On the Genesis of Species
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St. George Mivart >> On the Genesis of Species
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As yet all the changes which have taken place in pigeons are of a few
definite kinds only, such as may be well conceived to be compatible with a
species possessed of a certain inherent capacity for considerable yet
definite variation, a capacity for the ready production of certain degrees
of abnormality, which then cannot be further increased.
Mr. Darwin himself has already acquiesced in the proposition here
maintained, inasmuch as he distinctly affirms the existence of a marked
internal barrier to change in certain cases. And if this is admitted in one
case, the _principle_ is conceded, and it immediately becomes probable that
such internal barriers exist in all, although enclosing a much larger {119}
field for variation in some cases than in others. Mr. Darwin abundantly
demonstrates the variability of dogs, horses, fowls, and pigeons, but he
none the less shows clearly the _very small_ extent to which the goose, the
peacock, and the guinea-fowl have varied.[110] Mr. Darwin attempts to
explain this fact as regards the goose by the animal being valued only for
food and feathers, and from no pleasure having been felt in it on other
accounts. He adds, however, at the end the striking remark,[111] which
concedes the whole position, "but the goose seems to have _a singularly
inflexible organization_." This is not the only place in which such
expressions are used. He elsewhere makes use of phrases which quite
harmonize with the conception of a normal specific constancy, but varying
greatly and suddenly at intervals. Thus he speaks[112] of a _whole
organization seeming to have become plastic, and tending to depart from the
parental type_. That different organisms should have different degrees of
variability, is only what might have been expected _a priori_ from the
existence of parallel differences in inorganic species, some of these
having but a single form, and others being polymorphic.
To return to the goose, however, it may be remarked that it is at least as
probable that its fixity of character is the cause of the neglect, as the
reverse. It is by no means unfair to assume that _had_ the goose shown a
tendency to vary similar in degree to the tendency to variation of the fowl
or pigeon, it would have received attention at once on that account.
As to the peacock it is excused on the pleas (1), that the individuals
maintained are so few in number, and (2) that its beauty is so great it can
hardly be improved. But the individuals maintained _have not been too few_
for the independent origin of the black-shouldered form, or for the
supplanting of the commoner one by it. As to any neglect in selection,{120}
it can hardly be imagined that with regard to this bird (kept as it is all
but exclusively for its beauty), any spontaneous beautiful variation in
colour or form would have been neglected. On the contrary, it would have
been seized upon with avidity and preserved with anxious care. Yet apart
from the black-shouldered and white varieties, no tendency to change has
been known to show itself. As to its being too beautiful for improvement,
that is a proposition which can hardly be maintained. Many consider the
Javan bird as much handsomer than the common peacock, and it would be easy
to suggest a score of improvements as regards either species.
The guinea-fowl is excused, as being "no general favourite, and scarcely
more common than the peacock;" but Mr. Darwin himself shows and admits that
it is a noteworthy instance of constancy under very varied conditions.
These instances alone (and there are yet others) seem sufficient to
establish the assertion, that degree of change is different in different
domestic animals. It is, then, somewhat unwarrantable in any Darwinian to
assume that _all_ wild animals have a capacity for change similar to that
existing in _some_ of the domestic ones. It seems more reasonable to assert
the opposite, namely, that if, as Mr. Darwin says, the capacity for change
is different in different domestic animals, it must surely be limited in
those which have it least, and _a fortiori_ limited in wild animals.
Indeed, it cannot be reasonably maintained that wild species certainly vary
as much as do domestic races; it is possible that they may do so, but at
least this has not been yet shown. Indeed, the much greater degree of
variation amongst domestic animals than amongst wild ones is asserted over
and over again by Mr. Darwin, and his assertions are supported by an
overwhelming mass of facts and instances.
Of course, it may be asserted that a tendency to indefinite change exists
in all cases, and that it is only the circumstances and conditions of {121}
life which modify the effects of this tendency to change so as to produce
such different results in different cases. But assertion is not proof, and
this assertion has not been proved. Indeed, it may be equally asserted (and
the statement is more consonant with some of the facts given), that
domestication in certain animals induces and occasions a capacity for
change which is wanting in wild animals--the introduction of new causes
occasioning new effects. For, though a certain degree of variability
(normally, in all probability, only oscillation) exists in all organisms,
yet domestic ones are exposed to new and different causes of variability,
resulting in such striking divergencies as have been observed. Not even in
this latter case, however, is it necessary to believe that the variability
is indefinite, but only that the small oscillations become in certain
instances intensified into large and conspicuous ones. Moreover, it is
possible that some of our domestic animals have been in part chosen and
domesticated through possessing variability in an eminent degree.
That each species exhibits certain oscillations of structure is admitted on
all hands. Mr. Darwin asserts that this is the exhibition of a tendency to
vary which is absolutely indefinite. If this indefinite variability _does_
exist, of course no more need be said. But we have seen that there are
arguments _a priori_ and _a posteriori_ against it, while the occurrence of
variations in certain domestic animals greater in degree than the
differences between many wild species, is no argument in favour of its
existence, until it can be shown that the causes of variability in the one
case are the same as in the other. An argument against it, however, may be
drawn from the fact, that certain animals, though placed under the
influence of those exceptional causes of variation to which domestic
animals are subject, have yet never been known to vary, even in a degree
equal to that in which certain wild kinds have been ascertained to vary.
In addition to this immutability of character in some animals, it is {122}
undeniable, that domestic varieties have little stability, and much
tendency to reversion, whatever be the true explanation of such phenomena.
In controverting the generally received opinion as to "reversion," Mr.
Darwin has shown that it is not all breeds which in a few years revert to
the original form; but he has shown no more. Thus, the feral rabbits of
Porto Santo, Jamaica, and the Falkland Islands, have not yet so reverted in
those several localities.[113] Nevertheless, a Porto Santo rabbit brought
to England reverted in a manner the most striking, recovering the proper
colour of its fur "in rather less than four years."[114] Again, the white
silk fowl, in our climate, "reverts to the ordinary colour of the common
fowl in its skin and bones, due care having been taken to prevent any
cross."[115] This reversion taking place in spite of careful selection, is
very remarkable.
Numerous other instances of reversion are given by Mr. Darwin, both as
regards plants and animals; amongst others, the singular fact of bud
reversion.[116] The curiously recurring development of black sheep, in
spite of the most careful breeding, may also be mentioned, though, perhaps,
reversion has no part in the phenomenon.
These facts seem certainly to tell in favour of limited variability, while
the cases of non-reversion do not contradict it, as it is not contended
that all species have the same tendency to revert, but rather that their
capacities in this respect, as well as for change, are different in
different kinds, so that often reversion may only show itself at the end of
very long periods indeed.
Yet some of the instances given as probable or possible causes of reversion
by Mr. Darwin, can hardly be such. He cites, for example, the occasional
presence of supernumerary digits in man.[117] For this notion, however, he
is not responsible, as he rests his remark on the authority of a {123}
passage published by Professor Owen. Again, he refers[118] to "the greater
frequency of a monster proboscis in the pig than in any other animal." But
with the exception of the peculiar muzzle of the Saiga (or European
antelope), the only known proboscidian Ungulates are the elephants and
tapirs, and to neither of these has the pig any close affinity. It is
rather in the horse than in the pig that we might look for the appearance
of a reversionary proboscis, as both the elephants and the tapirs have the
toes of the hind foot of an odd number. It is true that the elephants are
generally considered to form a group apart from both the odd and the
even-toed Ungulata. But of the two, their affinities with the odd-toed
division are more marked.[119]
Another argument in favour of the, at least intermitting, constancy of
specific forms and of sudden modification, may be drawn from the absence of
minute transitional forms, but this will be considered in the next chapter.
It remains now to notice in favour of specific stability, that the
objection drawn from physiological difference between "species" and "races"
still exists unrefuted.
Mr. Darwin freely admits difficulties regarding the sterility of different
species when crossed, and shows satisfactorily that it could never have
arisen from the action of "Natural Selection." He remarks[120] also: "With
some few exceptions, in the case of plants, domesticated varieties, such as
those of the dog, fowl, pigeon, several fruit trees, and culinary
vegetables, which differ from each other in external characters more than
many species, are perfectly fertile when crossed, or even fertile in {124}
excess, whilst closely allied species are almost invariably in some degree
sterile."
Again, after speaking of "the general law of good being, derived from the
intercrossing of distinct individuals of the same species," and the
evidence that the pollen of a distinct _variety_ or race is prepotent over
a flower's own pollen, adds the very significant remark,[121] "When
distinct _species_ are crossed, the case is directly the reverse, for a
plant's own pollen is almost always prepotent over foreign pollen."
Again he adds:[122] "I believe from observations communicated to me by Mr.
Hewitt, who has had great experience in hybridizing pheasants and fowls,
that the early death of the embryo is a very frequent cause of sterility in
first crosses. Mr. Salter has recently given the results of an examination
of about 500 eggs produced from various crosses between three species of
Gallus and their hybrids. The majority of these eggs had been fertilized,
and in the majority of the fertilized eggs the embryos either had been
partially developed and had then aborted, or had become nearly mature, but
the young chickens had been unable to break through the shell. Of the
chickens which were born, more than four-fifths died within the first few
days, or at latest weeks, 'without any obvious cause, apparently from mere
inability to live,' so that from 500 eggs only twelve chickens were reared.
The early death of hybrid embryos probably occurs in like manner with
plants, at least it is known that hybrids raised from very distinct species
are sometimes weak and dwarfed, and perish at an early age, of which fact
Max Wichura has recently given some striking cases with hybrid willows."
Mr. Darwin objects to the notion that there is any special sterility
imposed to check specific intermixture and change, saying,[123] "To grant
to species the special power of producing hybrids, and then to stop {125}
their further propagation by different degrees of sterility, not strictly
related to the facility of the first union between their parents, seems a
strange arrangement."
But this only amounts to saying that the author himself would not have so
acted had he been the Creator. A "strange arrangement" must be admitted
anyhow, and all who acknowledge teleology at all, must admit that the
strange arrangement was designed. Mr. Darwin says, as to the sterility of
species, that the cause lies exclusively in their sexual constitution; but
all that need be affirmed is that sterility is brought about somehow, and
it is undeniable that "crossing" _is_ checked. All that is contended for is
that there _is_ a bar to the intermixture of _species_, but not of
_breeds_; and if the conditions of the generative products are that bar, it
is enough for the argument, no special kind of barring action being
contended for.
He, however, attempts to account for the modification of the sexual
products of species as compared with those of varieties, by the exposure of
the former to more uniform conditions during longer periods of time than
those to which varieties are exposed, and that as wild animals, when
captured, are often rendered sterile by captivity, so the influence of
union with another species may produce a similar effect. It seems to the
author an unwarrantable assumption that a cross with what, on the Darwinian
theory, can only be a slightly diverging descendant of a common parent,
should produce an effect equal to that of captivity, and consequent change
of habit, as well as considerable modification of food.
No clear case has been given by Mr. Darwin in which mongrel animals,
descended from the same undoubted species, have been persistently infertile
_inter se_; nor any clear case in which hybrids between animals, generally
admitted to be distinct species, have been continuously fertile _inter se_.
It is true that facts are brought forward tending to establish the
probability of the doctrine of Pallas, that species may sometimes be {126}
rendered fertile by domestication. But even if this were true, it would be
no approximation towards proving the converse, _i.e._ that races and
varieties may become sterile when wild. And whatever may be the preference
occasionally shown by certain breeds to mate with their own variety, no
sterility is recorded as resulting from unions with other varieties.
Indeed, Mr. Darwin remarks,[124] "With respect to sterility from the
crossing of domestic races, I know of no well-ascertained case with
animals. This fact (seeing the great difference in structure between some
breeds of pigeons, fowls, pigs, dogs, &c.) is extraordinary when contrasted
with the sterility, of many closely-allied natural species when crossed."
It has been alleged that the domestic and wild guinea-pig do not breed
together, but the specific identity of these forms is very problematical.
Mr. A. D. Bartlett, superintendent of the Zoological Gardens, whose
experience is so great, and observation so quick, believes them to be
decidedly distinct species.
Thus, then, it seems that a certain normal specific stability in species,
accompanied by occasional sudden and considerable modifications, might be
expected _a priori_ from what we know of crystalline inorganic forms and
from what we may anticipate with regard to the lowest organic ones. This
presumption is strengthened by the knowledge of the increasing difficulties
which beset any attempt to indefinitely intensify any race characteristics.
The obstacles to this indefinite intensification, as well as to certain
lines of variation in certain cases, appear to be not only external, but to
depend on internal causes or an internal cause. We have seen that Mr.
Darwin himself implicitly admits the principle of specific stability in
asserting the singular inflexibility of the organization of the goose. We
have also seen that it is not fair to conclude that all wild races can vary
as much as the most variable domestic ones. It has also been shown {127}
that there are grounds for believing in a tendency to reversion generally,
as it is distinctly present in certain instances. Also that specific
stability is confirmed by the physiological obstacles which oppose
themselves to any considerable or continued intermixture of species, while
no such barriers oppose themselves to the blending of varieties. All these
considerations taken together may fairly be considered as strengthening the
belief that specific manifestations are relatively stable. At the same time
the view advocated in this book does not depend upon, and is not identified
with, any such stability. All that the Author contends for is that specific
manifestation takes place along certain lines, and according to law, and
not in an exceedingly minute, indefinite, and fortuitous manner. Finally,
he cannot but feel justified, from all that has been brought forward, in
reiterating the opening assertion of this chapter that something is still
to be said for the view which maintains that species are stable, at least
in the intervals of their comparatively rapid successive {128}
manifestations.
* * * * *
CHAPTER VI.
SPECIES AND TIME.
Two relations of species to time.--No evidence of past existence of
minutely intermediate forms when such might be expected _a
priori_.--Bats, Pterodactyles, Dinosauria, and Birds.--Ichthyosauria,
Chelonia, and Anoura.--Horse ancestry.--Labyrinthodonts and
Trilobites.--Two subdivisions of the second relation of species to
time.--Sir William Thomson's views.---Probable period required for
ultimate specific evolution from primitive ancestral
forms.--Geometrical increase of time required for rapidly multiplying
increase of structural differences.--Proboscis monkey.--Time required
for deposition of strata necessary for Darwinian evolution.--High
organization of Silurian forms of life.--Absence of fossils in oldest
rocks.--Summary and conclusion.
Two considerations present themselves with regard to the necessary relation
of species to time if the theory of "Natural Selection" is valid and
sufficient.
The first is with regard to the evidences of the past existence of
intermediate form, their duration and succession.
The second is with regard to the total amount of time required for the
evolution of all organic forms from a few original ones, and the bearing of
other sciences on this question of time.
As to the first consideration, evidence is as yet against the modification
of species by "Natural Selection" alone, because not only are minutely
transitional forms generally absent, but they are absent in cases where we
might certainly _a priori_ have expected them to be present. [Page 129]
Now it has been said:[125] "If Mr. Darwin's theory be true, the number of
varieties differing one from another a very little must have been
indefinitely great, so great indeed as probably far to exceed the number of
individuals which have existed of any one variety. If this be true, it
would be more probable that no two specimens preserved as fossils should be
of one variety than that we should find a great many specimens collected
from a very few varieties, provided, of course, the chances of preservation
are equal for all individuals." "It is really strange that vast numbers of
perfectly similar specimens should be found, the chances against their
perpetuation as fossils are so great; but it is also very strange that the
specimens should be so exactly alike as they are, if, in fact, they came
and vanished by a gradual change."
Mr. Darwin attempts[126] to show cause why we should believe _a priori_
that intermediate varieties would exist in lesser numbers than the more
extreme forms; but though they would doubtless do so sometimes, it seems
too much to assert that they would do so generally, still less universally.
Now little less than universal and very marked inferiority in numbers would
account for the absence of certain series of minutely intermediate fossil
specimens. The mass of palaeontological evidence is indeed overwhelmingly
against minute and gradual modification. It is true that when once an
animal has obtained powers of flight its means of diffusion are
indefinitely increased, and we might expect to find many relics of an
aerial form and few of its antecedent state--with nascent wings just
commencing their suspensory power. Yet had such a slow mode of origin, as
Darwinians contend for, operated exclusively in all cases, it is absolutely
incredible that birds, bats, and pterodactyles should have left the remains
they have, and yet not a single relic be preserved in any one instance{130}
of any of these different forms of wing in their incipient and relatively
imperfect functional condition!
[Illustration: WING-BONES OF PTERODACTYLE, BAT, AND BIRD.]
Whenever the remains of bats have been found they have presented the exact
type of existing forms, and there is as yet no indication of the conditions
of an incipient elevation from the ground.
The pterodactyles, again, though a numerous group, are all true and perfect
pterodactyles, though surely _some_ of the many incipient forms, which on
the Darwinian theory have existed, must have had a good chance of
preservation.
As to birds, the only notable instance in which discoveries recently made
appear to fill up an important hiatus, is the interpretation given by
Professor Huxley[127] to the remains of Dinosaurian reptiles, and which
were noticed in the third chapter of this work. The learned Professor has
(as also has Professor Cope in America) shown that in very important {131}
and significant points the skeletons of the Iguanodon and of its allies
approach very closely to that existing in the ostrich, emeu, rhea, &c. He
has given weighty reasons for thinking that the line of affinity between
birds and reptiles passes to the birds last named from the Dinosauria
rather than from the Pterodactyles, through Archeopteryx-like forms to the
ordinary birds. Finally, he has thrown out the suggestion that the
celebrated footsteps left by some extinct three-toed creatures on the very
ancient sandstone of Connecticut were made, not, as hitherto supposed, by
true birds, but by more or less ornithic reptiles. But even supposing all
that is asserted or inferred on this subject to be fully proved, it would
not approach to a demonstration of specific origin by _minute_
modification. And though it harmonizes well with "Natural Selection," it is
equally consistent with the rapid and sudden development of new specific
forms of life. Indeed, Professor Huxley, with a laudable caution and
moderation too little observed by some Teutonic Darwinians, guarded himself
carefully from any imputation of asserting dogmatically the theory of
"Natural Selection," while upholding fully the doctrine of evolution.
But, after all, it is by no means certain, though very probable, that the
Connecticut footsteps were made by very ornithic reptiles, or extremely
sauroid birds. And it must not be forgotten that a completely carinate[128]
bird (the Archeopteryx) existed at a time, when, as yet, we have no
evidence of some of the Dinosauria having come into being. Moreover, if the
remarkable and minute similarity of the coracoid of a pterodactyle to that
of a bird be merely the result of function and no sign of genetic affinity,
it is not inconceivable that pelvic and leg resemblances of Dinosauria to
birds may be functional likewise, though such an explanation is, of {132}
course, by no means necessary to support the view maintained in this book.
[Illustration: THE ARCHEOPTERYX (OF THE OOLITE STRATA).]
[Illustration: SKELETON OF AN ICHTHYOSAURUS.]
But the number of forms represented by many individuals, yet by _no
transitional ones_, is so great that only two or three can be selected as
examples. Thus those remarkable fossil reptiles, the Ichthyosauria and
Plesiosauria, extended, through the secondary period, probably over the
greater part of the globe. Yet no single transitional form has yet been met
with in spite of the multitudinous individuals preserved. Again, with their
modern representatives the Cetacea, one or two aberrant forms alone {133}
have been found, but no series of transitional ones indicating minutely the
line of descent. This group, the whales, is a very marked one, and it is
curious, on Darwinian principles, that so few instances tending to indicate
its mode of origin should have presented themselves. Here, as in the bats,
we might surely expect that some relics of unquestionably incipient stages
of its development would have been left.
[Illustration: SKELETON OF A PLESIOSAURUS.]
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