Behavioral Ecology
Behav Ecol Sociobiol (1988) 22:37 47
Patch use as an indicator of habitat preference,
predation risk, and competition
Joel S. Brown*
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA
Received January 28, 1987 / Accepted September 25, 1987
Summary. A technique for using patch giving up
densities to investigate habitat preferences, preda-
tion risk, and interspecific competitive relation-
ships is theoretically analyzed and empirically in-
vestigated. Giving up densities, the density of re-
sources within a patch at which an individual
ceases foraging, provide considerably more infor-
mation than simply the amount of resources har-
vested. The giving up density of a forager, which
is behaving optimally, should correspond to a har-
vest rate that just balances the metabolic costs of
foraging, the predation cost of foraging, and the
missed opportunity cost of not engaging in alterna-
tive activities. In addition, changes in giving up
densities in response to climatic factors, predation
risk, and missed opportunities can be used to test
the model and to examine the consistency of the
foragers' behavior. The technique was applied to
a community of four Arizonan granivorous ro-
dents (Perognathus amplus, Dipodomys merriami,
Ammospermophilus harrisii, and Spermophilus tere-
ticaudus). Aluminum trays filled with 3 grams of
millet seeds mixed into 3 liters of sifted soil pro-
vided resource patches. The seeds remaining fol-
lowing a night or day of foraging were used to
determine the giving up density, and footprints in
the sifted sand indicated the identity of the forager.
Giving up densities consistently differed in re-
sponse to forager species, microhabitat (bush ver-
sus open), date, and station. The data also provide
useful information regarding the relative foraging
efficiencies and microhabitat preferences of the
coexisting rodent species.
Introduction
As first suggested by MacArthur and Pianka
(~ 966) and Emlen (1966), optimal foraging theory
* Current address: Department of Biological Sciences, Universi-
ty of Illinois at Chicago, Chicago, IL 60680, USA
and Sociobiology
9 Springer-Verlag 1988
can be used to investigate the properties of commu-
nities. A recent example of this includes Rosen-
zweig's (1979 et seq.) theory of habitat selection
where the behavior of individuals behaving opti-
mally is used to predict the outcome of intra- and
interspecific interactions (Rosenzweig 1981, 1985;
Pimm and Rosenzweig 1981; Pimm et al. 1985).
Other examples include short-term apparent com-
petition (Holt and Kotler 1987), moose foraging
dynamics (Belovsky 1978 et seq.), and resource
theory (Tilman 1982, 1985).
Testing models of population interactions
based upon the foraging behavior of individuals
often requires measuring habitat or patch use, hab-
itat preferences, and the rejection or acceptance
of patches. In this paper, I present a measuring
technique to accomplish these goals. The technique
uses artificial or manipulated resource patches to
measure a forager's giving up density. The tech-
nique is applicable to communities of active for-
agers seeking comparatively immobile prey.
I begin by discussing the theory which provides
an interpretation of patch giving up densities. The
theory is an extension of Charnov's (1976) mar-
ginal value theorem and it uses what, in economics,
is called the "marginal rate of substitution" (see
Russell and Wilkinson 1979) to include the effects
of predation risk and alternative activities on patch
use. I then give some preliminary results from ap-
plying this method to a community of four desert
granivorous rodent species: Arizona pocket mouse
(Perognathus amplus), Merriam's kangaroo rat
(Dipodomys merriami), round-tailed ground squir-
rel (Spermophilus tereticaudus), and Harris's ante-
lope ground squirrel (Ammospermophilus harrisii).
Optimal patch use
Theoretical treatments of optimal habitat use are
applicable to systems where foragers are able to
identify and direct their foraging efforts to subsets
38
of the environment (i.e. patches) that on average
yield higher harvest rates or benefits than the envi-
ronment at large. Once a habitat patch is located,
a forager must decide whether to accept the oppor-
tunity to harvest the patch (Rosenzweig 1974), and
it must decide how much time or effort to devote
to those patches it accepts for harvesting (Charnov
1976). By varying the foraging time allotted to each
patch, a forager can vary the total, average, mar-
ginal or net reward from each patch.
In the simplest case of habitat utilization, an
individual can only forage and has no alternative
activities. Assume that resources are distributed in
discrete patches and that foragers deplete the re-
sources as they harvest a patch. However, assume
also, that the distribution of resources among
patches remains fixed. In other words, although
the resources of a given patch are depleted by a
forager while it is in the patch, the resources of
the entire environment are not. Under these as-
sumptions, the marginal value theorem (Charnov
1976) states that the rate of energy gain to a forager
is maximized, and by assumption its fitness is also
maximized, when the forager's quitting harvest
rate in a patch equals its average harvest.
The above assumptions, which lead to the mar-
ginal value theorem, are restrictive. More realisti-
cally, foraging in patches will affect not only ener-
gy gain, but also other aspects of fitness such as
predation risk. Empirical work suggests that for-
agers balance the benefits of energetic reward and
the cost of predation when making foraging de-
cisions (Milinski and Heller 1978; Sih 1980; Grubb
and Greenwald 1982; Werner etal. 1983; Lima
et al. 1985). Furthermore, most organisms can en-
gage in other fitness-determining activities besides
foraging such as territorial defense, mate finding,
resting, dormancy, grooming, and nest mainte-
nance. Finally, foraging activity of individuals may
not only depress the resources of a given patch
but also, at least for a time, depress the resources
of the entire environment. For example, following
sunrise, hummingbirds and nectarivorous insects
often deplete their nectar resources (Feinsinger
1976; Schaffer et al. 1979; Brown et al. 1981).
In what follows, I relax these restrictive as-
sumptions (1) by assuming that foraging activity
affects both net energy gain and predation risk;
(2) by permitting the foragers to engage in alterna-
tive activities; and (3) by permitting the depletion
of resources over the entire environment.
Let fitness, denoted by G, be a function of net
energy gain from foraging, e, the probability of
surviving predation while foraging, p, and a vector
of inputs into fitness from engaging in any and
all alternative activites, a. I assume that the realiza-
tion of fitness is discrete and occurs following a
fixed amount of time T. If the individual is preyed
upon during this interval its fitness is zero. The
goal of the forager is to divide its time between
foraging and alternative activities so as to maxi-
mize fitness.
Net energy gain, e, and the probability of sur-
viving predation, p, are functions of time spent
foraging. Assume that time spent foraging is di-
vided over m resource patches. Resource patches
may vary with regard to initial resource density
and predation risk. Harvest rate is assumed to be
an increasing function of patch resource density.
As a forager devotes harvest time to the patch,
its resources are depleted and the harvest rate de-
clines. Let t I = ( 6 , " , tin) denote the vector of times
allotted to each of the m resource patches. Assume
that there are s alternative activities that contribute
to the other inputs into fitness, a. Let t " = ( t m + b ,
tm+s) be the vector of times devoted to each alter-
native activity. Note that alternative activities may
also expend energy and incur predation risk. For
accounting purposes, in what follows, I will include
these in the vector of inputs into fitness from alter-
native activities. Let
G [e (ts), p (t~), a (ta)] = p (t y) .f[e (ts), a (ta)]
denote the expected per capita rate of growth (fit-
ness) subject to the constraint that the sum of the
tj's and t~'s equals T where j = 1,..., m and i = m +
1,..., m + s. Futhermore, 0 < p (t s) _<1 is the proba-
bility of surviving to realize fitness as a function
of times exposed to predators while foraging, and
F[e(ti), a(t")]>0 is the expected per capita rate
of growth given the individual survives predation
while foraging.
I assume that energy enhances fitness (i.e. ~ G~
~e> 0) and that spending time foraging decreases
the probability of surviving predation to realize
fitness (i.e. ~p/~tj<O where j = 1,..., m). If patch
j has a lower risk of predation than patch k then
~p/~ t; > ~p/~tk.
The optimal values of t~ and ta can be found
through the technique of Lagrange multipliers
(Chiang 1974). I will consider those patches and
alternative activities such that the optimal values
for tj and ti are greater than zero for j = 1,..., m,
and i = m + 1,...m+s. Setting the necessary condi-
tion for the optimal tj equal to the necessary condi-
tion for the optimal ti yields for all j = 1,.--,m and
i = m + l,...,m+s:
[~ G (')/~ el [~e (.)/8 tj] + [~ G (-)/~p] [~p (")/~tj]
= [~ G (')/~ a] [ea (.)/eh]