It is certain that in some aerial insects the olfactory sense has dwindled to a minimum, e. g., in those species in which the male recognises and follows the female exclusively by means of the eyes, as in the Odonata (dragon-flies). To insects with such habits an olfactory sense would be almost superfluous. Here, too, the antennæ have dwindled to diminutive dimensions.

But there are insects whose antennæ are immovable and quite unable to touch objects. This is the case in most Diptera (flies). Still these antennæ are often highly developed and present striking dilatations densely beset with olfactory papillæ. By experiment I have demonstrated the existence of an olfactory sense in such Dipteran antennæ, and I have been able to show that, e. g., in Sarcophaga vivipara and other carrion flies, the egg-laying instinct is absolutely dependent on the sensation of the odor of carrion and the presence of the antennæ. In these cases the contact-odor sense is undoubtedly absent. More or less of a topochemical odor-sense at long range must, of course, be present, since the antennæ are external, but the precision of the spatial image must be very imperfect, owing to the immobility of the antennæ. Nevertheless, flies move about so rapidly in the air that they must be able by means of their antennæ to distinguish very quickly the direction from which odors are being wafted. These insects do, in fact, find the concealed source of odors with great assurance. But this is no great art, for even we ourselves are able to do the same by sniffing or going to and fro. But the flies find their way through the air with their eyes and not at all by means of their sense of smell. Hence their olfactory powers probably constitute a closer psychological approximation to those of mammals than to the topochemical odor-sense of ants, for they can hardly furnish any constant and definite space-relations.

Even in many insects with movable antennæ and of less ærial habits, e. g., the chafers and bombycid moths, the antennal olfactory sense is evidently much better adapted to function at a distance, i. e., to the perception of odors from distant objects, than to the perception of space and trails. Such insects find their way by means of their eyes, but fly in the direction whence their antennæ perceive an odor that is being sought.

A genuine topochemical antennal sense is, therefore, probably best developed in all arthropods, whose antennæ are not only movable in the atmosphere, but adapted to feeling of objects. In these cases the still imperfect topochemical odor-sense for distances can be momentarily controlled by the contact-odor-sense and definitively fixed topographically, i. e., topochemically, as we see so extensively practised in the ants.

It would be possible to meet this view with the objection that a contact-odor sense could not accomplish much more than the tactile sense. I have made this objection to myself. But in the first place it is necessary to reckon with the facts. Now it is a fact that insects in touching objects with their antennæ mainly perceive and distinguish the chemical constitution of the objects touched and heed these very much more than they do the mechanical impacts also perceived at the same time. Secondly, the tactile sense gives only resistance and through this, form. On the other hand, the multiplicity of odors is enormous, and it is possible to demonstrate, as I have done for the ants, and Von Buttel-Reepen for the bees, that these animals in distinguishing their different nest-mates and their enemies, betray nothing beyond the perception of extremely delicate and numerous gradations in the qualities of odors.

In combination with topochemical space-perception, these numerous odor-qualities must constitute a spatial sense which is vastly superior to the tactile sense. The whole biology of the social Hymenoptera furnishes the objective proof of this assertion.

It would certainly be well worth while to investigate this matter in other groups of arthropods which possess complex instincts.

In conclusion I will cite an example, which I have myself observed, for the purpose of illustrating the capacity of the topochemical olfactory sense.

The American genus Eciton comprises predatory ants that build temporary nests from which they undertake expeditions for the purpose of preying on all kinds of insects. The Ecitons follow one another in files, like geese, and are very quick to detect new hunting grounds. As “ants of visitation,” like the Africo-Indian species of Dorylus, they often take possession of human dwellings, ferret about in all the crevices of the walls and rooms for spiders, roaches, mice, and even rats, attack and tear to pieces all such vermin in the course of a few hours and then carry the booty home. They can convert a mouse into a clean skeleton. They also attack other ants and plunder their nests.

Now all the workers of the African species of Dorylus and of many of the species of Eciton are totally blind, so that they must orient themselves exclusively by means of their antennal sense.