_ON THE ORIGIN OF INSECTS._
"Personne," says Carl Vogt, "en Europe au moins, n'ose plus soutenir la
Création indépendante et de toutes pièces des espèces," and though this
statement is perhaps not strictly correct, still it is no doubt true,
that the Doctrine of Evolution, in some form or other, is accepted by
most, if not by all, the greatest naturalists of Europe. Yet it is
surprising how much, in spite of all that has been written, Mr. Darwin's
views are still misunderstood. Thus Browning, in one of his recent
poems, says:--
"That mass man sprang from was a jelly lump
Once on a time; he kept an after course
Through fish and insect, reptile, bird, and beast,
Till he attained to be an ape at last,
Or last but one."[53]
This theory, though it would be regarded by many as a fair statement of
his views, is one which Mr. Darwin would entirely repudiate. Whether
fish and insect, reptile, bird and beast, are derived from one original
stock or not, they are certainly not links in one sequence. I do not,
however, propose to discuss the question of Natural Selection, but may
observe that it is one thing to acknowledge that in Natural Selection,
or the survival of the fittest, Mr. Darwin has called attention to a
_vera causa_, has pointed out the true explanation of certain phenomena;
but it is quite another thing to maintain that all animals are descended
from some primordial source.
For my own part, I am satisfied that Natural Selection is a true cause,
and, whatever may be the final result of our present inquiries--whether
animated nature be derived from one ancestral source, or from many--the
publication of the Origin of Species will none the less have constituted
an epoch in the History of Biology. But, how far the present condition
of living beings is due to that cause; how far, on the other hand, the
action of Natural Selection has been modified and checked by other
natural laws--by the unalterability of types, by atavism, &c.; how many
types of life originally came into being; and whether they arose
simultaneously or successively,--these and many other similar questions
remain unsolved, even admitting the theory of Natural Selection. All
this has indeed been clearly pointed out by Mr. Darwin himself, and
would not need repetition but for the careless criticism by which in too
many cases the true question has been obscured. Without, however,
discussing the argument for and against Mr. Darwin's conclusions, so
often do we meet with travesties of it like that which I have just
quoted, that it is well worth while to consider the stages through which
some group, say for instance that of insects, have probably come to be
what they are, assuming them to have developed under natural laws from
simpler organisms. The question is one of great difficulty. It is hardly
necessary to say that insects cannot have passed through all the lower
forms of animal life, and naturalists do not at present agree as to the
actual line of their development.
In the case of insects, the gradual course of evolution through which
the present condition of the group has probably been reached, has been
discussed by Mr. Darwin, by Fritz Müller, Haeckel, Brauer, myself and
others.
In other instances Palæontology throws much light on this question.
Leidy has shown that the milk-teeth of the genus _Equus_ resemble the
permanent teeth of the ancient _Anchitherium_, while the milk-teeth of
_Anchitherium_ again approximate to the dental system of the still
earlier _Merychippus_. Rütimeyer, while calling attention to this
interesting observation, adds that the milk-teeth of _Equus caballus_ in
the same way, and still more those of _E. fossilis_, resemble the
permanent teeth of _Hipparion_.
"If we were not acquainted with the horse," says Flower,[54] "we could
scarcely conceive of an animal whose only support was the tip of a
single toe on each extremity, to say nothing of the singular
conformation of its teeth and other organs. So striking have these
characters appeared to many zoologists, that the animals possessing them
have been reckoned as an order apart, called Solidungula; but
palæontology has revealed that in the structure of its skull, its teeth,
its limbs, the horse is nothing more than a modified _Palæotherium_; and
though still with gaps in certain places, many of the intermediate
stages of these modifications are already known to us, being the
_Palæotherium_, _Anchitherium_, _Merychippus_, and _Hipparion_."
"All Echinoids," says A. Agassiz,[55] "pass, in their early stages,
through a condition which recalls to us the first Echinoids which made
their appearance in geological ages." On embryological grounds, he
observes, we should "place true Echini lowest, then the Clypeastroids,
next the Echinolamps, and finally the Spatangoids." Now among the
Echinoids of the Trias there are no Clypeastroids, Echinolamps, or
Spatangoids. The Clypeastroids make their appearance in the Lias, the
Echinolamps in the Jurassic, while the Spatangoids commence in the
Cretaceous period.
Again[56] "in the Radiates, the Acalephs in their first stages of
growth, that is, in their Hydroid condition, remind us of the adult
forms among Polyps, showing the structural rank of the Acalephs to be
the highest, since they pass beyond a stage which is permanent with the
Polyps; while the Adult forms of the Acalephs have in their turn a
certain resemblance to the embryonic phases of the class next above
them, the Echinoderms; within the limits of the classes, the same
correspondence exists as between the different orders; the embryonic
forms of the highest Polyps recall the adult forms of the lower ones,
and the same is true of the Acalephs as far as these phenomena have been
followed and compared among them." Indeed, the accomplished authors from
whom I have taken the above quotation, do not hesitate to say[57] that
"whenever such comparisons have been successfully carried out, the
result is always the same; the present representatives of the fossil
types recall in their embryonic condition the ancient forms, and often
explain their true position in the animal kingdom."
Fossil insects are unfortunately rare, there being but few strata in
which the remains of this group are well preserved. Moreover,
well-characterized Orthoptera and Neuroptera occur as early as the
Devonian strata; Coleoptera and Hemiptera in the Coal-measures;
Hymenoptera and Diptera in the Jurassic; Lepidoptera, on the contrary,
not until the Tertiary. But although it appears from these facts that,
as far as our present information goes, the Orthoptera and Neuroptera
are the most ancient orders, it is not, I think, conceivable that the
latter should have been derived from any known species of the former; on
the other hand, the earliest known Neuroptera and Orthoptera, though in
some respects less specialized than existing forms, are as truly, and as
well characterized, Insects, as any now existing; nor are we acquainted
with any earlier forms, which in any way tend to bridge over the gap
between them and lower groups, though, as we shall see, there are types
yet existing which throw much light on the subject.
In the consideration then of this question, we must rely principally on
Embryology and Development. I have already referred to the cases in
which species, very unlike in their mature condition, are very
similar one to another when young. Haeckel, in his "Naturliche
Schöpfungsgeschichte," gives a diagram which illustrates this very well
as regards Crustacea. Pls. 1-4 show the same to be the case with
Insects.
The Stag-beetle, the Dragon-fly, the Moth, the Bee, the Ant, the Gnat,
the Grasshopper,--these and other less familiar types seem at first to
have little in common. They differ in size, in form, in colour, in
habits, and modes of life. Yet the researches of entomologists,
following the clue supplied by the illustrious Savigny, have proved, not
only that while differing greatly in details, they are constructed on
one common plan; but also that other groups, as for instance, Crustacea
(Lobsters, Crabs, &c.) and Arachnida (Spiders and Mites), can be shown
to be fundamentally similar. In Pl. 4 I have figured the larvæ of an
_Ephemera_ (Fig. 1), of a _Meloë_ (Fig. 2), of a Dragon-fly (Fig. 3), of a
Sitaris (Fig. 4), of a _Campodea_ (Fig. 5), of a _Dyticus_ (Fig. 6), of a
Termite (Fig. 7), of a _Stylops_ (Fig. 8), and of a _Thrips_ (Fig. 9). All
these larvæ possess many characters in common. The mature forms are
represented in the corresponding figures of Plate 3, and it will at once
be seen how considerably they differ from one another. The same fact is
also illustrated in Figs. 48-55, where Figs. 48-51 represent the larval
states of the mature forms represented in Figs. 52-55. Fig. 48 is the
larva of a moth, _Agrotis suffusa_ (Fig. 52); Fig. 49 of a beetle,
_Haltica_ (Fig. 53); Fig. 50 of a Saw-fly, _Cimbex_ (Fig. 54); and Fig.
51 of a Centipede, _Julus_ (Fig. 55).
[Illustration: FIG. 48, Larva of Moth (_Agrotis suffusa_), after
Packard. 49, Larva of Beetle (_Haltica_), after Westwood. 50, Larva of
Saw-fly (_Cimbex_), Brischke and Zaddach. Beob. ub d. arten. der Blatt
und Holzwespen, Fig. 8. 51, Larva of _Julus_. Newport, Philos.
Transactions, 1841.]
Thus, then, although it can be demonstrated that perfect insects,
however much they differ in appearance, are yet reducible to one type,
the fact becomes much more evident if we compare the larvæ. M.
Brauer[58] and I[59] have pointed out that two types of larvæ, which I
have proposed to call _Campodea_-form and _Lindia_-form, and which Packard
has named Leptiform and Eruciform, run through the principal groups of
insects. This is obviously a fact of great importance: as all individual
_Meloës_ are derived from a form resembling Pl. 2, Fig. 2, it is surely no
rash hypothesis to suggest that the genus itself may have been so.
[Illustration: FIG. 52, _Agrotis suffusa_ (after Packard). 53, _Haltica_
(after Westwood).]
[Illustration: FIG. 54, _Cimbex_, Brischae and Zaddach. l.c. T. 2, Fig.
9.]
[Illustration: FIG. 55. _Julus_ (after Gervais).]
Firstly, however, let me say a word as to the general Insect type. It
may be described shortly as consisting of animals possessing a head,
with mouth parts, eyes and antennæ; a many segmented body, with three
pairs of legs on the segments immediately following the head; with, when
mature, either one or two pairs of wings, generally with caudal
appendages I will not now enter into a description of their internal
anatomy. It will be seen that, except as regards the wings, Pl. 4, Fig.
4, representing the larva of a small beetle named _Sitaris_, answers very
well to this description. Many other Beetles are developed from larvæ
closely resembling those of _Meloë_ (Pl. 4, Fig. 2), and Sitaris (Pl. 4,
Fig. 4); in fact--except those species the larvæ of which, as, for
instance of the Weevils (Pl. 2, Fig. 6), are internal feeders, and do
not require legs--we may say that the Coleoptera generally are derived
from larvæ of this type.
I will now pass to a second order, the Neuroptera. Pl. 4, Fig. 1,
represents the larva of _Chloëon_, a species the metamorphoses of which I
described some years ago in the Linnean Transactions,[60] and it is
obvious that in essential points it closely resembles the form to which
I have just alluded.
The Orthoptera, again, the order to which Grasshoppers, Crickets,
Locusts, &c. belong, commence life in a similar condition; and the same
may also be said of the Trichoptera.
The larvæ of Bees when they quit the egg are entirely legless, but in an
earlier stage they possess well-marked rudiments of thoracic legs,
showing, as it seems to me, that their apodal condition is an adaptation
to their circumstances. Other Hymenopterous larvæ, those for example of
_Sirex_ (Fig. 9), and of the Saw-flies (Fig. 50) have well-developed
thoracic legs.
From the difference in external form, and especially from the large
comparative size of the abdomen, these larvæ, as well as those of
Lepidoptera (Fig. 48), have generally been classed with the maggots of
Flies, Weevils, &c., rather than with the more active form of larva just
adverted to. This seems to me, as I have already pointed out,[61] to be
a mistake. The caterpillar type differs, no doubt, in its general
appearance, owing to its greater clumsiness, but still essentially
agrees with that already described.
No Dipterous larva, so far as I know, belongs truly to this type; in
fact, the early stages of the pupa in the Diptera seem in some respects
to correspond to the larvæ of other Insect orders. The Development of
the Diptera is, however, as Weissman[62] has shown, very abnormal in
other respects.
Thus, then, we find in many of the principal groups of insects that,
greatly as they differ from one another in their mature condition, when
they leave the egg they more nearly resemble the typical insect type;
consisting of a head; a three-segmented thorax, with three pairs of
legs; and a many-jointed abdomen, often with anal appendages. Now, is
there any mature animal which answers to this description? We need not
have been surprised if this type, through which it would appear that
insects must have passed so many ages since (for winged Neuroptera have
been found in the carboniferous strata) had long ago become extinct. Yet
it is not so. The interesting genus _Campodea_ (Pl. 3, Fig. 5) still
lives; it inhabits damp earth, and closely resembles the larva of
_Chloëon_ (Pl. 2, Fig. 1), constituting, indeed, a type which, as shown in
Pl. 4, occurs in many orders of insects. It is true that the
mouth-parts of _Campodea_ do not resemble either the strongly mandibulate
form which prevails among the larvæ of Coleoptera, Orthoptera,
Neuroptera, Hymenoptera, Lepidoptera; or the suctorial type of the
Homoptera and Heteroptera. It is, however, not the less interesting or
significant on that account, since, as I have elsewhere[63] pointed out,
its mouth-parts are intermediate between the mandibulate and haustellate
types; a fact which seems to me most suggestive.
It appears, then, that there are good grounds for considering that the
various types of insects are descended from ancestors more or less
resembling the genus _Campodea_, with a body divided into head, thorax,
and abdomen: the head provided with mouth-parts, eyes, and one pair of
antennæ; the thorax with three pairs of legs; and the abdomen, in all
probability, with caudal appendages.
If these views are correct, the genus _Campodea_ must be regarded as a
form of remarkable interest since it is the living representative of a
primæval type, from which not only the Collembola and Thysanura, but the
other great orders of insects have derived their origin.
From what lower group the _Campodea_ type was itself derived is a question
of great difficulty. Fritz Müller indeed says,[64] "if all the classes
of Arthropoda (Crustacea, Insecta, Myriopoda, and Arachnida) are indeed
all branches of a common stem (and of this there can scarcely be a
doubt), it is evident that the water-inhabiting and water-breathing
Crustacea must be regarded as the original stem from which the other
terrestrial classes, with their tracheal respiration, have branched
off." Haeckel, moreover, is of the opinion that the Tracheata are
developed from the Crustacea, and probably from the Zoëpoda. For my own
part, though I feel very great diffidence in expressing an opinion at
variance with that of such high authorities, I am rather disposed to
suggest that the _Campodea_ type may possibly have been derived from a
less highly developed one, resembling the modern Tardigrade,[65] a (Fig.
56) smaller and much less highly organized being than _Campodea_. It
possesses two eyes, three anterior pairs of legs, and one at the
posterior end of the body, giving it a curious resemblance to some
Lepidopterous larvæ.
[Illustration: FIG. 56, Tardigrade (after Dujardin).]
These legs, however, as will be seen, are reduced to mere projections.
But for them, the Tardigrada would closely resemble the vermiform larva
so common among insects. Among Trichoptera the larva early acquires
three pairs of legs, but as Zaddach has shown,[66] there is a stage,
though it is quickly passed through, in which the divisions of the body
are indicated, but no trace of legs is yet present. Indeed, there appear
to be reasons for considering that while among Crustacea the appendages
appear before the segments, in Insects the segments precede the
appendages, although this stage of development is very transitory, and
apparently, in some cases, altogether suppressed. I say "apparently,"
because, as I have already mentioned, I am not yet satisfied that it
will not eventually be found to be so in all cases. Zaddach, in his
careful observations of the embryology of _Phryganea_, only once found a
specimen in this stage, which also, according to the researches of
Huxley,[67] seems to be little more than indicated in _Aphis_. It is
therefore possible that in other cases, when no such stage has been
observed, it not really may be absent, but, from its transitoriness, may
have hitherto escaped attention.
Fritz Müller has expressed the opinion[68] that this vermiform type is
of comparatively recent origin. He says: "The ancient insects approached
more nearly to the existing Orthoptera, and perhaps to the wingless
Blattidæ, than to any other order, and the complete metamorphosis of the
Beetles, Lepidoptera, &c., is of later origin." "There were," he adds,
"perfect insects before larvæ and pupæ." This opinion has been adopted
by Mr. Packard[69] in his "Embryological Studies on Hexapodous Insects."
M. Brauer[70] also considers that the vermiform larva is a more recent
type than the Hexapod form, and is to be regarded not as a developmental
form, but as an adaptational modification of the earlier active hexapod
type. In proof of this he quotes the case of _Sitaris_.
Considering, however, the peculiar habits of this genus, to which I have
already referred, and also that the vermiform type is altogether lower
in organization and less differentiated than the _Campodea_ form, I cannot
but regard this case as exceptional; one in which the development has
been, as it were, to use an expression of Fritz Müller's, "falsified" by
the struggle for existence, and which therefore does not truly indicate
the successive stages of evolution. On the whole, the facts seem to me
to point to the conclusion that, though the grub-like larvæ of Coleoptera
and some other insects, owe their present form mainly to the influence
of external circumstances, and partially also to atavism, still the
_Campodea_ type is itself derived from earlier vermiform ancestors.
Nicolas Wagner has shown in the case of a small gnat, allied to
_Cecidomyia_, that even now, in some instances, the vermiform larvæ
possess the power of reproduction. Such a larva (as, for instance, Fig.
57) very closely resembles some of the Rotatoria, such for instance as
_Albertia_ or _Notommata_, which however possess vibratile cilia. There is,
indeed, one genus--_Lindia_ (Fig. 58)--in which these ciliæ are altogether
absent, and which, though resembling _Macrobiotus_ in many respects,
differs from that genus in being entirely destitute of legs. I have
never met with it myself, but it is described by Dujardin, who found it
in a ditch near Paris, as being oblong, vermiform, divided into rings,
and terminating posteriorly in two short conical appendages. The jaws
are not unlike those of the larvæ of Flies, and indeed many naturalists
meeting with such a creature would, I am sure, regard it as a small
Dipterous larva; yet Dujardin figures a specimen containing an egg, and
seems to have no doubt that it is a mature form.[71]
For the next descending stage we must, I think, look among the
Infusoria, through such genera as _Chætonotus_ or _Ichthydium_. Other
forms of the Rotatoria, such for instance as _Rattulus_, and still more
the very remarkable species discovered in 1871 by Mr. Hudson,[72] and
described under the name of _Pedalion mira_, seem to lead to the
Crustacea through the Nauplius form. Dr. Cobbold tells me that he
regards the _Gordii_ as the lowest of the Scolecida; Mr. E. Ray
Lankester considers some of the Turbellaria, such genera as
_Mesostomum_, _Vortex_, &c., to be the lowest of existing worms;
excluding the parasitic groups. Haeckel[73] also regards the Turbellaria
as forming the nearest approach to the Infusoria. The true worms seem,
however, to constitute a separate branch of the animal kingdom.
We may take, as an illustration of the lower worms, the genus
Prorhynchus (Fig. 59), which consists of a hollow cylindrical body,
containing a straight simple tube, the digestive organ.
But however simple such a creature as this may be, there are others
which are far less complex, far less differentiated; which therefore, on
Mr. Darwin's principles, may be considered still more closely to
represent the primæval ancestor from which these more highly-developed
types have been derived, and which, in spite of their great
antiquity--in spite of, or perhaps in consequence of, their simplicity,
still maintain themselves almost unaltered.
Thus the form which Haeckel has described[74] under the name
_Protamoeba primitiva_, Pl. 5, Fig. 1-5, consists of a homogeneous and
structureless substance, which continually alters its form; putting out
and drawing in again more or less elongated processes, and creeping
about like a true _Amoeba_, from which, however, _Protamoeba_ differs,
in the absence of a nucleus. It seems difficult to imagine anything
simpler; indeed, as described, it appears to be an illustration of
properties without structure. It takes into itself any suitable particle
with which it comes in contact, absorbs that which is nutritious, and
rejects the rest. From time to time a constriction appears at the centre
(Pl. 5, Fig. 2), its form approximates more and more to that of an
hour-glass (Pl. 5, Fig. 3), and at length the two halves separate, and
each commences an independent existence (Pl. 5, Fig. 5).
[Illustration: FIG. 59, _Prorhynchus stagnaus_.[75]]
[Illustration: PLATE V.
FIGS. 1-5, _Protamoeba_; 6-9, _Protamyxa Aurantiaca_,
Haeckel, Beit. zur Monog. der Moneren, pl. 1; 10-18, _Magosphæra planula_,
Haeckel, loc. cit. pl. 5.]
In the true _Amoebas_, on the contrary, we find a differentiation
between the exterior and the interior: the body being more or less
distinctly divisible into an outer layer and an inner parenchyme. In the
_Amoebas_, as in _Protamoeba_, multiplication takes place by
self-division, and nothing corresponding to sexual reproduction has yet
been discovered.
Somewhat more advanced, but still of great simplicity, is the _Protomyxa
aurantiaca_ (Pl. 5, Fig. 8), discovered by Haeckel[76] on dead shells of
_Spirula_, where it appears as a minute orange speck, which shows well
against the clear white of the _Spirula_. Examined with a microscope, the
speck is seen to be a spherical mass of orange-coloured, homogeneous,
albuminous matter, surrounded by a delicate, structureless membrane. It
is obvious from this description that these bodies closely resemble
eggs, for which indeed Haeckel at first mistook them. Gradually,
however, the yellow sphere broke itself up into smaller spherules (Pl.
5, Fig. 9), after which the containing membrane burst, and the separate
spherules, losing their globular form, crept out as small _Amoebæ_ (Pl.
5, Fig. 6), or amoeboid bodies. These little bodies moved about,
assimilated the minute particles of organic matter, with which they came
in contact, and gradually increased in size (Pl. 5, Fig. 7) with more or
less rapidity according to the amount of nourishment they were able to
obtain. They threw out arms in various directions, and if divided each
section maintained its individual existence. After a while their
movements ceased, they contracted into a ball, and again secreted round
themselves a clear structureless envelope.
This completes their life history as observed by Haeckel, who found it
easy to retain them in his glasses in perfect health, and who watched
them closely.
As another illustration I may take the _Magosphæra planula_, discovered
by Haeckel on the coast of Norway.
In one stage of its existence (Pl. 5, Fig. 10) it is a minute mass of
gelatinous matter, which continually alters its form, moves about,
feeds, and in fact behaves altogether like the _Amoeba_ just described.
It does not, however, remain always in this condition. After a while it
contracts into a spherical form (Pl. 5, Fig. ii), and secretes round
itself a structureless envelope, which, with the nucleus, gives it a
very close resemblance to a minute egg.
Gradually the nucleus divides, and the protoplasm also separates into
two spherules (Pl. 5, Fig. 12); these two subdivide into four (Pl. 5,
Fig. 13), and so on (Pl. 5, Fig 14), until at length thirty-two are
present, compressed into a more or less polygonal form (Pl. 5, Fig. 15).
Here this process ends. The separate spherules now begin to lose their
smooth outline, to throw out processes, and to show amoeboid movements
like those of the creatures just described. The processes or pseudopods
grow gradually longer, thinner, and more pointed. Their movements become
more active, until at length they take the form of ciliæ. The spherical
_Magosphæra_, the upper surface of which has thus become covered with
ciliæ, now begins to rotate within the cyst or envelope, which at length
gives way and sets free the contained sphere, which then swims about
freely in the water (Pl. 5, Fig. 16), thus closely resembling _Synura_, or
one of the Volvocineæ. After swimming about in this condition for a
certain time, the sphere breaks up into the separate cells of which it
is composed (Pl. 5, Fig. 17). As long as the individual cells remained
together, they had undergone no changes of form, but after separating
they show considerable contractility, and gradually alter their form,
until they become undistinguishable from true _Amoebæ_ (Pl. 5, Fig. 18).
Finally, according to Haeckel, these amoeboid bodies, after living for
a certain time in this condition, return to a state of rest, again
contract into a spherical form, and secrete round themselves a
structureless envelope. The life history of some other low organisms, as
for instance _Gregarina_, is of a similar character.
It may be said, and said truly, that the difference between such beings
as these and the _Campodea_, or Tardigrade, is immense. But if it be
considered incredible that even during the long lapse of geological time
such great changes should have taken place as are implied in the belief
that there is genetic connection between them and these lower groups,
let us consider what happens under our eyes in the development of each
one of these little creatures in the proverbially short space of their
individual life.
I will take for instance the first stages, and for the sake of brevity
only the first stages, of the life-history of a Tardigrade.[77] As shown
in Fig. 60, the egg is at first a round body or cell, with a clear
central nucleus--the germinal vesicle; it increases in size, and after
a while the yolk and the germinal vesicle divide into two (Fig. 61),
then into four (Fig. 62), and so on, just as we have seen to be the case
in _Magosphæra_. From the minute cells (Fig. 63) arising through this
process of yolk-segmentation, the body of the Tardigrade is then built
up.[78]
[Illustration: FIG. 60, Egg of Tardigrade, Kaufmann, Zeit f. Wiss.
Zool. 1851, Pl. 1. 61, Egg of Tardigrade after the yolk has subdivided.
62, Egg of Tardigrade in the next stage. 63, Egg of Tardigrade more
advanced.]
Though I will not now attempt to point out the full bearing of these
facts on the study of embryology generally, yet I cannot resist calling
attention to the similarity of the development of _Magosphæra_ with the
first stages of development of other animals, because it appears to me
to possess a significance, the importance of which it would be difficult
to overestimate.
Among the Zoophytes Prof. Allman thus describes[79] the process in
_Laomedea_, as representing the Hydroids (Pl. 6, Fig. 1, represents the
young egg):--"The first step observable in the segmentation-process is
the cleavage of the yolk into two segments (Pl. 6, Fig. 2), immediately
followed by the cleavage of these into other two, so that the vitellus
is now composed of four cleavage spheres (Pl. 6, Fig. 3)." These spheres
again divide (Pl. 6, Fig. 4) and subdivide, thus at length forming
minute cells, of which the body of the embryo is built up.
In Pl. 6, Figs. 5-9 represent the corresponding stages in the
development of a small parasitic worm--the _Filaria mustelarum_--as
given by Van Beneden.[80] The first process is that within the egg,
which represents, so to say, the encysted condition of _Magosphæra_, the
yolk divides itself into two balls (Pl. 6, Fig. 6), then into four,
eight, and so on, the cells thus constituted finally forming the young
worm. I have myself observed the same stages in the eggs of the very
remarkable and abnormal _Sphærularia bombi_.[81]
Among the Echinoderms M. Derbès thus describes the first stages (Pl. 6,
Figs. 10-13) in the development of the egg of an _Echinus_ (_Echinus
esculentus_):--"Le jaune commence à se segmenter, d'abord en deux, puis
en quatre et ainsi de suite, chacune des nouvelles cellules se
partageant à son tour en deux."[82] Sars has observed the same thing in
the starfish.[83]
[Illustration: PLATE. 6.]
In the Rotatoria, as shown by Huxley in _Lacinularia_,[84] and by
Williamson in _Melicerta_,[85] the yolk is at first a single globular
mass, the first changes which take place in it being as follows:--"The
central nucleus becomes drawn out and subdivides into two, this division
being followed by a corresponding segmentation of the yolk. The same
process is repeated again and again, until at length the entire yolk is
converted into a mass of minute cells." Among the Crustacea the total
segmentation of the yolk occurs among the Copepoda, Rhizocephala, and
Cirripedia. Sars has described the same process in one of the
nudibranchiate mollusca[86] (_Tritonia_), Müller in Entochocha,[87]
Haeckel in Ascidia,[88] Lacaze Duthiers in _Dentalium_.[89] Figures 18 to
21, Pl. 6, are taken from Koren and Danielssen's[90] memoir on the
development of _Purpura lapillus_.
Figs. 22-24 show the same stages in a fish (_Amphioxus_) as given by
Haeckel, and it is unnecessary to point out the great similarity.
Lastly, figures 25 to 29, Pl. 6, are given by Dr. Allen Thomson,[91] as
illustrating the first stages in the development of the vertebrata.
I might have given many other examples, but the above are probably
sufficient, and will show that the processes which constitute the
life-history of the lowest organized beings very closely resemble the
first stages in the development of more advanced groups; that as Allen
Thomson has truly observed,[92] "the occurrence of segmentation and the
regularity of its phenomena are so constant that we may regard it as one
of the best established series of facts in organic nature."
It is true that normal yolk-segmentation is not universal in the animal
kingdom; that there are great groups in which the yolk does not divide
in this manner,--perhaps owing to some difference in its relation to the
germinal vesicle, or perhaps because one of the suppressed stages in
embryological development, many examples might be given, not only in
zoology, but, as I may state on the authority of Dr. Hooker, in botany
also. But, however, this may be, it is surely not uninteresting, nor
without significance, to find that changes which constitute the
life-history of the lowest creatures for the initial stages even of the
highest.
Returning, in conclusion, to the immediate subject of this work, I have
pointed out that many beetles and other insects are derived from larvæ
closely resembling _Campodea_.
Since, then, individual insects are certainly in many cases developed
from larvæ closely resembling the genus _Campodea_, why should it be
regarded as incredible that insects as a group have gone through similar
stages? That the ancestors of beetles under the influence of varying
external conditions, and in the lapse of geological ages, should have
undergone changes which the individual beetle passes through under our
own eyes and in the space of a few days, is surely no wild or
extravagant hypothesis. Again, other insects come from vermiform larvæ
much resembling the genus _Lindia_, and it has been also repeatedly shown
that in many particulars the embryo of the more specialized forms
resembles the full-grown representatives of lower types. I conclude,
therefore, that the Insecta generally are descended from ancestors
resembling the existing genus _Campodea_, and that these again have arisen
from others belonging to a type represented more or less closely by the
existing genus _Lindia_.
Of course it may be argued that these facts have not really the
significance which they seem to me to possess. It may be said that when
Divine power created insects, they were created with these remarkable
developmental processes. By such arguments the conclusions of geologists
were long disputed. When God made the rocks, it was tersely said, He
made the fossils in them. No one, I suppose, would now be found to
maintain such a theory; and I believe the time will come when it will be
generally admitted that the structure of the embryo, and its
developmental changes, indicate as truly the course of organic
development in ancient times as the contents of rocks and their sequence
teach us the past history of the earth itself.
FOOTNOTES:
[1] Darwin's "Researches into the Geology and Natural History of the
Countries visited by H.M.S. _Beagle_," p. 326.
[2] Introduction to Entomology, vi. p. 50.
[3] Manual of Entomology, p. 30.
[4] Linnean Journal, vol. xi.
[5] Introduction to the Modern Classification of Insects, p. 17.
[6] Linnean Transactions, 1863--"On the Development of _Chloëon_."
[7] The figures on the first four plates are principally borrowed from
Mr. Westwood's excellent "Introduction to the Modern Classification of
Insects."
[8] "Sur la Domestication des _Clavigers_ par les Fourmis." Bull. de la
Soc. d'Anthropologie de Paris, 1868, p. 315.
[9] Westwood's Introduction, vol. i. p. 36.
[10] Westwood's Introduction, vol. ii. p. 52.
[11] Die Fortpflanzung und Entwickelung der Pupiparen. Von Dr. R.
Leuckart. Halle. 1848.
[12] Ann. des Sci. Nat., sér. 4, tome vii. See also _Natural History
Review_, April 1862.
[13] Ann. and Mag. of Nat. Hist. 1852.
[14] Zeits. für Wiss. Zool. 1869.
[15] Transactions of the Linnean Society, 1863.
[16] Lectures on the Anatomy, &c. of the Invertebrate Animals.
[17] Untersuchungen über die Entwickelung und den Bau der Gliederthiere,
1854.
[18] Linnean Transactions, vol. xxii. 1858.
[19] "Embryological Studies on Hexapodous Insects." Peabody Academy of
Science. Third Memoir.
[20] Mém. de l'Acad. Imp. des Sci. de St. Pétersbourg. 1869.
[21] Observationes de Prima Insectorum Genesi, p. 14.
[22] Mém. de l'Acad. Imp. des Sci. de St. Pétersbourg. tome xvi. 1871,
p. 35.
[23] Recherches sur l'Evolution des Araignées.
[24] Philosophical Transactions, 1841.
[25] Monog. of the Gymnoblastic or Tubularian Hydroids. See also Hincks,
British Hydroid Zoophytes. Pl. x.
[26] Loc. cit. p. 315.
[27] Philosophical Transactions, 1859, p. 589.
[28] "Facts for Darwin," Eng. Trans. p. 127.
[29] Rolleston, "Forms of Animal Life," p. 146.
[30] A. Agassiz, "Embryology of the Starfish," p. 25; "Embryology of
Echinoderms." Mem. of Am. Ac. of Arts and Sciences N.S. vol. ix. p. 9.
[31] Ueber die Gattungen der Seeigellarven. Siebente Abhandlung. Kön.
Akad. d. Wiss. zu Berlin. Von Joh. Müller, 1855, Pl. iii. fig. 3.
[32] Huxley, Introduction to the Classification of Animals, p. 45.
[33] Philosophical Transactions, 1865 and 1866.
[34] Loc. cit. Zweit. Abh. Pl. i., figs. 8 and 9.
[35] Thomson, on the Embryology of the Echinodermata, _Natural History
Review_, 1863, p. 415. See also Agassiz, "Embryology of the Starfish,"
p. 62.
[36] A. Agassiz, Embryology of Echinoderms, p. 18.
[37] Hincks. British Hydroid Zoophytes, pp. 120-147.
[38] Zeits. für Wiss. Zool. 1864, p. 228.
[39] Introduction to Entomology, 6th ed. vol. i. p. 61.
[40] Métamorphoses de l'Homme et des Animaux, p. 133. See also
Carpenter, Principles of Physiology. 1851, p. 389.
[41] Darwin, Origin of Species, 4th ed. p. 532.
[42] Principles of Biology, vi. p. 349.
[43] For differences in larva consequent on variation in the external
condition, see _ante_, p. 61.
[44] See Hincks. British Hydroid Zoophytes, P. lxii. Agassiz, Sea-side
Studies, p. 43.
[45] See Newport, Phil. Trans., 1832.
[46] Linnean Transactions, 1862.
[47] Origin of Species, 4th ed., pp. 14 and 97.
[48] On the Alternation of Generations. By J. J. Steenstrup. Trans. by
C. Busk, Esq. Ray Society. 1842.
[49] Zeit. für Wiss. Zool. 1863.
[50] Mém. de l'Acad. Imp. de St. Pétersbourg. vol. xv. 1870.
[51] Of course all animals in which the sexes are distinct are in one
sense dimorphic.
[52] "There is no such thing as a true case of 'alternation of
generations in the animal kingdom;' there is only an alternation of true
generation with the totally distinct process of gemmation or
fission."--HUXLEY _on Animal Individuality_, Ann. and Mag. of Nat. Hist.
June 1852.
[53] Prince Hohenstiel Schwangau, p. 68.
[54] Journal of the Royal Institution. April 1873.
[55] "Embryology of Echinoderms," l. c. p. 15.
[56] Mr. and Mrs. Agassiz: "Sea-side Studies," p. 139.
[57] l. c. p. 138.
[58] Wien. Zool. Bot. Gesells, 1869.
[59] Linnean Transactions, 1863.
[60] Linnean Transactions, 1866, vol. xxv.
[61] Linnean Transactions, vol. xxiv. p. 65.
[62] Siebold und Kolliker's Zeitschr. f. Wiss. Zool., 1864.
[63] Linnean Journal, vol. xi.
[64] Facts for Darwin, p. 120.
[65] A still nearer approach is afforded by the genus _Peripatus_, which
since the above was written has been carefully described, especially by
Moseley and Hutton. There are several species, scattered over the
southern hemisphere. In general appearance they look like a link between
a caterpillar and a centipede. They have a pair of antennæ, two pairs of
jaws, and (according to the species) from fourteen to thirty-three pairs
of legs. They breathe by means of tracheæ, which open diffusely all over
the body.
[66] Unters. üb. die Entwick, und den Bau der Gliederthiere, p. 73.
[67] Linnean Transactions, v. xxii.
[68] Facts for Darwin, trans. by Dallas, p. 118. See also Darwin,
"Origin of Species," p. 530. 4th ed.
[69] Mem. Peabody Academy of Science, v. I. No, 3.
[70] Wien. Zool. Bott. Gesells. 1869, p. 310.
[71] See also the descriptions given by Dujardin (Ann. des Sci. Nat.
1851, v. xv.) and Claparède (Anat. und Entwickl. der Wirbel osen Thiere)
of the interesting genus _Echinoderes_, which these two eminent
naturalists unite in regarding as intermediate between the Annelides and
the Crustacea.
[72] "On a New Rotifer." _Monthly Microscopical Journal_, Sept. 1871.
[73] Generelle Morphologie, vol. ii. p. 79.
[74] Monographie der Moneren, p. 43.
[75] Gegenbaur. Grund. d. Vergleich. Anat. p. 210. See also Dr. M. S.
Schultze, Beiträge zur Naturg. der. Turbellarien. 1851. Pl. vi. fig. 1.
[76] Monographieder Moneren, p. 10.
[77] See Kauffmann, Ueber die Entwickelung and systematische Stellung
der Tardigraden. Zeits. f. Wiss. Zool. 1851, p. 220.
[78] It is true that among the Insecta generally the first stages of
development differ in appearance considerably from those above
described; those of _Platygaster_, as figured by Ganin (ante Figs. 17-22),
being very exceptional.
[79] Monograph of the Gymnoblastic or Tubularian Hydroids, by G. J.
Allman, Ray Soc. 1871, p. 86.
[80] Mém. sur les Vers Intestinaux, 1858.
[81] Natural History Review, 1861, p. 44.
[82] Ann. des Sci. Nat. 1847, p. 90.
[83] Fauna littoralis Norvegiæ, pl. viii.
[84] Trans. of the Microsc. Soc. of London, 1851.
[85] Quarterly Journal of Microsc. Science, 1853.
[86] Wiegmann's Archiv., 1840, p. 196.
[87] Ueber die Erzeugung von Schnecken in Holothurier. Berlin, Bericht,
1851. Ann. Nat. Hist. 1852, v. ix. Müller's Archiv., 1852.
[88] Natürliche Schöpfungsgeschichte, pl. x.
[89] Ann. des Sci. Nat. 1853, p. 89.
[90] Ann. des Sci. Nat. 1857, pl. vi.
[91] Cyclopædia of Anatomy and Physiology. Art. Ovum, p. 4.
[92] Thomson, loc. cit. Article, Ovum, p. 139.
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