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Studies in the Theory of Descent, Volume I
August Weismann

Studies in the Theory of Descent, Volume I
August Weismann

August Weismann

Studies in the Theory of Descent, Volume I


The present work by Professor Weismann, well known for his profound embryological investigations on the Diptera, will appear, I believe, to every naturalist extremely interesting and well deserving of careful study. Any one looking at the longitudinal and oblique stripes, often of various and bright colours, on the caterpillars of Sphinx-moths, would naturally be inclined to doubt whether these could be of the least use to the insect; in the olden time they would have been called freaks of Nature. But the present book shows that in most cases the colouring can hardly fail to be of high importance as a protection. This indeed was proved experimentally in one of the most curious instances described, in which the thickened anterior end of the caterpillar bears two large ocelli or eye-like spots, which give to the creature so formidable an appearance that birds were frightened away. But the mere explanation of the colouring of these caterpillars is but a very small part of the merit of the work. This mainly consists in the light thrown on the laws of variation and of inheritance by the facts given and discussed. There is also a valuable discussion on classification, as founded on characters displayed at different ages by animals belonging to the same group. Several distinguished naturalists maintain with much confidence that organic beings tend to vary and to rise in the scale, independently of the conditions to which they and their progenitors have been exposed; whilst others maintain that all variation is due to such exposure, though the manner in which the environment acts is as yet quite unknown. At the present time there is hardly any question in biology of more importance than this of the nature and causes of variability, and the reader will find in the present work an able discussion on the whole subject, which will probably lead him to pause before he admits the existence of an innate tendency to perfectibility. Finally, whoever compares the discussions in this volume with those published twenty years ago on any branch of Natural History, will see how wide and rich a field for study has been opened up through the principle of Evolution; and such fields, without the light shed on them by this principle, would for long or for ever have remained barren.

    Charles Darwin.


In offering to English readers this translation of Professor Weismann’s well-known “Studies in the Theory of Descent,” the main part of which is devoted to entomological subjects, I have been actuated by the desire of placing in the hands of English naturalists one of the most complete of recent contributions to the theory of Evolution as applied to the elucidation of certain interesting groups of facts offered by the insect world. Although many, if not most, working naturalists are already familiar with the results of Dr. Weismann’s researches, of which abstracts have from time to time appeared in English and American scientific journals, I nevertheless believe that a study of the complete work, by enabling the reader to follow closely the detailed lines of reasoning and methods of experiment employed by the author, will be found to be of considerable value to those biologists who have not been able to follow the somewhat difficult phraseology of the original. It is not my intention, nor would it be becoming in me to discuss here the merits of the results arrived at by the minute and laborious investigations with which Dr. Weismann has for many years occupied himself. I may however point out that before the appearance of the present work the author, in addition to his well-known papers on the embryology and development of insects, had published two valuable contributions to the theory of descent, viz. one entitled “Über die Berechtigung der Darwin’schen Theorie” (1868), and another “Über den Einfluss der Isolirung auf die Artbildung” (1872). These works, which are perhaps not so well known in this country as could be desired, might be advantageously studied in connection with the present volume wherein they are frequently referred to.

Since every new contribution to science is a fresh starting-point for future work, I may venture without any great breach of propriety to dwell briefly upon one or two of the main points which appear to me to be suggested by Prof. Weismann’s investigations.

Although the causes of Glacial Epochs is a subject which has much occupied the attention of geologists and physiographers, the question is one of such great complexity that it cannot yet be regarded as finally settled. But apart from the question of causes – a most able discussion of which is given by the author of “Island Life” – there is not the least doubt that at no very distant geological period there occurred such an epoch, which, although intermittent, was of considerable duration. The last great geological event which our globe experienced was in fact this Ice Age, and the pure naturalist has not hitherto attributed in my opinion sufficient importance to the direct modifying effects of this prolonged period of cold. It is scarcely possible that such a vast climatic change as that which came on at the close of the Pliocene Period should have left no permanent effect upon our present fauna and flora, all the species of which have survived from the glacial age. The great principle of Natural Selection leads us to see how pre-glacial forms may have become adapted to the new climatic conditions (which came on gradually) by the “survival of the fittest” or “indirect equilibration.” The influence of the last Glacial Epoch as a factor in determining the present geographical distribution of animals and plants has already been amply treated of by many writers since the broad paths were traced out by Darwin, Lyell, and Wallace. The last-named author has indeed quite recently discussed this branch of the subject most exhaustively in his work on “Island Life” above mentioned. The reference of a particular group of phenomena – the seasonal dimorphism of butterflies – to the direct action of the Glacial Period and the subsequent influence of the ameliorating climate, was however the first step taken in this neglected field by the author of the present work in 1875. It is possible, and indeed probable, that future researches will show that other characters among existing species can be traced to the same causes.

The great generalizations of embryology, which science owes so largely to the researches of Karl Ernst von Baer, bear to the theory of descent the same relations that Kepler’s laws bear to the theory of gravitation. These last-named laws are nothing more than generalized statements of the motions of the planets, which were devoid of meaning till the enunciation of the theory of gravitation. Similarly the generalized facts of embryology are meaningless except in the light of the theory of descent. It has now become a recognized principle in biology that animals in the course of their development from the ovum recapitulate more or less completely the phases through which their ancestors have passed. The practical application of this principle to the determination of the line of descent of any species or group of species is surrounded by difficulties, but attempts have been made of late years – as by Haeckel in his Gastrula theory – to push the law to its legitimate consequences. In this country Sir John Lubbock, in 1874, appealed to the embryonic characters of larvæ in support of his views on the origin of insects. To the author of this work (1876) is due the first application of the principle of Ontogeny as revealing the origin of the markings of caterpillars. A most valuable method of research is thus opened up, and entomologists should not be long in availing themselves of it. Our knowledge of the subject of larval development in Lepidoptera is still most imperfect, and it cannot as yet be foreseen to what extent the existing notions of classification in this much-studied order may have to be modified when a minute study of the Comparative Ontogeny of larval characters, worked out as completely as possible for each family, has enabled a true genealogical system to be drawn up. The extent to which such a larval genealogy would coincide with our present classification cannot now be decided, but he who approaches this fruitful line of inquiry in the true spirit of an investigator, will derive much instruction from Prof. Weismann’s remarks on “Phyletic Parallelism in Metamorphic Species.” The affinities of the larger groups among Lepidoptera would most probably be made out once and for ever if systematists would devote more time to observation in this field, and to the co-ordination and working up of the numerous data scattered throughout the vast number of entomological publications.

The doctrine of development by no means implies, as has sometimes been maintained, a continuous advancement in organization. Although the scale of organic nature has continued to rise as a whole, cases may occasionally occur where a lower grade of organization is better adapted to certain conditions of life. This principle of “degeneration” was recognized by Darwin as early as in the first edition of the “Origin of Species;” it was soon perceived to be applicable to the phenomenon of parasitism, and was first definitely formulated by Dr. Anton Dohrn in 1875. In a lecture delivered before the British Association at Sheffield in 1879, Prof. E. Ray Lankester ascribed to “degeneration” a distinct and well-defined function in the theory of descent. Dr. Weismann’s explanation of the transformation of Axolotl given in the fourth essay of this work, may be regarded as a special contribution to this phase of Darwinism. Whilst refuting the idea held by certain naturalists, that such cases are arguments against the origin of species by the accumulation of minute variations, and prove the possibility of development per saltum, the theory here advanced (that Siredon at a former period existed at a higher stage of development as Amblystoma, and that the observed cases of metamorphosis are but reversions to this lost higher stage) suggests the question whether there may not still be in existence many other degenerated forms quite unsuspected by naturalists.

Many of the opponents of Evolution have from time to time denounced this doctrine as leading to “pure materialism,” a denunciation which may appear somewhat alarming to the uninitiated, but which may not seem fraught with any serious consequences to those who have followed the course of philosophical speculation during the last few years. Those who attack the doctrine on this ground will however do well to consider Prof. Weismann’s views set forth in the last essay in this volume, before hastily assuming that the much dreaded “materialism” is incompatible with any other conception of Nature.

The small amount of leisure time which I have been able to devote to the translation of this volume has delayed its completion considerably beyond the anticipated time, and it was with a view to meeting this difficulty that I departed from the original form of the German edition and issued it in parts. Owing to the extremely idiomatic character of the German text, I have throughout endeavoured to preserve only the author’s meaning, regardless of literal translation or of the construction of the original. In some few cases, however, I have intentionally adopted literal translations of certain technical expressions which might, I think, be advantageously introduced into our biological vocabularies. Some alterations have been made in the original text by the author for the present edition, and many new notes have been added. For those bearing my initials I am alone responsible.

It gives me much pleasure in conclusion to express my thanks to Dr. Weismann, not only for the readily given permission to publish an English translation of his work, but also for much valuable assistance during the execution of the task. The author has been good enough to superintend the drawing of the plates for this edition, and he has also read through the greater part of the manuscript. From Mr. Darwin also I have received much kindly encouragement, and among entomologists I am especially indebted to Mr. W. H. Edwards of West Virginia, for his valuable additions to the first part. To my friends Mr. A. G. Butler, Mr. Roland Trimen, and Mr. F. Moore, I owe acknowledgments for much useful information concerning the caterpillars of exotic Sphingidæ, which I have incorporated in the notes and appendices, and Mr. W. S. Simpson has given me occasional advice in the translation of some of the more difficult passages.

    R. M.

London, November, 1881.


With the appearance of Charles Darwin’s work “On the Origin of Species,” in the year 1858, there commenced a new era in biology. Weary of the philosophical speculations which, at the beginning of this century, had at first been started with moderation but had afterwards been pushed to excess, biologists had entirely let drop all general questions and confined themselves to special investigations. The consideration even of general questions had quite fallen into disuse, and the investigation of mere details had led to a state of intellectual shortsightedness, interest being shown only for that which was immediately in view. Immense numbers of detailed facts were thus accumulated, but they could not possibly be mastered; the intellectual bond which should have bound them together was wanting.

But all this was changed in a short time. At first only single and mostly the younger naturalists fell in with the new theory of development proclaimed by Darwin, but the conviction soon became general that this was the only scientifically justifiable hypothesis of the origin of the organic world.

The materials accumulated in all the provinces of biology now for the first time acquired a deeper meaning and significance; unexpected inter-relations revealed themselves as though spontaneously, and what formerly appeared as unanswerable enigmas now became clear and comprehensible. Since that time what a vast modification has the subject of animal embryology undergone; how full of meaning appear the youngest developmental stages, how important the larvæ; how significant are rudimentary organs; what department of biology has not in some measure become affected by the modifying influence of the new ideas!

But the doctrine of development not only enabled us to understand the facts already existing; it gave at the same time an impetus to the acquisition of unforeseen new ones. If at the present day we glance back at the development of the biological sciences within the last twenty years, we must be astonished both at the enormous array of new facts which have been evoked by the theory of development, and by the immense series of special investigations which have been called forth by this doctrine.

But while the development theory for by far the greater majority of these investigations served as a light which more and more illuminated the darkness of ignorance, there appeared at the same time some other researches in which this doctrine itself became the object of investigation, and which were undertaken with a view to establish it more securely.

To this latter class of work belong the “Studies” in the present volume.

It will perhaps be objected that the theory of descent has already been sufficiently established by Darwin and Wallace. It is true that their newly-discovered principle of selection is of the very greatest importance, since it solves the riddle as to how that which is useful can arise in a purely mechanical way. Nor can the transforming influence of direct action, as upheld by Lamarck, be called in question, although its extent cannot as yet be estimated with any certainty. The secondary modifications which Darwin regards as the consequence of a change in some other organ must also be conceded. But are these three factors actually competent to explain the complete transformation of one species into another? Can they transform more than mere single characters or groups of characters? Can we consider them as the sole causes of the regular phenomena of the development of the races of animals and plants? Is there not perhaps an unknown force underlying these numberless developmental series as the true motor power – a “developmental force” urging species to vary in certain directions and thus calling into existence the chief types and sub-types of the animal and vegetable kingdoms?

At the time these “Studies” first appeared (1875) they had been preceded by a whole series of attempts to introduce into science such an unknown power. The botanists, Nägeli and Askenasy, had designated it the “perfecting principle” or the “fixed direction of variation;” Kolliker as the “law of creation;” the philosophers, Von Hartmann and Huber, as the “law of organic development,” and also “the universal principle of organic nature.”

It was thus not entirely superfluous to test the capabilities of the known factors of transformation. We had here before us a question of the highest importance – a question which entered deeply into all our general notions, not only of the organic world, but of the universe as a whole.

This question – does there exist a special “developmental force”? – obviously cannot be decided by mere speculation; it must also be attempted to approach it by the inductive method.

The five essays in this volume are attempts to arrive, from various sides, somewhat nearer at a solution of the problem indicated.

The first essay on the “Seasonal Dimorphism of Butterflies” is certainly but indirectly connected with the question; it is therein attempted to discover the causes of this remarkable dimorphism, and by this means to indicate at the same time the extent of one of the transforming factors with reference to a definite case. The experiments upon which I base my views are not as numerous as I could desire, and if I were now able to repeat them they would be carried out more exactly than was possible at that time, when an experimental basis had first to be established. In spite of this, the conclusions to which I was led appear to be on the whole correct. That admirable and most conscientious observer of the North American butterflies, Mr. W. H. Edwards, has for many years experimented with American species in a manner similar to that which I employed for European species, and his results, which are published here in Appendix II (#litres_trial_promo). 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 - A most minute and exact description of the newly hatched larva of Chionobas Aëllo is given by the American entomologist, Samuel H. Scudder. Ann. Soc. Ent. de Belgique, xvi., 1873.] 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 with some quite ordinary phenomenon which requires no explanation, now go too far towards the other extreme. The term “larval reproduction” is an expression, but no explanation; we have therefore to attempt to find out the true interpretation, but whether the one which I have given is correct must be judged of by others.

These four essays lead up to a fifth and concluding one “On the Mechanical Conception of Nature.” Whilst the results obtained are here summed up, it is attempted to form them into a philosophical conception of Nature and of the Universe. It will be thought by many that this should have been left to professed philosophers, and I readily admit that I made this attempt with some misgiving. Two considerations, however, induced me to express here my own views. The first was that the facts of science are frequently misunderstood, or at any rate not estimated at their true value, by philosophers;[2 - I am aware that this certainly cannot be said of philosophers like Lotze or Herbert Spencer; but these are at the same time both naturalists and philosophers.] the second consideration was, that even certain naturalists and certainly very many non-naturalists, turn distrustfully from the results of science, because they fear that these would infallibly lead to a view of the Universe which is to them unacceptable, viz. the materialistic view. With regard to the former I wished to show that the views of the development of organic Nature inaugurated by Darwin and defended in this work are certainly correctly designated mechanical; with reference to the latter I wished to prove that such a mechanical conception of the organic world and of Nature in general, by no means leads merely to one single philosophical conception of Nature, viz. to Materialism, but that on the contrary it rather admits of legitimate development in a quite different manner.

Thus in these last four essays much that appears heterogeneous will be found in close association, viz. scientific details and general philosophical ideas. In truth, however, these are most intimately connected, and the one cannot dispense with the other. As the detailed investigations of the three essays find their highest value in the general considerations of the fourth, and were indeed only possible by constantly keeping this end in view, so the general conclusions could only grow out of the results of the special investigations as out of a solid foundation. Had the new materials here brought together been already known, the reader would certainly have been spared the trouble of going into the details of special scientific research. But as matters stood it was indispensable that the facts should be examined into and established even down to the most trifling details. The essay “On the Origin of the Markings of Caterpillars” especially, had obviously to commence with the sifting and compilation of extensive morphological materials.

    August Weismann.

Freiburg in Baden,

November, 1881.


I. The Origin and Significance of Seasonal Dimorphism
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