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A comparative study of models for predation and parasitism

A comparative study of models for predation and parasitism

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24<br />

This problem will not be discussed any further in this paper.<br />

The solution <strong>of</strong> simultaneous eqs. (4a. la) <strong>and</strong> (4a. lb) under the assumption <strong>of</strong> discrete<br />

generations was not considered by the original authors. The solution, as I will<br />

show in the next section, is in fact possible <strong>and</strong> is related to the NICHOLSON-BAILEY<br />

model.<br />

b).<br />

The NICHOLSON-BAILEY model<br />

This model is known as the 'Competition model'. It is, primarily, constructed <strong>for</strong><br />

the purpose <strong>of</strong> demonstrating NICHOLSON'S hypothesis that animal populations are in<br />

the state <strong>of</strong> balance fluctuating around a steady density <strong>of</strong> each species concerned,<br />

<strong>and</strong> that this steady density (or steady state) is brought about by competition among<br />

the members <strong>of</strong> the parasite species (NICHOLSON 1933). NICHOLSON with the collabo-<br />

ration <strong>of</strong> a mathematician, BAILEY (NICHOLSON <strong>and</strong> BAILEY 1935), intended to con-<br />

struct a model on the assumption <strong>of</strong> a very simple, idealized situation, concerning a<br />

theoretical relationship in densities between host <strong>and</strong> parasite species. By altering<br />

conditions in this simple model, they drew numerous conclusions about the mode <strong>of</strong><br />

existence <strong>of</strong> steady states.<br />

Whether or not NICHOLSON'S basic philosophy that animal populations are in the<br />

state <strong>of</strong> balance is a useful one, is not <strong>of</strong> concern here.<br />

It is more important to<br />

determine whether the basic premises in the NICHOLSON-BAILEY theory can produce<br />

a reasonable model <strong>for</strong> <strong>parasitism</strong>, so that a comparison between the model <strong>and</strong><br />

observation can provide any useful direction.<br />

original authors.<br />

The following is the reasoning by the<br />

Let x0 be the number <strong>of</strong> objects (hosts) originally present in a unit area, <strong>and</strong> let<br />

x be the number left undiscovered after an area s has been traversed (by all parasites<br />

concerned). Then the number <strong>of</strong> previously undiscovered objects discovered in a<br />

fraction <strong>of</strong> area traversed, i.e. ds, is xds. This must be equal to the decrease, -dx,<br />

<strong>of</strong> the number <strong>of</strong> undiscovered objects per unit area, i.e.<br />

-dx--xds (4b. 1),<br />

<strong>and</strong> since x=xo when s=0, integrating eq. (4b. 1) <strong>for</strong> the range (0, s), <strong>and</strong> hence<br />

(x0, x), we obtain<br />

X ~Xoe-*<br />

from which we have<br />

z/xo: 1- e-' (4b. 2)<br />

where z=xo-x. Factor s is called by the authors the 'area traversed', which is the<br />

area that is searched effectively by all parasites involved <strong>and</strong> includes possible overlaps.<br />

In passing, the average area traversed by each individual parasite is called the 'area <strong>of</strong><br />

discovery'. As against the area traversed, the net total area searched by all parasites<br />

concerned is called the 'area covered', which excludes areas already searched. Thus,<br />

the right-h<strong>and</strong> side <strong>of</strong> eq.<br />

(4b. 2) shows that the proportion <strong>of</strong> the 'area covered'<br />

increases only asymptotically as the 'area traversed' increases, <strong>and</strong> that there<strong>for</strong>e the

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