Studies in the Theory of Descent, Volume II

We may therefore fairly assert that the structure of larvæ is on the whole remarkably uniform, in consequence of the uniformity in the conditions of life. Notwithstanding the great variety of external aspects, the general structure of caterpillars does not become changed – it is only their outward garb which varies, sometimes in one direction, and sometimes in another, and which, starting from inherited characters, becomes adapted to the various special conditions of life in the best possible manner.

All this is quite different in the case of the imagines, where we meet with very important differences in the conditions of life. The butterflies, which live under the influence of direct sunlight and a much higher temperature, and which are on the wing for a much longer period during the day, must evidently be differently equipped to the moths in their motor organs (wings), degree of hairiness, and in the development of their eyes and other organs of sense. It is true that we are not at present in a condition to furnish special proofs that the individual organs of butterflies are exactly adapted to a diurnal life, but we may safely draw this general conclusion from the circumstance that no butterfly is of nocturnal habits.25 It cannot be stated in objection that there are many moths which fly by day. It certainly appears that no great structural change is necessary to confer upon a Lepidopteron organized for nocturnal life the power of also flying by day; but this proves nothing against the view that the structure of the butterflies depends upon adaptation to a diurnal life. Analogous cases are known to occur in many other groups of animals. Thus, the decapodous Crustacea are obviously organized for an aquatic life; but there are some crabs which take long journeys by land. Fish appear no less to be exclusively adapted to live in water; nevertheless the “climbing-perch” (Anabas) can live for hours on land.

It is not the circumstance that some of the moths fly by day which is extraordinary and demands a special explanation, but the reverse fact just mentioned, that no known butterfly flies by night. We may conclude from this that the organization of the latter is not adapted to a nocturnal life.

If we assume26 that the Lepidopterous family adapted to a diurnal life gives rise in the course of time to a nocturnal family, there can be no doubt but that the transformation of structure would be far greater on the part of the imagines than on that of the larvæ. The latter would not remain quite unchanged – not because their imagines had taken to a nocturnal life which for the larva would be quite immaterial, but because this change could only occur very gradually in the course of a large number of generations, and during this long period the conditions of life would necessarily often change with respect to the larvæ. It has been shown above that within the period of time necessary for the formation of a new species impulses to change occur on both sides; how much more numerous therefore must these be in the case of a group of much higher rank, for the establishment of which a considerably longer period is required. In the case assumed, therefore, the larvæ would also change, but they would suffer much smaller transformations than the imagines. Whilst in the latter almost all the typical portions of the body would undergo deep changes in consequence of the entirely different conditions of life, the larvæ would perhaps only change in marking, hairs, bristles, or other external characters, the typical parts experiencing only unimportant modifications.

In this manner it can easily be understood why the larvæ of a family of Noctuæ do not differ to a greater extent from those of a family of butterflies than do the latter from some other Rhopalocerous family, or why the imagines of a Rhopalocerous and a Heterocerous family present much greater form-divergences than their larvæ. At the same time is therefore explained the unequal value that must be attributed to any single family of butterflies in its larvæ and in its imagines. The unequal form-divergences coincide exactly with the inequalities in the conditions of life.

When whole families of butterflies show the same structure in their typical parts (antennæ, wings, &c.), and, what is of more importance, can be separated as a systematic group of a higher order (i. e. as a section or sub-order) from the other Lepidoptera whilst their larval families do not appear to be connected by any common character, the cause of this incongruence lies simply in the circumstance that the imagines live under some peculiar conditions which are common to them all, but which do not recur in other Lepidopterous groups. Their larvæ live in precisely the same manner as those of all the other families of Lepidoptera – they do not differ in their mode of life from those of the Heterocerous families to a greater extent than they do from one another.

We therefore see here a community of form within the same compass as that in which there is community in the conditions of life. In all butterflies such community is found in their diurnal habits, and in accordance with this we find that these only, and not their larvæ, can be formed into a group having common characters.

In the larvæ also we only find agreement in the conditions of life within a much wider compass, viz. within the whole order. Between the limits of the order Lepidoptera the conditions of life in the caterpillars are, as has just been shown, on the whole very uniform, and the structure of the larvæ accordingly agrees almost exactly in all Lepidopterous families in every essential, i. e. typical, part.

In this way is explained the hitherto incomprehensible phenomenon that the sub-ordinal group Rhopalocera cannot be based on the larvæ, but that Lepidopterous caterpillars can as a whole be associated into a higher group (order); they constitute altogether families and an order, but not the intermediate group of a sub-order. By this means we at the same time reply to an objection that may be raised, viz. that larval forms cannot be formed into high systematic groups because of their “low and undeveloped” organization.

To this form of incongruence, viz. to the formation of systematic groups of unequal value and magnitude, I must attach the greatest weight with respect to theoretical considerations. I maintain that this, as I have already briefly indicated above, is wholly incompatible with the admission of a phyletic force. How is it conceivable that such a power could work in the same organism in two entirely different directions – that it should in the same species lead to the constitution of quite different systems for the larvæ and for the imagines, or that it should lead only to the formation of families in the larvæ and to sub-orders in the imagines? If an internal force existed which had a tendency to call into existence certain groups of animal forms of such a nature that these constituted one harmonious whole of which the components bore to one another fixed morphological relationships, it would certainly have been an easy matter for such a power to have given to the larvæ of butterflies some small character which would have distinguished them as such, and which would in some measure have impressed them with the stamp of “Rhopalocera.” Of such a character we find no trace however; on the contrary, everything goes to show that the transformations of the organic world result entirely from external influences.

III. Incongruences in other Orders of Insects

Although the order Lepidoptera is for many reasons especially favourable for an investigation such as that undertaken in the previous section, it will nevertheless be advantageous to inquire into the form-relationships of the two chief stages in some other orders of metamorphic insects, and to investigate whether in these cases the formation of systematic groups also coincides with common conditions of life.

Hymenoptera

In this order there cannot be the least doubt as to the form-relationship of the imagines. The characteristic combination of the pro- and meso-thorax, the number and venation of the wings, and the mouth-organs formed for biting and licking, are found throughout the whole order, and leave no doubt that the Hymenoptera are well based on their imaginal characters.

But it is quite different with the larvæ. It may be boldly asserted that the order would never have been founded if the larvæ only had been known. Two distinct larval types here occur, the one – caterpillar-like – possessing a distinct horny head provided with the typical masticatory organs of insects, and a body having thirteen segments, to which, in addition to a variable number of abdominal legs, there are always attached three pairs of horny thoracic legs: the other type is maggot-shaped, without the horny head, and is entirely destitute of mouth-organs, or at least of the three pairs of typical insect jaws, and is also without abdominal and thoracic legs. The number of segments is extremely variable; the larvæ of the saw-flies have thirteen besides the head, the maggot-shaped larvæ of bees possess fourteen segments altogether, and the gall-flies and ichneumons only twelve or ten. We should be much mistaken also if we expected to find connecting characters in the internal organs. The intestine is quite different in the two types of larvæ, the posterior opening being absent in the maggot-like grubs; at most only the tracheal and nervous systems show a certain agreement, but this is not complete.

The order Hymenoptera, precisely speaking and conceived only morphologically, exists therefore but in the imagines; in the larvæ there exist only the caterpillar- and maggot-formed groups. The former shows a great resemblance to Lepidopterous larvæ, and in the absence of all knowledge of the further development it might be attempted to unite them with these into one group. The two certainly differ in certain details of structure in the mouth-organs and in the number of segments, abdominal legs, &c., to a sufficient extent to warrant their being considered as two sub-orders of one larval order; but they would in any case be regarded as much more nearly related in form than the caterpillar- and maggot-like types of the Hymenopterous larvæ.

Is it not conceivable, however, that the imagines of the Hymenoptera – that ichneumons and wasps may be only accidentally alike, and that they have in fact arisen from quite distinct ancestral forms, the one having proceeded with the Lepidopterous caterpillars from one root, and the other with the grub-like Dipterous larvæ from another root?

This is certainly not the case; the common characters are too deep-seated to allow the supposition that the resemblance is here only superficial. From the structure of the imagines alone the common origin of all the Hymenoptera may be inferred with great probability. This would be raised into a certainty if we could demonstrate the phyletic development of the maggot-formed out of the caterpillar-formed Hymenopterous larvæ by means of the ontogeny of the former. From the beautiful investigations of Bütschli on the embryonic development of bees27 we know that the embryo of the grub possesses a complete head, consisting of four segments and provided with the three typical pairs of jaws. These head segments do not subsequently become formed into a true horny head, but shrivel up; whilst the jaws disappear with the exception of the first pair, which are retained in the form of soft processes with small horny points. We know also that from the three foremost segments of the embryo the three typical pairs of legs are developed in the form of round buds, just as they first appear in all insects.28 These rudimentary limbs undergo complete degeneration before the birth of the larva, as also do those of the whole29 of the remaining segments, which, even in this primitive condition, show a small difference to the three foremost rudimentary legs.

The grub-like larvæ of the Hymenoptera have therefore descended from forms which possessed a horny head with antennæ and three pairs of gnathites and a 13-segmented body, of which the three foremost segments were provided with legs differing somewhat from those of the other segments; that is to say, they have descended from larvæ which possessed a structure generally similar to that of the existing saw-fly larvæ. The common derivation of all the Hymenoptera from one source is thus established with certainty.30

But upon what does this great inequality in the form-relationship of the larvæ and imagines depend? The existing maggot-like grubs are without doubt much further removed from the active caterpillar-like larvæ than are the saw-flies from the Aculeate Hymenoptera. Whilst these two groups differ only through various modifications of the typical parts (limbs, &c.), their larvæ are separable by much deeper-seated distinctions; limbs of typical importance entirely vanish in the one group, but in the other attain to complete development.

In the Hymenoptera there exists therefore a very considerable incongruence in the systems based morphologically, i. e. on the pure form-relationships of the larvæ and of the imagines. The reason of this is not difficult to find: the conditions of life differ much less in the case of the imagines than in that of the larvæ. In the former the conditions of life are similar in their broad features. Hymenoptera live chiefly in the air and fly by day, and in their mode of obtaining food do not present any considerable differences. Their larvæ, on the other hand, live under almost diametrically opposite conditions. Those of the saw-flies live after the manner of caterpillars upon or in plants, in both cases their peculiar locomotion being adapted for the acquisition and their masticatory organs for the reduction of food. The larvæ of the other Hymenoptera, however, do not as a rule require any means of locomotion for reaching nor any organs of mastication for swallowing their food, since they are fed in cells, like the bees and wasps, or grow up in plant galls of which they suck the juice, or are parasitic on other insects by whose blood they are nourished. We can readily comprehend that in the whole of this last group the legs should disappear, that the jaws should likewise vanish or should become diminished to one pair retained in a much reduced condition, that the horny casing of the head, the surface of attachment of the muscles of the jaws, should consequently be lost, and that even the segments of the head itself should become more or less shrivelled up as the organs of sense therein located became suppressed.

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1

[The slight variability in the colour of this pupa, opens up the interesting question of the photographic sensitiveness of this and other species, which is stated to cause them to assimilate in colour to the surface on which the larva undergoes its final ecdysis. Some experiments upon this subject have been recorded by Mr. T. W. Wood, Proc. Ent. Soc. 1867, p. xcix, but the field is still almost unexplored. R.M.]

2

“Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872, p. 20.

3

In some instances Deilephila Lineata has also been seen by day hovering over flowers.

4

It is true that I only reared one brood, but from this fifty specimens were obtained. It would be interesting to know whether this variety of the caterpillar is distributed over the whole of Southern Europe.

5

In this sense Lubbock says: – “It is evident that creatures which, like the majority of insects, live during the successive periods of their existence in very different circumstances, may undergo considerable changes in their larval organization in consequence of forces acting on them while in that condition; not, indeed, without affecting, but certainly without affecting to any corresponding extent, their ultimate form.” – “Origin and Metamorphoses of Insects,” London, 1874, p. 39.

6

“Grundzüge der Zoologie,” 1875.

7

[Lepidopterists are of course aware that even these distinctions are not absolute, as no single character can be named which does not also appear in certain moths. The definition in this case, as in that of most other groups of animals and plants, is only a general one. See, for instance, Westwood’s “Introduction to the Classification of Insects,” vol. ii. pp. 330–332. Also some remarks by C. V. Riley in his “Eighth Annual Report” on the insects of Missouri, 1876, p. 170. With reference to the antennæ as a distinguishing character, see Mr. A. G. Butler’s article in “Science for All,” 1880, part xxvii. p. 65. R.M.]

8

The genus of Morphinæ, Discophora, possesses hairs very similar to those of the genus Cnethocampa belonging to the Bombycidæ.

9

[The larvæ of genera 14, Phyciodes, and 35, Crenis, are likewise spiny. See Edwards’ “Butt. of N. Amer.” vol. ii. for figures of the caterpillar of Phyc. Tharos: for notes on the larvæ of Crenis Natalensis and C. Boisduvali see a paper by W. D. Gooch, “Entomologist,” vol. xiv. p. 36. The larvæ of genus 55, Ageronia, are also spiny. (See Burmeister’s figure of A. Arethusa, “Lép. Rép. Arg.” Pl. V. Fig. 4). The larvæ of genus 98, Aganisthos, also appear to be somewhat spiny (see Burmeister’s figure of A. Orion, loc. cit. Pl. V. Fig. 6), and this raises the question as to whether the genus is correctly located in its present position. The larvæ of the following genera figured in Moore’s “Lepidoptera of Ceylon,” parts i. and ii., are all spiny: – 6, Cirrochroa (Pl. XXXII.); 7, Cynthia (Pl. XXVI.); 27, Kallima (Pl. XIX.); and 74, Parthenos (Pl. XXIV.). Many species of caterpillars which are spiny when adult appear to be spineless, or only slightly hairy when young. See Edwards’ figures of Melitæa Phaeton, Argynnis Diana, and Phyc. Tharos (loc. cit.) and his description of the larva of Arg. Cybele, “Canad. Entom.” vol. xii. p. 141. The spiny covering thus appears to be a character acquired at a comparatively recent period in the phyletic development. R.M.]

10

[The larvæ of the 110th genus, Paphia, Fabr. (Anæa, Hübn.) are also smoothed-skinned. See Edwards’ figure (loc. cit. vol. i. Pl. XLVI.) of P. Glycerium. Also C. V. Riley’s “Second Annual Report” on the insects of Missouri, 1870, p. 125. Burmeister figures the larva of a species of Prepona (genus 99) which is smooth (P. Demophon, loc. cit. Pl. V. Fig. 1). The horns on the head of Apatura, &c., may possibly be a survival from a former spiny condition. R.M.]

11

“Synopsis of the described Lepidoptera of North America.” Washington, 1862.

12

“Catalog der Lepidopteren des Europäischen Faunengebietes.” Dresden, 1871.

13

This group of moths (“Schwärmer”) is regarded as of very different extents by systematists; when I here comprise under it only the Sphingidæ proper and the Sesiidæ, I by no means ignore the grounds which favour a greater extension of the group; the latter is not rigidly limited. [The affinities of the Sesiidæ (Ægeriidæ) are by no means clearly made out: it appears probable that they are not related to the Sphingidæ. See note 160, p. 370. R.M.]

14

[For Mr. A. G. Butler’s observations on the genus Acronycta, see “Trans. Ent. Soc.” 1879, p. 313; and note 68, p. 169, of the present volume. R.M.]

15

[The following characters are given in Stainton’s “Manual of British Butterflies and Moths,” vol. i. p. 114: – “Larva of very variable form: at one extreme we find the singular Cerura larvæ, with only fourteen legs, and two long projecting tails from the last segment; at the other extreme we have larvæ with sixteen legs and no peculiarity of form, such as Chaonia and Bucephala; most have, however, the peculiarity of holding the hind segment of the body erect when in repose; generally quite naked, though downy in Bucephala and rather hairy in Curtulu; very frequently there are projections on the back of the twelfth segment.” R.M.]

16

Encyl. Meth. ix. p. 310.

17

[The genus Vanessa (in the wide sense) appears to be in a remarkable condition of what may be called phyletic preservation. Thus, the group of species allied to V. C. – album passes by almost insensible steps into the group of butterflies typified by our “Tortoiseshells.” The following is a list of some of the intermediate species in their transitional order: —I. – album, V. – album, Faunus, Comma, California, Dryas, Polychloros, Xanthomelas, Cashmirensis, Urticæ, Milberti, &c. Similarly, our Atalanta and Cardui are connected by a number of intermediate forms, showing a complete transition from the one to the other. The following is the order of the species so far as I am acquainted with them: —Atalanta, Dejeanii, Callirhoë, Tammeamea, Myrinna, Huntera, Terpsichore, Carye, Kershawii, and Cardui. R.M.]

18

“Prodromus Systematis Lepidopterorum.” Regensburg, 1864.

19

[The larva of Acherontia Morta, figured by Butler (see note 121, p. 262), possesses the characteristically recurved horn; that of Ach. Medusa figured by the same author, does not appear to possess this character in any marked degree. R.M.]

20

[See note 97, p. 233. R.M.]

21

Loc. cit. Pl. XXV. [This species is referred by Butler to the genus Paonias, Hübn. R.M.]

22

Abbot and Smith, Pl. XXIX. [Placed by Butler in the genus Cressonia, Grote and Robinson. Abbot and Smith state that this larva is sometimes green. According to Mr. Herman Strecker (Lepidop. Rhopal. and Hetero, Reading, Pa. 1874, p. 54) it feeds upon black walnut (Juglans Nigra), hickory (Carya Alba), and ironwood (Ostrya Virginica). Of the North American species of Smerinthus, the following, in addition to Excæcatus, closely resemble our Ocellatus: —S. (Calasymbolus) Geminatus, Say; (C.) Cerisii, Kirby; and Ophthalmicus, Boisd. In addition to S. (Cressonia) Juglandis, S. (Triptogon) Modesta much resembles our Populi. The larva of Geminatus, according to Strecker, is “pale green, lightest above, with yellow lateral granulated stripes; caudal horn violet; stigmata red. It feeds on the willow.” R.M.]

23

Cat. Brit. Mus.

24

[This lengthening of the true legs is mimetic according to Hermann Müller, and causes the anterior portion of the caterpillar to resemble a spider. See note 129, p. 290. R.M.]

25

[Certain butterflies appear to be crepuscular, if not nocturnal in their habits. Thus in his “Notes on the Lepidoptera of Natal,” Mr. W. D. Gooch states that he never saw Melanitis, Leda, or Gnophodes Parmeno on the wing by day, but generally during the hour after sunset. He adds: – “My sugar always attracted them freely, even up to 10 or 11 p.m.” Many species of Hesperidæ are also stated to be of crepuscular habits by this same observer. See “Entomologist,” vol xvi. pp. 38 and 40. R.M.]

26

I only make this assumption for the sake of simplicity, and not because I am convinced that the existing Rhopalocera are actually the oldest Lepidopterous group.