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(usually as juveniles) than before the reduction. Such compensation is rarely complete,
so that the equilibrium in the presence of a constant offtake of animals will be somewhat
lower than the equilibrium in the absence of any hunting. But as long as offtake does
not exceed the population's ability to compensate in some way, an equilibrium will be
reached, and continued hunting will not result in a chronic decline. 11
Further, in the case of trophy hunting, managers quickly discover that sustainable
offtake rates are conservative ones. Unlike in models underlying the classic theory of
sustainable yield, the objective in trophy hunting is to produce large males, not merely
a specific yield of animal bodies. To become large, one must become at least somewhat
old, and to be able to grow old, mortality rates among young males must be tempered
lest all die before they attain trophy size. In any case, females are generally protected in
a population managed for trophy hunting, and it is the female side of things that actually
controls population growth. Thus, researchers have rarely been terribly concerned about
the demographic issue in trophy hunting.
A second concern is that a population subjected to hunting only of mature males could
decline if there remain too few to inseminate females. If so, some females that would
otherwise produce young in the spring following the hunting season might fail to do so,
thus reducing the reproductive (and growth) rate of the population. Again keeping in mind
the lack of detailed studies for any of the Chinese species involved, there is little reason
for concern that this would occur in a population managed for trophy hunts. Studies of
ecologically similar species in North America and Europe, even in cases where almost all
of the oldest age-classes of males are removed, have generally shown that younger males
are more than willing to step up and breed in their stead. 12 It may not be the same males
who are doing the breeding as in an unhunted population, but the imperative to produce
the next generation is strong; it takes an unusually heavy removal of males to reduce the
normal proportion of adult females impregnated. 13
A third concern—one that might be applicable particularly to argali—is that removing
the oldest, most dominant males exacts a cost in survival for the remaining, younger males.
This effect has been suggested as potentially present in species that feature hierarchical
mating systems in which males physically contest with each other for dominance, and the
dominant males then obtain the majority of copulations. 14 That is, the very contesting of
dominance exacts a price: considerable energy is expended in constantly defending one's
high place in the hierarchy, and animals that focus their energy on such competition also
reduce their food intake. Younger males, which have little chance of becoming dominant
in any case, will usually expend relatively little energy even trying. Easily dissuaded from
their ambition, they will generally resort to alternative tactics to attempt copulations,
waiting until they grow large enough to seriously contest for dominance. 15
Thus, we generally find that the survival rate among young males in such hierarchically
structured species is higher than among the dominant males. The risk in trophy hunting is
that by removing most or all of the older, dominant males (which suffer a higher mortal-
ity rate in any case), the energetically costly competition for dominance then devolves to
younger males who otherwise would be spared it. This line of reasoning acknowledges
that females will be bred in any case, whether by older males or younger ones, but the
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