Animal Behaviour

by C. Lloyd Morgan

From the nature of the rock surfaces the removal of a limpet to a distance of two feet almost invariably involved placing them on the further side of an angle. And though some returned over such an angle, the majority did not.

In most cases the individuals which failed to return to their respective scars took up new positions; and in several instances, when they were subsequently removed to a distance of a few inches from this new position, they returned to it. Their return to the scar was watched in many cases, and the course was fairly, but not quite direct. One limpet covered a distance of ten inches, over a somewhat curved course, in a little under twenty minutes. In another case the limpet on its return journey had to pass between two others, which necessitated the lifting of the shell to some height so as to clear one of them. On reaching the scar they twist and turn about so as to fit down in the normal position which is constant. When they come up the wrong way round they rotate pretty rapidly through the 180 degrees to get into position. One was observed to make a short excursion from and return to its scar under stillish water. But as a rule they seem to remain fixed when they are submerged, moving for the most part when the tide has just receded.

The greatest distance I have watched a limpet reach from its home was twenty-two inches. But I have found them at a distance of three feet from their scars—that is to say, from those to which they fitted perfectly. This was on a large flat surface.

When they move, the tentacles are projected out beyond the shell, and keep on touching and slightly adhering to the rock. On reaching the scar they carefully feel round it with the tentacles. By excision of these feelers Professor Davis was led to conclude that it is not through their instrumentality that the limpet finds its way back to its particular scar. But I am inclined to question these results. At any rate, further observations and experiments are needed to settle the point.

Snails will also return to special dark hollows or crannies in the wall after their foraging excursions. Such behaviour in molluscs affords evidence of something more than instinct. In popular speech, we should say that there is memory of the locality. And in any case it is difficult to interpret the facts without the assumption that the animals are conscious, and that re-presentative states are evoked through the mediation of presentative sense-impressions. But how difficult, if not impossible, it is to form anything like a satisfactory conception of the rudimentary mental processes of a limpet!

The most highly developed molluscs are the cephalopods. They have long sensitive mobile arms with which they feel for and capture their prey. “Now Schneider observed,” writes Dr. Stout,[68] “a very young octopus seize a hermit-crab. The hermit-crab covers the shell in which it takes up its abode with stinging zoophytes. Stung by these, the octopus immediately recoiled and let its prey escape. Subsequently it was observed to avoid hermit-crabs. Older animals of the same species managed cleverly to pull the crab out of its house without being stung.” Such cases afford evidence of profiting by experience through the exercise of intelligence.

Darwin’s careful observations on the manner in which earthworms drag leaves into their burrows seem to show that these annelids act intelligently, and deal with leaves of different shapes in different ways. The leaves of Pine trees, consisting of two needles arising from a common base, were almost invariably drawn down by seizing this basal point of junction; while the leaves of the Lime were, in 79 per cent. of the cases examined, drawn down by the apex; in only 4 per cent. by the base; and in the remaining 17 per cent. by seizing some intermediate portion. On the other hand, the leaves of the Rhododendron, in which the basal part of the blade is often narrower than the apical part, were in 66 per cent. of the observations drawn down by the narrower base. Triangles of paper were in the majority of cases seized by the apex. Commenting upon his observations, carried out with great care under experimental conditions, Darwin says,[69] “As worms are not guided by special instincts in each particular case, though possessing a general instinct to plug up their burrows, and as chance is excluded, the next most probable conclusion seems to be that they try in different ways to draw in objects, and at last succeed in some one way;” that is to say, they profit by experience based on the method of trial and failure. But Darwin adds that the evidence he obtained shows “that worms do not habitually try to draw objects into their burrows in many different ways.” And he seems to attribute to them an almost rational power of dealing with the circumstances in the light of general conceptions. “If worms,” he says, “are able to judge, either before drawing or after having drawn an object close to the mouths of their burrows, how best to drag it in, they must acquire some notion of its general shape. This they probably acquire by touching it in many places with the anterior extremity of their bodies, which acts as a tactual organ. It may be well to remember how perfect the sense of touch becomes in a man when born blind and deaf, as are worms. If worms have the power of acquiring some notion, however rude, of the shape of an object and of their burrows, as seems to be the case, they deserve to be called intelligent; for they then act in nearly the same manner as would a man under similar circumstances.”

Such power of perceiving the relation of the shape of a leaf or other object to the form of the burrow is presumably beyond the reach of an earthworm. It may be regarded as more probable that the earthworm inherits an instinctive tendency to draw down objects in special ways, and that this is subject to some modification under the play of experience, without the formation of anything so psychologically complex as a general notion, however rude. In any case the behaviour of earthworms in closing their burrows seems to afford indications of something more than instinct—of that profiting by the results of experience which characterizes intelligent procedure. More than this we cannot say.

Professor Whitman[70] has made some interesting observations on the leech Clepsine. “Place the animal,” he says, “in a shallow, flat-bottomed dish, and leave it for a few hours or a day, in order to give it time to get accustomed to the place, and come to rest on the bottom. Then, taking the utmost care not to jar the dish or breathe upon the surface of the water, look at the Clepsine through a low magnifying lens, and see what happens when the surface of the water is touched with the point of a needle held vertically above the animal’s back. If the experiment is properly carried out, it will be seen that the respiratory undulations (if such movements happen to be going on) suddenly cease, and that the animal slightly expands its body and hugs the glass. Wait a few moments until the animal, recovering its normal composure, again resumes its respiratory movements. Then let the needle descend through the water until the point rests on the bottom of the dish at a little distance from the edge of the body. Again the movements will cease, and the animal will hug the glass with its body somewhat expanded. Now push the needle slowly along towards the leech, and notice as the needle comes almost in contact with the thin margin of the body, that the part nearest the needle begins to retreat slowly before it. This behaviour shows a surprising keenness of tactile sensibility, the least touch of the water with a needle-point being felt at once.... If its back were rubbed with a brush or the handle of a dissecting needle, in order to test its sensitiveness to touch, the appearance would probably be that of insensibility and indifference to the treatment. Closer examination, however, would show that the flesh of the animal was more rigid than usual, and that the surface was covered with numerous stiff, conical elevations, the dermal papillæ or warts, which are so low and blunt in the normal state of rest as to be scarcely visible. It would be seen that the animal, although motionless, was in a state of active resistance to attack.... Clepsine has another and entirely different method of keeping quiet. The animal rolls itself up (head first and ventral side innermost) into a hard ball, outwardly passive, free to roll or fall whithersoever gravity or currents of water may direct it.... If by chance the animal has eggs, it will not desert them to escape in this way.... This species, then, has two quite distinct and peculiar ways of keeping quiet, and thus avoiding its enemies. If the animal has no eggs, or if it has young, it may adopt either mode of escape, while if it has eggs it has no choice but to remain quiet over them.... The act of rolling up into a passive ball may be performed (a) under compulsion, as when it is her last resort in self-defence; (b) under a milder provocation, as one of three courses of behaviour, as when the resting-place is turned up to light, and the choice is offered between remaining quiet in place, creeping away at leisure, or rolling into a ball and dropping to the bottom; (c) or finally, under no special external stimulus, but rather from internal motive, the normal demand for rest and shady seclusion, presumably very strong in Clepsine after gorging itself with the blood of its turtle host.”