APPENDIX II.
The following paper by Dr. Fritz Müller[222] forms the third of a series of communications on Brazilian butterflies published in “Kosmos,” and as it bears upon the investigations made known in the third essay of the present work, I will here give a translation, by permission of the publisher, Herr Karl Alberts.
“Acræa and the Maracujá Butterflies as Larvæ, Pupæ, and Imagines.
“In a thoughtful essay on ‘Phyletic Parallelism in Metamorphic Species,’ Weismann has shown that in the case of Lepidoptera the developmental stages of larva, pupa, and imago vary independently, and that a change occurring in one stage is without influence upon the preceding and succeeding stages, so that the course which has been followed by the individual stages in their developmental history has not been in all cases identical. This want of agreement may manifest itself both by unequal divergence of form-relationship, and by unequal group formation. With respect to unequal form-divergence the caterpillars are sometimes more closely related in form than their imagines, and at other times the reverse is the case. With respect to unequal group formation again, two cases are possible; the larvæ and imagines may form groups of unequal value, the one stage forming higher or lower groups than the other, or they may form groups of unequal size, i.e., groups which do not coincide but which overlap. Form-relationship and blood-relationship do not therefore always agree; the resemblances among the caterpillars would lead to a quite different arrangement to that resulting from the resemblances among the imagines, and it is probable that neither of these arrangements would correspond with the actual relationships.
“Starting from this fact, which he establishes by numerous examples, Weismann proceeds to show most convincingly that an innate power of development or of transformation, such as has been assumed under various names by many adherents of the development theory, has no existence, but that every modification and advancement in species has been called forth by external influences.
“A most beautiful illustration of the want of ‘phyletic parallelism,’ as Weismann designates the different form-relationships of the larvæ, pupæ, and imagines, is furnished by the five genera Acræa, Heliconius, Eueides, Colænis, and Dione (= Agraulis). This instance seems to me to be of especial value, because it offers the rare case of pupæ showing greater differences than the larvæ and imagines.
“The species of which I observed the larvæ and pupæ are Acræa Thalia and Alalia, Heliconius Eucrate, Eueides Isabella, Colænis Dido and Julia, Dione Vanillæ and Juno; besides these I noticed the pupa of Eueides Aliphera.
“The following remarks apply only to these species, although we may suppose with great probability that the whole of the congeneric forms—excepting perhaps the widely ranging species of Acræa—would display similar characters to their Brazilian representatives.
“The imagines of the five genera mentioned form two sharply defined families, the Acræidæ and the butterflies of the Maracujá group.[223] The latter comprises the three genera Heliconius, Eueides, and Colænis, which differ only in very unimportant characters; Eueides is distinguished from Heliconius by its shorter antennæ, and Colænis differs from Eueides in having the discoidal cell of the hind-wings open. The genus Dione is further removed by the different structure of the legs, and the silvery spots on the underside of the wings. Certain species resemble those of other genera in a most striking manner, and much more closely both in colour and marking, and even in the form of their wings, than they do their own congeners. This is the case with Acræa Thalia and Eueides Pavana, with Heliconius Eucrate and Eueides Isabella, and with Eueides Aliphera and Colænis Julia, which are deceptively alike, and the last two are connected with Dione Juno, at least by the upper side of the wings. The difficulty of judging of the relationships of the single species is thus much aggravated; it cannot be said how much of this resemblance is to be attributed to blood-relationship, and how much to deceptive imitation.
“As larvæ all the Brazilian species must be placed in one genus, as they agree exactly in the number and arrangement of their spines (4 spines, not in a transverse row, on segments 2 and 3; 6 spines, in a transverse row, on segments 4–11; 4 spines, not in a transverse row, on the last (12th) segment). They differ from one another much less in this respect than do the German species of Vanessa, such, for instance, as V. Io or Antiopa from V. Polychloros, Urticæ, and Atalanta.[224] The larvæ of Acræa Thalia are certainly without the two spines on the head which the others possess, and, on the other hand, they have a well-developed pair of spines on the first segment, which, in most of the other species, are completely absent; but this does not justify their separation, since the head spines of Heliconius, Eueides, and Colænis Dido, which are of a considerable length, are shorter than those of the next segment in Colænis Julia, and Dione Vanillæ, and in Dione Juno they dwindle down to two minute points, this last species also bearing a short pair on the first segment. The larva of Dione Juno is thus as closely related to that of Acræa Thalia as it is to that of its congener Dione Vanillæ.
“If it were desired to form two distinct larval groups this could not be effected on the basis of their differences in form, but could only be based on their food-plants. The larvæ of Heliconius, Eueides, Colænis, and Dione live on species of Maracujá (Passiflora); those of Acræa Thalia and Alalia on Compositæ (Mikania and Veronia). These larval groups would agree with those founded on the form-relationships of the imagines, but unlike the imaginal groups, which can be formed into families, they would scarcely possess a generic value.
“If we arrange the single species of caterpillars according to their resemblances, this arrangement does not agree with that based on the resemblances of the imagines, even if we disregard the different values of the groups. The result is somewhat as follows:—
IMAGINES.
(Nymphalideous butterflies with tufts on wing-veins.)
/---------------------------------------------------\
(Families.) MARACUJÁ-GROUP. ACRÆIDÆ.
/---------------------------\ |
/-------------------\ /-------------------\ |
(Genera.) Heliconius. Eueides. Colænis. Dione. Acræa.
| /-------\ /-------\ |
(Species.) Eucrate. Isabella. Dido. Julia. Vanillæ. Juno. Thalia. | | | | | | |
\----------+---------/ \--------/ \------/
\--------------------------------------/
LARVÆ.”
* * * * *
[Here follow the remarks on the habits of the larvæ in connection with their colours, &c., which have already been quoted in illustration of the use of the spiny protection (note [133], p. [293]). From these facts the author draws the conclusion that the form-relationships of the caterpillars depend rather upon their mode of life than upon their blood-relationships, assuming the latter to be correctly expressed by the arrangement of the imagines at present adopted.]
Figs. 1–4. Pupæ of Acræa Thalia; Heliconius Eucrate; Eueides Isabella, and Colænis Dido; life size.
“A glance at the above figures of the pupæ of Heliconius Eucrate (Fig. 2), Eueides Isabella (Fig. 3), and Colænis Dido (Fig. 4), will show how great are the differences between these pupæ as compared with the close form-relationship of all the Maracujá butterflies, and with the no less close resemblance of their larvæ. A family which comprised three such dissimilar pupæ would also be capable of including that of Acræa Thalia (Fig. 1).
“The pupa of this last species has nothing peculiar in its general appearance, but possesses the ordinary pupal form; it is tolerably rounded, without any great elevations or depressions; a minute pointed projection is situated on the head over each eye-cover, and a similar process projects from the roots of the wings. Its distinguishing characters are five pairs of spines on the back of the abdominal segments. These spines are found also in Acræa Alalia, but appear to be absent in other species, e.g. in the Indian A. Violæ. Last summer, among some batches of Thalia larvæ—each batch being the progeny from one lot of eggs—I found certain individuals which differed from the others in having much shorter spines, and these changed into pupæ in which the five pairs of spines were proportionally shorter than usual, thus being an exception to the rule that changes in one stage of development are without influence on the other stages. I may remark, by the way, that this law, enunciated by Weismann, can only be applied to imagines and pupæ with certain restrictions. The skin of the pupa forms a sheath or cover for the eyes, antennæ, trunk, legs, and wings of the imago, and if these parts undergo any considerable modification in the latter, corresponding changes must appear in the pupa. This is shown, for instance, by many ‘Skippers’ (Hesperidæ), in which the extraordinarily long trunk necessitates a sheath of a corresponding length. The colour of the pupa of Acræa Thalia is whitish, the wing-veins with some other markings and the spines are black; metallic spots are absent.
“In the pupa of Heliconius Eucrate the laterally compressed region of the wings is raised into a large projection, the antennal sheaths lying on the edges of the wings are serrated and beset with short pointed spines; instead of the minute projections of Acræa Thalia, the head bears two large humped processes; the body is raised on each side into a foliaceous border carrying five spines of different lengths, the foremost pair, directed towards the head, being the longest. The pupa is brown, and ornamented with four pairs of brilliant metallic spots, one pair close behind the antennæ, and three pairs, almost coalescent, on the back before the longest pair of spines. A short spine projects from the middle of each of the latter somewhat arched metallic patches.
“In the pupa of Colænis Dido (which resembles that of Colænis Julia, and to which may be added those of Dione Vanillæ and Juno) the spines are absent, the wing region is but moderately arched, and the antennæ marked only by small elevations; instead of the leaf-like border, there are on each side of the back five knotty or humped processes. The metallic spots are similar in number and position to those of Heliconius Eucrate; those on the back have a wart-like process in the middle, instead of a spine.
“The pupæ of Heliconius and Colænis when moving their posterior segments rapidly, as they do whenever they are disturbed, produce a very perceptible hissing noise by the friction of these segments, this sound, which is especially noticeable in the case of Heliconius Eucrate, perhaps serving to terrify small foes. (So loud is the sound produced in this manner by the pupæ of Epicalia Numilia, that my children have named them ‘Schreipuppen.’)
“The pupæ of Heliconius and Colænis thus differ to a much greater extent than the imagines or larvæ, and the same holds good for Eueides in a much higher degree as compared with its above-mentioned allies. The larvæ of Eueides have no distinctive characters, and even the generic rank of the imagines is doubtful; as pupæ, on the other hand, they are far removed (even by their mode of suspension) not only from the remainder of the Maracujá group and from the whole of the great Nymphalideous group (Danainæ, Satyrinæ, Elymniinæ, Brassolinæ, Morphinæ, Acræinæ and Nymphalinæ), but from almost all other butterflies. The larva pupates on the underside of a leaf; the pupa is fastened by the tail, but does not hang down like the pupæ of the other Nymphalidæ,—its last segments are so curved that the breast of the chrysalis is in contact with the underside of the leaf. I am not acquainted with any other pupa among those not suspended by a girdle which assumes such a position. Something similar occurs, however, in the pupa of Stalachtis, which is without a girdle, and according to Bates, is ‘kept in an inclined position by the fastening of the tail.’ By this peculiarity Bates distinguishes the Stalachtinæ from the Libytheæ with pupæ ‘freely suspended by the tail.’
“Besides through this peculiar position of the body, the pupa of Eueides Isabella is distinguished by short hooked and long narrow sabre-like pairs of processes on the back and head. Its colour is whitish, yellowish, or sordid yellowish-grey; in the last variety both the four long dorsal processes and the surrounding portions, as well as the points of the other processes, remain white or yellowish. The pupa Eueides Aliphera is very similar, only all the processes are somewhat shorter, the four longest (dorsal) and some other markings being black.
“Now if, as Weismann has attempted to show for larvæ and imagines, the form-divergence always ‘corresponds exactly with the divergence in the mode of life,’ the question arises as to what difference in the conditions of life has brought about such a considerable form-divergence between the pupæ of such closely allied species as the Maracujá butterflies. In pupæ which do not eat or drink, and which have neither to seek in courtship nor to care for progeny, it is only protection from foes that can concern us. But in the pupæ of nearly allied species of which the larvæ feed on kindred plants in the same districts at the same periods of the year, can the enemies be so different as to produce such a considerable divergence in form? One might answer this question in the negative with some confidence, and affirm that in this case the difference in the pupæ does not result from the ‘divergence in the mode of life,’ or from the difference in the external conditions, but is accidental, i.e. a consequence of some fortunate variation induced by some external cause, which variation afforded protection against common foes—to one species in one way, and to the other species in some other way; this course, once entered upon, having been urged on by natural selection, until at length the wide divergence now shown is attained. How in the case of any of the species the peculiarity in colour or form can actually serve as a protection, I must confess myself at fault in answering. Only in the case of the pupa of Eueides Isabella will I venture to offer a supposition. That it is not green like other pupæ which suspend themselves among foliage (Siderone, Epicalia, Callidryas, &c.), but contrasts more or less brightly with the dark green of the leaves, precludes the idea of concealment; on the other hand its colour is too dull to serve as a conspicuous sign of distastefulness. In either case the meaning of the wonderful processes of the pupa would remain unexplained.
“We are thus compelled to seek another possibility in mimicry, by which foes would be deceived by deceptive resemblance. But what is the object imitated? Dead insects overgrown by fungi are often found on leaves, the whitish or yellowish fungi growing from their bodies in various fantastic forms. Such insects of course no longer serve as tempting morsels. The processes of the pupa of Eueides suggest such fungoid growths, although I certainly cannot assert that to our eyes in broad daylight the resemblance is very striking. But the pupæ hang among the shadows of the leaves, and a less perfect imitation may deceive foes that are not so sharp-sighted; protective resemblance must commence moreover with an imperfect degree of imitation.”