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A comparative study of models for predation and parasitism

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39<br />

particles.<br />

As the average number <strong>of</strong> particles within the ring is 7rR2X <strong>and</strong> the<br />

number <strong>of</strong> tosses in time ts is kt,, where k is the frequency <strong>of</strong> tosses per unit time,<br />

the total number caught by the ring will be<br />

n~ =~R~kXt~.<br />

Although the right-h<strong>and</strong> side <strong>of</strong> the above <strong>for</strong>mula is the same as in eq. (4c. 4)<br />

describing HOLLING'S experiment, notation n, in the left-h<strong>and</strong> side is not n as in eq.<br />

(4c. 4). It was assumed in HOLLING'S model that all the discs tapped were taken,<br />

<strong>and</strong> it was a justifiable assumption since only one disc was discovered at a time as<br />

there was no overlap <strong>of</strong> the discs. In the present model, however, more than two<br />

particles may occur within the ring at one toss. Thus, if the predator is allowed to<br />

catch only one prey individual within its recognition zone at a time, the total number<br />

captured (i. e. n) will be the number <strong>of</strong> successful tosses rather than ns, the total<br />

number discovered.<br />

In the present model, the distribution <strong>of</strong> the prey individuals <strong>and</strong> the tosses <strong>of</strong><br />

the ring are ideally at r<strong>and</strong>om, <strong>and</strong> so, provided that R is not too large as compared<br />

with the table, the frequency <strong>of</strong> tosses in which a given number <strong>of</strong> particles falls<br />

must follow a POISSON series with its mean equal to the average number <strong>of</strong> particles<br />

in the ring at one toss, i.e. rcR~X. Hence, as the frequency with which no particle<br />

occurs within the ring in the period t, will be kt,e -~R*X the total frequency <strong>of</strong><br />

successful tosses in t, will be<br />

n =kt, (l-e- ~R~X ).<br />

In the above model, only one ring was assumed to be tossed at the frequency <strong>of</strong> k<br />

per unit time per table.<br />

If there are Y rings per unit area tossed independently,<br />

each at the average frequency <strong>of</strong> k per unit time, the total frequency is kYts, <strong>and</strong><br />

so the above equation becomes<br />

n=kYt~(1-e -'~R~X ) (4d. 4).<br />

If kYt~ in the above equation is set equal to b, <strong>and</strong> rcR 2 to a, we have IVLEV'S eq.<br />

(4d. 2"). Thus, we find that IVLEV'S equation, in terms <strong>of</strong> the toss-a-ring simulation,<br />

is a generalized version <strong>of</strong> HOLLING'S experiment, since eq. (4d. 4) is a generalization<br />

<strong>of</strong> eq. (4c. 5). That is, recognition zones can overlap, <strong>and</strong> so there is no restriction<br />

to the range <strong>of</strong> variation in the prey density X (remembering that HOLLING'S eq.<br />

(4c. 5) holds in the disc experiment only <strong>for</strong> X

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