Studies in the Theory of Descent, Volume I. Weismann August

I employed for European species, and his results, which are published here in Appendix II. to the first essay, contain nothing as far as I can see which is not in harmony with my views. Many new questions suggest themselves, however, and it would be a grateful task if some entomologist would go further into these investigations.

      The second essay directly attacks the main problem above indicated. It treats of the “Origin of the Markings of Caterpillars,” and is to some extent a test of the correctness and capabilities of the Darwinian principles; it attempts to trace the differences in form in a definite although small group entirely to known factors.

      Why the markings of caterpillars have particularly been chosen for this purpose will appear for two reasons.

      The action of Natural Selection, on account of the nature of this agency, can only be exerted on those characters which are of biological importance. As it was to be tested whether, besides Natural Selection and the direct action of external conditions, together with the correlative results of these two factors, there might not lie concealed in the organism some other unknown transforming power, it was desirable to select for the investigation a group of forms which, if not absolutely excluding, nevertheless appeared possibly to restrict, the action of one of the two known factors of transformation, that of Natural Selection; a group of forms consisting essentially of so-called “purely morphological” characters, and not of those the utility of which was obvious, and of which the origin by means of Natural Selection was both possible and probable ab initio. Now, although the colouring can readily be seen to be of value to the life of its possessors, this is not the case with the quite independent markings of caterpillars; excepting perhaps those occasional forms of marking which have been regarded as special cases of protective resemblance. The markings of caterpillars must in general be considered as “purely morphological” characters, i. e. as characters which we do not know to be of any importance to the life of the species, and which cannot therefore be referred to Natural Selection. The most plausible explanation of these markings might have been that they were to be regarded as ornaments, but this view precludes the possibility of referring them either to Natural Selection or to the influence of direct changes in the environment.

      The markings of caterpillars offered also another advantage which cannot be lightly estimated; they precluded from the first any attempt at an explanation by means of Sexual Selection. Although I am strongly convinced of the activity and great importance of this last process of selection, its effects cannot be estimated in any particular case, and the origin of a cycle of forms could never be clearly traced to its various factors, if Sexual Selection had also to be taken into consideration. Thus, we may fairly suppose that many features in the markings of butterflies owe their origin to Sexual Selection, but we are, at least at present, quite in the dark as to how many and which of these characters can be traced to this factor.

      An investigation such as that which has been kept in view in this second essay would have been impracticable in the case of butterflies, as well as in the analogous case of the colouring and marking of birds, because it would have always been doubtful whether a character which did not appear to be attributable to any of the other transforming factors, should not be referred to Sexual Selection. It would have been impossible either to exclude or to infer an unknown developmental force, since we should have had to deal with two unknowns which could in no way be kept separate.

      We escape this dilemma in the markings of caterpillars, because the latter do not propagate in this state. If the phenomena are not here entirely referable to Natural Selection and the direct action of the environment – if there remains an inexplicable residue, this cannot be referred to Sexual Selection, but to some as yet unknown power.

      But it is not only in this respect that caterpillars offer especial advantages. If it is to be attempted to trace transformations in form to the action of the environment, an exact knowledge of this environment is in the first place necessary, i. e. a precise acquaintance with the conditions of life under the influence of which the species concerned exist. With respect to caterpillars, our knowledge of the life conditions is certainly by no means as complete as might be supposed, when we consider that hundreds of Lepidopterists have constantly bred and observed them during a most extended period. Much may have been observed, but it has not been thought worthy of publication; much has also been published, but so scattered and disconnected and at the same time of such unequal credibility, that a lifetime would be required to sift and collect it. A comprehensive biology of caterpillars, based on a broad ground, is as yet wanting, although such a labour would be both most interesting and valuable. Nevertheless, we know considerably more of the life of caterpillars than of any other larvæ, and as we are also acquainted with an immense number of species and are able to compare their life and the phenomena of their development, the subject of the markings of caterpillars must from this side also appear as the most favourable for the problem set before us.

      To this must be added as a last, though not as the least, valuable circumstance, that we have here preserved to us in the development of the individual a fragment of the history of the species, so that we thus have at hand a means of following the course which the characters to be traced to their causes – the forms of marking – have taken during the lapse of thousands of years.

      If with reference to the question as to the precise conditions of life in caterpillars I was frequently driven to my own observations, it was because I found as good as no previous work bearing upon this subject. It was well known generally that many caterpillars were differently marked and coloured when young to what they were when old; in some very striking cases brief notices of this fact are to be found in the works,1 more especially, of the older writers, and principally in that of the excellent observer Rösel von Rosenhof, the Nuremberg naturalist and miniature painter. In no single case, however, do the available materials suffice when we have to draw conclusions respecting the phyletic development. We distinctly see here how doubtful is the value of those observations which are made, so to speak, at random, i. e. without some definite object in view. Many of these observations may be both good and correct, but they are frequently wanting precisely in that which would make them available for scientific purposes. Thus everything had to be established de novo, and for this reason the investigations were extended over a considerable number of years, and had to be restricted to a small and as sharply defined a group as possible – a group which was easily surveyed, viz. that of the Hawk-moths or Sphinges.

      Since the appearance of the German edition of this work many new observations respecting the markings of caterpillars have been published, such, for example, as those of W. H. Edwards and Fritz Müller. I have, however, made but little use of them here, as I had no intention of giving anything like a complete ontogeny of the markings in all caterpillars: larval markings were with me but means to an end, and I wished only to bring together such a number of facts as were necessary for drawing certain general conclusions. It would indeed be most interesting to extend such observations to other groups of Lepidoptera.

      The third essay also, for similar reasons, is based essentially upon the same materials, viz. the Lepidoptera. It is therein attempted to approach the general problem – does there or does there not exist an internal transforming force? – from a quite different and, I may say, opposite point of view. The form-relationships of Lepidoptera in their two chief stages of development, imago and larva, are therein analysed, and by an examination of the respective forms it has been attempted to discover the nature of the causes which have led thereto.

      I may be permitted to say that the fact here disclosed of a different morphological, with the same genealogical relationship, appears to me to be of decided importance. The agreement of the conclusions following therefrom with the results of the former investigation has, at least in my own mind, removed the last doubts as to the correctness of the latter.

      The fourth and shortest essay on the “Transformation of the Axolotl into Amblystoma,” starts primarily with the intention of showing that cases of sudden transformation are no proof of per saltum development. When this essay first appeared the view was still widely entertained that we had here a case proving per saltum development. That this explanation was erroneous is now generally admitted, but I believe that those who suppose that we have here to deal