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70 II. DE ARACHNIDIS<br />
using a scanning electron microscope, has seen on the upper surface of<br />
the abdomen knobbed setae peculiar to female Lycosidae. These afford<br />
a firm grasp to the undermost layer of the brood in their precarious<br />
situation. The knobbed setae are not innervated, but long, smooth setae<br />
are in contact with nerves and probably play a part in determining<br />
maternal behaviour.<br />
The characteristic of all these methods is the evident bond between<br />
the spider and her cocoon. Although we cannot know whether the<br />
cocoon appears to her as an object of affection or of value, we are soon<br />
aware of the facts that she resists any attempts to remove it, that if it is<br />
taken away by force her behaviour is conspicuously changed, and that<br />
if it is offered to her again she at once accepts and replaces it, reacting<br />
not to its appearance but to its touch. Investigators have shown that a<br />
deprived spider will accept the cocoon of another spider or any one of a<br />
wide variety of substitutes, such as a pith ball or a pellet of cotton wool<br />
or blotting paper; and in nature such spiders have been found carrying<br />
small shells.<br />
The memory of a bereaved spider for her cocoon varies from a few<br />
hours to a few days, after ·which she will accept neither her own cocoon<br />
nor any substitute. It is believed that her behaviour is dependent on the<br />
state of her ovaries; if the arrival of a new cocoon is near, her memory<br />
is short, although she has carried it for longer than if the arrival of the<br />
new cocoon was in the distant future.<br />
No male arachnid ever shows any parental responsibility and no male<br />
wolf spider has ever been recorded carrying a cocoon.<br />
The time of courtship is but one of the occasions when <strong>Arachnida</strong><br />
are in communication with one another, and it is more easily recognized<br />
by ourselves because the movements involved are unusual and conspicuous.<br />
The study of intercommunication in a wider sense is receiving<br />
closer attention, and Rovner ( 1967b) has found that <strong>Arachnida</strong> can<br />
affect one another by drumming or scraping on the ground, by a display<br />
of colours, and by particular waving of the legs or pal pi. These are<br />
additional to the acts of stridulation, found among scorpions, spiders<br />
and in other orders.<br />
Several of the sounds made by spiders have been recorded, and playback<br />
experiments have proved that spiders can hear air-borne sounds.<br />
8<br />
Ethology: Behaviour<br />
The descriptions of the daily habits of <strong>Arachnida</strong> given in the last<br />
chapter form an example of >vhat may be called the simple or natural<br />
history method of studying animal behaviour. They attempt to provide<br />
answers to the question: how do these animals behave? They thus<br />
supply material on which to base the more valuable discussions which<br />
attempt to answer the more difficult question: what makes these<br />
animals behave as they do? To do this the different types of activity<br />
must be analysed, classified, and their characteristics defined.<br />
The simplest complete reaction of the nervous system of an animal<br />
is the form of response known as areflex-"aneuro-muscularadjustment<br />
due to the inherited mechanism of the nervous system". The familiar<br />
examples of reflex action, such as the blinking of an eyelid, are often<br />
described with an emphasis on the speed with which the whole system<br />
carries out its function of appropriate response. Thus there is a tendency<br />
to neglect the fact that many important reflexes have to persist for<br />
hours at a time, an uninterrupted series of unit actions. Important<br />
examples of this are the reflexes of posture or of habitual attitude, the<br />
maintenance of which is their chief function. These are called tonic<br />
reflexes. The majority of reflex actions are quickly fatigued and quickly<br />
recover, but the tonic reflexes of posture do not tire in this way. On the<br />
other hand, they are the most easily dispossessed or over-ridden of all,<br />
as if they paid for the continuous use of the nerve-paths by an exaggerated<br />
readiness to make way for other traffic, and in this we can perceive<br />
their nature and function. They form a perpetual substratum of<br />
nervous activity, maintaining the body in a state of dynamic equilibrium.<br />
It is clearly important that this state of equilibrium shall be<br />
upset easily, so that there shall arise that agility of response to the<br />
changing circumstances of life which produces efficiency and successful<br />
activity in the living animal.<br />
A very large proportion of the life of an arachnid is spent in waiting,<br />
and during these long periods of inactivity the tonic reflexes are in sole