Chapter IX: Preface (6)

Another unanswerable question asked by the Boston reviewers is, Why, when structure and instinct or habit vary-- as they must have varied, on Darwin's hypothesis--they vary together and harmoniously, instead of vaguely? We cannot tell, because we cannot tell why either varies at all. Yet, as they both do vary in successive generations--as is seen under domestication--and are correlated, we can only adduce the fact. Darwin may be precluded from our answer, but we may say that they vary together because designed to do so. A reviewer says that the chance of their varying together is inconceivably small; yet, if they do not, the variant individuals must all perish. Then it is well that it is not left to chance. To refer to a parallel case: before we were born, nourishment and the equivalent to respiration took place in a certain way. But the moment we were ushered into this breathing world, our actions promptly conformed, both as to respiration and nourishment, to the before unused structure and to the new surroundings.

"Now," says the Examiner, "suppose, for instance, the gills of an aquatic animal converted into lungs, while instinct still compelled a continuance under water, would not drowning ensue?" No doubt. But--simply contemplating the facts, instead of theorizing--we notice that young frogs do not keep their heads under water after ceasing to be tadpoles. The instinct promptly changes with the structure, without supernatural interposition--just as Darwin would have it, if the development of a variety or incipient species, though rare, were as natural as a metamorphosis.

"Or if a quadruped, not yet furnished with wings, were suddenly inspired with the instinct of a bird, and precipitated itself from a cliff, would not the descent be hazardously rapid?" Doubtless the animal would be no better supported than the objection. But Darwin makes very little indeed of voluntary efforts as a cause of change, and even poor Lamarck need not be caricatured. He never supposed that an elephant would take such a notion into his wise head, or that a squirrel would begin with other than short and easy leaps; yet might not the length of the leap be increased by practice?

The North American reviewer's position, that the higher brute animals have comparatively little instinct and no intelligence, is a heavy blow and great discouragement to dogs, horses, elephants, and monkeys. Thus stripped of their all, and left to shift for themselves as they may in this hard world, their pursuit and seeming attainment of knowledge under such peculiar difficulties are interesting to contemplate. However, we are not so sure as is the critic that instinct regularly increases downward and decreases upward in the scale of being. Now that the case of the bee is reduced to moderate proportions,[III-19] we know of nothing in instinct surpassing that of an animal so high as a bird, the talegal, the male of which plumes himself upon making a hot-bed in which to batch his partner's eggs--which he tends and regulates the beat of about as carefully and skillfully as the unplumed biped does an eccaleobion.[III-20]

As to the real intelligence of the higher brutes, it has been ably defended by a far more competent observer, Mr. Agassiz, to whose conclusions we yield a general assent, although we cannot quite place the best of dogs "in that respect upon a level with a considerable proportion of poor humanity," nor indulge the hope, or indeed the desire, of a renewed acquaintance with the whole animal kingdom in a future life.

The assertion that acquired habitudes or instincts, and acquired structures, are not heritable, any breeder or good observer can refute.

That "the human mind has become what it is out of a developed instinct," is a statement which Mr. Darwin nowhere makes, and, we presume, would not accept. That he would have us believe that individual animals acquire their instincts gradually,[III-21] is a statement which must have been penned in inadvertence both of the very definition of instinct, and of everything we know of in Mr. Darwin's book.

It has been attempted to destroy the very foundation of Darwin's hypothesis by denying that there are any wild varieties, to speak of, for natural selection to operate upon. We cannot gravely sit down to prove that wild varieties abound. We should think it just as necessary to prove that snow falls in winter. That variation among plants cannot be largely due to hybridism, and that their variation in Nature is not essentially different from much that occurs in domestication, and, in the long-run, probably hardly less in amount, we could show if our space permitted.

As to the sterility of hybrids, that can no longer be insisted upon as absolutely true, nor be practically used as a test between species and varieties, unless we allow that hares and rabbits are of one species. That such sterility, whether total or partial, subserves a purpose in keeping species apart, and was so designed, we do not doubt. But the critics fail to perceive that this sterility proves nothing whatever against the derivative origin of the actual species; for it may as well have been intended to keep separate those forms which have reached a certain amount of divergence, as those which were always thus distinct.

The argument for the permanence of species, drawn from the identity with those now living of cats, birds, and other animals preserved in Egyptian catacombs, was good enough as used by Cuvier against St.-Hilaire, that is, against the supposition that time brings about a gradual alteration of whole species; but it goes for little against Darwin, unless it be proved that species never vary, or that the perpetuation of a variety necessitates the extinction of the parent breed. For Darwin clearly maintains--what the facts warrant--that the mass of a species remains fixed so long as it exists at all, though it may set off a variety now and then. The variety may finally supersede the parent form, or it may coexist with it; yet it does not in the least hinder the unvaried stock from continuing true to the breed, unless it crosses with it. The common law of inheritance may be expected to keep both the original and the variety mainly true as long as they last, and none the less so because they have given rise to occasional varieties. The tailless Manx cats, like the curtailed fox in the fable, have not induced the normal breeds to dispense with their tails, nor have the Dorkings (apparently known to Pliny) affected the permanence of the common sort of fowl.

As to the objection that the lower forms of life ought, on Darwin's theory, to have been long ago improved out of existence, and replaced by higher forms, the objectors forget what a vacuum that would leave below, and what a vast field there is to which a simple organization is best adapted, and where an advance would be no improvement, but the contrary. To accumulate the greatest amount of being upon a given space, and to provide as much enjoyment of life as can be under the conditions, is what Nature seems to aim at; and this is effected by diversification.

Finally, we advise nobody to accept Darwin's or any other derivative theory as true. The time has not come for that, and perhaps never will. We also advise against a similar credulity on the other side, in a blind faith that species--that the manifold sorts and forms of existing animals and vegetables--"have no secondary cause." The contrary is already not unlikely, and we suppose will hereafter become more and more probable. But we are confident that, if a derivative hypothesis ever is established, it will be so on a solid theistic ground.

Meanwhile an inevitable and legitimate hypothesis is on trial--an hypothesis thus far not untenable--a trial just now very useful to science, and, we conclude, not harmful to religion, unless injudicious assailants temporarily make it so.

One good effect is already manifest; its enabling the advocates of the hypothesis of a multiplicity of human species to perceive the double insecurity of their ground. When the races of men are admitted to be of one species, the corollary, that they are of one origin, may be expected to follow. Those who allow them to be of one species must admit an actual diversification into strongly-marked and persistent varieties, and so admit the basis of fact upon which the Darwinian hypothesis is built; while those, on the other hand, who recognize several or numerous human species, will hardly be able to maintain that such species were primordial and supernatural in the ordinary sense of the word.

The English mind is prone to positivism and kindred forms of materialistic philosophy, and we must expect the derivative theory to be taken up in that interest. We have no predilection for that school, but the contrary. If we had, we might have looked complacently upon a line of criticism which would indirectly, but effectively, play into the hands of positivists and materialistic atheists generally. The wiser and stronger ground to take is, that the derivative hypothesis leaves the argument for design, and therefore for a designer, as valid as it ever was; that to do any work by an instrument must require, and therefore presuppose, the exertion rather of more than of less power than to do it directly; that whoever would be a consistent theist should believe that Design in the natural world is coextensive with Providence, and hold as firmly to the one as he does to the other, in spite of the wholly similar and apparently insuperable difficulties which the mind encounters whenever it endeavors to develop the idea into a system, either in the material and organic, or in the moral world. It is enough, in the way of obviating objections, to show that the philosophical difficulties of the one are the same, and only the same, as of the other. IV

SPECIES AS TO VARIATION,

GEOGRAPHICAL

DISTRIBUTION,

AND SUCCESSION

(American Journal of Science and Arts, May, 1863)

Etude sur l'Espece, a l'Occasion d'une Revision de la Famille des Cupuliferes, par M. ALPHONSE DE CANDOLLE.-- This is the title of a paper by M. Alph. De Candolle, growing out of his study of the oaks. It was published in the November number of the Bibliotheque Universelle, and separately issued as a pamphlet. A less inspiring task could hardly be assigned to a botanist than the systematic elaboration of the genus Quercus and its allies. The vast materials assembled under De Candolle's hands, while disheartening for their bulk, offered small hope of novelty. The subject was both extremely trite and extremely difficult. Happily it occurred to De Candolle that an interest might be imparted to an onerous undertaking, and a work of necessity be turned to good account for science, by studying the oaks in view of the question of species. What this term species means, or should mean, in natural history, what the limits of species, inter se or chronologically, or in geographical distribution, their modifications, actual or probable, their origin, and their destiny--these are questions which surge up from time to time; and now and then in the progress of science they come to assume a new and hopeful interest. Botany and zoology, geology, and what our author, feeling the want of a new term proposes to name epiontology, [IV-1] all lead up to and converge into this class of questions, while recent theories shape and point the discussion So we look with eager interest to see what light the study of oaks by a very careful experienced and conservative botanist, particularly conversant with the geographical relations of plants may throw upon the subject.

The course of investigation in this instance does not differ from that ordinarily pursued by working botanists nor, in deed are the theoretical conclusions other than those to which a similar study of other orders might not have equally led. The oaks afford a very good occasion for the discussion of questions which press upon our attention, and perhaps they offer peculiarly good materials on account of the number of fossil species.

Preconceived notions about species being laid aside, the specimens in hand were distributed, according to their obvious resemblances, into groups of apparently identical or nearly identical forms, which were severally examined and compared. Where specimens were few, as from countries little explored, the work was easy, but the conclusions, as will be seen, of small value. The fewer the materials, the smaller the likelihood of forms intermediate between any two, and--what does not appear being treated upon the old law-maxim as non-existent--species are readily enough defined. Where, however, specimens abound, as in the case of the oaks of Europe, of the Orient, and of the United States, of which the specimens amounted to hundreds, collected at different ages, in varied localities, by botanists of all sorts of views and predilections--here alone were data fit to draw useful conclusions from. Here, as De Candolle remarks, he had every advantage, being furnished with materials more complete than any one person could have procured from his own herborizations, more varied than if he had observed a hundred times over the same forms in the same district, and more impartial than if they had all been amassed by one person with his own ideas or predispositions. So that vast herbaria, into which contributions from every source have flowed for years, furnish the best possible data--at least are far better than any practicable amount of personal herborization--or the comparative study of related forms occurring over wide tracts of territory. But as the materials increase, so do the difficulties. Forms, which appeared totally distinct, approach or blend through intermediate gradations; characters, stable in a limited number of instances or in a limited district, prove unstable occasionally, or when observed over a wider area; and the practical question is forced upon the investigator, What here is probably fixed and specific, and what is variant, pertaining to individual, variety, or race?

In the examination of these rich materials, certain characters were found to vary upon the same branch, or upon the same tree, sometimes according to age or development, sometimes irrespective of such relations or of any assignable reasons. Such characters, of course, are not specific, although many of them are such as would have been expected to be constant in the same species, and are such as generally enter into specific definitions. Variations of this sort, De Candolle, with his usual painstaking, classifies and tabulates, and even expresses numerically their frequency in certain species. The results are brought well to view in a systematic enumeration:

1. Of characters which frequently vary upon the same branch: over a dozen
such are mentioned.

2. Of those which sometimes vary upon the same branch: a smaller number of
these are mentioned.

3. Those so rare that they might be called monstrosities. Then he enumerates characters, ten in number, which he has never found to vary on the same branch, and which, therefore, may better claim to be employed as specific. But, as among them he includes the duration of the leaves, the size of the cupule, and the form and size of its scales, which are by no means quite uniform in different trees of the same species, even these characters must be taken with allowance. In fact, having first brought together, as groups of the lowest order, those forms which varied upon the same stock, he next had to combine similarly various forms which, though not found associated upon the same branch, were thoroughly blended by intermediate degrees:

"The lower groups (varieties or races) being thus constituted, I have given the rank of species to the groups next above these, which differ in other respects, i.e., either in characters which were not found united upon certain individuals, or in those which do not show transitions from one individual to another. For the oaks of regions sufficiently known, the species thus formed rest upon satisfactory bases, of which the proof can be furnished. It is quite otherwise with those which are represented in our herbaria by single or few specimens. These are provisional species--species which may hereafter fall to the rank of simple varieties. I have not been inclined to prejudge such questions; indeed, in this regard, I am not disposed to follow those authors whose tendency is, as they say, to reunite species. I never reunite them without proof in each particular case; while the botanists to whom I refer do so on the ground of analogous variations or transitions occurring in the same genus or in the same family. For example resting on the fact that Quercus hex, Q. coccifera, Q. acutifolia, etc., have the leaves sometimes entire and sometimes toothed upon the same branch, or present transitions from one tree to another, I might readily have united my Q. Tlapuxahuensis to Q. Sartorii of Liebmann, since these two differ only in their entire or their toothed leaves. From the fact that the length of the peduncle varies in Q. Robur and many other oaks, I might have combined Q. Seemannii Liebm. with Q. salicifolia Nee. I have not admitted these inductions, but have demanded visible proof in each particular case. Many species are thus left as provisional; but, in proceeding thus, the progress of the science will be more regular, and the synonymy less dependent upon the caprice or the theoretical opinions of each author."

This is safe and to a certain degree judicious, no doubt, as respects published species. Once admitted, they may stand until they are put down by evidence, direct or circumstantial. Doubtless a species may rightfully be condemned on good circumstantial evidence. But what course does De Candolle pursue in the case--of every-day occurrence to most working botanists, having to elaborate collections from countries not so well explored as Europe--when the forms in question, or one of the two, are as yet unnamed? Does he introduce as a new species every form which he cannot connect by ocular proof with a near relative, from which it differs only in particulars which he sees are inconstant in better known species of the same group? We suppose not. But, if he does, little improvement for the future upon the state of things revealed in the following quotation can be expected:

"In the actual state of our knowledge, after having seen nearly all the original specimens, and in some species as many as two hundred representatives from different localities, I estimate that, out of the three hundred species of Cupuliferae which will be enumerated in the Prodromus, two-thirds at least are provisional species. In general, when we consider what a multitude of species were described from a single specimen, or from the forms of a single locality, of a single country, or are badly described, it is difficult to believe that above one-third of the actual species in botanical works will remain unchanged."

Such being the results of the want of adequate knowledge, how is it likely to be when our knowledge is largely increased? The judgment of so practised a botanist as De Candolle is important in this regard, and it accords with that of other botanists of equal experience.

"They are mistaken," he pointedly asserts, "who repeat that the greater part of our species are clearly limited, and that the doubtful species are in a feeble minority. This seemed to be true, so long as a genus was imperfectly known, and its species were founded upon few specimens, that is to say, were provisional. Just as we come to know them better, intermediate forms flow in, and doubts as to specific limits augment."

De Candolle insists, indeed, in this connection, that the higher the rank of the groups the more definite their limitation, or, in other terms, the fewer the ambiguous or doubtful forms, that genera are more strictly limited than species tribes than genera, orders than tribes, etc. We are not convinced of this Often where it has appeared to be so, advancing discovery has brought intermediate forms to light, perplexing to the systematist. "They are mistaken, we think more than one systematic botanist will say, "who repeat that the greater part of our natural orders and tribes are absolutely limited," however we may agree that we will limit them. Provisional genera we suppose are proportionally hardly less common than provisional species; and hundreds of genera are kept up on considerations of general propriety or general convenience, although well known to shade off into adjacent ones by complete gradations. Somewhat of this greater fixity of higher groups, therefore, is rather apparent than real. On the other hand, that varieties should be less definite than species, follows from the very terms employed. They are ranked as varieties, rather than species, just because of their less definiteness.

Singular as it may appear, we have heard it denied that spontaneous varieties occur. De Candolle makes the important announcement that, in the oak genus, the best known species are just those which present the greatest number of spontaneous varieties and sub-varieties. The maximum is found in Q. Robur, with twenty-eight varieties, all spontaneous. Of Q. Lusitanica eleven varieties are enumerated, of Q. Calliprinos ten, of Q. coccifera eight, * etc. And he significantly adds that "these very species which offer such numerous modifications are themselves ordinarily surrounded by other forms, provisionally called species, because of the absence of known transitions or variations, but to which some of these will probably have to be joined hereafter." The inference is natural, if not inevitable, that the difference between such species and such varieties is only one of degree, either as to amount of divergence, or of hereditary fixity, or as to the frequency or rarity at the present time of intermediate forms.

This brings us to the second section of De Candolle's article, in which he passes on, from the observation of the present forms and affinities of cupuliferous plants, to the consideration of their probable history and origin. Suffice it to say, that he frankly accepts the inferences derived from the whole course of observation, and contemplates a probable historical connection between congeneric species. He accepts and, by various considerations drawn from the geographical distribution of European Cupuliferae, fortifies the conclusion--long ago arrived at by Edward Forbes--that the present species, and even some of their varieties, date back to about the close of the Tertiary epoch, since which time they have been subject to frequent and great changes of habitation or limitation, but without appreciable change of specific form or character; that is, without profounder changes than those within which a species at the present time is known to vary. Moreover, he is careful to state that he is far from concluding that the time of the appearance of a species in Europe at all indicates the time of its origin. Looking back still further into the Tertiary epoch, of which the vegetable remains indicate many analogous, but few, if any, identical forms, he concludes, with Heer and others, that specific changes of form, as well as changes of station, are to be presumed; and, finally, that "the theory of a succession of forms through the deviation of anterior forms is the most natural hypothesis, and the most accordant with the known facts in palaeontology, geographical botany and zoology, of anatomical structure and classification: but direct proof of it is wanting, and moreover, if true, it must have taken place very slowly; so slowly, indeed, that its effects are discernible only after a lapse of time far longer than our historic epoch." In contemplating the present state of the species of Cupuliferae in Europe, De Candolle comes to the conclusion that, while the beech is increasing, and extending its limits southward and westward (at the expense of Coniferae and birches), the common oak, to some extent, and the Turkey oak decidedly, are diminishing and retreating, and this wholly irrespective of man's agency. This is inferred of the Turkey oak from the great gaps found in its present geographical area, which are otherwise inexplicable, and which he regards as plain indications of a partial extinction. Community of descent of all the individuals of species is of course implied in these and all similar reasonings.

An obvious result of such partial extinction is clearly enough brought to view The European oaks (like the American species) greatly tend to vary that is they manifest an active disposition to produce new forms Every form tends to become hereditary and so to pass from the state of mere variation to that of race and of these competing incipient races some only will survive. Quercus Robur offers a familiar illustration of the manner in which one form may in the course of time become separated into two or more distinct ones.

To Linnaeus this common oak of Europe was all of one species. But of late years the greater number of European botanists have regarded it as including three species, Q. pedunculata, Q. sessiliflora, and Q. pubescens. De Candolle looks with satisfaction to the independent conclusion which he reached from a long and patient study of the forms (and which Webb, Gay, Bentham, and others, had equally reached), that the view of Linnaeus was correct, inasmuch as it goes to show that the idea and the practical application of the term species have remained unchanged during the century which has elapsed since the publication of the "Species Plantarum." But, the idea remaining unchanged, the facts might appear under a different aspect, and the conclusion be different, under a slight and very supposable change of circumstances. Of the twenty-eight spontaneous varieties of Q. Robur, which De Candolle recognizes, all but six, he remarks, fall naturally under the three sub-species, pedunculata, sessiliflora, and pubescens, and are therefore forms grouped around these as centres; and, moreover, the few connecting forms are by no means the most common. Were these to die out, it is clear that the three forms which have already been so frequently taken for species would be what the group of four or five provisionally admitted species which closely surround Q. Robur now are. The best example of such a case, as having in all probability occurred through geographical segregation and partial extinction, is that of the cedar, thus separated into the Deodar, the Lebanon, and the Atlantic cedars--a case admirably worked out by Dr. Hooker two or three years ago. [IV-2] A special advantage of the Cupuliferae for determining the probable antiquity of existing species in Europe, De Candolle finds in the size and character of their fruits. However it may be with other plants (and he comes to the conclusion generally that marine currents and all other means of distant transport have played only a very small part in the actual dispersion of species), the transport of acorns and chestnuts by natural causes across an arm of the sea in a condition to germinate, and much more the spontaneous establishment of a forest of oaks or chestnuts in this way, De Candolle conceives to be fairly impossible in itself, and contrary to all experience. From such considerations, i.e., from the actual dispersion of the existing species (with occasional aid from post-tertiary deposits), it is thought to be shown that the principal Cupuliferae of the Old World attained their actual extension before the present separation of Sicily, Sardinia and Corsica, and of Britain, from the European Continent.

This view once adopted, and this course once entered upon, has to be pursued farther. Quercus Robur of Europe with its bevy of admitted derivatives, and its attending species only provisionally admitted to that rank, is very closely related to certain species of Eastern Asia, and of Oregon and California--so closely that "a view of the specimens by no means forbids the idea that they have all originated from Q. Robur, or have originated, with the latter, from one or more preceding forms so like the present ones that a naturalist could hardly know whether to call them species or varieties." Moreover, there are fossil leaves from diluvian deposits in Italy, figured by Gaudin, which are hardly distinguishable from those of Q. Robur on the one hand, and from those of Q. Douglasii, etc., of California, on the other. No such leaves are found in any tertiary deposit in Europe; but such are found of that age, it appears, in Northwest America, where their remote descendants still flourish. So that the probable genealogy of Q. Robur, traceable in Europe up to the commencement of the present epoch, looks eastward and far into the past on far-distant shores.

Quercus Ilex, the evergreen oak of Southern Europe and Northern Africa, reveals a similar archaeology; but its presence in Algeria leads De Candolle to regard it as a much more ancient denizen of Europe than Q. Robur; and a Tertiary oak, Q. ilicoides, from a very old Miocene bed in Switzerland, is thought to be one of its ancestral forms. This high antiquity once established, it follows almost of course that the very nearly-related species in Central Asia, in Japan, in California, and even our own live-oak with its Mexican relatives, may probably enough be regarded as early offshoots from the same stock with Q. hex.

In brief--not to continue these abstracts and remarks, and without reference to Darwin's particular theory (which De Candolle at the close very fairly considers)--if existing species, or many of them, are as ancient as they are now generally thought to be, and were subject to the physical and geographical changes (among them the coming and the going of the glacial epoch) which this antiquity implies; if in former times they were as liable to variation as they now are; and if the individuals of the same species may claim a common local origin, then we cannot wonder that "the theory of a succession of forms by deviations of anterior forms" should be regarded as "the most natural hypothesis," nor at the general advance made toward its acceptance.

The question being, not, how plants and animals originated, but, how came the existing animals and plants to be just where they are and what they are, it is plain that naturalists interested in such inquiries are mostly looking for the answer in one direction. The general drift of opinion, or at least of expectation, is exemplified by this essay of De Candolle; and the set and force of the current are seen by noticing how it carries along naturalists of widely different views and prepossessions--some faster and farther than others--but all in one way. The tendency is, we may say, to extend the law of continuity, or something analogous to it, from inorganic to organic Nature, and in the latter to connect the present with the past in some sort of material connection. The generalization may indeed be expressed so as not to assert that the connection is genetic, as in Mr. Wallace's formula: "Every species has come into existence coincident both in time and space with preexisting closely-allied species." Edward Forbes, who may be called the originator of this whole line of inquiry, long ago expressed a similar view. But the only material sequence we know, or can clearly conceive, in plants and animals, is that from parent to progeny; and, as De Candolle implies, the origin of species and that of races can hardly be much unlike, nor governed by other than the same laws, whatever these may be.

The progress of opinion upon this subject in one generation is not badly represented by that of De Candolle himself, who is by no means prone to adopt new views without much consideration. In an elementary treatise published in the year 1835, he adopted and, if we rightly remember, vigorously maintained, Schouw's idea of the double or multiple origin of species, at least of some species--a view which has been carried out to its ultimate development only perhaps by Agassiz, in the denial of any necessary genetic connection among the individuals of the same species, or of any original localization more restricted than the area now occupied by the species. But in i855, in his "Geographic Botanique," the multiple hypothesis, although in principle not abandoned, loses its point, in view of the probable high antiquity of existing species. The actual vegetation of the world being now regarded as a continuation, through numerous geological, geographical, and more recently historical changes, of anterior vegetations, the actual distribution of plants is seen to be a consequence of preceding conditions; and geological considerations, and these alone, may be expected to explain all the facts--many of them so curious and extraordinary--of the actual geographical distribution of the species. In the present essay, not only the distribution but the origin of congeneric species is regarded as something derivative; whether derived by slow and very gradual changes in the course of ages, according to Darwin, or by a sudden, inexplicable change of their tertiary ancestors, as conceived by Heer, De Candolle hazards no opinion. It may, however, be inferred that he looks upon "natural selection" as a real, but insufficient cause; while some curious remarks upon the number of monstrosities annually produced, and the possibility of their enduring, may be regarded as favorable to Heer's view.

As an index to the progress of opinion in the direction referred to, it will be interesting to compare Sir Charles Lyell's well-known chapters of twenty or thirty years ago, in which the permanence of species was ably maintained, with his treatment of the same subject in a work just issued in England, which, however, has not yet reached us.

A belief of the derivation of species may be maintained along with a conviction of great persistence of specific characters. This is the idea of the excellent Swiss vegetable palaeontologist, Heer, who imagines a sudden change of specific type at certain periods, and perhaps is that of Pictet. Falconer adheres to somewhat similar views in his elaborate paper on elephants, living and fossil, in the Natural History Review for January last. Noting that "there is clear evidence of the true mammoth having existed in America long after the period of the northern drift, when the surface of the country had settled down into its present form, and also in Europe so late as to have been a contemporary of the Irish elk, and on the other hand that it existed in England so far back as before the deposition of the bowlder clay; also that four well-defined species of fossil elephant are known to have existed in Europe; that "a vast number of the remains of three of these species have been exhumed over a large area in Europe; and, even in the geological sense, an enormous interval of time has elapsed between the formation of the most ancient and the most recent of these deposits, quite sufficient to test the persistence of specific characters in an elephant," he presents the question, "Do, then, the successive elephants occurring in these strata show any signs of a passage from the older form into the newer?"

To which the reply is: "If there is one fact which is impressed on the conviction of the observer with more force than any other, it is the persistence and uniformity of the characters of the molar teeth in the earliest known mammoth and his most modern successor . . . Assuming the observation to be correct, what strong proof does it not afford of the persistence and constancy, throughout vast intervals of time, of the distinctive characters of those organs which arc most concerned in the existence and habits of the species? If we cast a glance back on the long vista of physical changes which our planet has undergone since the Neozoic epoch, we can nowhere detect signs of a revolution more sudden and pronounced, or more important in its results, than the intercalation and sudden disappearance of the glacial period. Yet the 'dicyclotherian' mammoth lived before it, and passed through the ordeal of all the hard extremities it involved, bearing his organs of locomotion and digestion all but unchanged. Taking the group of four European fossil species above enumerated, do they show any signs in the successive deposits of a transition from the one form into the other? Here again the result of my observation, in so far as it has extended over the European area, is, that the specific characters of the molars are constant in each, within a moderate range of variation, and that we nowhere meet with intermediate forms." . . .

Dr. Falconer continues (page 80):

"The inferences which I draw from these facts are not opposed to one of the leading propositions of Darwin's theory. With him, I have no faith in the opinion that the mammoth and other extinct elephants made their appearance suddenly, after the type in which their fossil remains are presented to us. The most rational view seems to be, that they are in some shape the modified descendants of earlier progenitors. But if the asserted facts be correct, they seem clearly to indicate that the older elephants of Europe, such as E. meridionalis and E. antiguus, were not the stocks from which the later species, E. primigenius and E. Africanus sprung, and that we must look elsewhere for their origin. The nearest affinity, and that a very close one, of the European E. meridionalis is with the Miocene E. planifrons of India; and of E. primigenius, with the existing India species.

"Another reflection is equally strong in my mind--that the means which have been adduced to explain the origin of the species by 'natural selection,' or a process of variation from external influences, are inadequate to account for the phenomena. The law of phyllotaxis, which governs the evolution of leaves around the axis of a plant, is as nearly constant in its manifestation as any of the physical laws connected with the material world. Each instance, however different from another, can be shown to be a term of some series of continued fractions. When this is coupled with the geometrical law governing the evolution of form, so manifest in some departments of the animal kingdom, e. g., the spiral shells of the Mollusca, it is difficult to believe that there is not, in Nature, a deeper-seated and innate principle, to the operation of which natural selection is merely an adjunct. The whole range of the Mammalia, fossil and recent, cannot furnish a species which has had a wider geographical distribution, and passed through a longer term of time, and through more extreme changes of climatal conditions, than the mammoth. If species are so unstable, and so susceptible of mutation through such influences, why does that extinct form stand out so signally a monument of stability? By his admirable researches and earnest writings, Darwin has, beyond all his contemporaries, given an impulse to the philosophical investigation of the most backward and obscure branch of the biological sciences of his day; he has laid the foundations of a great edifice; but he need not be surprised if, in the progress of erection, the superstructure is altered by his successors, like the Duomo of Milan from the Roman to a different style of architecture."

Entertaining ourselves the opinion that something more than natural selection is requisite to account for the orderly production and succession of species, we offer two incidental remarks upon the above extract.

1. We find in it--in the phrase "natural selection, or a process of variation from external influences"--an example of the very common confusion of two distinct things, viz., variation and natural selection. The former has never yet been shown to have its cause in "external influences," nor to occur at random. As we have elsewhere insisted, if not inexplicable, it has never been explained; all we can yet say is, that plants and animals are prone to vary, and that some conditions favor variation. Perhaps in this Dr. Falconer may yet find what he seeks: for "it is difficult to believe that there is not in nature a deeper-seated and innate principle, to the operation of which natural selection is merely an adjunct." The latter, which is the ensemble of the external influences, including the competition of the individuals them selves, picks out certain variations as they arise, but in no proper sense can be said to originate them

2. Although we are not quite sure how Dr Falconer in tends to apply the law of phyllotaxis to illustrate his idea, we fancy that a pertinent illustration may be drawn from it in this way. There are two species of phyllotaxis, perfectly distinct, and we suppose, not mathematically reducible the one to the other, viz.: (1.) That of alternate leaves, with its vane ties and (2.) That of verticillate leaves, of which opposite leaves present the simplest case That although generally constant a change from one variety of alternate phyllotaxis to an other should occur on the same axis, or on successive axes, is not surprising, the different sorts being terms of a regular series--although indeed we have not the least idea as to how the change from the one to the other comes to pass But it is interesting and in this connection perhaps instructive, to remark that while some dicotyledonous plants hold to the verticillate, i.e., opposite-leaved phyllotaxis throughout, a larger number--through the operation of some deep seated and innate principle which we cannot fathom--change abruptly into the other species at the second or third node, and change back again in the flower, or else effect a synthesis of the two species in a manner which is puzzling to understand. Here is a change from one fixed law to another, as unaccountable, if not as great, as from one specific form to another.

An elaborate paper on the vegetation of the Tertiary period in the southeast of France, by Count Gaston de Saporta, published in the Annales des Sciences Naturelles in 1862, vol. xvi., pp. 309-344--which we have not space to analyze--is worthy of attention from the general inquirer, on account of its analysis of the Tertiary flora into its separate types, Cretaceous, Austral, Tropical, and Boreal, each of which has its separate and different history--and for the announcement that "the hiatus, which, in the idea of most geologists, intervened between the close of the Cretaceous and the beginning of the Tertiary, appears to have had no existence, so far as concerns the vegetation; that in general it was not by means of a total overthrow, followed by a complete new emission of species, that the flora has been renewed at each successive period; and that while the plants of Southern Europe inherited from the Cretaceous period more or less rapidly disappeared, as also the austral forms, and later the tropical types (except the laurel, the myrtle, and the Chamaerops humilis), the boreal types, coming later, survived all the others, and now compose, either in Europe, or in the north of Asia, or in North America, the basis of the actual arborescent vegetation. Especially "a very considerable number of forms nearly identical with tertiary forms now exist in America, where they have found, more easily than in our soil--less vast and less extended southward--refuge from ulterior revolutions," The extinction of species is attributed to two kinds of causes; the one material or physical, whether slow or rapid; the other inherent in the nature of organic beings, incessant, but slow, in a manner latent, but somehow assigning to the species, as to the individuals, a limited period of existence, and, in some equally mysterious but wholly natural way, connected with the development of organic types: "By type meaning a collection of vegetable forms constructed upon the same plan of organization, of which they reproduce the essential lineaments with certain secondary modifications, and which appear to run back to a common point of departure."

In this community of types, no less than in the community of certain existing species, Saporta recognizes a prolonged material union between North America and Europe in former times. Most naturalists and geologists reason in the same way--some more cautiously than others--yet perhaps most of them seem not to perceive how far such inferences imply the doctrine of the common origin of related species.

For obvious reasons such doctrines are likely to find more favor with botanists than with zoologists. But with both the advance in this direction is seen to have been rapid and great; yet to us not unexpected. We note, also, an evident disposition, notwithstanding some endeavors to the contrary, to allow derivative hypotheses to stand or fall upon their own merits--to have indeed upon philosophical grounds certain presumptions in their favor--and to be, perhaps, quite as capable of being turned to good account as to bad account in natural theology.[IV-3]

Among the leading naturalists, indeed, such views--taken in the widest sense--have one and, so far as we are now aware, only one thoroughgoing and thoroughly consistent opponent, viz., Mr. Agassiz.

Most naturalists take into their very conception of a species, explicitly or by implication, the notion of a material connection resulting from the descent of the individuals composing it from a common stock, of local origin. Agassiz wholly eliminates community of descent from his idea of species, and even conceives a species to have been as numerous in individuals and as wide-spread over space, or as segregated in discontinuous spaces, from the first as at the later period.

The station which it inhabits, therefore, is with other naturalists in no wise essential to the species, and may not have been the region of its origin. In Agassiz's view the habitat is supposed to mark the origin, and to be a part of the character of the species. The habitat is not merely the place where it is, but a part of what it is.

Most naturalists recognize varieties of species; and many, like De Candolle, have come to conclude that varieties of the highest grade, or races, so far partake of the characteristics of species, and are so far governed by the same laws, that it is often very difficult to draw a clear and certain distinction between the two. Agassiz will not allow that varieties or races exist in Nature, apart from man's agency.

Most naturalists believe that the origin of species is supernatural, their dispersion or particular geographical area, natural, and their extinction, when they disappear, also the result of physical causes. In the view of Agassiz, if rightly understood, all three are equally independent of physical cause and effect, are equally supernatural.

In comparing preceding periods with the present and with each other, most naturalists and palaeontologists now appear to recognize a certain number of species as having survived from one epoch to the next, or even through more than one formation, especially from the Tertiary into the post-Tertiary period, and from that to the present age. Agassiz is understood to believe in total extinctions and total new creations at each successive epoch, and even to recognize no existing species as ever contemporary with extinct ones, except in the case of recent exterminations.

These peculiar views if sustained will effectually dispose of every form of derivative hypothesis.

Returning for a moment to De Candolle's article, we are disposed to notice his criticism of Linnaeus's "definition" of the term species (Philosophia Botanica, No. 157): "Species tot numeramus quot diversae formae in principio sunt creatae"-- which he declares illogical, inapplicable, and the worst that has been propounded. "So, to determine if a form is specific, it is necessary to go back to its origin which is impossible A definition by a character which can never be verified is no definition at all."

Now as Linnaeus practically applied the idea of species with a sagacity which has never been surpassed and rarely equaled and indeed may be said to have fixed its received meaning in natural history, it may well be inferred that in the phrase above cited he did not so much undertake to frame a logical definition, as to set forth the idea which, in his opinion, lay at the foundation of species; on which basis A.L. Jussieu did construct a logical definition--"Nunc rectius definitur perennis individuorum similium successio continuata generatione renascentium." The fundamental idea of species, we would still maintain, is that of a chain of which genetically-connected individuals are the links. That, in the practical recognition of species, the essential characteristic has to be inferred, is no great objection--the general fact that like engenders like being an induction from a vast number of instances, and the only assumption being that of the uniformity of Nature. The idea of gravitation, that of the atomic constitution of matter, and the like, equally have to be verified inferentially. If we still hold to the idea of Linnaeus, and of Agassiz, that existing species were created independently and essentially all at once at the beginning of the present era, we could not better the propositions of Linnaeus and of Jussieu. If; on the other hand, the time has come in which we may accept, with De Candolle, their successive origination, at the commencement of the present era or before, and even by derivation from other forms, then the "in principio" of Linnaeus will refer to that time, whenever it was, and his proposition be as sound and wise as ever.

In his "Geographie Botanique" (ii., 1068-1077) De Candolle discusses this subject at length, and in the same interest. Remarking that of the two great facts of species, viz., likeness among the individuals, and genealogical connection, zoologists have generally preferred the latter,[IV-4] while botanists have been divided in opinion, he pronounces for the former as the essential thing, in the following argumentative statement:

"Quant a moi, j'ai ete conduit, dans ma definition de l'espece, a mettre decidement la ressemblance au-dessus de caracteres de succession. Ce n'est pas seulement a cause des circonstances propres au regne vegetal, dont je m'occupe exclusivement; ce n'est pas non plus afin de sortir ma definition des theories et de la rendre le plus possible utile aux naturalistes descripteurs et nomenclateurs, c'est aussi par un motif philosophique. En toute chose il faut aller au fond des questions, quand on le peut. Or, pourquoi la reproduction est-elle possible, habituelle, feconde indefiniment, entre des etres organises que nous dirons de la meme espece? Parce qu'ils se ressemblent et uniquement a cause de cela. Lorsque deux especes ne peuvent, ou, s'il s'agit d'animaux superieurs, ne peuvent et ne veulent se croiser, c'est qu'elles sont tres differentes. Si l'on obtient des croisements, c'est que les individus sont analogues; si ces croisements donnent des produits feconds, c'est que les individus etaient plus analogues; si ces produits euxmemes sont feconds, c'est que la ressemblance etait plus grande; s'ils sont fecond habituellement et indefiniment, c'est que la ressemblance interieure et exterieure etait tres grande. Ainsi le degre de ressemblance est le fond; la reproduction en est seulement la manifestation et la mesure, et il est logique de placer la cause au-dessus de l'effet."

We are not yet convinced. We still hold that genealogical connection, rather than mutual resemblance, is the fundamental thing--first on the ground of fact, and then from the philosophy of the case. Practically, no botanist can say what amount of dissimilarity is compatible with unity of species; in wild plants it is sometimes very great, in cultivated races often enormous. De Candolle himself informs us that the different variations which the same oak-tree exhibits arc significant indications of a disposition to set up separate varieties, which becoming hereditary may constitute a race; he evidently looks upon the extreme forms, say of Quercus Robur, as having thus originated; and on this ground, inferred from transitional forms, and not from their mutual resemblance, he includes them in that species. This will be more apparent should the discovery of transitions, which he leads us to expect, hereafter cause the four provisional species which attend Q. Robur to be merged in that species. It may rightly be replied that this conclusion would be arrived at from the likeness step by step in the series of forms; but the cause of the likeness here is obvious. And this brings in our "motif philosophique."

Not to insist that the likeness is after all the variable, not the constant, element--to learn which is the essential thing, resemblance among individuals or their genetic connection--we have only to ask which can be the cause of the other.

In hermaphrodite plants (the normal case), and even as the question is ingeniously put by De Candolle in the above extract, the former surely cannot be the cause of the latter, though it may, in case of crossing, offer occasion. But, on the ground of the most fundamental of all things in the constitution of plants and animals--the fact incapable of further analysis, that individuals reproduce their like, that characteristics are inheritable--the likeness is a direct natural consequence of the genetic succession; "and it is logical to place the cause above the effect."

We are equally disposed to combat a proposition of De Candolle's about genera, elaborately argued in the "Geographie Botanique," and incidentally reaffirmed in his present article, viz., that genera are more natural than species, and more correctly distinguished by people in general, as is shown by vernacular names. But we have no space left in which to present some evidence to the contrary.

V

SEQUOIA AND ITS HISTORY

THE RELATIONS OF NORTH AMERICAN

TO NORTHEAST ASIAN AND TO

TERTIARY VEGETATION

(A Presidential Address to the American Association for the Advancement of Science, at Dubuque, August, 1872)