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A comparative study of models for predation and parasitism

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54<br />

effect <strong>of</strong> local depletion <strong>of</strong> prey upon the hunting efficiency will be less at higher<br />

values <strong>of</strong> x0. If a cross-section parallel to the z-xo<br />

circumstances, it should again be wavy.<br />

plane is observed under these<br />

A similar tendency to periodic deviation from a r<strong>and</strong>om distribution in accordance<br />

with changes in the host density was shown by SIMMONDS (1943, see also citation by<br />

WILLIAMS 1964). SIMMONDS, <strong>and</strong><br />

odic deviation in relation to the<br />

SIMMONDS' experiments, however,<br />

WILLIAMS as well, presented <strong>and</strong> analysed the peri-<br />

parasite/host ratio, rather than to the density. In<br />

parasite density was kept constant <strong>and</strong> host density<br />

changed <strong>for</strong> ratios between 1/200 <strong>and</strong> 1/25, but <strong>for</strong> those between 2/25 <strong>and</strong> 10/25 the<br />

parasite density was changed <strong>and</strong> the host density kept constant.<br />

There<strong>for</strong>e, the<br />

<strong>for</strong>mer corresponds to a cross-section parallel to the z-xo plane in the hunting surface,<br />

but the latter is a cross-section parallel to the z-Yt<br />

parable even if the ratio declines continually.<br />

plane, <strong>and</strong> so they are not com-<br />

Keeping this point in mind, one can<br />

compare my simulation model <strong>and</strong> SIMMONDS' observation, <strong>and</strong> find again a close simi-<br />

larity between them. It should be borne in mind that from the st<strong>and</strong>point <strong>of</strong> my<br />

simulation model, it is not the parasite/host ratio which is essential: it is the geo-<br />

metric properties <strong>of</strong> parasites' searching activity, changing as the densities change,<br />

that results in the pattern described above.<br />

Now, a question is posed as to whether the wavy trend in the hunting surface is<br />

inherent in <strong>predation</strong> <strong>and</strong> <strong>parasitism</strong>. As already shown, a hunting curve comes closer<br />

to the one expected in the r<strong>and</strong>om encounters between prey <strong>and</strong> predators, <strong>and</strong><br />

between hosts <strong>and</strong> parasites, when the predators, or parasites, shift their hunting area<br />

frequently.<br />

Of course, an approach to the curve expected in r<strong>and</strong>om encounters<br />

suggests a rise in the average hunting efficiency. This is, however, just an apparent<br />

relationship, because the time needed to h<strong>and</strong>le a victim has been excluded in the<br />

above <strong>models</strong>. If, in reality, a hunter shifts its hunting area much too <strong>of</strong>ten, the time<br />

involved in travelling from place to place will also increase. Consequently, the advan-<br />

tage <strong>of</strong> shifting will eventually be cancelled by the disadvantage.<br />

Hence, there must<br />

be an optimal frequency <strong>of</strong> shifts <strong>and</strong> an optimal distance (average) <strong>of</strong> travel that<br />

result in the highest hunting efficiency. There<strong>for</strong>e, a perfectly smooth hunting surface<br />

would not at any rate be expected.<br />

If, however, the prey or host individuals also<br />

move around independently <strong>of</strong> each other <strong>and</strong> <strong>of</strong> the predators or parasites, then<br />

r<strong>and</strong>om encounters might again be expected.<br />

It should be mentioned that the wavy pattern <strong>of</strong> a hunting curve (i. e. z plotted<br />

against n) is caused entirely by uninterrupted searching in the first simulation experi-<br />

ment, i.e. the predator's path is continuously increased with time. If the experi-<br />

ment was designed, however, so that the paths <strong>of</strong> some predators consisted <strong>of</strong> a<br />

number <strong>of</strong> short ones, as in the second simulation experiment, the wavy trend will<br />

be less pronounced. If the predator's path was completely discontinuous, that is, if n<br />

was varied entirely by Y, <strong>and</strong> t was extremely short, the waves would eventually<br />

disappear. However, as long as the path <strong>of</strong> each predator is allowed to be continuous

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