For the reader who can accurately picture to himself the external structure of a flea and of the typical insects belonging to other orders, a few words may be said on the probable ancestry of fleas and their relationship to other living insects. This vexed and much debated question is still, as the older naturalists would have said, tremendum mysterium. Very little light has yet been thrown upon it, and the most divergent views have been expressed by learned and competent entomologists. A historic survey of the various opinions that have been held since the days of Linnæus would fill many pages; but a short summary of the different orders to which fleas have been referred by different zoologists will suffice.
The older authors, Linnæus, Geoffroy, Cuvier and Duméril, and Gervais placed them among the Aptera because they were wingless. Kircher regarded them as Orthoptera, an order which includes grasshoppers and crickets; but he has had few followers. By Fabricius and by Illiger they were treated as Hemiptera or bugs. Lameere, a Belgian, has recently expressed a decided view that fleas are really a family of Coleoptera or beetles. Those who have held the once orthodox opinion that they belonged to the Diptera or flies are Roesel, Oken, Straus-Dürkheim, Burmeister, Newman, Walker, von Siebold and Wagner.
The structure of an adult flea, however, differs from that of an adult fly in the following noteworthy respects: the mouth-parts are differently constructed, the head of the flea is closely joined to its thorax, the three divisions of the thorax are not joined and fused, the flea is wingless, the eyes of fleas are simple ocelli, and there are differences of lesser importance in the stigmata, which give access to the tracheal system by which all insects breathe.
The number of those who have regarded fleas as belonging to a distinct order of insects is considerable: they are Lamarck, De Geer, Latreille, Kirby and Spence, MacLeay, Leach, Dugès, Bouché, van der Hoeven, Westwood, Landois, Brauer, Kraepelin, and Taschenberg. Modern opinion is all but unanimous on this point.
There remains, however, a second question. Even if it be agreed that there must be a distinct order for Suctoria, Aphaniptera, Siphonaptera, or fleas; where ought that order to be placed? In which other order of insects must we look for the nearest relations of fleas? For a time after the acceptance of the fact that insect forms have been evolved, and not separately created, the ancestors of fleas were searched for among some species of fly.
Then Kraepelin rejected the view that flies were as closely related to fleas as most entomologists thought and his followers could only find points of difference and no points of resemblance. Dahl (1899), a German, then took up the cudgels for the fly theory. Dahl pointed out the resemblance between fleas and a group of flies called Phoridæ also parasitic on warm-blooded animals. During the ensuing years the debate was resumed afresh with much liveliness and sometimes with a little acrimony.
The fleas were placed by MacLeay and by Balbiani between the Diptera and Hemiptera; by Leach between the Hemiptera and Lepidoptera; by Dugès between the Hymenoptera and Diptera; by Brauer between the Diptera and Coleoptera. Handlirsch thinks that fleas have no connection at all with beetles and Gross can find no signs of relationship with either Coleoptera or Diptera.
Embryology and the study of larval forms have thrown so much light on the ancestry of many animals, that it was hoped that a microscopic examination of the larvæ of fleas, in various stages of development, would produce some facts of importance. In this hope entomologists have, to a great extent, been disappointed. There seems to be much similarity between the embryos of beetles, moths, flies, wasps and fleas. Those who have dwelt on the likeness of the larval flea to the maggot of a fly seem to forget that the resemblance to an embryo beetle is nearly as strong.
The young larva of the flea is very transparent and the digestive canal, heart and nervous system are easily recognised. The egg-shell breaker is an interesting example of the development of a temporary larval structure and it is the only known instance of such a structure in an insect. There are no traces of eyes. The antennæ are three-jointed. They are rather long and slender, being about one-third as long as the head. The head is well-developed and the larva has no feet.