Although our knowledge of the history of the development of the individual species is still so fragmentary, we may conclude with certainty that the development of the markings has been uniform throughout—that it has proceeded in the same manner in all species. All the species appear to be making for the same goal, and the question thus arises whether there may not be an innate force urging their phyletic development. The rigorous examination of this conception must be reserved for a later section. Here, as we are only occupied essentially in establishing facts, it must be remarked that retrogression has never been observed. The young larval forms of a species never show the markings of a later phyletic stage than the older larval forms; the development takes the same course in all species, only making a greater advance in the same direction in some than in others.
Thus, Nicæa and Euphorbiæ have advanced to the seventh phyletic stage, Zygophylli and Hippophaës only to the third, and some specimens of Zygophylli to the fourth. But at whatever phyletic stage the ontogeny of a species may terminate, the young larval stages always display the older phyletic stages. Thus, Galii in its last ontogenetic stage reaches the sixth phyletic stage; in its penultimate stage it reaches the fifth phyletic stage; and in its third stage; the fourth phyletic stage is represented, so that little imagination is required to anticipate that in the second stage the third or second phyletic stage would be pictured.
If we tabulate the development of the various species, indicating the ontogenetic stages by Arabic numerals, and the stages of the phylogeny which are reached in each stage of the ontogeny by Roman numerals, we obtain a useful synopsis of the series of developments, and, at the same time, it shows how many gaps still remain to be filled up in order to complete our knowledge even of this small group of species.
Table of Development of the Species of Deilephila.
| Deilephila. | Ontogeny Stage 1. | Ontogeny Stage 2. | Ontogeny Stage 3. | Ontogeny Stage 4. | Ontogeny Stage 5. |
| 1. Hippophaës | ? | ? | ? | III. | III.-IV. |
| 2. Zygophylli | ? | ? | ? | ? | III.-IV. |
| 3. Lineata | ? | ? | ? | ? | IV. |
| 4. Livornica | ? | ? | ? | IV. | V. |
| 5. Galii | ? | ? | IV. | V. | VI. |
| 6. Vespertilio | ? | II. (?) | IV. | V. | VI. |
| 7. Mauritanica | ? | ? | ? | ? | VI. |
| 8. Dahlii | I. | ? | VI. | VII. | VII. |
| 9. Euphorbiæ | I. | V. | VI. | VII. | VII. |
| 10. Nicæa | ? | ? | VII. | VII. | VII. |
From this very incomplete table we perceive that, in certain instances, the stages can be represented as a continuous series of phyletic steps, as in the case of D. Galii; that in others certain steps may be omitted, as with D. Euphorbiæ, in which grade I. of stage 1 is immediately followed by grade V. in stage 2. In reality the gap caused by this omission is still greater than would appear, as grade V. is only indicated, and not actually reached, the subdorsal not being present as a sharply-defined line, but only as a faint stripe. The suppression of phyletic steps increases with the advancement in phyletic development. The higher the step to which a species finally attains, the greater is the tendency of the initial stages to be compressed, or omitted altogether.
From what has thus far been seen with respect to the development of D. Hippophaës, there may be drawn what to me appears to be a very important conclusion, viz. that the ring-spots of Deilephila first originated on the segment bearing the caudal horn, and were then gradually transferred as secondary spots to the preceding segments. Complete certainty would be given to this conclusion by a knowledge of the young forms of other phyletically retarded species, especially those of the American D. Lineata, and perhaps also those of Zygophylli and Livornica. The few observations on the development of D. Galii already recorded give support to this view, since the absence of ring-spots on the three front segments in the young caterpillar (one instance), or their less perfect formation on these segments (second instance), indicates a forward transference of the spots.
If the foregoing view be accepted, there follows from it a fundamental difference between the development of the genera Chærocampa and Deilephila. In the former the formation of the eye-spots proceeds from a subdorsal line, but they first appear on two of the front segments, and are then transferred to the posterior segments. In Deilephila, on the other hand, a single ring-spot is formed on the penultimate segment bearing the caudal horn, and this is repeated on the anterior segments by secondary transference. With respect to the origination of the ring-spot also, there is a distinction between this genus and Chærocampa, inasmuch as the first step towards the eye-formation in the latter consists in the separation of a curved portion of the subdorsal line, whilst in Deilephila the nuclear spot first seems to originate and the separation of the mirror-spot from the subdorsal line appears to occur secondarily. It is difficult here to draw further conclusions, since the first appearance of the primary ring-spot has not yet been observed, and no more certain inference respecting the history of the formation of the primary ring-spots can be drawn from the manner in which the secondary ring-spots are formed. Because in Hippophaës the formation of the secondary ring-spots begins with the red coloration of one or two shagreen-dots, it does not follow that the primary spot on the eleventh segment also originated in this manner; and this is not without importance when we are concerned with the causes which underlie the formation of ring-spots. In Chærocampa also, the formation of the primary eye-spots appears to differ from that of the secondary—in the latter the black “ground-area” first appearing, and in the former the “mirror-spot.” The secondary eye-spots certainly remain rudimentary in this last genus, so that the evidence in support of this conclusion is thus much weakened; but it must be admitted that we are here on ground still too uncertain to admit of wider conclusions being based thereon.
As a final result of the investigation, we may advance the opinion that the existing species of the genus Deilephila have reached five different phyletic stages, and that their very different external appearance is explained by their different phyletic ages; the appearance from these caterpillars of moths so extremely similar, can otherwise be scarcely understood.
It may appear almost unnecessary to bring forward additional proofs in support of this interpretation of the facts, but in a field where the data are so scanty, no argument which can be drawn from them should be considered as superfluous. The variations which occasionally occur in the larvæ, however, to a certain extent furnish a proof of the correctness of the theoretical interpretation offered.