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A comparative study of models for predation and parasitism

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66<br />

(4g. 8). Then f(X) <strong>for</strong> a st<strong>and</strong>ard X is the efficiency <strong>of</strong> the species concerned.<br />

further discussion, see the appendix to w 4i).<br />

It should be pointed out here that, on the whole, the review <strong>of</strong> <strong>models</strong> by WATT<br />

(1959) is invalid, firstly because his mathematics is <strong>of</strong>ten wrong, <strong>and</strong> secondly because<br />

he was confused between<br />

(For<br />

instantaneous <strong>and</strong> overall functions, between <strong>parasitism</strong><br />

<strong>and</strong> <strong>predation</strong>, <strong>and</strong> between the Z-Xo <strong>and</strong> z-Y relationships. It should also be noticed<br />

that the criticism against the assumption f(x)=ax invalidating the NICHOLSON-BAILEY<br />

<strong>predation</strong> equation does not invalidate THOMPSON'S <strong>parasitism</strong> equation, since the latter<br />

does not assume f(x)=ax.<br />

h).<br />

The HASSELL-VARLEY model <strong>of</strong> social interference in parasites<br />

Although this model is called by the authors (HASSELL <strong>and</strong> VARLEY 1969) 'a<br />

new model' based on the NICHOLSON-BAILEY competition equation (see w it is in<br />

fact a special case <strong>of</strong> the generalized THOMPSON'S model <strong>for</strong> indiscriminate parasites,<br />

eq. (4g. 8), in which the instantaneous hunting function is a modified NICHOLSON-<br />

BAILEY linear function. As already pointed out, THOMPSON'S equation <strong>for</strong> <strong>parasitism</strong><br />

takes the same <strong>for</strong>m as the NICHOLSON-BAILEY 'competition equation' <strong>for</strong> <strong>predation</strong><br />

if the instantaneous hunting function f(X) is assumed to be a linear function <strong>of</strong> X,<br />

i.e. f(X) =aX, in which the coefficient a is the 'effective area <strong>of</strong> recognition per<br />

unit time'. Under these circumstances, the value <strong>of</strong> /~, as defined by eq. (4b. 8), becomes<br />

at, a constant.<br />

It has been shown in w 4e, however, that the /~ cannot be constant, but is at least<br />

a function <strong>of</strong> X, host density. HASSELL <strong>and</strong> VARLEY, however, found that in some<br />

published data the value <strong>of</strong> ~ was not independent <strong>of</strong> Y, parasite density. These data<br />

are shown graphically in Fig. 11; this is a reproduction <strong>of</strong> figure 1 in HASSELL <strong>and</strong><br />

VARLEY (1969) with a slightly different arrangement. These data show that the<br />

value <strong>of</strong> In ii tends to decrease as the value <strong>of</strong> In Y increases. The interpretation <strong>of</strong><br />

these relationships by HASSELL <strong>and</strong> VARLEY is that the parasites interfered with each<br />

other more strongly as their density increased, <strong>and</strong> hence the reduction in "the area<br />

<strong>of</strong> discovery", i.e. /f. The authors stated that "the striking feature <strong>of</strong> the relationships<br />

in (Fig. 11) is that they are linear over several orders <strong>of</strong> magnitude", <strong>and</strong> that "the<br />

data <strong>for</strong> Chelonus texanus CRESS. [curve (c)] cover a narrow range <strong>of</strong> parasite densities<br />

but seem to imply a curvilinear relationship".<br />

Thus, it was concluded that the relationships were described by the following<br />

<strong>for</strong>mula<br />

In ii=ln Q-m In Y (4h. 1)<br />

or<br />

= O Y-~ (4h. 1'),<br />

in which the factor Q is called "the quest constant" <strong>and</strong> m "the mutual interference<br />

constant".<br />

If the above relationships are incorporated into the NICHOLSON-BAILEY model

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