January 1999 REPORTS 337

Ecology, 80(1), 1999, pp. 337–340

q 1999 by the Ecological Society of America

ENERGETIC CONSTRAINTS AND THE RELATIONSHIP BETWEEN BODY

SIZE AND HOME RANGE AREA IN MAMMALS
DOUGLAS A. KELT AND DIRK VAN VUREN
Department of Wildlife, Fish, and Conservation Biology, University of California, Davis, California 95616 USA

Abstract. Recent theoretical developments to explain the unimodal and asymmetric

distribution of body sizes among species in higher taxa have yielded predictions for related
demographic and life history traits. In particular, it has been predicted that there is an
energetically optimal body size (M*) for terrestrial mammals at ;100 g, and that the
relationships of many biological characteristics will change slope or even sign at this point.
We reanalyze a well-known data set on the relationship between home range size and body
size in mammals. If the distribution of home range sizes as a function of body size is
energetically constrained, then it is reasonable to assume that the lower size limit of a home
range for a given body mass will also be constrained. To evaluate this hypothesis, we
predicted that the relationship between minimal home range size and body size is nonlinear,
and that the smallest home ranges should correspond to species in the vicinity of M*. Our
data tentatively support both hypotheses but constitute a clear call for more comprehensive
analyses with larger data sets.
Key words: body size; energetic constraints; home range area; mammals; optimal body size.

INTRODUCTION Maurer 1987, 1989). The very largest geographic rang-

A recent review on the relationship between home
range size and body size in mammals concluded that
‘‘we presently know less about the scaling of home
range area than about the dependence on size of almost
any other ecological, anatomical or behavioral vari-
able’’ (Reiss 1988:86). Until recently, most authors as-
sumed that allometric relationships with body size were
linear across all species. Many characteristics of ani-
mals, including basal metabolic rate, brain size, heart
size and rate of blood circulation, litter size and size
of offspring at birth, etc., scale linearly over most body
sizes when plotted on logarithmic axes (Peters 1983,
Calder 1984). However, there is increasing evidence
that nonlinear relationships may better describe many
variables, ranging from maximum life-span to median
dispersal distances to the frequency distribution of
body sizes (e.g., Calder 1989, Brown et al. 1993, Fin-
dley 1993, Van Vuren 1998).
Such nonlinearity in allometric relationships may re-
sult from energetic constraints. Many variables appear
to be constrained in the range of variation that is al-
lowable for a given body size. For example, the rela-
tionship between geographic range size and adult body
size in North American birds and mammals is con-
strained by at least two separate dynamics (Brown and
Manuscript received 12 January 1998; revised and accepted
23 March 1998.
es are limited by the size of the North American con-
tinent, and the smallest geographic ranges are con-
strained by the energetic requirements of small body
size. As a result of the latter constraint, very small
geographic ranges are not possible for animals of ex-
treme body sizes (Brown and Maurer 1987, 1989). Re-
cently, Brown et al. (1993) provided a model to explain
such nonlinear constraints involving body size. They
argued that there is an optimal body size (M*) for mam-
mals, and that this was ;100 g. Because energetic con-
straints increase with both increasing and decreasing
size away from M*, Brown et al. (1993) predicted that
scaling relations would change in magnitude, or even
change sign, in the vicinity of M*, and they offered
examples of some ecological variables that exhibited
such nonlinear scaling relationships.
Home range size varies with body size and is influ-
enced by energetic requirements (McNab 1963, Hare-
stad and Bunnell 1979, Lindstedt et al. 1986, Swihart
et al. 1988). If there is an energetically based optimal
body size, such that energetic constraints increase with
both increasing and decreasing size from M*, then
these constraints should influence the lower limit of
home range area when plotted as a function of body
size. In particular, species whose mass approximates
M* should have the smallest minimum home ranges,
and the minimum home range that is possible should
increase for species whose mass is progressively fur-
 338 REPORTS Ecology, Vol. 80, No. 1

ther from M*, resulting in a ‘‘u-shaped’’ constraint

space. Our objective was to determine if home ranges
of mammals demonstrate the predicted pattern. To do
so, we reanalyzed a well-known data set on home range
area and body size in mammals (Harestad and Bunnell
1979).
METHODS
Harestad and Bunnell (1979) compiled data from the
literature on 48 species and subspecies of mammals,
averaging male and female adult masses. Their data
were taken from a variety of sources and included home
range areas based on direct observation, radioteleme-
try, and trapping. Because data for the shrews Sorex
vagrans and S. obscurus (the latter is currently syn-
onymized with S. monticolus; see Wilson and Reeder
1993) presented in Harestad and Bunnell (1979) were
unpublished, we obtained the subsequently published
data (Hawes 1977), and have calculated home range
size and body mass (obtained from Brown and Nico-
letto 1991) for these species, averaged across sexes
(Fig. 1a).
To determine if energetics produce a nonlinear con-
straint space that is least in the region of M*, we eval-
uated the lower limit of the body size–home range re-
lationship by focusing on the distribution of minimal
FIG. 1. Plot of the distribution of home range size, mea-
home range size. We divided the mass axis according sured in hectares, against body mass, measured in grams.
to 1/4 log10 segments, extracted the smallest home Panel (a) presents original data from Harestad and Bunnell
range in each segment, and plotted log10(smallest home (1979), with trophic groups denoted by different symbols (m,
range) against log10(body size). We applied a linear herbivores; v, omnivores; m, carnivores). In panel (b), we
regression to the data points below 100 g and a separate plot only the minimum home ranges in log10 body size cat-
egories vs. log10 body mass (linear regression statistics are
regression to data points above 100 g. The existence presented). Brown et al. (1993) calculated a range of possible
of a u-shaped constraint space is supported if the slope optimal body sizes (M*), from ;80 g to 250 g; this region
of the lower regression is significantly negative, and is denoted by the light vertical lines.
that of the upper regression is significantly positive.
It is worth reiterating that we are not evaluating
whether or not the overall relationship between body Most notably, minimal home ranges are smallest for
size and home range size is linear; rather, we are ad- mammals in the region of M*, ;100 g. These data
dressing the shape of the constraints that restrict the therefore are consistent with the hypothesis that con-
range of variation in this relationship. Additionally, straints determining minimal home range area in mam-
because our objectives are to evaluate a pattern that mals are nonlinear, and that they are greater for species
characterizes the entire class Mammalia, we have not with masses that are progressively farther from M*.
segregated taxa according to secondary features such Thus, the arguments proposed by Brown and Maurer
as trophic category, mode of locomotion, degree of (1989) to explain the existence and shape of constraint
sociality, etc. Although patterns at these lower scales lines for geographic ranges may also apply to home
of analysis are interesting in their own right, our ob- ranges: very large and very small species are energet-
jective is to evaluate the spatial implications of a hy- ically constrained to use larger foraging areas, as re-
pothesis based on underlying energetic constraints op- flected in Fig. 1b.
erating on all mammals. An alternative explanation for the distribution of data
points in Fig. 1 is that it merely reflects the superpos-
RESULTS AND DISCUSSION ition of simple linear relationships for the three trophic
The slopes of both regressions are significantly dif- groups (carnivores, omnivores, and herbivores), with
ferent from zero (Fig. 1b). Additionally, the lower re- similar slopes but with different intercepts. However,
gression has a significantly negative slope, whereas the this argument fails to account for the absence of species
upper regression has a significantly positive slope. from the lower left corner of the figure. Such species
 January 1999 REPORTS
would be very small (K100 g) and would have very
small home ranges. The absence of such species argues
against this explanation and for an energetic constraint
hypothesis, which states that this portion of bivariate
space is energetically unavailable to mammals, regard-
less of their trophic position.
It has been demonstrated elsewhere (Silva and
Downing 1995, Silva et al. 1997) that population den-
sity is nonlinearly related to body size, such that density
decreases with increasing distance from M*. Our anal-
ysis suggests that this lower density may partially be
a response to increasing home range requirements away
from M*. Damuth (1981) argued that the home ranges
of larger mammals have greater overlap with conspe-
cifics than do those of smaller mammals. If this gen-
erality can be extended to a relationship between home
range area and home range overlap, then the real re-
lationship between body size and home range area may
be less distinct than suggested by Fig. 1b. Unfortu-
nately, we are not aware of data that could either sup-
port or refute this hypothesis. Although such a rela-
tionship might reduce the slope of the constraint space
below M*, it would be unlikely to change the results
qualitatively; a constraint space evidently precludes
species from occurring in either the extreme lower right
or left-hand corners of this bivariate space.
It is worth noting that the smallest home ranges in
these data correspond to two species with a fossorial
habit: the northern pocket gopher (Thomomys talpoi-
des; M 5 140 g, A 5 0.02 ha, where M is body mass
and A is home range area) and Townsend’s mole ( Sca-
panus townsendii; M 5 142 g, A 5 0.10 ha). It might
be argued that the home ranges of these species are
constrained more by their environment than by their
body size. An alternative explanation is that their near-
ly optimal body sizes have allowed them to exploit a
subterranean environment, and that reduced home rang-
es are one facet of the selective regime to which these
species have adapted. Apart from this, fossorial species
have lower metabolic rates than nonfossorial taxa of
similar body sizes, and correspondingly lower ener-
getic requirements (McNab 1966, 1979, Contreras and
McNab 1990). Thus, they are predicted to have smaller
home ranges than their nonfossorial relatives. However,
the real question to ask is why we do not see any smaller
fossorial species with smaller home ranges. The con-
straint hypothesis predicts that smaller fossorial species
would also have larger home ranges. Again, only with
a larger data set will this question be resolved.
We would expect that these constraints on minimum
home range size would also hold true within major
trophic groups. Both theoretical considerations (see
Brown et al. 1993) and our analyses predict that the
relationship between body size and minimum home
range area should be nonlinear within different trophic
339
groups. The data used here are too coarse to test this
idea, but there is a hint that this may occur: the smallest
home ranges for both carnivores and herbivores occur
within the region that Brown et al. (1993) considered
energetically optimal (Fig. 1a).
Minimal home range area for North American mam-
mals appears to be nonlinearly related to body size.
Although this analysis does not fully resolve many de-
tails of this relationship, it does demonstrate that the
lower limit to home range size is nonlinearly related
to body size in mammals, and underscores the impor-
tance of nonlinear relations in ecology. Although ad-
ditional data, perhaps especially for nonfossorial mam-
mals in the vicinity of M*, will be required to fully
resolve this issue, we hope that this will encourage
other ecologists to consider nonlinear relations when
evaluating seemingly straightforward patterns.
ACKNOWLEDGMENTS
Although they may not agree with all of our interpretations,
Brent Danielson, Tim Caro, John Eadie, the Steve’s Pizza
seminar group, and three anonymous reviewers provided use-
ful criticisms on earlier drafts.
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