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Alfred Russel Wallace : Alfred Wallace : A. R. Wallace :
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Two Darwinian Essays (S324: 1880)

Editor Charles H. Smith's Note: A review of two works: Studies in the Theory of Descent, Part I. On the Seasonal Dimorphism of Butterflies, with Two Coloured Plates, by August Weismann, and Degeneration. A Chapter in Darwinism., by E. Ray Lankester. Printed in the Nature issue of 17 June 1880. Original pagination indicated within double brackets. To link directly to this page connect with: http://people.wku.edu/charles.smith/wallace/S324.htm

    [[p. 141]] The first of Dr. Weismann's "Studies," of which Mr. Meldola has given us an excellent translation, with the author's latest notes and additions, is devoted to a thorough examination of the well-known but hitherto little understood phenomenon of the seasonal forms of butterflies. For the benefit of those unacquainted with entomology we may state, that many butterflies have two, or even three broods in a year. One brood appears in spring, their larvæ having fed during the preceding autumn and passed the winter in the pupa state, while the others appear later in the year, having passed rapidly through all their transformations and thus never having been exposed to the cold of winter. In most cases the insects produced under these opposite conditions present little or no perceptible difference; but in others there is a constant variation, and sometimes this is so great that the two forms have been described as distinct species. The most remarkable case among European butterflies is that of Araschnia prorsa, the winter or spring form of which was formerly considered to be a distinct species and named Araschnia levana. The two insects differ considerably in both sexes, in markings, in colour, and even in the form of the wings, so that till they were bred and found to be alternate broods of the same species (about the year 1830) no one doubted their being altogether distinct.

    In order to learn something of the origin and nature of this curious phenomenon Dr. Weismann has for many years carried on a variety of experiments, breeding the species in large numbers and subjecting the pupæ to artificial heat or cold for the purpose of hastening or retarding the transformation. The result of these experiments is, that by subjecting the summer brood to severe artificial cold in the pupa state, it may be made to produce perfect insects the great majority of which are of the winter form; but, on the other hand, no change of conditions that have yet been tried have any effect in changing the winter to the summer form. Taking this result in connection with the fact that in high latitudes where there is only one brood a year it is always the winter form, Dr. Weismann was led to the hypothesis that this winter form was the original type of the species, and that the summer form has been produced gradually, since the glacial epoch, by the summer becoming longer and thus admitting of the production of a second or summer brood. This explains why the production of the winter form (A. levana) from summer larvæ is easy, it being a reversion to the ancestral type; while the production of the summer form (A. prorsa) from autumnal larvæ is impossible, because that form is the result of gradual development; and processes of development which have taken thousands of years to bring about cannot be artificially reproduced in a single season.

    This hypothesis was supported by experiments with another two-brooded species, Pieris napi, with similar results, the winter form being produced with certainty by the application of cold to summer pupæ; and Mr. Edwards, in America, has made similar experiments with the various forms of Papilii ajax, finding that the summer broods can be changed into the winter form by the application of cold, while the winter broods can never be made to assume the summer form by hastening the process of transformation. In the Arctic regions and in the high Alps there is only one form of Pieris napi, which very closely resembles the winter form of the rest of Europe, and this could never be the least changed by rapidly developing the pupæ under the influence of heat.

    Another curious case is that of one of the Lycænidæ (Plebeius agestis) which exhibits three forms, which may be designated as A, B, and C. The first two, A and B, are alternate broods (winter and summer) in Germany, while in Italy the corresponding forms are B and C, so that B is the summer form in Germany and the winter form in Italy. Here we see climatic varieties in process of formation in a very curious way.

    That temperature during the pupa stage is a very powerful agent in modifying the characters of butterflies, is well shown by the case of Polyommatus phlæas. The two broods of this insect are alike in Germany, while in Italy the summer brood has the wings dusky instead of copper-coloured. The period of development is exactly the same in both countries, so that the change must, it is argued, be attributed to the higher temperature of the Italian summer. It has been noticed that in Italy a large number of species of butterflies are thus seasonally dimorphic which are not so in Central and Northern Europe.

    Dr. Weismann lays great stress on the varied effects of temperature in modifying allied species or the two sexes of the same species, from which he argues that the essential cause of all these changes is to be found in peculiarities of physical constitution, which cause different species, varieties, or sexes to respond differently to the same change of temperature; and he thinks that many sexual differences can be traced to this cause alone without calling in the aid of sexual selection. The general result arrived at by the laborious investigation of these phenomena is, that--"a species is only caused to change through the influence of changing external conditions of life, this change being in a fixed direction which entirely depends on the physical nature of the varying organism, and is different in different species, or even in the two sexes of the same species;" and he adds:--"According to my view, transmutation by purely internal causes is not to be entertained. If we could absolutely suspend the changes of the external conditions of life, existing species would remain stationary. The action of external inciting causes, in the widest sense of the word, is alone able to produce modifications; and even the never-failing 'individual variations,' together with the inherited dissimilarity [[p. 142]] of constitution, appear to me to depend upon unlike external influences, the inherited constitution itself being dissimilar because the individuals have been at all times exposed to somewhat varying external influences." The present writer has arrived at almost exactly similar conclusions to these, from a study of the geographical distribution and specific variation of animal forms, as stated in an article on "The Origin of Species and Genera," which appeared in the Nineteenth Century of January last, and it is gratifying to find them supported by the results of a very different line of inquiry, and by the authority of so eminent and original an observer as Dr. Weismann.

    The second work referred to in our heading, is Prof. Lankester's British Association evening lecture last year at Sheffield, now republished with illustrations as one of the useful little volumes of the "Nature Series." It discusses the little-known phenomena of "Degeneration" as a phase of development much more general, and of far greater importance than is usually supposed. Degeneration causes an organism to become more simple in structure, in adaptation to less varied and less complex conditions of life. "Any new set of conditions occurring to an animal which render its food and safety very easily attained, seem to lead as a rule to degeneration; just as an active healthy man sometimes degenerates when he becomes suddenly possessed of a fortune; or as Rome degenerated when possessed of the riches of the ancient world. The habit of parasitism clearly acts upon animal organisation in this way. Let the parasitic life once be secured, and away go legs, jaws, eyes, and ears; the active and highly-gifted crab, insect, or annelid may become a mere sac, absorbing nourishment and laying eggs."

    We see incipient cases of degeneration in the loss of limbs of the serpentiform lizards and the pisciform mammals; the loss of eyes in the inhabitants of caverns and in some earth-burrowers; the loss of wings in the Apteryx and of toes in the horse; and, still more curious, the loss of the power of feeding themselves in some slave-holding ants. More pronounced cases are those of the barnacles--degenerated crustacea, and the mites--degenerate spiders; while we reach the climax of the process in Ascidians--degenerate vertebrates, and such mere living sacs as the parasitic Sacculina and Lernæocera, which are degenerated crustaceans. Not only such lesser groups as the above, but whole orders may be the result of degeneration. Such are the headless bivalve mollusca known as Lamellibranchs, which are believed to have degenerated from the head-bearing active cuttle-fish type; while the Polyzoa or Moss-polyps stand in the same relation to the higher Mollusca as do the Ascidians to the higher Vertebrates.

    While discarding the hypothesis that all savages are the descendants of more civilised races, Prof. Lankester yet admits the application of his principle to explain the condition of some of the most barbarous races--"such as the Fuegians, the Bushmen, and even the Australians. They exhibit evidence of being descended from ancestors more cultivated than themselves." He even applies it to the higher races in intellectual matters, and asks: "Does the reason of the average man of civilised Europe stand out clearly as an evidence of progress when compared with that of the men of bygone ages? Are all the inventions and figments of human superstition and folly, the self-inflicted torturing of mind, the reiterated substitution of wrong for right, and of falsehood for truth, which disfigure our modern civilisation--are these evidence of progress? In such respects we have at least reason to fear that we may be degenerate. It is possible for us--just as the Ascidian throws away its tail and its eye and sinks into a quiescent state of inferiority--to reject the good gift of reason with which every child is born, and to degenerate into a contented life of material enjoyment accompanied by ignorance and superstition."

    This is very suggestive; but we may, I think, draw a yet higher and deeper teaching from the phenomena of degeneration. We seem to learn from it the absolute necessity of labour and effort, of struggle and difficulty, of discomfort and pain, as the condition of all progress, whether physical or mental, and that the lower the organism the more need there is of these ever-present stimuli, not only to effect progress, but to avoid retrogression. And if so, does not this afford us the nearest attainable solution of the great problem of the origin of evil? What we call evil is the essential condition of progress in the lower stages of the development of conscious organisms, and will only cease when the mind has become so thoroughly healthy, so well balanced, and so highly organised, that the happiness derived from mental activity, moral harmony, and the social affections, will itself be a sufficient stimulus to higher progress and to the attainment of a more perfect life.

    For numerous instructive details connected with degenerated animals we refer our readers to the work itself--truly a small book on a great subject, and one which discusses matters of the deepest interest, alike to the naturalist and the philosopher.

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