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82 II. DE ARACH::-.;IDIS<br />
declined to zero just before dawn. The content of 5-HT showed the<br />
same peak at 2.00 a.m. The distinctive feature, which agrees with that<br />
found in other animals, is that the occurrence, rather than the duration<br />
of activity, is constant.<br />
Spiders and harvestmen are not exceptional among arachnids in this<br />
respect, for a similar rhythm is detectable in most, if not in all the other<br />
orders. As an example, one may quote the observation of \Veygoldt<br />
( 1971) that the Amblypygi in his laboratory showed three peaks of<br />
activity during the day, the first and most pronounced just after sunset,<br />
a second before dawn, and the third and weakest at about noon.<br />
Similarly, Cloudsley-Thompson ( 1973) has described the activity of the<br />
scorpion Butlwtus minax, which rises to two maxima during each day,<br />
by reacting to the change from light to dark and, in daylight, to rising<br />
temperatures.<br />
The activity of many animals is similarly regular, but the mechanism<br />
that controls them is often unknown. The rather abrupt changes that<br />
are to be observed suggest that a series of neurophysiological events<br />
must be involved. In any case the similarities between vertebrate and<br />
invertebrate behaviour, taking the form of response to the internal<br />
environment, is extremely interesting.<br />
9<br />
Zoogeography: Distribution<br />
The distribution on the earth of any group of animals is the result of a<br />
combination of the place of origin, the means of travel and the nature<br />
of the obstacles encountered en route. The zoogeography of <strong>Arachnida</strong><br />
is no less interesting than is that of any other class.<br />
Early considerations suggest that four categories can be recognized.<br />
(i) <strong>Arachnida</strong> which, though confined to the hot tropical or very<br />
warm sub-tropical belt are, however, more or less continuously<br />
represented therein-scorpions, theraphosid spiders, Amblypygi and<br />
Uropygi.<br />
(ii) <strong>Arachnida</strong> which are also tropical or sub-tropical, but which<br />
are sporadically and discontinuously distributed therein-Palpigradi,<br />
Ricinulei, Schizomida and liphistiid spiders.<br />
(iii) <strong>Arachnida</strong> which spread to the limits of the temperate zones<br />
Opiliones, Pseudoscorpiones, Acari and most spiders.<br />
(iv) <strong>Arachnida</strong> which do, in fact, spread into the polar regionscertain<br />
spiders, Opiliones and mites.<br />
These may be taken in turn.<br />
It is, in fact, very obvious that in the tropics the <strong>Arachnida</strong> are<br />
represented by some of their largest members. The scorpions, Solifugae<br />
and theraphosid spiders are groups which average more than 2 cm in<br />
body length, a great contrast to all the other orders, in which the body<br />
length averages less than a centimetre. This generalization is in striking<br />
contrast with that usually known as Alien's rule, which states that in<br />
any one group the polar members are larger than the tropical ones.<br />
This rule, however, was based on observations on warm-blooded<br />
mammals. The ratio area : volume is necessarily smaller for a large<br />
animal than for a small one, so that the former loses heat less rapidly<br />
than the latter. It is clear that for cold-blooded animals like <strong>Arachnida</strong><br />
the rule cannot be accepted.<br />
The most remarkable of the tropical <strong>Arachnida</strong> are those that are<br />
able to survive in the hostile conditions of the deserts that cover so