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|10 - On The Geological Succession of Organic Beings||10-01 - On the slow and successive appearance of new species||10||
Let us now see whether the several facts and rules relating to the geological succession of organic beings, better accord with the common view of the immutability of species, or with that of their slow and gradual modification, through descent and natural selection.
New species have appeared very slowly, one after another, both on the land and in the waters.
Lyell has shown that it is hardly possible to resist the evidence on this head in the case of the several tertiary stages; and every year tends to fill up the blanks between them, and to make the percentage system of lost and new forms more gradual.
In some of the most recent beds, though undoubtedly of high antiquity if measured by years, only one or two species are lost forms, and only one or two are new forms, having here appeared for the first time, either locally, or, as far as we know, on the face of the earth.
If we may trust the observations of Philippi in Sicily, the successive changes in the marine inhabitants of that island have been many and most gradual.
The secondary formations are more broken; but, as Bronn has remarked, neither the appearance nor disappearance of their many now extinct species has been simultaneous in each separate formation.
|10 - On The Geological Succession of Organic Beings||10-07 - On the affinities of extinct species to each other and to living species||10||
Let us now look to the mutual affinities of extinct and living species.
They all fall into one grand natural system; and this fact is at once explained on the principle of descent.
The more ancient any form is, the more, as a general rule, it differs from living forms.
But, as Buckland long ago remarked, all fossils can be classed either in still existing groups, or between them.
That the extinct forms of life help to fill up the wide intervals between existing genera, families, and orders, cannot be disputed.
For if we confine our attention either to the living or to the extinct alone, the series is far less perfect than if we combine both into one general system.
With respect to the Vertebrata, whole pages could be filled with striking illustrations from our great palaeontologist, Owen, showing how extinct animals fall in between existing groups.
Cuvier ranked the Ruminants and Pachyderms, as the two most distinct orders of mammals; but Owen has discovered so many fossil links, that he has had to alter the whole classification of these two orders; and has placed certain pachyderms in the same sub-order with ruminants: for example, he dissolves by fine gradations the apparently wide difference between the pig and the camel.
In regard to the Invertebrata, Barrande, and a higher authority could not be named, asserts that he is every day taught that palaeozoic animals, though belonging to the same orders, families, or genera with those living at the present day, were not at this early epoch limited in such distinct groups as they now are.
Some writers have objected to any extinct species or group of species being considered as intermediate between living species or groups.
If by this term it is meant that an extinct form is directly intermediate in all its characters between two living forms, the objection is probably valid.
But I apprehend that in a perfectly natural classification many fossil species would have to stand between living species, and some extinct genera between living genera, even between genera belonging to distinct families.
The most common case, especially with respect to very distinct groups, such as fish and reptiles, seems to be, that supposing them to be distinguished at the present day from each other by a dozen characters, the ancient members of the same two groups would be distinguished by a somewhat lesser number of characters, so that the two groups, though formerly quite distinct, at that period made some small approach to each other.
It is a common belief that the more ancient a form is, by so much the more it tends to connect by some of its characters groups now widely separated from each other.
This remark no doubt must be restricted to those groups which have undergone much change in the course of geological ages; and it would be difficult to prove the truth of the proposition, for every now and then even a living animal, as the Lepidosiren, is discovered having affinities directed towards very distinct groups.
Yet if we compare the older Reptiles and Batrachians, the older Fish, the older Cephalopods, and the eocene Mammals, with the more recent members of the same classes, we must admit that there is some truth in the remark.
Let us see how far these several facts and inferences accord with the theory of descent with modification.
As the subject is somewhat complex, I must request the reader to turn to the diagram in the fourth chapter.
We may suppose that the numbered letters represent genera, and the dotted lines diverging from them the species in each genus.
The diagram is much too simple, too few genera and too few species being given, but this is unimportant for us.
The horizontal lines may represent successive geological formations, and all the forms beneath the uppermost line may be considered as extinct.
The three existing genera, a14, q14, p14, will form a small family; b14 and f14 a closely allied family or sub-family; and o14, e14, m14, a third family.
These three families, together with the many extinct genera on the several lines of descent diverging from the parent-form A, will form an order; for all will have inherited something in common from their ancient and common progenitor.
On the principle of the continued tendency to divergence of character, which was formerly illustrated by this diagram, the more recent any form is, the more it will generally differ from its ancient progenitor.
Hence we can understand the rule that the most ancient fossils differ most from existing forms.
We must not, however, assume that divergence of character is a necessary contingency; it depends solely on the descendants from a species being thus enabled to seize on many and different places in the economy of nature.
Therefore it is quite possible, as we have seen in the case of some Silurian forms, that a species might go on being slightly modified in relation to its slightly altered conditions of life, and yet retain throughout a vast period the same general characteristics.
This is represented in the diagram by the letter F14.
All the many forms, extinct and recent, descended from A, make, as before remarked, one order; and this order, from the continued effects of extinction and divergence of character, has become divided into several sub-families and families, some of which are supposed to have perished at different periods, and some to have endured to the present day.
By looking at the diagram we can see that if many of the extinct forms, supposed to be embedded in the successive formations, were discovered at several points low down in the series, the three existing families on the uppermost line would be rendered less distinct from each other.
If, for instance, the genera a1, a5, a10, m3, m6, m9 were disinterred, these three families would be so closely linked together that they probably would have to be united into one great family, in nearly the same manner as has occurred with ruminants and pachyderms.
Yet he who objected to call the extinct genera, which thus linked the living genera of three families together, intermediate in character, would be justified, as they are intermediate, not directly, but only by a long and circuitous course through many widely different forms.
If many extinct forms were to be discovered above one of the middle horizontal lines or geological formations for instance, above No. VI. but none from beneath this line, then only the two families on the left hand (namely, a14, &c., and b14, &c.) would have to be united into one family; and the two other families (namely, a14 to f14 now including five genera, and o14 to m14) would yet remain distinct.
These two families, however, would be less distinct from each other than they were before the discovery of the fossils.
If, for instance, we suppose the existing genera of the two families to differ from each other by a dozen characters, in this case the genera, at the early period marked VI., would differ by a lesser number of characters; for at this early stage of descent they have not diverged in character from the common progenitor of the order, nearly so much as they subsequently diverged.
Thus it comes that ancient and extinct genera are often in some slight degree intermediate in character between their modified descendants, or between their collateral relations.
In nature the case will be far more complicated than is represented in the diagram; for the groups will have been more numerous, they will have endured for extremely unequal lengths of time, and will have been modified in various degrees.
As we possess only the last volume of the geological record, and that in a very broken condition, we have no right to expect, except in very rare cases, to fill up wide intervals in the natural system, and thus unite distinct families or orders.
All that we have a right to expect, is that those groups, which have within known geological periods undergone much modification, should in the older formations make some slight approach to each other; so that the older members should differ less from each other in some of their characters than do the existing members of the same groups; and this by the concurrent evidence of our best palaeontologists seems frequently to be the case.
Thus, on the theory of descent with modification, the main facts with respect to the mutual affinities of the extinct forms of life to each other and to living forms, seem to me explained in a satisfactory manner.
And they are wholly inexplicable on any other view.
On this same theory, it is evident that the fauna of any great period in the earth's history will be intermediate in general character between that which preceded and that which succeeded it.
Thus, the species which lived at the sixth great stage of descent in the diagram are the modified offspring of those which lived at the fifth stage, and are the parents of those which became still more modified at the seventh stage; hence they could hardly fail to be nearly intermediate in character between the forms of life above and below.
We must, however, allow for the entire extinction of some preceding forms, and for the coming in of quite new forms by immigration, and for a large amount of modification, during the long and blank intervals between the successive formations.
Subject to these allowances, the fauna of each geological period undoubtedly is intermediate in character, between the preceding and succeeding faunas.
I need give only one instance, namely, the manner in which the fossils of the Devonian system, when this system was first discovered, were at once recognised by palaeontologists as intermediate in character between those of the overlying carboniferous, and underlying Silurian system.
But each fauna is not necessarily exactly intermediate, as unequal intervals of time have elapsed between consecutive formations.
It is no real objection to the truth of the statement, that the fauna of each period as a whole is nearly intermediate in character between the preceding and succeeding faunas, that certain genera offer exceptions to the rule.
For instance, mastodons and elephants, when arranged by Dr Falconer in two series, first according to their mutual affinities and then according to their periods of existence, do not accord in arrangement.
The species extreme in character are not the oldest, or the most recent; nor are those which are intermediate in character, intermediate in age.
But supposing for an instant, in this and other such cases, that the record of the first appearance and disappearance of the species was perfect, we have no reason to believe that forms successively produced necessarily endure for corresponding lengths of time: a very ancient form might occasionally last much longer than a form elsewhere subsequently produced, especially in the case of terrestrial productions inhabiting separated districts.
To compare small things with great: if the principal living and extinct races of the domestic pigeon were arranged as well as they could be in serial affinity, this arrangement would not closely accord with the order in time of their production, and still less with the order of their disappearance; for the parent rock-pigeon now lives; and many varieties between the rock-pigeon and the carrier have become extinct; and carriers which are extreme in the important character of length of beak originated earlier than short-beaked tumblers, which are at the opposite end of the series in this same respect.
Closely connected with the statement, that the organic remains from an intermediate formation are in some degree intermediate in character, is the fact, insisted on by all palaeontologists, that fossils from two consecutive formations are far more closely related to each other, than are the fossils from two remote formations.
Pictet gives as a well-known instance, the general resemblance of the organic remains from the several stages of the chalk formation, though the species are distinct in each stage.
This fact alone, from its generality, seems to have shaken Professor Pictet in his firm belief in the immutability of species.
He who is acquainted with the distribution of existing species over the globe, will not attempt to account for the close resemblance of the distinct species in closely consecutive formations, by the physical conditions of the ancient areas having remained nearly the same.
Let it be remembered that the forms of life, at least those inhabiting the sea, have changed almost simultaneously throughout the world, and therefore under the most different climates and conditions.
Consider the prodigious vicissitudes of climate during the pleistocene period, which includes the whole glacial period, and note how little the specific forms of the inhabitants of the sea have been affected.
On the theory of descent, the full meaning of the fact of fossil remains from closely consecutive formations, though ranked as distinct species, being closely related, is obvious.
As the accumulation of each formation has often been interrupted, and as long blank intervals have intervened between successive formations, we ought not to expect to find, as I attempted to show in the last chapter, in any one or two formations all the intermediate varieties between the species which appeared at the commencement and close of these periods; but we ought to find after intervals, very long as measured by years, but only moderately long as measured geologically, closely allied forms, or, as they have been called by some authors, representative species; and these we assuredly do find.
We find, in short, such evidence of the slow and scarcely sensible mutation of specific forms, as we have a just right to expect to find.
|13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or||13-03 - Rules and difficulties in classification, explained on the theory of descent with modification||10||
Let us now consider the rules followed in classification, and the difficulties which are encountered on the view that classification either gives some unknown plan of creation, or is simply a scheme for enunciating general propositions and of placing together the forms most like each other.
It might have been thought (and was in ancient times thought) that those parts of the structure which determined the habits of life, and the general place of each being in the economy of nature, would be of very high importance in classification.
Nothing can be more false.
No one regards the external similarity of a mouse to a shrew, of a dugong to a whale, of a whale to a fish, as of any importance.
|01 - Variations Under Domestication||01-09 - Principles of Selection anciently followed, and their Effects||10||
Let us now briefly consider the steps by which domestic races have been produced, either from one or from several allied species.
Some effect may be attributed to the direct and definite action of the external conditions of life, and some to habit; but he would be a bold man who would account by such agencies for the differences between a dray- and race-horse, a greyhound and bloodhound, a carrier and tumbler pigeon.
One of the most remarkable features in our domesticated races is that we see in them adaptation, not indeed to the animal's or plant's own good, but to man's use or fancy.
Some variations useful to him have probably arisen suddenly, or by one step; many botanists, for instance, believe that the fuller's teasel, with its hooks, which cannot be rivalled by any mechanical contrivance, is only a variety of the wild Dipsacus; and this amount of change may have suddenly arisen in a seedling.
So it has probably been with the turnspit dog; and this is known to have been the case with the ancon sheep.
But when we compare the dray-horse and race-horse, the dromedary and camel, the various breeds of sheep fitted either for cultivated land or mountain pasture, with the wool of one breed good for one purpose, and that of another breed for another purpose; when we compare the many breeds of dogs, each good for man in different ways; when we compare the game-cock, so pertinacious in battle, with other breeds so little quarrelsome, with "everlasting layers" which never desire to sit, and with the bantam so small and elegant; when we compare the host of agricultural, culinary, orchard, and flower-garden races of plants, most useful to man at different seasons and for different purposes, or so beautiful in his eyes, we must, I think, look further than to mere variability.
We cannot suppose that all the breeds were suddenly produced as perfect and as useful as we now see them; indeed, in many cases, we know that this has not been their history. The key is man's power of accumulative selection: nature gives successive variations; man adds them up in certain directions useful to him. In this sense he may be said to have made for himself useful breeds.