Not only by their structure and embryology, as well as their metamorphosis, do the myriopods and insects stand apart from the Arachnida and other arthropods, but it seems probable that they have had a different ancestry, the arthropods being apparently polyphyletic.
There are two animals which appear to connect the insects with the worms, and which indicate a separate line of descent from the worms independent of that of the other classes. These are the singular Peripatus, which serves as a connecting link between arthropods and worms, and Scolopendrella (Symphyla). These two animals are guide-posts, pointing out, though vaguely to be sure, the way probably trod by the forms, now extinct, which led up to the insects.
Relations of Peripatus to Insects.—We will first recount the characteristics of this monotypic class. Peripatus (Fig. 4) stands alone, with no forms intermediate between itself and the worms on the one hand, and the true Arthropoda on the other. Originally supposed to be a worm, it is now referred to a class by itself, the Malacopoda of Blainville, or Protracheata of Haeckel. It lives in the tropics, in damp places under decaying wood. In general appearance it somewhat resembles a caterpillar, but the head is soft and worm-like, though it bears a pair of antenna-like tentacles. It may be said rather to superficially resemble a leech with clawed legs, the skin and its wrinkles being like those of a leech. There is a pair of horny jaws in the mouth, but these are more like the pharyngeal teeth of worms than the jaws of arthropods. The numerous legs end each in a pair of claws. The ladder-like nervous system is unlike that of annelid worms or arthropods, but rather recalls that of certain molluscs (Chiton, etc.), as well as that of certain flat and nemertine worms. Its annelid features are the large number of segmentally arranged true nephridia, and the nature of the integument. Its arthropodan features, which appear to take it out of the group of worms, are the presence of tracheæ, of true salivary and slime glands, of a pair of coxal glands (Fig. 4, C, cd) as well as the claws at the end of the legs. The tracheæ, which are by no means the only arthropodan features, are evidently modified dermal glands. The heart is arthropodan, being a dorsal tube lying in a pericardial sinus, with many openings. This assemblage of characters is not to be found in any marine or terrestrial worm.
The tracheæ (Fig. 4, D, tr) are unbranched fine tubes, without a “spiral thread,” and are arranged in tufts, in P. edwardsii opening by simple orifices or pores (“stigmata”) scattered irregularly over the surface of the body; but in another species (P. capensis) some of the stigmata are arranged more definitely in longitudinal rows,—on each side two, one dorsally and one ventrally. “The stigmata in a longitudinal row are, however, more numerous than the pairs of legs.” (Lang.)
The salivary glands, opening by a short common duct into the under side of the mouth, in the same general position as in insects, are evidently, as the embryology of the animal proves, transformed nephridia, and being of the arthropodan type explain the origin and morphology of those of insects. It is so with the slime glands; these, with the coxal glands, being transformed and very large dermal glands. Those of insects arose in the same manner, and are evidently their homologues, while those of Peripatus were probably originally derived from the setiparous glands in the appendages (parapodia) of annelid worms.
Fig. 4.—A, Peripatus novæ zealandiæ.—After Sedgwick, from Lang. B, Peripatus capensis, side view, enlarged about twice the natural size.—After Moseley, from Balfour. C, Anatomy of Peripatus capensis. The enteric canal behind the pharynx has been removed. g, brain; a, antenna; op, oral or slime papillæ; sd, slime gland; sr, slime reservoir, which at the same time acts as a duct to the gland; so4, so5, so6, so9, nephridia of the 4th, 5th, 6th, and 9th pairs of limbs; cd, elongated coxal gland of the last pair of feet; go, genital aperture; an, anus; ph, pharynx; n, longitudinal trunk of the nervous system.—After Balfour, from Lang. D, Portion of the body of Peripatus capensis opened to show the scattered tufts of tracheæ (tr); v, v, ventral nerve cords.—After Moseley.
The genital glands and ducts are paired, but it is to be observed that the outlets are single and situated at the end of the body. In the male the ejaculatory duct is single; in its base a spermatophore is formed. It will be seen, then, that Peripatus is not only a composite type, and a connecting link between worms and tracheate arthropods, but that it may reasonably be regarded, if not itself the ancestor, as resembling the probable progenitor of myriopods and insects, though of course there is a very wide gap between Peripatus and the other antennate, air-breathing Arthropoda.
Fig. 4.—E, Peripatus edwardsii, head from the under side: a, base of antenna; op, oral papilla; the figure also shows the papillæ around the mouth, and the four jaws.—After Balfour, from Lang. F, Anterior end of Peripatus capensis, ventral side, laid open: a, antenna; z, tongue; k, jaw; sd, salivary gland; gs, union of the two salivary glands; ph, pharynx; œ, œsophagus; l, lip papillæ around the mouth; op, oral or slime papilla; sld, duct or reservoir of the slime gland.—After Balfour, from Lang.