy requirements.
Multiday home-range size of mongooses varied little among
sex—age classes regardless of the method used to estimate
home-range size. These observations suggest that individual
variation was greater than that observed between age—sex
classes. Individual variation could well be produced by
—
different body masses, as supported by the relationships found
between home-range size and body mass and between
observed and predicted (by Lindstedt et al.’s (1986) equation)
home-range sizes.
As previously shown by other authors through simulations
or the use of data gathered from other species (e.g., Swihart
and Slade 1985a; Boulanger and White 1990). sample size
influenced estimates of home-range size in Egyptian
mongooses. This justified an assessment of the effect of
sample size on home-range size and the use of the corrected
MCP method to correlate home-range size with body mass.
However, the use of complete data sets in other analyses to
more closely assess the true home-range sizes of mongooses
was preferred, so useful biological information was not
missed.
A negative relation between home-range size and body mass
was observed in females. A hypothesis to explain the female
pattern could be that bigger individuals may be able to find
and defend areas with richer resources (i.e.. to be socially
dominant), and, therefore, are able to support themselves in
smaller areas. The proportion of foraging habitat inside
females’ core areas was greater than in. the rest of the multiday
home-range area (Palomares and Delibes 1993a). Furthermore, mongoose core areas were small (they represented only
17—28% of whole multiday hone-range size), suggesting high
habitat productivity (prey density) in the study area. This
seems to be the case because mongooses devote only a small
percentage of their daily time (25%) to activity (Palomares
and Delibes l992c). Because females’ core areas were
exclusive (Paloinares and Delibes 1993a), this proposed
hypothesis seems reasonable. Although there is no stu4y that
specifically relates body size to social dominance in
mongooses, it has been shown in the dwarf mongoose,
Helogale parvukz, that the socially dominant individuals are
the oldest ones (Rood 1990), who in its turn may be the largest
ones (e.g., see Fig. 1 of Creel et al. 1991). Why was the same
pattern not observed in male Egyptian mongooses, though?
The number of core areas used by males was significantly
greater than those used by females and young. Males were
territorial and therefore increased the number of females
monopolized (including more females’ core areas) as their
home-range sizes increased (Paloniares and Delibes 1993a),
so females rather than food niny be the limiting resource for
males, even outside of the mating season (see below). If the
females’ home-range size is determined by food, the body
mass of the two sexes can again be used to predict the
home-range size required by males from an energetic
standpoint (Sandell 1989). For the males sampled, the ratios
of the expected home-range size for males (see above) to the
observed mean home-range size for females were 1.2, 1.0,
1.1, and 1.2, However, actual ratios were 3.8, 0.5, 0.8, and
2.4, respectively. Actual ratios were much higher than
expected for heavier males and lower for smaller males,
suggesting that home-range sizes, as a rule, could not be
explained in terms of food resources alone. Sandell (1986)
also found two different spacing behaviours in male stoats,
Mustela erininea, which were related to age and size of
individuals.
Larger home-ranges for males would be predicted during
the mating season if females are their limiting resource
(Sandeli 1989). Even though there were insufficient individuals with long tracking periods to confidently test the
t
PALOMARES
seasonal hypothesis, available data for two males appear to
indicate that seasonal variations in range size were not
important. These individuals had home-range sizes (using the
MCP4O method and excluding 5% outlier fixes) of 3.9, 4.5,
and 4.5 kin2 for autumn, winter, and spring and 1.9 and
1.3 km2 for winter and spring, respectively (F. Palomares,
unpublished data). This observation does not invalidate the
proposed hypothesis that females are the main resources for
males, because in fact males increased the frequency of
contacts with the females inhabiting their territories during
the mating season (Palomares 1990).
Acknowledgements
The DirecciOn General de Investigacidn CientIfica y
TecnolOgica (project PB87-0405) and Consejo Superior de
lnvestigaciou.es Cientificas provided financial su.pport to the
author for this research. 1 am grateful to E. Collado for his work
and patience with computer simulations, and to M. Delibes,
E. Collado, tiC. White, T.M. Caro, J.W. Laundré, and two
anonymous reviewers for providing useful comments on earlier
versions of the manuscript. K. Nelson reviewed the English
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