"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.