376 GAME APPLICATIONS (Ch. 30)
(i) If the proportion of Hawks is slightly greater than E,whichstrat–
egy does better? Dove. If the proportion of Hawks is slightly less
than E, which strategy does better? Hawk. If the more profitable
strategy tends to be adopted more frequently in future plays, then if the
strategy proportions are out of equilibrium, will changes tend to move the
proportions back toward equilibrium or further away from equilibrium?
30.9 (2) Economic ideas and equilibrium analysis have many fascinating
applications in biology. Popular discussions of natural selection and bio-
logical fitness often take it for granted that animal traits are selected for
the benefit of the species. Modern thinking in biology emphasizes that
individuals (or strictly speaking, genes) are the unit of selection. A mu-
tant gene that induces an animal to behave in such a way as to help the
species at the expense of the individuals that carry that gene will soon
be eliminated, no matter how beneficial that behavior is to the species.
A good illustration is a paper in the Journal of Theoretical Biology,
1979, by H. J. Brockmann, A. Grafen, and R. Dawkins, called “Evo-
lutionarily Stable Nesting Strategy in a Digger Wasp.” They maintain
that natural selection results in behavioral strategies that maximize an
individual animal’s expected rate of reproduction over the course of its
lifetime. According to the authors, “Time is the currency which an animal
spends.”
Females of the digger wasp Sphex ichneumoneus nest in underground
burrows. Some of these wasps dig their own burrows. After she has dug
her burrow, a wasp goes out to the fields and hunts katydids. These
she stores in her burrow to be used as food for her offspring when they
hatch. When she has accumulated several katydids, she lays a single egg
in the burrow, closes off the food chamber, and starts the process over
again. But digging burrows and catching katydids is time-consuming. An
alternative strategy for a female wasp is to sneak into somebody else’s
burrow while she is out hunting katydids. This happens frequently in
digger wasp colonies. A wasp will enter a burrow that has been dug by
another wasp and partially stocked with katydids. The invader will start
catching katydids, herself, to add to the stock. When the founder and
the invader finally meet, they fight. The loser of the fight goes away and
never comes back. The winner gets to lay her egg in the nest.
Since some wasps dig their own burrows and some invade burrows
begun by others, it is likely that we are observing a biological equilibrium
in which each strategy is as effective a way for a wasp to use its time for
producing offspring as the other. If one strategy were more effective than
the other, then we would expect that a gene that led wasps to behave
in the more effective way would prosper at the expense of genes that led
them to behave in a less effective way.
Suppose the average nesting episode takes 5 days for a wasp that
digs its own burrow and tries to stock it with katydids. Suppose that the