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(Brent) #1
As a natural consequence of these multispecies effects on feeding rates, optimal
patterns of diet choice can have important implications for predator–prey inter-
actions. An adaptive predator would shift its attention away from some species
as they become more and more rare. Such behavior can have a stabilizing influence
on predator–prey dynamics, reducing the degree of variability over time of cyclical
predator–prey systems (Gleeson and Wilson 1986; Fryxell and Lundberg 1994;
Krivan 1996). This is especially likely when the growth rate of the predator is poor
on alternative prey, when the forager exhibits partial preferences, or when alterna-
tive prey do not have overlapping spatial distributions (Fryxell and Lundberg 1997).

An alternative way to model diet choice employs a technique called linear programming
to identify the optimal solution to a requirement influenced by several constraints
(Belovsky 1978; Belovsky et al. 1989). When applied to optimal foraging, this allows
researchers the means to explore more subtle hypotheses. Linear programming can
be used to predict the optimal diet for a forager which is trying to maximize its intake
of energy, while at the same ensuring that it obtains sufficient intake of a scarce
nutrient to meet its metabolic requirements. When conducted for pairwise com-
binations of alternative foods, linear programming can be readily understood from
simple graphs (Fig. 5.5).
Belovsky (1978) used linear programming to predict the optimal choice of aquatic
versus terrestrial plants by moose, based on parameters for moose living on Isle Royale,
a small island in Lake Superior. One constraint is that moose must obtain a daily
intake of 2.57 g of sodium in order to meet their metabolic requirements. Terrestrial
plants in this system are deficient in sodium, whereas aquatic plants have much higher
concentrations. Like many other herbivores, moose have limits on the amount of food
material that can be processed each day through the digestive tract. The total daily
consumption of aquatic and terrestrial plants eaten by a moose cannot exceed this
processing rate, which we call the digestive constraint. Moose also have limits on
the number of hours that can be devoted to cropping food items. Finally, each food
type has different profitability (ratio of digestible energy content to cropping time).
Thus, time spent cropping energetically poor food items (such as aquatic plants) reduces
the opportunity to look for energetically richer items (terrestrial plants). In other words,
a moose might waste valuable time eating poor food that could be spent looking for
better food.

THE ECOLOGY OF BEHAVIOR 65

5.2.3Optimal diet
selection: linear
programming

4000

3000

2000

1000

0
0 1000 2000 3000 4000 5000
Intake of terrestrial plants

Intake of aquatic plants

Minimum energy requirement

Sodium requirement

Digestive constraint

Fig. 5.5Linear
programming model of
diet selection in moose,
based on Belovsky
(1978). The lines
represent constraints
that must be met. The
range of diets that fall
within these constraints
is enclosed in the
triangle in the middle
of the diagram.

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