It is now thirty years since I gave an analysis in the first edition of my "Principles of Geology" (volume 2 1832) of the views which had been put forth by Lamarck, in the beginning of the century, on this subject. In that interval the progress made in zoology and botany, both in augmenting the number of known animals and plants, and in studying their physiology and geographical distribution and above all in examining and describing fossil species, is so vast that the additions made to our knowledge probably exceed all that was previously known; and what Lamarck then foretold has come to pass; the more new forms have been multiplied, the less are we able to decide what we mean by a variety, and what by a species. In fact, zoologists and botanists are not only more at a loss than ever how to define a species, but even to determine whether it has any real existence in nature, or is a mere abstraction of the human intellect, some contending that it is constant within certain narrow and impassable limits of variability, others that it is capable of indefinite and endless modification.
Before I attempt to explain a great step, which has recently been made by Mr. Darwin and his fellow-labourers in this field of inquiry, I think it useful to recapitulate in this place some of the leading features of Lamarck's system, without attempting to adjust the claims of some of his contemporaries (Geoffroy St. Hilaire in particular) to share in the credit of some of his original speculations.
From the time of Linnaeus to the commencement of the present century, it seemed a sufficient definition of the term species to say that "a species consisted of individuals all resembling each other, and reproducing their like by generation." But Lamarck after having first studied botany with success, had then turned his attention to conchology, and soon became aware that in the newer (or Tertiary) strata of the earth's crust there were a multitude of fossil species of shells, some of them identical with living ones, others simply varieties of the living, and which as such were entitled to be designated, according to the ordinary rules of classification, by the same names. He also observed that other shells were so nearly allied to living forms that it was difficult not to suspect that they had been connected by a common bond of descent. He therefore proposed that the element of time should enter into the definition of a species, and that it should run thus: "A species consists of individuals all resembling each other, and reproducing their like by generation, SO LONG AS THE SURROUNDING CONDITIONS DO NOT UNDERGO CHANGES SUFFICIENT TO CAUSE THEIR HABITS, CHARACTERS, AND FORMS TO VARY." He came at last to the conclusion that none of the animals and plants now existing were primordial creations, but were all derived from pre-existing forms, which, after they may have gone on for indefinite ages reproducing their like, had at length, by the influence of alterations in climate and in the animate world been made to vary gradually, and adapt themselves to new circumstances, some of them deviating in the course of ages so far from their original type as to have claims to be regarded as new species.
In support of these views, he referred to wild and cultivated plants and to wild and domesticated animals, pointing out how their colour, form, structure, physiological attributes and even instincts were gradually modified by exposure to new soils and climates, new enemies, modes of subsistence, and kinds of food.
Nor did he omit to notice that the newly acquired peculiarities may be inherited by the offspring for an indefinite series of generations, whether they be brought about naturally—as when a species, on the extreme verge of its geographical range, comes into competition with new antagonists and is subjected to new physical conditions; or artificially—as when by the act of the breeder or horticulturist peculiar varieties of form or disposition are selected.
But Lamarck taught not only that species had been constantly undergoing changes from one geological period to another, but that there also had been a progressive advance of the organic world from the earliest to the latest times, from beings of the simplest to those of more and more complex structure, and from the lowest instincts up to the highest, and finally from brute intelligence to the reasoning powers of Man. The improvement in the grade of being had been slow and continuous, and the human race itself was at length evolved out of the most highly organised and endowed of the inferior mammalia.
In order to explain how, after an indefinite lapse of ages, so many of the lowest grades of animal or plant still abounded, he imagined that the germs or rudiments of living things, which he called monads, were continually coming into the world and that there were different kinds of these monads for each primary division of the animal and vegetable kingdoms. This last hypothesis does not seem essentially different from the old doctrine of equivocal or spontaneous generation; it is wholly unsupported by any modern experiments or observation, and therefore affords us no aid whatever in speculating on the commencement of vital phenomena on the earth.
Some of the laws which govern the appearance of new varieties were clearly pointed out by Lamarck. He remarked, for example, that as the muscles of the arm become strengthened by exercise or enfeebled by disuse, some organs may in this way, in the course of time, become entirely obsolete, and others previously weak become strong and play a new or more leading part in the organisation of a species. And so with instincts, where animals experience new dangers they become more cautious and cunning, and transmit these acquired faculties to their posterity. But not satisfied with such legitimate speculations, the French philosopher conceived that by repeated acts of volition animals might acquire new organs and attributes, and that in plants, which could not exert a will of their own, certain subtle fluids or organising forces might operate so as to work out analogous effects.
After commenting on these purely imaginary causes, I pointed out in 1832, as the two great flaws in Lamarck's attempt to explain the origin of species, first, that he had failed to adduce a single instance of the initiation of a new organ in any species of animal or plant; and secondly, that variation, whether taking place in the course of nature or assisted artificially by the breeder and horticulturist, had never yet gone so far as to produce two races sufficiently remote from each other in physiological constitution as to be sterile when intermarried, or, if fertile, only capable of producing sterile hybrids, etc.* (* "Principles of Geology" 1st edition volume 2 chapter 2.)
To this objection Lamarck would, no doubt, have answered that there had not been time for bringing about so great an amount of variation; for when Cuvier and some other of his contemporaries appealed to the embalmed animals and plants taken from Egyptian tombs, some of them 3000 years old, which had not experienced in that long period the slightest modification in their specific characters, he replied that the climate and soil of the valley of the Nile had not varied in the interval, and that there was therefore no reason for expecting that we should be able to detect any change in the fauna and flora. "But if," he went on to say, "the physical geography, temperature, and other conditions of life had been altered in Egypt as much as we know from geology has happened in other regions, some of the same animals and plants would have deviated so far from their pristine types as to be thought entitled to take rank as new and distinct species."
Although I cited this answer of Lamarck in my account of his theory,* (* Ibid. page 587.) I did not at the time fully appreciate the deep conviction which it displays of the slow manner in which geological changes have taken place and the insignificance of thirty or forty centuries in the history of a species, and that, too, at a period when very narrow views were entertained of the extent of past time by most of the ablest geologists, and when great revolutions of the earth's crust, and its inhabitants, were generally attributed to sudden and violent catastrophes.
While in 1832 I argued against Lamarck's doctrine of the gradual transmutation of one species into another, I agreed with him in believing that the system of changes now in progress in the organic world would afford, when fully understood, a complete key to the interpretation of all the vicissitudes of the living creation in past ages. I contended against the doctrine, then very popular, of the sudden destruction of vast multitudes of species and the abrupt ushering into the world of new batches of plants and animals.
I endeavoured to sketch out (and it was, I believe, the first systematic attempt to accomplish such a task) the laws which govern the extinction of species, with a view of showing that the slow but ceaseless variations now in progress in physical geography, together with the migration of plants and animals into new regions, must in the course of ages give rise to the occasional loss of some of them and eventually cause an entire fauna and flora to die out; also that we must infer from geological data that the places thus left vacant from time to time are filled up without delay by new forms adapted to new conditions, sometimes by immigration from adjoining provinces, sometimes by new creations. Among the many causes of extinction enumerated by me were the power of hostile species, diminution of food, mutations in climate, the conversion of land into sea and of sea into land, etc. I firmly opposed Brocchi's hypothesis of a decline in the vital energy of each species;* (* "Principles of Geology" 1st edition volume 2 chapter 8; and 9th edition page 668.) maintaining that there was every reason to believe that the reproductive powers of the last surviving representatives of a species were as vigorous as those of their predecessors, and that they were as capable, under favourable circumstances, of repeopling the earth with their kind. The manner in which some species are now becoming scarce and dying out, one after the other, appeared to me to favour the doctrine of the fixity of the specific character, showing a want of pliancy and capability of varying, which ensured their annihilation whenever changes adverse to their well-being occurred; time not being allowed for such a transformation as might be conceived capable of adapting them to the new circumstances, and of converting them into what naturalists would call new species.* (* Laws of Extinction, "Principles of Geology" 1st edition 1832 volume 2 chapters 5 to 11 inclusive; and 9th edition chapters 37 to 42 inclusive 1853.)
But while rejecting transmutation, I was equally opposed to the popular theory that the creative power had diminished in energy, or that it had been in abeyance ever since Man had entered upon the scene. That a renovating force which had been in full operation for millions of years should cease to act while the causes of extinction were still in full activity, or even intensified by the accession of Man's destroying power, seemed to me in the highest degree improbable. The only point on which I doubted was whether the force might not be intermittent instead of being, as Lamarck supposed, in ceaseless operation. Might not the births of new species, like the deaths of old ones, be sudden? Might they not still escape our observation? If the coming in of one new species, and the loss of one other which had endured for ages, should take place annually, still, assuming that there are a million of animals and plants living on the globe, it would require, I observed, a million of years to bring about a complete revolution in the fauna and flora. In that case, I imagined that, although the first appearance of a new form might be as abrupt as the disappearance of an old one, yet naturalists might never yet have witnessed the first entrance on the stage of a large and conspicuous animal or plant, and as to the smaller kinds, many of them may be conceived to have stolen in unseen, and to have spread gradually over a wide area, like species migrating into new provinces.* (* "Principles of Geology" 1st edition 1832 volume 2 chapter 11; and 9th edition page 706.)
It may now be useful to offer some remarks on the very different reception which the twin branches of Lamarck's development theory, namely, progression and transmutation, have met with, and to inquire into the causes of the popularity of the one and the great unpopularity of the other. We usually test the value of a scientific hypothesis by the number and variety of the phenomena of which it offers a fair or plausible explanation. If transmutation, when thus tested, has decidedly the advantage over progression and yet is comparatively in disfavour, we may reasonably suspect that its reception is retarded, not so much by its own inherent demerits, as by some apprehended consequences which it is supposed to involve and which run counter to our preconceived opinions.
THEORY OF PROGRESSION.
In treating of this question, I shall begin with the doctrine of progression, a concise statement of which, so far as it relates to the animal kingdom, was thus given twelve years ago by Professor Sedgwick, in the preface to his "Discourse on the Studies of the University of Cambridge."
"There are traces," he says, "among the old deposits of the earth of an organic progression among the successive forms of life. They are to be seen in the absence of mammalia in the older, and their very rare appearance in the newer Secondary groups; in the diffusion of warm-blooded quadrupeds (frequently of unknown genera) in the older Tertiary system, and in their great abundance (and frequently of known genera) in the upper portions of the same series; and lastly, in the recent appearance of Man on the surface of the earth."
"This historical development," continues the same author, of the forms and functions of organic life during successive epochs, "seems to mark a gradual evolution of creative power, manifested by a gradual ascent towards a higher type of being." "But the elevation of the fauna of successive periods was not made by transmutation, but by creative additions; and it is by watching these additions that we get some insight into Nature's true historical progress, and learn that there was a time when Cephalopoda were the highest types of animal life, the primates of this world; that Fishes next took the lead, then Reptiles; and that during the secondary period they were anatomically raised far above any forms of the reptile class now living in the world. Mammals were added next, until Nature became what she now is, by the addition of Man."* (* Professor Sedgwick's "Discourse on the Studies of the University of Cambridge" Preface to 5th edition pages 44, 154, 216, 1850.)
Although in the half century which has elapsed between the time of Lamarck and the publication of the above summary, new discoveries have caused geologists to assign a higher antiquity both to Man and the oldest fossil mammalia, fish, and reptiles than formerly, yet the generalisation, as laid down by the Woodwardian Professor, as to progression, still holds good in all essential particulars.
The progressive theory was propounded in the following terms by the late Hugh Miller in his "Footprints of the Creator."
"It is of itself an extraordinary fact without reference to other considerations, that the order adopted by Cuvier in his "Animal Kingdom," as that in which the four great classes of vertebrate animals, when marshalled according to their rank and standing, naturally range, should be also that in which they occur in order of time. The brain, which bears an average proportion to the spinal cord of not more than two to one, comes first—it is the brain of the fish; that which bears to the spinal cord an average proportion of two and a half to one succeeded it—it is the brain of the reptile; then came the brain averaging as three to one—it is that of the bird. Next in succession came the brain that averages as four to one—it is that of the mammal; and last of all there appeared a brain that averages as twenty-three to one—reasoning, calculating Man had come upon the scene."* (* "Footprints of the Creator" Edinburgh 1849 page 283.)
M. Agassiz, in his "Essay on Classification," has devoted a chapter to the "Parallelism between the Geological Succession of Animals and Plants and their present relative Standing;" in which he has expressed a decided opinion that within the limits of the orders of each great class there is a coincidence between their relative rank in organisation and the order of succession of their representatives in time.* (* "Contributions to the Natural History of the United States" Part 1.—Essay on Classification page 108.)
Professor Owen, in his Palaeontology, has advanced similar views, and has remarked, in regard to the vertebrata that there is much positive as well as negative evidence in support of the doctrine of an advance in the scale of being, from ancient to more modern geological periods. We observe, for example, in the Triassic, Oolitic, and Cretaceous strata, not only an absence of placental mammalia, but the presence of innumerable reptiles, some of large size, terrestrial and aquatic, herbivorous and predaceous, fitted to perform the functions now discharged by the mammalia.
The late Professor Bronn, of Heidelberg, after passing in review more than 24,000 fossil animals and plants, which he had classified and referred each to their geological position in his "Index Palaeontologicus," came to the conclusion that, in the course of time, there had been introduced into the earth more and more highly organised types of animal and vegetable life; the modern species being, on the whole, more specialised, i.e. having separate organs, or parts of the body, to perform different functions, which, in the earlier periods and in beings of simpler structure, were discharged in common by a single part or organ.
Professor Adolphe Brongniart, in an essay published in 1849 on the botanical classification and geological distribution of the genera of fossil plants,* (* Tableau des Genres de Vegetaux fossiles, etc. "Dictionnaire Universel d'Histoire Naturelle" Paris 1849.) arrives at similar results as to the progress of the vegetable world from the earliest periods to the present. He does not pretend to trace an exact historical series from the sea-weed to the fern, or from the fern again to the conifers and cycads, and lastly from those families to the palms and oaks, but he, nevertheless, points out that the cryptogamic forms, especially the acrogens, predominate among the fossils of the primary formations, the Carboniferous especially, while the gymnosperms or coniferous and cycadeous plants abound in all the strata, from the Trias to the Wealden inclusive; and lastly, the more highly developed angiosperms, both monocotyledonous and dicotyledonous, do not become abundant until the Tertiary period. It is a remarkable fact, as he justly observes, that the angiospermous exogens, which comprise four-fifths of living plants—a division to which all our native European trees, except the Coniferae, belong, and which embrace all the Compositae, Leguminosae, Umbelliferae, Cruciferae, Heaths, and so many other families—are wholly unrepresented by any fossils hitherto discovered in the Primary and Secondary formations from the Silurian to the Oolitic inclusive. It is not till we arrive at the Cretaceous period that they begin to appear, sparingly at first, and only playing a conspicuous part, together with the palms and other endogens, in the Tertiary epoch.
When commenting on the eagerness with which the doctrine of progression was embraced from the close of the last century to the time when I first attempted, in 1830, to give some account of the prevailing theories in geology, I observed that far too much reliance was commonly placed on the received dates of the first appearances of certain orders or classes of animals or plants, such dates being determined by the age of the stratum in which we then happened to have discovered the earliest memorials of such types. At that time (1830), it was taken for granted that Man had not co-existed with the mammoth and other extinct mammalia, yet now that we have traced back the signs of his existence to the Pleistocene era, and may anticipate the finding of his remains on some future day in the Pliocene period, the theory of progression is not shaken; for we cannot expect to meet with human bones in the Miocene formations, where all the species and nearly all the genera of mammalia belong to types widely differing from those now living; and had some other rational being, representing Man, then flourished, some signs of his existence could hardly have escaped unnoticed, in the shape of implements of stone or metal, more frequent and more durable than the osseous remains of any of the mammalia.
In the beginning of this century it was one of the canons of the popular geological creed that the first warm-blooded quadrupeds which had inhabited this planet were those derived from the Eocene gypsum of Montmartre in the suburbs of Paris, almost all of which Cuvier had shown to belong to extinct genera. This dogma continued in force for more than a quarter of a century, in spite of the discovery in 1818 of a marsupial quadruped in the Stonesfield strata, a member of the Lower Oolite, near Oxford. Some disputed the authority of Cuvier himself as to the mammalian character of the fossil; others, the accuracy of those who had assigned to it so ancient a place in the chronological series of rocks. In 1832 I pointed out that the occurrence of this single fossil in the Oolite was "fatal to the theory of successive development" as then propounded.* (* "Principles of Geology" 2nd edition 1 173.) Since that period great additions have been made to our knowledge of the existence of land quadrupeds in the olden times. We have ascertained that, in Eocene strata older than the gypsum of Paris, no less than four distinct sets of placental mammalia have flourished; namely, first, those of the Headon series in the Isle of Wight, from which fourteen species have been procured; secondly, those of the antecedent Bagshot and Bracklesham beds, which have yielded, together with the contemporaneous "calcaire grossier" of Paris, twenty species; thirdly, the still older beds of Kyson, near Ipswich, and those of Herne Bay, at the mouth of the Thames, in which seven species have been found; and fourthly, the Woolwich and Reading beds, which have supplied ten species.* (* Lyell's supplement to 5th edition of "Elements" 1857.)
We can scarcely doubt that we should already have traced back the evidence of this class of fossils much farther had not our inquiries been arrested, first by the vast gap between the Tertiary and Secondary formations, and then by the marine nature of the Cretaceous rocks.
The mammalia next in antiquity, of which we have any cognisance, are those of the Upper Oolite of Purbeck, discovered between the years 1854 and 1857, and comprising no less than fourteen species, referable to eight or nine genera; one of them, Plagiaulax, considered by Dr. Falconer to have been a herbivorous marsupial. The whole assemblage appear, from the joint observations of Professor Owen and Dr. Falconer, to indicate a low grade of quadruped, probably of the marsupial type. They were, for the most part, diminutive, the two largest not much exceeding our common hedgehog and polecat in size.
Next anterior in age are the mammalia of the Lower Oolite of Stonesfield, of which four species are known, also very small and probably marsupial, with one exception, the Stereognathus ooliticus, which, according to Professor Owen's conjecture, may have been a hoofed quadruped and placental, though, as we have only half of the lower jaw with teeth, and the molars are unlike any living type, such an opinion is of course hazarded with due caution.
Still older than the above are some fossil quadrupeds of small size, found in the Upper Trias of Stuttgart in Germany, and more lately by Mr. C. Moore in beds of corresponding age near Frome, which are also of a very low grade, like the living Myrmecobius of Australia. Beyond this limit our knowledge of the highest class of vertebrata does not as yet extend into the past, but the frequent shifting back of the old landmarks, nearly all of them once supposed in their turn to indicate the date of the first appearance of warm-blooded quadrupeds on this planet, should serve as a warning to us not to consider the goal at present reached by palaeontology as one beyond which they who come after us are never destined to pass.
On the other hand, it may be truly said in favour of progression that after all these discoveries the doctrine is not gainsaid, for the less advanced marsupials precede the more perfect placental mammalia in the order of their appearance on the earth.
If the three localities where the most ancient mammalia have been found—Purbeck, Stonesfield, and Stuttgart—had belonged all of them to formations of the same age, we might well have imagined so limited an area to have been peopled exclusively with pouched quadrupeds, just as Australia now is, while other parts of the globe were inhabited by placentals, for Australia now supports one hundred and sixty species of marsupials, while the rest of the continents and islands are tenanted by about seventeen hundred species of mammalia, of which only forty-six are marsupial, namely, the opossums of North and South America. But the great difference of age of the strata in each of these three localities seems to indicate the predominance throughout a vast lapse of time (from the era of the Upper Trias to that of the Purbeck beds) of a low grade of quadrupeds; and this persistency of similar generic and ordinal types in Europe while the species were changing, and while the fish, reptiles, and mollusca were undergoing vast modifications, raises a strong presumption that there was also a vast extension in space of the same marsupial forms during that portion of the Secondary epoch which has been termed "the age of reptiles."
As to the class Reptilia, some of the orders which prevailed when the Secondary rocks were formed are confessedly much higher in their organisation than any of the same class now living. If the less perfect ophidians, or snakes, which now abound on the earth had taken the lead in those ancient days among the land reptiles, and the Deinosaurians had been contemporary with Man, there can be no doubt that the progressionist would have seized upon this fact with unfeigned satisfaction as confirmatory of his views. Now that the order of succession is precisely reversed, and that the age of the Iguanodon was long anterior to that of the Eocene Palaeophis and living boa, while the crocodile is in our own times the highest representative of its class, a retrograde movement in this important division of the vertebrata must be admitted. It may perhaps be accounted for by the power acquired by the placental mammalia, when they became dominant, a power before which the class of vertebrata next below them, as coming most directly in competition with them, may more than any other have given way.
For no less than thirty-four years it had been a received axiom in palaeontology that reptiles had never existed before the Permian or Magnesian Limestone period, when at length in 1844 this supposed barrier was thrown down, and Carboniferous reptiles, terrestrial and aquatic, of several genera were brought to light; and discussions are now going on as to whether some remains of an Enaliosaur (perhaps a large Labyrinthodon) have not been detected in the coal of Nova Scotia, and whether certain sandstones near Elgin in Scotland, containing the bones of lacertian, crocodilian, and rhynchosaurian reptiles, may not be referable to the "Old Red" or Devonian group. Still, no traces of this class have yet been detected in rocks as ancient as those in which the oldest fish have been found. [Note 38.]
As to fossil representatives of the ichthyic type, the most ancient were not supposed before 1838 to be of a date anterior to the Coal, but they have since been traced back, first to the Devonian, and then to the Silurian rocks. No remains, however, of them or of any vertebrate animal have yet been discovered in the Ordovician strata, rich as these are in invertebrate fossils, nor in the still older Cambrian; so that we seem authorised to conclude, though not without considerable reserve, that the vertebrate type was extremely scarce, if not wholly wanting, in those epochs often spoken of as "primitive," but which, if the Development Theory be true, were probably the last of a long series of antecedent ages in which living beings flourished.
As to the Mollusca, which afford the most unbroken series of geological medals, the highest of that class, the Cephalopoda, abounded in older Silurian times, comprising several hundred species of chambered univalves. Had there been strong prepossessions against the progressive theory, it would probably have been argued that when these cephalopods abounded, and the siphonated gasteropods were absent, a higher order of zoophagous mollusca discharged the functions afterwards performed by an inferior order in the Secondary, Tertiary, and Post-Tertiary seas. But I have never seen this view suggested as adverse to the doctrine of progress, although much stress has been laid on the fact that the Silurian Brachiopoda, creatures of a lower grade, formerly discharged the functions of the existing lamellibranchiate bivalves, which are higher in the scale.
It is said truly that the Ammonite, Orthoceras, and Nautilus of these ancient rocks were of the tetrabranchiate division, and none of them so highly organised as the Belemnite and other dibranchiate cephalopods which afterwards appeared, and some of which now flourish in our seas. Therefore, we may infer that the simplest forms of the Cephalopoda took precedence of the more complex in time. But if we embrace this view, we must not forget that there are living Cephalopoda, such as the Octopods, which are devoid of any hard parts, whether external or internal, and which could leave behind them no fossil memorials of their existence, so that we must make a somewhat arbitrary assumption, namely, that at a remote era, no such Dibranchiata were in being, in order to avail ourselves of this argument in favour of progression. On the other hand, it is true that in the Lower Cambrian not even the shell-bearing tetrabranchiates have yet been discovered.
In regard to plants, although the generalisation above cited of M. Adolphe Brongniart is probably true, there has been a tendency in the advocates of progression to push the inferences deducible from known facts, in support of their favourite dogma, somewhat beyond the limits which the evidence justifies. Dr. Hooker observes, in his recent "Introductory Essay to the Flora of Australia," that it is impossible to establish a parallel between the successive appearances of vegetable forms in time, and their complexity of structure or specialisation of organs as represented by the successively higher groups in the natural method of classification. He also adds that the earliest recognisable Cryptogams are not only the highest now existing, but have more highly differentiated vegetative organs than any subsequently appearing, and that the dicotyledonous embryo and perfect exogenous wood, with the highest specialised tissue known (the coniferous with glandular tissue), preceded the monocotyledonous embryo and endogenous wood in date of appearance on the globe—facts wholly opposed to the doctrine of progression, and which can only be set aside on the supposition that they are fragmentary evidence of a time farther removed from the origin of vegetation than from the present day.* (* "Introductory Essay to the Flora of Australia," page 31 London 1859. Published separately.) [Note 39.]
It would be an easy task to multiply objections to the theory now under consideration; but from this I refrain, as I regard it not only as a useful, but rather in the present state of science as an indispensable hypothesis, and one which though destined hereafter to undergo many and great modifications will never be overthrown.
It may be thought almost paradoxical that writers who are most in favour of transmutation (Mr. C. Darwin and Dr. J. Hooker, for example) are nevertheless among those who are most cautious, and one would say timid, in their mode of espousing the doctrine of progression; while, on the other hand, the most zealous advocates of progression are oftener than not very vehement opponents of transmutation. We might have anticipated a contrary leaning on the part of both, for to what does the theory of progression point? It supposes a gradual elevation in grade of the vertebrate type in the course of ages from the most simple ichthyic form to that of the placental mammalia and the coming upon the stage last in the order of time of the most anthropomorphous mammalia, followed by the human race—this last thus appearing as an integral part of the same continuous series of acts of development, one link in the same chain, the crowning operation as it were of one and the same series of manifestations of creative power. If the dangers apprehended from transmutation arise from the too intimate connection which it tends to establish between the human and merely animal natures, it might have been expected that the progressive development of organisation, instinct, and intelligence might have been unpopular, as likely to pioneer the way for the reception of the less favoured doctrine. But the true explanation of the seeming anomaly is this, that no one can believe in transmutation who is not profoundly convinced that all we know in palaeontology is as nothing compared with what we have yet to learn, and they who regard the record as so fragmentary, and our acquaintance with the fragments which are extant as so rudimentary, are apt to be astounded at the confidence placed by the progressionists in data which must be defective in the extreme. But exactly in proportion as the completeness of the record and our knowledge of it are overrated, in that same degree are many progressionists unconscious of the goal towards which they are drifting. Their faith in the fullness of the annals leads them to regard all breaks in the series of organic existence, or in the sequence of the fossiliferous rocks, as proofs of original chasms and leaps in the course of nature—signs of the intermittent action of the creational force, or of catastrophes which devastated the habitable surface. They do not doubt that there is a continuity of plan, but they believe that it exists in the Divine mind alone, and they are therefore without apprehension that any facts will be discovered which would imply a material connection between the outgoing organisms and the incoming ones.
ON THE ORIGIN OF SPECIES BY VARIATION AND NATURAL SELECTION.
Mr. Darwin's Theory of the Origin of Species by Natural Selection. Memoir by Mr. Wallace. Manner in which favoured Races prevail in the Struggle for Existence. Formation of new Races by breeding. Hypotheses of definite and indefinite Modifiability equally arbitrary. Competition and Extinction of Races. Progression not a necessary Accompaniment of Variation. Distinct Classes of Phenomena which Natural Selection explains. Unity of Type, Rudimentary Organs, Geographical Distribution, Relation of the extinct to the living Fauna and Flora, and mutual Relations of successive Groups of Fossil Forms. Light thrown on Embryological Development by Natural Selection. Why large Genera have more variable Species than small ones. Dr. Hooker on the Evidence afforded by the Vegetable Kingdom in favour of Creation by Variation. Steenstrup on alternation of Generations. How far the Doctrine of Independent Creation is opposed to the Laws now governing the Migration of Species.
For many years after the promulgation of Lamarck's doctrine of progressive development, geologists were much occupied with the question whether the past changes in the animate and inanimate world were brought about by sudden and paroxysmal action, or gradually and continuously, by causes differing neither in kind nor degree from those now in operation.
The anonymous author of "The Vestiges of Creation" published in 1844 a treatise, written in a clear and attractive style, which made the English public familiar with the leading views of Lamarck on transmutation and progression, but brought no new facts or original line of argument to support those views, or to combat the principal objections which the scientific world entertained against them.
No decided step in this direction was made until the publication in 1858 of two papers, one by Mr. Darwin and another by Mr. Wallace, followed in 1859 by Mr. Darwin's celebrated work on "The Origin of Species by Means of Natural Selection; or, the Preservation of favoured Races in the Struggle for Life." The author of this treatise had for twenty previous years strongly inclined to believe that variation and the ordinary laws of reproduction were among the secondary causes always employed by the Author of nature, in the introduction from time to time of new species into the world, and he had devoted himself patiently to the collecting of facts and making of experiments in zoology and botany, with a view of testing the soundness of the theory of transmutation. Part of the manuscript of his projected work was read to Dr. Hooker as early as 1844 and some of the principal results were communicated to me on several occasions. [Note 40.] Dr. Hooker and I had repeatedly urged him to publish without delay, but in vain, as he was always unwilling to interrupt the course of his investigations; until at length Mr. Alfred R. Wallace, who had been engaged for years in collecting and studying the animals of the East Indian archipelago, thought out independently for himself one of the most novel and important of Mr. Darwin's theories. This he embodied in an essay "On the Tendency of Varieties to depart indefinitely from the original Type." It was written at Ternate in February 1858, and sent to Mr. Darwin with a request that it might be shown to me if thought sufficiently novel and interesting. Dr. Hooker and I were of opinion that it should be immediately printed, and we succeeded in persuading Mr. Darwin to allow one of the manuscript chapters of his "Origin of Species," entitled "On the Tendency of Species to form Varieties, and on the Perpetuation of Species and Varieties by natural Means of Selection," to appear at the same time.* (* See "Proceedings of the Linnaean Society" 1858.)
By reference to these memoirs it will be seen that both writers begin by applying to the animal and vegetable worlds the Malthusian doctrine of population, or its tendency to increase in a geometrical ratio, while food can only be made to augment even locally in an arithmetical one. There being therefore no room or means of subsistence for a large proportion of the plants and animals which are born into the world, a great number must annually perish. Hence there is a constant struggle for existence among the individuals which represent each species and the vast majority can never reach the adult state, to say nothing of the multitudes of ova and seeds which are never hatched or allowed to germinate. Of birds it is estimated that the number of those which die every year equals the aggregate number by which the species to which they respectively belong is on the average permanently represented.
The trial of strength which must decide what individuals are to survive and what to succumb occurs in the season when the means of subsistence are fewest, or enemies most numerous, or when the individuals are enfeebled by climate or other causes; and it is then that those varieties which have any, even the slightest, advantage over others come off victorious. They may often owe their safety to what would seem to a casual observer a trifling difference, such as a darker or lighter shade of colour rendering them less visible to a species which preys upon them, or sometimes to attributes more obviously advantageous, such as greater cunning or superior powers of flight or swiftness of foot. These peculiar qualities and faculties, bodily and instinctive, may enable them to outlive their less favoured rivals, and being transmitted by the force of inheritance to their offspring will constitute new races, or what Mr. Darwin calls "incipient species." If one variety, being in other respects just equal to its competitors, happens to be more prolific, some of its offspring will stand a greater chance of being among those which will escape destruction, and their descendants, being in like manner very fertile, will continue to multiply at the expense of all less prolific varieties.
As breeders of domestic animals, when they choose certain varieties in preference to others to breed from, speak technically of their method as that of "selecting," Mr. Darwin calls the combination of natural causes, which may enable certain varieties of wild animals or plants to prevail over others of the same species, "natural selection."
A breeder finds that a new race of cattle with short horns or without horns may be formed in the course of several generations by choosing varieties having the most stunted horns as his stock from which to breed; so nature, by altering in the course of ages, the conditions of life, the geographical features of a country, its climate, the associated plants and animals, and consequently the food and enemies of a species and its mode of life, may be said, by this means to select certain varieties best adapted for the new state of things. Such new races may often supplant the original type from which they have diverged, although that type may have been perpetuated without modification for countless anterior ages in the same region, so long as it was in harmony with the surrounding conditions then prevailing.
Lamarck, when speculating on the origin of the long neck of the giraffe, imagined that quadruped to have stretched himself up in order to reach the boughs of lofty trees, until by continued efforts and longing to reach higher he obtained an elongated neck. Mr. Darwin and Mr. Wallace simply suppose that, in a season of scarcity, a longer-necked variety, having the advantage in this respect over most of the herd, as being able to browse on foliage out of their reach, survived them and transmitted its peculiarity of cervical conformation to its successors.
By the multiplying of slight modifications in the course of thousands of generations and by the handing down of the newly-acquired peculiarities by inheritance, a greater and greater divergence from the original standard is supposed to be effected, until what may be called a new species, or in a greater lapse of time a new genus will be the result.
Every naturalist admits that there is a general tendency in animals and plants to vary; but it is usually taken for granted, though he have no means of proving the assumption to be true, that there are certain limits beyond which each species cannot pass under any circumstances or in any number of generations. Mr. Darwin and Mr. Wallace say that the opposite hypothesis, which assumes that every species is capable of varying indefinitely from its original type, is not a whit more arbitrary, and has this manifest claim to be preferred, that it will account for a multitude of phenomena which the ordinary theory is incapable of explaining.
We have no right, they say, to assume, should we find that a variable species can no longer be made to vary in a certain direction, that it has reached the utmost limit to which it might under more favourable conditions or if more time were allowed be made to diverge from the parent type.
Hybridisation is not considered by Mr. Darwin as a cause of new species, but rather as tending to keep variation within bounds. Varieties which are nearly allied cross readily with each other, and with the parent stock, and such crossing tends to keep the species true to its type, while forms which are less nearly related, although they may intermarry, produce no mule offspring capable of perpetuating their kind.
The competition of races and species, observes Mr. Darwin, is always most severe between those which are most closely allied and which fill nearly the same place in the economy of nature. Hence when the conditions of existence are modified the original stock runs great risk of being superseded by some one of its modified offshoots. The new race or species may not be absolutely superior in the sum of its powers and endowments to the parent stock, and may even be more simple in structure and of a lower grade of intelligence, as well as of organisation, provided on the whole it happens to have some slight advantage over its rivals. Progression, therefore, is not a necessary accompaniment of variation and natural selection, though when a higher organisation happens to be coincident with superior fitness to new conditions, the new species will have greater power and a greater chance of permanently maintaining and extending its ground. One of the principal claims of Mr. Darwin's theory to acceptance is that it enables us to dispense with a law of progression as a necessary accompaniment of variation. It will account equally well for what is called degradation, or a retrograde movement towards a simpler structure, and does not require Lamarck's continual creation of monads; for this was a necessary part of his system, in order to explain how, after the progressive power had been at work for myriads of ages, there were as many beings of the simplest structure in existence as ever.
Mr. Darwin argues, and with no small success, that all true classification in zoology and botany is in fact genealogical, and that community of descent is the hidden bond which naturalists have been unconsciously seeking, while they often imagined that they were looking for some unknown plan of creation.
As the "Origin of Species"* (* "Origin of Species" page 121.) is in itself a condensed abstract of a much larger work not yet published [Note 41] I could not easily give an analysis of its contents within narrower limits than those of the original, but it may be useful to enumerate briefly some of the principal classes of phenomena on which the theory of "natural selection" would throw light.
In the first place it would explain, says Mr. Darwin, the unity of type which runs through the whole organic world, and why there is sometimes a fundamental agreement in structure in the same class of beings which is quite independent of their habits of life, for such structure, derived by inheritance from a remote progenitor, has been modified in the course of ages in different ways according to the conditions of existence. It would also explain why all living and extinct beings are united, by complex radiating and circuitous lines of affinity with one another into one grand system;* (* "Origin" page 498.) also, there having been a continued extinction of old races and species in progress and a formation of new ones by variation, why in some genera which are largely represented, or to which a great many species belong, many of these are closely but unequally related; also, why there are distinct geographical provinces of species of animals and plants, for after long isolation by physical barriers each fauna and flora by varying continually must become distinct from its ancestral type, and from the new forms assumed by other descendants which have diverged from the same stock.
The theory of indefinite modification would also explain why rudimentary organs are so useful in classification, being the remnants preserved by inheritance of organs which the present species once used—as in the case of the rudiments of eyes in insects and reptiles inhabiting dark caverns, or of the wings of birds and beetles which have lost all power of flight. In such cases the affinities of species are often more readily discerned by reference to these imperfect structures than by others of much more physiological importance to the individuals themselves.
The same hypothesis would explain why there are no mammalia in islands far from continents, except bats, which can reach them by flying; and also why the birds, insects, plants, and other inhabitants of islands, even when specifically unlike, usually agree generically with those of the nearest continent, it being assumed that the original stock of such species came by migration from the nearest land.
Variation and natural selection would also afford a key to a multitude of geological facts otherwise wholly unaccounted for, as for example why there is generally an intimate connection between the living animals and plants of each great division of the globe and the extinct fauna and flora of the Post-Tertiary or Tertiary formations of the same region; as, for example, in North America, where we not only find among the living mollusca peculiar forms foreign to Europe, such as Gnathodon and Fulgur (a subgenus of Fusus), but meet also with extinct species of those same genera in the Tertiary fauna of the same part of the world. In like manner, among the mammalia we find in Australia not only living kangaroos and wombats, but fossil individuals of extinct species of the same genera. So also there are recent and fossil sloths, armadilloes and other Edentata in South America, and living and extinct species of elephant, rhinoceros, tiger, and bear in the great Europeo-Asiatic continent. The theory of the origin of new species by variation will also explain why a species which has once died out never reappears and why the fossil fauna and flora recede farther and farther from the living type in proportion as we trace them back to remoter ages. It would also account for the fact that when we have to intercalate a new set of fossiliferous strata between two groups previously known, the newly discovered fossils serve to fill up gaps between specific or generic types previously familiar to us, supplying often the missing links of the chain, which, if transmutation is accepted, must once have been continuous.
One of the most original speculations in Mr. Darwin's work is derived from the fact that, in the breeding of animals, it is often observed that at whatever age any variation first appears in the parent, it tends to reappear at a corresponding age in the offspring. Hence the young individuals of two races which have sprung from the same parent stock are usually more like each other than the adults. Thus the puppies of the greyhound and bull-dog are much more nearly alike in their proportions than the grown-up dogs, and in like manner the foals of the carthorse and racehorse than the adult individuals. For the same reason we may understand why the species of the same genus, or genera of the same family, resemble each other more nearly in their embryonic than in their more fully developed state, or how it is that in the eyes of most naturalists the structure of the embryo is even more important in classification than that of the adult, "for the embryo is the animal in its less modified state, and in so far it reveals the structure of its progenitor. In two groups of animals, however much they may at present differ from each other in structure and habits, if they pass through the same or similar embryonic stages, we may feel assured that they have both descended from the same or nearly similar parents, and are therefore in that degree closely related. Thus community in embryonic structure reveals community of descent, however much the structure of the adult may have been modified."* (* Darwin, "Origin" etc. page 448.)
If then there had been a system of progressive development, the successive changes through which the embryo of a species of a high class, a mammifer for example, now passes, may be expected to present us with a picture of the stages through which, in the course of ages, that class of animals has successively passed in advancing from a lower to a higher grade. Hence the embryonic states exhibited one after the other by the human individual bear a certain amount of resemblance to those of the fish, reptile, and bird before assuming those of the highest division of the vertebrata.
Mr. Darwin, after making a laborious analysis of many floras, found that those genera which are represented by a large number of species contain a greater number of variable species, relatively speaking, than the smaller genera or those less numerously represented. This fact he adduces in support of his opinion that varieties are incipient species, for he observes that the existence of the larger genera implies that the manufacturing of species has been active in the period immediately preceding our own, in which case we ought generally to find the same forces still in full activity, more especially as we have every reason to believe the process by which new species are produced is a slow one.* (* "Origin of Species" chapter 2 page 56.)
Dr. Hooker tells us that he was long disposed to doubt this result, as he was acquainted with so many variable small genera, but after examining Mr. Darwin's data, he was compelled to acquiesce in his generalisation.* (* "Introductory Essay to the Flora of Australia" page 6.)
It is one of those conclusions, to verify which requires the investigation of many thousands of species, and to which exceptions may easily be adduced both in the animal and vegetable kingdoms, so that it will be long before we can expect it to be thoroughly tested, and if true, fairly appreciated. Among the most striking exceptions will be some genera still large, but which are beginning to decrease, the conditions favourable to their former predominance having already begun to change. To many, this doctrine of "natural selection," or "the preservation of favoured races in the struggle for life," seems so simple, when once clearly stated, and so consonant with known facts and received principles, that they have difficulty in conceiving how it can constitute a great step in the progress of science. Such is often the case with important discoveries, but in order to assure ourselves that the doctrine was by no means obvious, we have only to refer back to the writings of skilful naturalists who attempted in the earlier part of the nineteenth century to theorise on this subject, before the invention of this new method of explaining how certain forms are supplanted by new ones and in what manner these last are selected out of innumerable varieties and rendered permanent.
DR. HOOKER ON THE THEORY OF "CREATION BY VARIATION" AS APPLIED TO THE VEGETABLE KINGDOM.
Of Dr. Hooker, whom I have often cited in this chapter, Mr. Darwin has spoken in the Introduction to his "Origin of Species," as one "who had, for fifteen years, aided him in every possible way, by his large stores of knowledge, and his excellent judgment." This distinguished botanist published his "Introductory Essay to the Flora of Australia" in December 1859, the year after the memoir on "Natural Selection" was communicated to the Linnaean Society, and a month after the appearance of the "Origin of Species."
Having, in the course of his extensive travels, studied the botany of arctic, temperate, and tropical regions, and written on the flora of India, which he had examined at all heights above the sea from the plains of Bengal to the limits of perpetual snow in the Himalaya, and having specially devoted his attention to "geographical varieties," or those changes of character which plants exhibit when traced over wide areas and seen under new conditions; being also practically versed in the description and classification of new plants, from various parts of the world, and having been called upon carefully to consider the claims of thousands of varieties to rank as species, no one was better qualified by observation and reflection to give an authoritative opinion on the question, whether the present vegetation of the globe is or is not in accordance with the theory which Mr. Darwin has proposed. We cannot but feel, therefore, deeply interested when we find him making the following declaration:
"The mutual relations of the plants of each great botanical province, and, in fact, of the world generally, is just such as would have resulted if variation had gone on operating throughout indefinite periods, in the same manner as we see it act in a limited number of centuries, so as gradually to give rise in the course of time, to the most widely divergent forms."
In the same essay, this author remarks, "The element of mutability pervades the whole Vegetable Kingdom; no class, nor order, nor genus of more than a few species claims absolute exemption from it, whilst the grand total of unstable forms, generally assumed to be species, probably exceeds that of the stable." Yet he contends that species are neither visionary, nor even arbitrary creations of the naturalist, but realities, though they may not remain true for ever. The majority of them, he remarks, are so far constant, "within the range of our experience," and their forms and characters so faithfully handed down through thousands of generations, that they admit of being treated as if they were permanent and immutable. But the range of "our experience" is so limited, that it will "not account for a single fact in the present geographical distribution, or origin of any one species of plant, nor for the amount of variation it has undergone, nor will it indicate the time when it first appeared, nor the form it had when created."* (* Hooker, "Introductory Essay to the Flora of Australia.")
To what an extent the limits of species are indefinable, is evinced, he says, by the singular fact that, among those botanists who believe them to be immutable, the number of flowering plants is by some assumed to be 80,000, and by others over 150,000. The general limitation of species to certain areas suggests the idea that each of them, with all their varieties, have sprung from a common parent and have spread in various directions from a common centre. The frequency also of the grouping of genera within certain geographical limits is in favour of the same law, although the migration of species may sometimes cause apparent exceptions to the rule and make the same types appear to have originated independently at different spots.* (* Ibid. page 13.)
Certain genera of plants, which, like the brambles, roses, and willows in Europe, consist of a continuous series of varieties between the terms of which no intermediate forms can be intercalated, may be supposed to be newer types and on the increase, and therefore undergoing much variation; whereas genera which present no such perplexing gradations may be of older date and may have been losing species and varieties by extinction. In this case, the annihilation of intermediate forms which once existed makes it an easy task to distinguish those which remain.
It had usually been supposed by the advocates of the immutability of species that domesticated races, if allowed to run wild, always revert to their parent type. Mr. Wallace had said in reply that a domesticated species, if it loses the protection of Man, can only stand its ground in a wild state by resuming those habits and recovering those attributes which it may have lost when under domestication. If these faculties are so much enfeebled as to be irrecoverable it will perish; if not and if it can adapt itself to the surrounding conditions, it will revert to the state in which Man first found it: for in one, two, or three thousand years, which may have elapsed since it was originally tamed, there will not have been time for such geographical, climatal, and organic changes as would only be suited to a new race or a new and allied species.
But in regard to plants Dr. Hooker questions the fact of reversion. According to him, species in general do not readily vary, but when they once begin to do so the new varieties, as every horticulturist knows, show a great inclination to go on departing more and more from the old stock. As the best marked varieties of a wild species occur on the confines of the area which it inhabits, so the best marked varieties of a cultivated plant are those last produced by the gardener. Cabbages, for example, wall fruits, and cereal, show no disposition, when neglected, to assume the characters of the wild states of these plants. Hence the difficulty of determining what are the true parent species of most of our cultivated plants. Thus the finer kinds of apples, if grown from seed, degenerate and become crabs, but in so doing they do not revert to the original wild crab-apple, but become crab states of the varieties to which they belong.* (* "Introductory Essay to the Flora of Australia" page 9.)
It would lead me into too long a digression were I to attempt to give a fuller analysis of this admirable essay; but I may add that none of the observations are more in point, as bearing on the doctrine of what Hooker terms "creation by variation," than the great extent to which the internal characters and properties of plants, or their physiological constitution, are capable of being modified, while they exhibit externally no visible departure from the normal form. Thus, in one region a species may possess peculiar medicinal qualities which it wants in another, or it may be hardier and better able to resist cold. The average range in altitude, says Hooker, of each species of flowering plant in the Himalayan Mountains, whether in the tropical, temperate, or Alpine region, is 4000 feet, which is equivalent to twelve degrees of isothermals of latitude. If an individual of any of these species be taken from the upper limits of its range and carried to England, it is found to be better able to stand our climate than those from the lower or warmer stations. When several of these internal or physiological modifications are accompanied by variation in size, habits of growth, colour of the flowers, and other external characters, and these are found to be constant in successive generations, botanists may well begin to differ in opinion as to whether they ought to regard them as distinct species or not.
ALTERNATION OF GENERATIONS.
Hitherto, no rival hypothesis has been proposed as a substitute for the doctrine of transmutation; for what we term "independent creation," or the direct intervention of the Supreme Cause, must simply be considered as an avowal that we deem the question to lie beyond the domain of science.
The discovery by Steenstrup of alternate generation enlarges our views of the range of metamorphosis through which a species may pass, so that some of its stages (as when a Sertularia and a Medusa interchange) deviate so far from others as to have been referred by able zoologists to distinct genera, or even families. But in all these cases the organism, after running through a certain cycle of change, returns to the exact point from which it set out, and no new form or species is thereby introduced into the world. The only secondary cause therefore which has as yet been even conjecturally brought forward, to explain how in the ordinary course of nature a new specific form may be generated is, as Lamarck declared, "variation," and this has been rendered a far more probable hypothesis by the way in which "natural selection" is shown to give intensity and permanency to certain varieties.
When I formerly advocated the doctrine that species were primordial creations and not derivative, I endeavoured to explain the manner of their geographical distribution, and the affinity of living forms to the fossil types nearest akin to them in the Tertiary strata of the same part of the globe, by supposing that the creative power, which originally adapts certain types to aquatic and others to terrestrial conditions, has at successive geological epochs introduced new forms best suited to each area and climate, so as to fill the places of those which may have died out.
In that case, although the new species would differ from the old (for these would not be revived, having been already proved by the fact of their extinction to be incapable of holding their ground), still they would resemble their predecessors generically. For, as Mr. Darwin states in regard to new races, those of a dominant type inherit the advantages which made their parent species flourish in the same country, and they likewise partake in those general advantages which made the genus to which the parent species belonged a large genus in its own country.
We might therefore, by parity of reasoning, have anticipated that the creative power, adapting the new types to the new combination of organic and inorganic conditions of a given region, such as its soil, climate, and inhabitants, would introduce new modifications of the old types—marsupials, for example, in Australia, new sloths and armadilloes in South America, new heaths at the Cape, new roses in the northern and new calceolarias in the southern hemisphere. But to this line of argument Mr. Darwin and Dr. Hooker reply that when animals or plants migrate into new countries, whether assisted by man or without his aid, the most successful colonisers appertain by no means to those types which are most allied to the old indigenous species. On the contrary it more frequently happens that members of genera, orders, or even classes, distinct and foreign to the invaded country, make their way most rapidly and become dominant at the expense of the endemic species. Such is the case with the placental quadrupeds in Australia, and with horses and many foreign plants in the pampas of South America, and numberless instances in the United States and elsewhere which might easily be enumerated. Hence the transmutationists infer that the reason why these foreign types, so peculiarly fitted for these regions, have never before been developed there is simply that they were excluded by natural barriers. But these barriers of sea or desert or mountain could never have been of the least avail had the creative force acted independently of material laws or had it not pleased the Author of Nature that the origin of new species should be governed by some secondary causes analogous to those which we see preside over the appearance of new varieties, which never appear except as the offspring of a parent stock very closely resembling them.
OBJECTIONS TO THE HYPOTHESIS OF TRANSMUTATION CONSIDERED.
Statement of Objections to the Hypothesis of Transmutation founded on the Absence of Intermediate Forms. Genera of which the Species are closely allied. Occasional Discovery of the missing Links in a Fossil State. Davidson's Monograph on the Brachiopoda. Why the Gradational Forms, when found, are not accepted as Evidence of Transmutation. Gaps caused by Extinction of Races and Species. Vast Tertiary Periods during which this Extinction has been going on in the Fauna and Flora now existing. Genealogical Bond between Miocene and Recent Plants and Insects. Fossils of Oeningen. Species of Insects in Britain and North America represented by distinct Varieties. Falconer's Monograph on living and fossil Elephants. Fossil Species and Genera of the Horse Tribe in North and South America. Relation of the Pliocene Mammalia of North America, Asia, and Europe. Species of Mammalia, though less persistent than the Mollusca, change slowly. Arguments for and against Transmutation derived from the Absence of Mammalia in Islands. Imperfection of the Geological Record. Intercalation of newly discovered Formation of intermediate Age in the chronological Series. Reference of the St. Cassian Beds to the Triassic Periods. Discovery of new organic Types. Feathered Archaeopteryx of the Oolite.
THEORY OF TRANSMUTATION—ABSENCE OF INTERMEDIATE LINKS.
The most obvious and popular of the objections urged against the theory of transmutation may be thus expressed: If the extinct species of plants and animals of the later geological periods were the progenitors of the living species, and gave origin to them by variation and natural selection, where are all the intermediate forms, fossil and living, through which the lost types must have passed during their conversion into the living ones? And why do we not find almost everywhere passages between the nearest allied species and genera, instead of such strong lines of demarcation and often wide intervening gaps?
We may consider this objection under two heads:—
First. To what extent are the gradational links really wanting in the living creation or in the fossil world, and how far may we expect to discover such as are missing by future research?
Secondly. Are the gaps more numerous than we ought to anticipate, allowing for the original defective state of the geological records, their subsequent dilapidation and our slight acquaintance with such parts of them as are extant, and allowing also for the rate of extinction of races and species now going on, and which has been going on since the commencement of the Tertiary period?
First. As to the alleged absence of intermediate varieties connecting one species with another, every zoologist and botanist who has engaged in the task of classification has been occasionally thrown into this dilemma—if I make more than one species in this group, I must, to be consistent, make a great many. Even in a limited region like the British Isles this embarrassment is continually felt.
Scarcely any two botanists, for example, can agree as to the number of roses, still less as to how many species of bramble we possess. Of the latter genus, Rubus, there is one set of forms respecting which it is still a question whether it ought to be regarded as constituting three species or thirty-seven. Mr. Bentham adopts the first alternative and Mr. Babington the second, in their well-known treatises on British plants.
We learn from Dr. Hooker that at the antipodes, both in New Zealand and Australia, this same genus Rubus is represented by several species rich in individuals and remarkable for their variability. When we consider how, as we extend our knowledge of the same plant over a wider area, new geographical varieties commonly present themselves, and then endeavour to imagine the number of forms of the genus Rubus which may now exist, or probably have existed, in Europe and in regions intervening between Europe and Australia, comprehending all which may have flourished in Tertiary and Post-Tertiary periods, we shall perceive how little stress should be laid on arguments founded on the assumed absence of missing links in the flora as it now exists.
If in the battle of life the competition is keenest between closely allied varieties and species, as Mr. Darwin contends, many forms can never be of long duration, nor have a wide range, and these must often pass away without leaving behind them any fossil memorials. In this manner we may account for many breaks in the series which no future researches will ever fill up.
DAVIDSON ON FOSSIL BRACHIOPODA.
It is from fossil conchology more than from any other department of the organic world that we may hope to derive traces of a transition from certain types to others, and fossil memorials of all the intermediate shades of form. We may especially hope to gain this information from the study of some of the lower groups, such as the Brachiopoda, which are persistent in type, so that the thread of our inquiry is less likely to be interrupted by breaks in the sequence of the fossiliferous rocks. The splendid monograph just concluded by Mr. Davidson on the British Brachiopoda, illustrates, in the first place, the tendency of certain generic forms in this division of the mollusca to be persistent throughout the whole range of geological time yet known to us; for the four genera, Rhynchonella, Crania, Discina, and Lingula, have been traced through the Silurian, Devonian, Carboniferous, Permian, Jurassic, Cretaceous, Tertiary, and Recent periods, and still retain in the existing seas the identical shape and character which they exhibited in the earliest formations. On the other hand, other Brachiopoda have gone through in shorter periods a vast series of transformations, so that distinct specific and even generic names have been given to the same varying form, according to the different aspects and characters it has put on in successive sets of strata.
In proportion as materials of comparison have accumulated, the necessity of uniting species previously regarded as distinct under one denomination has become more and more apparent. Mr. Davidson, accordingly, after studying not less than 260 reputed species from the British Carboniferous rocks, has been obliged to reduce that number to 100, to which he has added 20 species either entirely new or new to the British strata; but he declares his conviction that, when our knowledge of these 120 Brachiopoda is more complete, a further reduction of species will take place.
Speaking of one of these forms, which he calls Spirifer trigonalis, he says that it is so dissimilar to another extreme of the series, S. crassa, that in the first part of his memoir (published some ten years ago) he described them as distinct, and the idea of confounding them together must, he admits, appear absurd to those who have never seen the intermediate links, such as are presented by S. bisulcata, and at least four others with their varieties, most of them shells formerly recognised as distinct by the most eminent palaeontologists, but respecting which these same authorities now agree with Mr. Davidson in uniting them into one species.* (* "Monograph on British Brachiopoda" Palaeontographical Society page 222.)
The same species has sometimes continued to exist under slightly modified forms throughout the whole of the Ordovician and Silurian as well as the entire Devonian and Carboniferous periods, as in the case of the shell generally known as Leptaena rhomboidalis, Wahlenberg. No less than fifteen commonly received species are demonstrated by Mr. Davidson by the aid of a long series of transitional forms, to appertain to this one type; and it is acknowledged by some of the best writers that they were induced on purely theoretical grounds to give distinct names to some of the varieties now suppressed, merely because they found them in rocks so widely remote in time that they deemed it contrary to analogy to suppose that the same species could have endured so long: a fallacious mode of reasoning, analogous to that which leads some zoologists and botanists to distinguish by specific names slight varieties of living plants and animals met with in very remote countries, as in Europe and Australia, for example; it being assumed that each species has had a single birthplace or area of creation, and that they could not by migration have gone from the northern to the southern hemisphere across the intervening tropics.
Examples are also given by Mr. Davidson of species which pass from the Devonian into the Carboniferous, and from that again into the Permian rocks. The vast longevity of such specific forms has not been generally recognised in consequence of the change of names which they have undergone when derived from such distant formations, as when Atrypa unguicularis assumes, when derived from a Carboniferous rock, the name of Spirifer Urei, besides several other synonyms, and then, when it reaches the Permian period, takes the name of Spirifer Clannyana, King; all of which forms the author of the monograph, now under consideration, asserts to be one and the same.
No geologist will deny that the distance of time which separates some of the eras above alluded to, or the dates of the earliest and latest appearances of some of the fossils above mentioned, must be reckoned by millions of years. According to Mr. Darwin's views, it is only by having at our command the records of such enormous periods that we can expect to be able to point out the gradations which unite very distinct specific forms. But the advocate of transmutation must not be disappointed if, when he has succeeded in obtaining some of the proofs which he was challenged to produce, they make no impression on the mind of his opponent. All that will be conceded is that specific variation in the Brachiopoda, at least, has a wider range than was formerly suspected. So long as several allied species were brought nearer and nearer to each other, considerable uneasiness might have been felt as to the reality of species in general, but when fifteen or more are once fairly merged in one group, constituting in the aggregate a single species, one and indivisible, and capable of being readily distinguished from every other group at present known, all misgivings are at an end. Implicit trust in the immutability of species is then restored, and the more insensible the shades from one extreme to the other, in a word, the more complete the evidence of transition, the more nugatory does the argument derived from it appear. It then simply resolves itself into one of those exceptional instances of what is called a protean form.
Thirty years ago a great London dealer in shells, himself an able naturalist, told me that there was nothing he had so much reason to dread, as tending to depreciate his stock in trade, as the appearance of a good monograph on some large genus of mollusca; for, in proportion as the work was executed in a philosophical spirit, it was sure to injure him, every reputed species pronounced to be a mere variety becoming from that time unsaleable. Fortunately, so much progress has since been made in England in estimating the true ends and aims of science, that specimens indicating a passage between forms usually separated by wide gaps, whether in the Recent or fossil fauna, are eagerly sought for, and often more prized than the mere normal or typical forms.
It is clear that the more ancient the existing mollusca, or the farther back into the past we can trace the remains of shells still living, the more easy it becomes to reconcile with the doctrine of transmutation the distinctness in character of the majority of living species. For, what we want is time, first, for the gradual formation, and then for the extinction of races and allied species, occasioning gaps between the survivors.
In the year 1830 I announced, on the authority of M. Deshayes, that about one-fifth of the mollusca of the Falunian or Upper Miocene strata of Europe, belonged to living species. Although the soundness of that conclusion was afterwards called in question by two or three eminent conchologists (and by the late M. Alcide d'Orbigny among others), it has since been confirmed by the majority of living naturalists and is well borne out by the copious evidence on the subject laid before the public in the magnificent work edited by Dr. Hoernes, and published under the auspices of the Austrian Government, "On the Fossil Shells of the Vienna Basin."
The collection of Tertiary shells from which those descriptions and beautiful figures were taken is almost unexampled for the fine state of preservation of the specimens, and the care with which all the varieties have been compared. It is now admitted that about one-third of these Miocene forms, univalves and bivalves included, agree specifically with living mollusca, so that much more than the enormous interval which divides the Miocene from the Recent period must be taken into our account when we speculate on the origin by transmutation of the shells now living, and the disappearance by extinction of intermediate varieties and species.
MIOCENE PLANTS AND INSECTS RELATED TO RECENT SPECIES.
Geologists were acquainted with about three hundred species of marine shells from the Falunian strata on the banks of the Loire, before they knew anything of the contemporary insects and plants. At length, as if to warn us against inferring from negative evidence the poverty of any ancient set of strata in organic remains proper to the land, a rich flora and entomological fauna was suddenly revealed to us characteristic of Central Europe during the Upper Miocene period. This result followed the determination of the true position of the Oeningen beds in Switzerland, and of certain formations of "Brown Coal" in Germany.
Professor Heer, who has described nearly five hundred species of fossil plants from Oeningen, besides many more from other Miocene localities in Switzerland,* (* Heer, "Flora tertiaria Helvetiae" 1859; and Gaudin's French translation, with additions, 1861.) estimates the phanerogamous species which must have flourished in Central Europe at that time at 3000, and the insects as having been more numerous in the same proportion as they now exceed the plants in all latitudes. This European Miocene flora was remarkable for the preponderance of arborescent and shrubby evergreens, and comprised many generic types no longer associated together in any existing flora or geographical province. Some genera, for example, which are at present restricted to America, co-existed in Switzerland with forms now peculiar to Asia, and with others at present confined to Australia.
Professor Heer has not ventured to identify any of this vast assemblage of Miocene plants and insects with living species, so far at least as to assign to them the same specific names, but he presents us with a list of what he terms homologous forms, which are so like the living ones that he supposes the one to have been derived genealogically from the others. He hesitates indeed as to the manner of the transformation or the precise nature of the relationship, "whether the changes were brought about by some influence exerted continually for ages, or whether at some given moment the old types were struck with a new image."
Among the homologous plants alluded to are forty species, of which both the leaves and fruits are preserved, and thirty others, known at present by their leaves only. In the first list we find many American types, such as the tulip tree (Liriodendron), the deciduous cypress (Taxodium), the red maple and others, together with Japanese forms, such as a cinnamon, which is very abundant. And what is worthy of notice, some of these fossils so closely allied to living plants occur not only in the Upper, but even some few of them as far back in time as the Lower Miocene formations of Switzerland and Germany, which are probably as distant from the Upper Miocene or Oeningen beds as are the latter from our own era.
Some of the fossil plants to which Professor Heer has given new names have been regarded as Recent species by other eminent naturalists. Thus, one of the trees allied to the elm Unger had called Planera Richardi, a species which now flourishes in the Caucasus and Crete. Professor Heer had attempted to distinguish it from the living tree by the greater size of its fruit, but this character he confessed did not hold good, when he had an opportunity (1861) of comparing all the varieties of the living Planera Richardi which Dr. Hooker laid before him in the rich herbarium of Kew.
As to the "homologous insects" of the Upper Miocene period in Switzerland, we find among them, mingled with genera now wholly foreign to Europe, some very familiar forms, such as the common glowworm, Lampyris noctiluca, Linn., the dung-beetle, Geotrupes stercorarius, Linn., the ladybird, Coccinella septempunctata, Linn., the ear-wig, Forficula auricularia, Linn., some of our common dragon-flies, as Libellula depressa, Linn., the honey-bee, Apis mellifera, Linn., the cuckoo spittle insect, Aphrophora spumaria, Linn., and a long catalogue of others, to all of which Professor Heer had given new names, but which some entomologists may regard as mere varieties until some stronger reasons are adduced for coming to a contrary opinion.
Several of the insects above enumerated, like the common ladybird, are well known at present to have a very wide range over nearly the whole of the Old World, for example, without varying, and might therefore be expected to have been persistent throughout many successive changes of the earth's surface and climate. Yet we may fairly anticipate that even the most constant types will have undergone some modifications in passing from the Miocene to the Recent epoch, since in the former period the geography and climate of Europe, the height of the Alps, and the general fauna and flora were so different from what they now are. But the deviation may not exceed that which would generally be expressed by what is called a well-marked variety.
Before I pass on to another topic, it may be well to answer a question which may have occurred to the reader; how it happens that we remained so long ignorant of the vegetation and insects of the Upper Miocene period in Europe? The answer may be instructive to those who are in the habit of underrating the former richness of the organic world wherever they happen to have no evidence of its condition. A large part of the Upper Miocene insects and plants alluded to have been met with at Oeningen, near the Lake of Constance, in two or three spots embedded in thinly laminated marls, the entire thickness of which scarcely exceeds 3 or 4 feet, and in two quarries of very limited dimensions. The rare combination of causes which seems to have led to the faithful preservation of so many treasures of a perishable nature in so small an area, appear to have been the following: first, a river flowing into a lake; secondly, storms of wind, by which leaves and sometimes the boughs of trees were torn off and floated by the stream into the lake; thirdly, mephitic gases rising from the lake, by which insects flying over its surface were occasionally killed: and fourthly, a constant supply of carbonate of lime in solution from mineral springs, the calcareous matter when precipitated to the bottom mingling with fine mud and thus forming the fossiliferous marls.
SPECIES OF INSECTS IN BRITAIN AND NORTH AMERICA, REPRESENTED BY DISTINCT VARIETIES.
If we compare the living British insects with those of the American continent, we frequently find that even those species which are considered to be identical, are nevertheless varieties of the European types. I have noticed this fact when speaking of the common English butterfly, Vanessa atalanta, or "red admiral," which I saw flying about the woods of Alabama in mid-winter. I was unable to detect any difference myself, but all the American specimens which I took to the British Museum were observed by Mr. Doubleday to exhibit a slight peculiarity in the colouring of a minute part of the anterior wing,* (* Lyell's "Second Visit to the United States" volume 2 page 293.) a character first detected by Mr. T.F. Stephens, who has also discovered that similar slight, but equally constant variations, distinguish other Lepidoptera now inhabiting the opposite sides of the Atlantic, insects which, nevertheless, he and Mr. Westwood and the late Mr. Kirby, have always agreed to regard as mere varieties of the same species.
Mr. T.V. Wollaston, in treating of the variation of insects in maritime situations and small islands, has shown how the colour, growth of the wings, and many other characters, undergo modification under the influence of local conditions, continued for long periods of time;* (* Wollaston, "On the Variation of Species" etc. London 1856.) and Mr. Brown has lately called our attention to the fact that the insects of the Shetland Isles present slight deviations from the corresponding types occurring in Great Britain, but far less marked than those which distinguish the American from the European varieties.* (* "Transactions of Northern Entomological Society" 1862.) In the case of Shetland, Mr. Brown remarks, a land communication may well be supposed to have prevailed with Scotland at a more modern era than that between Europe and America. In fact, we have seen that Shetland can hardly fail to have been united with Scotland after the commencement of the glacial period (see map, Figure 41); whereas a communication between the north of Europe by Iceland and Greenland (which, as before stated, once enjoyed a genial climate) must have been anterior to the glacial epoch. A much larger isolation, and the impossibility of varieties formed in the two separated areas crossing with each other, would account, according to Mr. Darwin's theory, for the much wider divergence observed in the specific types of the two regions.
The reader will remember that at the commencement of the Glacial period there was scarcely any appreciable difference between the molluscous fauna and that now living. When therefore the events of the Glacial period, as described in the earlier part of this volume, are duly pondered on, and when we reflect that in the Upper Miocene period the living species of mollusca constitute only one-third of the whole fauna, we see clearly by how high a figure we must multiply the time in order to express the distance between the Miocene period and our own days.
SPECIES OF MAMMALIA RECENT AND FOSSIL—PROBOSCIDIANS.
But it may perhaps be said that the mammalia afford more conspicuous examples than do the mollusca, insects, or plants of the wide gaps which separate species and genera, and that if in this higher class such a multitude of transitional forms had ever existed as would be required to unite the Tertiary and Recent species into one series or net-work of allied or transitional forms, they could not so entirely have escaped observation whether in the fossil or living fauna. A zoologist who entertains such an opinion would do well to devote himself to the study of some one genus of mammalia, such as the elephant, rhinoceros, hippopotamus, bear, horse, ox, or deer; and after collecting all the materials he can get together respecting the extinct and Recent species, decide for himself whether the present state of science justifies his assuming that the chain could never have been continuous, the number of the missing links being so great.
Among the extinct species formerly contemporary with man, no fossil quadruped has so often been alluded to in this work as the mammoth, Elephas primigenius. From a monograph on the proboscidians by Dr. Falconer, it appears that this species represents one extreme of a type of which the Pliocene Mastodon borsoni represents the other. Between these extremes there are already enumerated by Dr. Falconer no less than twenty-six species, some of them ranging as far back in time as the Miocene period, others still living, like the Indian and African forms. Two of these species, however, he has always considered as doubtful, Stegodon ganesa, probably a mere variety of one of the others, and Elephas priscus of Goldfuss, founded partly on specimens of the African elephant, assumed by mistake to be fossil, and partly on some aberrant forms of E. antiquus.
The first effect of the intercalation of so many intermediate forms between the two most divergent types, has been to break down almost entirely the generic distinction between Mastodon and Elephas. Dr. Falconer, indeed, observes that Stegodon (one of several subgenera which he has founded) constitutes an intermediate group, from which the other species diverge through their dental characters, on the one side into the mastodons, and on the other into the Elephants.* (* "Quarterly Journal of the Geological Society" volume 13 1857 page 314.) The next result is to diminish the distance between the several members of each of these groups.
Dr. Falconer has discovered that no less than four species of elephant were formerly confounded together under the title of Elephas primigenius, whence its supposed ubiquity in Pleistocene times, or its wide range over half the habitable globe. But even when this form has been thus restricted in its specific characters, it has still its geographical varieties; for the mammoth's teeth brought from America may in most instances, according to Dr. Falconer, be distinguished from those proper to Europe. On this American variety Dr. Leidy has conferred the name of E. americanus. Another race of the same mammoth (as determined by Dr. Falconer) existed, as we have seen, before the Glacial period, or at the time when the buried forest of Cromer and the Norfolk cliffs was deposited; and the Swiss geologists have lately found remains of the mammoth in their country, both in pre-glacial and post-glacial formations.
Since the publication of Dr. Falconer's monograph, two other species of elephant, F. mirificus, Leidy, and F. imperator, have been obtained from the Pliocene formations of the Niobrara Valley in Nebraska, one of which, however, may possibly be found hereafter to be the same as E. columbi, Falc. A remarkable dwarf species also (Elephas melitensis) has been discovered, belonging, like the existing E. africanus, to the group Loxodon. This species has been established by Dr. Falconer on remains found by Captain Spratt R.N. in a cave in Malta.* (* "Proceedings of the Geological Society" London 1862.)
How much the difficulty of discriminating between the fossil representatives of this genus may hereafter augment, when all the species with their respective geographical varieties are known, may be inferred from the following fact—Professor H. Schlegel, in a recently published memoir, endeavours to show that the living elephant of Sumatra agrees with that of Ceylon, but is a distinct species from that of Continental India, being distinguishable by the number of its dorsal vertebrae and ribs, the form of its teeth, and other characteristics.* (* Schlegel, "Natural History Review" Number 5 1862 page 72.) Dr. Falconer, on the other hand, considers these two living species as mere geographical varieties, the characters referred to not being constant, as he has ascertained, on comparing different individuals of E. indicus in different parts of Bengal in which the ribs vary from nineteen to twenty, and different varieties of E. africanus in which they vary from twenty to twenty-one.
An inquiry into the various species of the genus Rhinoceros, recent and fossil, has led Dr. Falconer to analogous results, as might be inferred from what was said in Chapter 10, and as a forthcoming memoir by the same writer will soon more fully demonstrate.
Among the fossils brought in 1858 by Mr. Hayden from the Niobrara Valley, Dr. Leidy describes a rhinoceros so like the Asiatic species, R. indicus, that he at first referred it to the same, and, what is most singular, he remarks generally of the Pliocene fauna of that part of North America that it is far more related in character to the Pleistocene and Recent fauna of Europe than to that now inhabiting the American continent.
It seems indeed more and more evident that when we speculate in future on the pedigree of any extinct quadruped which abounds in the drift or caverns of Europe, we shall have to look to North and South America as a principal source of information. Thirty years ago, if we had been searching for fossil types which might fill up a gap between two species or genera of the horse tribe (or great family of the Solipedes), we might have thought it sufficient to have got together as ample materials as we could obtain from the continents of Europe, Africa, and Asia. We might have presumed that as no living representative of the equine family, whether horse, ass, zebra, or quagga, had been furnished by North or South America when those regions were first explored by Europeans, a search in the transatlantic world for fossil species might be dispensed with. But how different is the prospect now opening before us! Mr. Darwin first detected the remains of a fossil horse during his visit to South America, since which two other species have been met with on the same continent, while in North America, in the valley of the Nebraska alone, Mr. Hayden, besides a species not distinguishable from the domestic horse, has obtained, according to Dr. Leidy, representatives of five other fossil genera of Solipedes. These he names, Hipparion, Protohippus, Merychippus, Hypohippus, and Parahippus. On the whole, no less than twelve equine species, belonging to seven genera (including the Miocene Anchitherium of Nebraska), being already detected in the Tertiary and Post-Tertiary formations of the United States.* (* "Proceedings of the Academy of Natural Science" Philadelphia for 1858 page 89.)
Professors Unger* (* "Die versunkene Insel Atlantis.") and Heer* (* "Flora tertiaria Helvetiae.") have advocated, on botanical grounds, the former existence of an Atlantic continent during some part of the Tertiary period, as affording the only plausible explanation that can be imagined, of the analogy between the Miocene flora of Central Europe and the existing flora of Eastern America. Professor Oliver, on the other hand, after showing how many of the American types found fossil in Europe are common to Japan, inclines to the theory, first advanced by Dr. Asa Gray, that the migration of species, to which the community of types in the eastern states of North America and the Miocene flora of Europe is due, took place when there was an overland communication from America to eastern Asia between the fiftieth and sixtieth parallels of latitude, or south of Behring Straits, following the direction of the Aleutian islands.* (* Oliver, Lecture at the Royal Institution, March 7, 1862.) By this course they may have made their way, at any epoch, Miocene, Pliocene, or Pleistocene, antecedently to the glacial epoch, to Mongolia, on the east coast of northern Asia.
We have already seen that a large proportion of the living quadrupeds of Mongolia (34 out of 48) are specifically identical with those at present inhabiting the continent of Western Europe and the British Isles.
A monograph on the hippopotamus, bear, ox, stag, or any other genus of mammalia common in the European drift or caverns, might equally well illustrate the defective state of the materials at present at our command. We are rarely in possession of one perfect skeleton of any extinct species, still less of skeletons of both sexes, and of different ages. We usually know nothing of the geographical varieties of the Pleistocene and Pliocene species, least of all, those successive changes of form which they must have undergone in the preglacial epoch between the Upper Miocene and Pleistocene eras. Such being the poverty of our palaeontological data, we cannot wonder that osteologists are at variance as to whether certain remains found in caverns are of the same species as those now living; whether, for example, the Talpa fossilis is really the common mole, the Meles morreni the common badger, Lutra antiqua the otter of Europe, Sciurus priscus the squirrel, Arctomys primigenia the marmot, Myoxus fossilis the dormouse, Schmerling's Felis engihoulensis the European lynx, or whether Ursus spelaeus and Ursus priscus are not extinct races of the living brown bear (Ursus arctos).