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62 11. DE ARACHNIDIS<br />
of the exoskeleton prevents expansion of the internal fluids, so that<br />
vessels and ducts are fatally closed by pressure. This is normally<br />
avoided by the adoption of a cryptozoic life, so that the animal, under<br />
stones, leaves or logs, is always in the shade and is active only at night.<br />
Scorpions, however, have been shown by Alexander ( 1958) to meet<br />
a rise of temperature by a remarkable habit which she has described<br />
as stilting. The scorpion straightens its legs, so that its body is raised<br />
above the ground and free circulation of air around it is made possible.<br />
Experiment made it clear that heat is the stimulus for this action and<br />
that control of the body temperature does, in fact, follow it.<br />
A widespread method of protection and of escape from predatory<br />
enemies, which is common among both <strong>Arachnida</strong> and Crustacea, is<br />
the quick dropping of a captured limb. This permits the animal to<br />
escape, purchasing life and freedom at the temporary expense of a<br />
limb. The habit was closely studied by Wood ( 1926), who divided the<br />
circumstances in which it occurs into the following.<br />
(i) Autotomy is the reflex self-mutilation or automatic severance of a<br />
limb from the body. This does not exist in the <strong>Arachnida</strong>, but is found<br />
among Crustacea.<br />
(ii) Autospasy is the casting of a limb when pulled by some outside<br />
agent, like the forceps of an investigator.<br />
(iii) Autotilly is the pulling off of a limb by the animal itself, as<br />
when an injured leg is seized in the chelicerae and severed from the<br />
body.<br />
(iv) Autophagy is the act of eating a part of the body after severance<br />
from the rest.<br />
In addition to these a fifth term, autosalizy, has been coined by<br />
Pieron ( 1924) with much the same meaning as autotilly, but apparently<br />
involving an element of choice or decision on the part of the animal.<br />
According to most views of arachnid behaviour, this phenomenon could<br />
not be exhibited by them. It is impossible to imagine that the arachnid,<br />
as it pulls on its imprisoned leg, shall come to a decision to run away on<br />
seven legs rather than perish.<br />
Wood's experiments showed conclusively that the shedding of a limb<br />
is not the result of a reflex action or of a special mechanism, as had<br />
previously been believed. It is simply the inability of the skeletal and<br />
muscular components of the leg to resist more than a certain force. The<br />
leg, when pulled, parts at its weakest point. Consequently, all spiders<br />
cast their legs at the coxa-trochanter joint, all "Pedipalpi" at the<br />
patella-tibia joint and Opiliones at the trochanter-femur joint. On the<br />
other hand, in the scorpions, king-crabs, and some of the mites there is<br />
no predetermined locus of fracture. The muscles are evenly arranged<br />
throughout the legs and the chitin is well provided with longitudinal<br />
7. BIONOMICS: GENERAL HABITS 63<br />
fibres at all the interarticular membranes. These factors, combined<br />
with a lack of response on stimulation, make autotomy and autospasy<br />
impossible and autotilly very unlikely.<br />
The spider Tidarrenfordum, however, provides a remarkable example<br />
of autotilly. The palpal organ of the mature male is large; it is almost<br />
halfthe size of the spider's small body, and Chamberlin and lvie (1933)<br />
found that while all young specimens of both sexes had two palpi, the<br />
mature and sub-adult males had all lost either the right or the left<br />
palpus. Bonnet ( 1935) subsequently discovered that during the last<br />
instar the sex organs inside the palp cause it to swell to a considerable<br />
size. To the small owner one such organ was an inconvenience, but two<br />
were intolerable, and the spider therefore pulled off, with its chelicerae,<br />
one of its overgrown appendages. It seemed to be a matter of indifference<br />
whether the right or the left was removed, but the self-mutilation<br />
was invariable. After the last ecdysis the remaining palpus was fully<br />
developed and functionaL It is clear that this autotilly is habitual in this<br />
species and the loss of one palp is not the result of an accident.<br />
A method of defence that is interesting because it is unusual is<br />
adopted by several species of the spider family Theraphosidae. Specialized<br />
"hairs" of four different kinds grow on the dorsal of the<br />
abdomen and from here they can be scraped off in numbers by the<br />
metatarsal spines of the spider's fourth legs. Under threat a shower of<br />
these hairs is directed towards the predator. They produce intense<br />
irritation of the human skin, and seem to be used chiefly against small<br />
mammals (Cooke, 1974).<br />
Among the Uropygi defence by secretion of a nauseous vapour is<br />
more fully developed than among the familiar Opiliones. Glands at the<br />
base of the flagellum can produce a spray of acidic droplets, which,<br />
owing to the mobility of the abdomen can be directed over a wide<br />
range. The method is said to be most against birds and<br />
mammals.<br />
An arachnid's constant occupation, like that of every other animal,<br />
is to provide itself with food.<br />
The average arachnid, if there is such a thing, is a predator, and the<br />
picture that rises in the mind as we think of its feeding is one of a<br />
nocturnal wanderer, which does not so much seek its prey as come upon<br />
it by chance, detecting it by the sense of touch and seizing it forthwith.<br />
No doubt this is broadly accurate for many and there is little in such<br />
casual collecting that is worth attention. It seems reasonable to regard<br />
it as the primitive method of obtaining food, to be contrasted with more<br />
specialized methods evolved later. The most obvious of these improvements<br />
is perhaps the acquisition of greater speed and strength, or, as in<br />
spiders, the highly specialized and unique method of web spinning.