Romanes[31] thought that the manner of stinging and paralyzing their prey might “be justly deemed the most remarkable instinct in the world.” Spiders, insects, and caterpillars are stung, he says, “in their chief nerve-centres, in consequence of which the victims are not killed outright, but rendered motionless; they are then conveyed to a burrow and, continuing to live in their paralyzed condition for several weeks, are then available as food for the larvæ when these are hatched. Of course the extraordinary fact which stands to be explained is that of the precise anatomical, not to say also physiological knowledge which appears to be displayed by the insect in stinging only the nerve-centres of its prey.” Eimer[32] thought that it “is absolutely impossible that the animal has arrived at its habit otherwise than by reflection upon the facts of experience.” “At the beginning,” he says, “she probably killed larvæ by stinging them anywhere, and then placed them in the cell. The bad results of this showed themselves; the larvæ putrified before they could serve as food for the larval wasps. In the mean time the mother wasp discovered that those larvæ which she had stung in particular parts of the body were motionless but still alive, and then she concluded that larvæ stung in this particular way could be kept for a longer time unchanged as living motionless food.”
Now, since these wasps, when they have stored their nests and laid an egg on one of the victims, close it up once and for all, and take no further interest in it or its contents, there seems no opportunity, at any rate in the existing state of matters, for the acquisition of that experience on which Eimer relied. But both his explanation and Romanes’s difficulty are based on the following assumptions: first, that the victims are instinctively or habitually stung in the chief nerve-centres; secondly, that when thus stung they are not killed but remain paralyzed for weeks; and thirdly, that the marvellously definite and delicate instinctive behaviour is in direct relation to the uniform result of prolonged paralysis and consequent preservation of the food in the fresh state. But Dr. Peckham’s careful observations and experiments show that, with the American wasps, the victims stored in the nests are quite as often dead as alive; that those which are only paralyzed live for a varying number of days, some more, some less; that wasp larvæ thrive just as well on dead victims, sometimes dried-up, sometimes undergoing decomposition, as on living and paralyzed prey; that the nerve-centres are not stung with the supposed uniformity; and that in some cases paralysis, in others death, follows when the victims are stung in parts far removed from any nerve-centre. “We believe,” he says, “that the primary purpose of the stinging is to overcome resistance, and to prevent the escape of the victims, and that incidentally some of them are killed and others are paralyzed.”
If, therefore, as will probably be shown to be the case, these conclusions are found to be generally true for this interesting group of insects, the mystery of “the precise anatomical, not to say also physiological knowledge which appears to be displayed” by these wasps turns out to be one of our own fabrication. It melts away in the light of fuller and more searching investigation.
Fig. 11.—Solitary Wasp stinging Caterpillar (after Peckham).
It must not be supposed, however, from what has been said, that the behaviour in the act of stinging is altogether indefinite. On the contrary, each species proceeds in a relatively definite manner with some variation or modification of method. Philanthus punctatus, for example, stings the bees, on which she preys, under the neck, and the thrust is at once fatal. Dr. Peckham further notes that he was only successful in getting the wasps to sting when they were hunting; those that had not yet begun to store the nests paid no attention to the bees. This is an example of that internal factor to which reference was made in the last section. Marchal observed that Cerceris ornata runs the end of her abdomen along the under surface of the thorax of the bee, and delivers her thrust at the division of the segments—that is, where the sting can enter. The action does not imply any physiological knowledge. In general she begins at the neck. Spiders are usually, but not always, stung on the ventral surface. To give but one more example, Dr. Peckham observed in three cases the procedure of Ammophila urnaria which preys on caterpillars, and often, after stinging, bites the neck in several places, this process being termed malaxation. In three observed captures, all the caterpillars being of the same species and alike in size, the thrusts were given on the ventral surface near the middle line, between the segments. In the first, seven stings were given at the extremities (there being thirteen segments), the middle segments being left untouched, and no malaxation was practised. In the second, seven stings were again given, but in the anterior and middle segments, followed by slight malaxation. In both these cases the first three thrusts were in definite order, behind the third, the second, and the first segments successively. In the case of the third caterpillar, only one thrust was given, between the third and fourth segments—that is to say, in the position of the first stab in the other cases,—and after this one thrust there was prolonged malaxation. Of fifteen stored caterpillars examined, some lived only three days, others a little longer, while a few showed signs of life at the end of a fortnight. In more than one instance the second of the two caterpillars stored in each nest died and became discoloured before the first one was entirely eaten. The larva under such circumstances ate it with good appetite, and then spun its cocoon as if nothing unpleasant had occurred.
Fig. 12.—Solitary Wasp dragging a Caterpillar to its Nest (after Peckham).