PAST AND PRESENT; THE MEANING OF THE STORY
In the previous chapter we recognised how the seasonal changes in
various species of butterflies as observable in two or three
generations, indicate changes in the history of the race as it might be
traced through innumerable generations. The endless variety in the form
and habits of insect-larvae and their adaptations to various modes of
life, which have been briefly sketched in this little book, suggest
vaster changes in the class of insects, as a whole, through the long
periods of geological time. Every student of life, influenced by the
teaching of Charles Darwin (1859) and his successors, now regards all
groups of animals from the evolutionary standpoint, and believes that
comparisons of facts of structure and life-history of orders and classes
evidently akin to each other, furnish at least some indications of the
course of development in the greater systematic divisions, even as the
facts of seasonal dimorphism, mentioned in the last chapter, give hints
as to the course of development in those restricted groups that we call
species or varieties. A brief discussion of the main outlines of the
life-story of insects in the wide, evolutionary sense may thus fitly
conclude this book.
In the first place we turn to the 'records' of those rocks, in whose
stratified layers[12] are entombed remains, often fragmentary and
obscure, of the insects of past ages of the earth's history. Compared
with the thousands of extinct types of hard-shelled marine animals, such
as the Mollusca, fossil insects are few, as could only be expected,
seeing that insects are terrestrial and aerial creatures with slight
chance of preservation in sediments formed under water. Yet a number of
insect remains are now known to naturalists, who are, in this
connection, more particularly indebted to the researches of S.H. Scudder
(1885), C. Brongniart (1894), and A. Handlirsch (1906).
[12] See Table of Geological Systems, p. 123.
We are now considering insects from the standpoint of their
life-histories, and the individual life-story of an insect of which we
possess but a few fragments of wings or body, entombed in a rock formed
possibly before the period of the Coal Measures, can only be a matter of
inference. Still it may safely be inferred that when the structure of
these remains clearly indicates affinity to some existing order or
family, the life-history of the extinct creature must have resembled, on
the whole, that of its nearest living allies. And all the fossil
insects known can be either referred to existing orders, or shown to
indicate definite relationship to some existing group.
Passing over some doubtful remains of Silurian age, we find in rocks
usually regarded as Devonian[13] the most ancient fossils that can be
certainly referred to the insects, while from beds of the succeeding
Carboniferous period, a number of insect remains have been disinterred.
These Palaeozoic insects were frequently of large size, and they show
distinct affinities with our recent may-flies, dragon-flies,
stone-flies, and cockroaches. In the Permian period, the latest of the
divisions of the Palaeozoic, lived Eugereon, an insect with hemipteroid
jaws and orthopteroid wings. All these insects must have been
exopterygote in their life-history, if we may trust the indications of
affinity furnished by their structure. In the Mesozoic period, however,
insects with complete transformations must have been fairly abundant.
Rocks of Triassic age have yielded beetles and lacewing-flies, while
from among Jurassic fossils specimens have been described as
representing most of our existing orders, including Lepidoptera,
Hymenoptera and Diptera. In Cainozoic rocks fossil insects of nearly six
thousand species have been found, which are easily referable to
existing families and often to existing genera. We may conclude then,
imperfect though our knowledge of extinct insects is, that some of the
most complex of insect life-stories were being worked out before the
dawn of the Cainozoic era. Some instructive hints as to differences in
the rate of change among different insect groups may be drawn from the
study of parasites. For example, V.L. Kellogg (1913) points out that an
identical species of the Mallophaga (Bird-lice) infests an Australian
Cassowary and two of the South American Rheas; while two species of the
same genus (Lipeurus) are common to the African Ostrich and a third kind
of South American Rhea. These parasites must have been inherited
unchanged by the various members of these three families of flightless
birds from their common ancestors, that is from early Cainozoic times at
latest. On the other hand, the various kinds of such highly specialised
parasites as the warble-flies of the oxen and deer, must have become
differentiated during those later stages of the Cainozoic period which
witnessed the evolution of their respective mammalian hosts.
[13] The 'Little River' beds of St John, New Brunswick, Canada, by some
modern geologists however considered as Carboniferous.
The foregoing brief outline of our knowledge of the geological
succession of insects shows that the exopterygote preceded, in time, the
endopterygote type of life-history. We have already seen that those
insects undergoing little change in the life-cycle, and with visible,
external wing-rudiments, are on the whole less specialised in structure
than those which pass through a complete transformation. These two
considerations, taken together, suggest strongly that in the evolution
of the insect class, the simpler life-history preceded the more complex.
Such a conclusion seems reasonable and what might have been expected,
but we are confronted with the difficulty that if the most highly
organised insects pass through the most profound transformations, then
insects present a remarkable and puzzling exception to the general rules
of development among animals, as has already been pointed out in the
first chapter of this volume (p. 7). A few students of insect
transformation have indeed supposed that the crawling caterpillar or
maggot must be regarded as a larval stage which recalls the worm-like
nature of the supposed far-off ancestors of insects generally. Even in
Poulton's classical memoir (1891, p. 190), this view finds some support,
and it may be hard to give up the seductive idea that the worm-like
insect-larva has some phylogenetic meaning. But the weight of evidence,
when we take a comprehensive survey of the life-story of insects, must
be pronounced to be strongly in favour of the view put forward by Brauer
(1869), and since supported by the great majority of naturalists who
have discussed the subject, that the caterpillar or the maggot is itself
a specialised product of the evolutionary process, adapted to its own
particular mode of larval life.
The explanation of insect transformation is, in brief, to be found in an
increasing amount of divergence between larva and imago. The most
profound metamorphosis is but a special type of growth, accompanied by
successive castings and renewings of the chitinous cuticle, which
envelopes all arthropods. In the simplest type of insect life-story,
there is no marked difference in form between the newly-hatched young
and the adult, and in such cases we find that the young insect lives in
the same way as the adult, has the same surroundings, eats the same
food. This is the rule (see Chapters II and III) with the Apterygota,
the Orthoptera, and most of the Hemiptera. In the last-named order,
however, we find in certain families marked divergence between larva and
imago, for example in the cicads, whose larvae live underground, while
in the coccids, whose males are highly specialised and females degraded,
there succeeds to the larva--very like the young stage in allied
families--a resting instar, which in the case of the male, suggests
comparison with the pupa of a moth or beetle.
Turning to the stone-flies, dragon-flies and may-flies, whose
life-stories have been sketched in Chapter IV, we find that the early
stages are passed in water, whence before the final moult, the insects
emerge to the upper air. Except for the possession of tufted gills,
adapting them to an aquatic life, the stone-fly nymphs differ but
slightly from the adults; the grubs of the dragon-flies and may-flies,
however, are markedly different from their parents. In connection with
these comparisons, it is to be noted that the dragon-flies and may-flies
are more highly specialised insects than stone-flies, divergent
specialisation of the adult and larva is therefore well illustrated in
these groups, which nevertheless have, like the Hemiptera and
Orthoptera, visible external wing-rudiments.
From the vast array of insects that show internal wing-growth and a true
pupal stage, a few larval types were chosen for description in Chapter
VI, and a review of these suggests again the thought of increasing
divergence between larva and imago. Reference has been made previously
to the many instances in which the former has become pre-eminently the
feeding, and the latter the breeding stage in the life-cycle. It seems
impossible to avoid the conclusion that the active, armoured
campodeiform grub differing less from its parent than an eruciform larva
differs from its parent, is as a larval type more primitive than the
caterpillar or maggot. A. Lameere has indeed, while admitting the
adaptive character of insect larvae generally, argued (1899) with much
ingenuity that the eruciform or vermiform type must have been primitive
among the Endopterygota, believing that the original environment of the
larvae of the ancestral stock of all these insects must have been the
interior of plant tissues. He is thus forced to the necessity of
suggesting that the campodeiform larvae of ground-beetles or lacewings
must be regarded as due to secondarily acquired adaptations; 'they
resemble Thysanura and the larvae of Heterometabola only as whales
resemble fishes.' There are two considerations which render these
theories untenable. The Neuroptera and Coleoptera among which
campodeiform larvae are common, are less specialised than Lepidoptera,
Hymenoptera, and Diptera, in which they are unknown. And among the
Coleoptera which as we have seen (pp. 50 _f._) display a most
interesting variety of larval structure, the legless, eruciform larva
characterises families in which the imago shows the greatest
specialisation, while in the same life-story, as in the case of the
oil-beetles (pp. 56-7), the newly-hatched grub may be campodeiform,
changing to the eruciform type as soon as it finds itself within reach
of its host's rich store of food.
A certain amount of difficulty may be felt with regard to the theory of
divergent evolution between imago and larva, in the case of those
insects with complete transformation whose grubs and adults live in much
the same conditions. By turning over stones the naturalist may find
ground-beetles in company with the larvae of their own species. On the
leaves of a willow tree he may observe leaf-beetles (Phyllodecta and
Galerucella) together with their grubs, all greedily eating the foliage;
or lady-bird beetles (Coccinella) and their larvae hunting and devouring
the 'greenfly.' All of these insects are, however, Coleoptera, and the
adult insects of this order are much more disposed to walk and crawl and
less disposed to fly than other endopterygote insects. Their heavily
armoured bodies and their firm shield-like forewings render them less
aerial than other insects; in many genera the power of flight has been
altogether lost. It is not surprising, therefore, that many beetles,
even when adult, should live as their larvae do; since the acquirement
of complete metamorphosis they have become modified towards the larval
condition, and an extreme case of such modification is afforded by the
wingless grub-like female Glow-worm (Lampyris).
With most insects, however, the larva must be regarded as the more
specially modified, even if degraded, stage. Miall (1895) has pointed
out that the insect grub is not a precociously hatched embryo, like the
larvae of multitudes of marine animals, but that it exhibits in a
modified form the essential characters of the adult. Comparison for
example can be readily made between the parts of the caterpillar and the
butterfly, whose story was sketched in the first chapter of this book,
widely different though caterpillar and butterfly may appear at a
superficial glance. And the survey of variety in form, food, and habit
of insect larvae given in Chapter VI enforces surely the conclusion that
the larva is eminently plastic, adaptable, capable of changing so as to
suit the most diverse surroundings. In a most suggestive recent
discussion on the transformation of insects P. Deegener (1909) has
claimed that the larva must be regarded as the more modified stage,
because while all the adult's structures are represented in the larva,
even if only as imaginal buds, there are commonly present in the larva
special adaptive organs not found in the imago, for example the pro-legs
of caterpillars or the skin-gills of midge-grubs. The correspondence of
parts in butterfly and caterpillar just referred to, may still be
traced, though less easily, in bluebottle and maggot. The latter is an
extreme example of degenerative evolution, and its contrast with the
elaborately organised two-winged fly marks the greatest divergence
observable between the larva and imago. With this divergence the resting
pupal stage, during which more or less dissolution and reconstruction of
organs goes on, becomes a necessity, and it has already been pointed out
how the amount of this reconstruction is greatest where the divergence
between the larval and perfect stages is most marked. Whatever
differences of opinion may prevail on points of detail, the general
explanation of insect metamorphosis as the result of divergent evolution
in the two active stages of the life-story must assuredly be accepted.
No other explanation accords with the increasing degree of divergence to
be observed as we pass from the lower to the higher insect orders.
The successive incidents of the life-story of most insects are largely
connected with the acquisition of wings. Wings, and the power of flight
wherewith they endow their possessors, are evidently beneficial to the
race in giving power of extending the range during the breeding period
and thus ensuring a wide distribution of the eggs. In no case are wings
fully developed until the closing stage of the insect's life, they are
always acquired after hatching or birth. We have already noticed (p. 40)
how Sharp (1899) has laid stress on the essential difference between the
exopterygote and endopterygote insects, the wing-rudiments of the former
growing outwards throughout life while those of the latter remain hidden
until the pupal instar. Sharp considers that there is some difficulty in
bridging, in thought, the gap between these two methods of wing-growth,
and has put forward an ingenious suggestion to meet it (1902). Reference
has already been made to insects of various orders in which one sex is
wingless, the Vapourer Moth (p. 96) for example, or all the individuals
of both sexes are wingless, as the aberrant cockroaches mentioned in
Chapter II (p. 15), or certain generations of virgin females are
wingless, for example aphids (pp. 18-19) and gall-flies (pp. 94-5).
Insects may thus become secondarily wingless, that is to say be
manifestly the offspring of winged parents, and such wingless forms may
on the other hand give rise to offspring or descendants with
well-developed wings. Frequently, as in the case of the aphids, many
wingless generations intervene between two winged generations. A
striking illustration of this fact is afforded by an aquatic bug, _Velia
currens_, commonly to be seen skating over the surface of running water.
The adults of Velia are nearly always wingless, but now and then the
naturalist meets with a specimen provided with functional wings, the
possession of which enables the insect to make its way to a fresh
stream. Moreover there are whole orders of parasitic insects, such as
the lice and fleas, which, showing clear affinity to orders of winged
insects, are believed to be secondarily wingless. These orders are
designated by Sharp 'Anapterygota.' And from the analogy of the periodic
loss and recovery of wings in various generations of the same species,
he has concluded that the gap between the exopterygote and the
endopterygote method of development may have been bridged by an
anapterygote condition; that the ancestors of those insects with
complete transformations were the wingless descendants of primitive
insects which grew their wings from visible external rudiments, and
that in later times re-acquiring wings, they developed these organs in a
new way, from inwardly directed rudiments or imaginal buds.
This theory of Sharp's is original, daring, and ingenious, but the loss
and re-acquisition of wings which it presupposes is difficult to imagine
in large groups during a prolonged evolutionary history, while the
sudden appearance of a totally new mode of wing-growth in the offspring
of wingless insects would be an extreme example of discontinuity in
development.
On the whole the most probable suggestion which can be made as to the
origin of 'complete' transformation in insects is that the instar in
which wings were first visible externally became later and later in the
course of the evolution of the more highly organised groups. In this way
a gradual transition from the exopterygote to the endopterygote type of
life-story is at least conceivable. It will be remembered that a may-fly
(p. 33) undergoes a moult after acquiring functional wings, emerging
into the air as a 'sub-imago.' In not a few endopterygote insects, the
pupa shows more or less activity, swimming through water intermittently
(gnats) or just before the imago has to emerge (caddis-flies); working
its way out of the ground (crane-flies) or coming half-way out of its
cocoon (many moths). The pupa of the higher insects almost certainly
corresponds with the may-fly's sub-imago, and the facts just recalled as
to remnants of pupal activity suggest that in the ancestors of
endopterygote insects what is now the pupal instar was represented by an
active nymphal or sub-imaginal stage, possibly indeed by more than one
stage, as Packard and other writers have stated that pupae of bees and
wasps undergo two or three moults before the final exposure of the
imago. Such an early pupal instar has been defined as a 'pro-nymph' or a
'semi-pupa.' Examples have been given of the exceptional passive
condition of the penultimate instar in Exopterygota. The instars
preceding this presumably had originally outward wing-rudiments in all
insect life-histories, and the endopterygote condition was attained by
the postponement of the outward appearance of these to successively
later stages. The leg and wing rudiments of the male coccid (pp. 20-1)
beneath the cuticle of the second instar are strictly comparable to
imaginal buds, and these are present in one instar of what is generally
regarded as an exopterygote life-history. The first instar in all
insects has no visible wing-rudiments, but when they grow outwardly from
the body, they necessarily become covered with cuticle, so that they
must be visible after the first moult. There is no supreme difficulty in
supposing that the important change was for these early rudiments to
become sunk into the body, so that the cuticle of the second, and,
later, of the third and succeeding instars, showed no outward sign of
their presence. This suggestion is confirmed by Heymons' (1896, 1907)
observation of the occasional appearance of outward wing-rudiments on
the thoracic segments of a mealworm, the larva of the beetle _Tenebrio
molitor_, and by F. Silvestri's discovery (1905) of a 'pro-nymph' stage
with short external wing-rudiments between the second larval and the
pupal instars of the small ground-beetle _Lebia scapularis_. Whatever
may be the exact explanation of these abnormalities, they show that in
the life-story of the higher insects outward wing-rudiments may even yet
appear before the pupal stage, confirming our belief that such
appearance is an ancestral character. The inward growth of these
wing-rudiments may well have been correlated with a difference in form
between the newly-hatched insect and its parent. As this difference
persisted until a constantly later stage, and the pre-imaginal instar
became necessarily a stage for reconstruction, the present condition of
complete metamorphosis in the more highly organised orders was finally
attained.
To explain satisfactorily these complex life-stories is however
admittedly a difficult task. The acquisition of wings is, as we have
seen, a dominating feature in them all, but if we try to go yet a step
farther back and speculate on the origin of wings in the most primitive
exopterygote insects, the task becomes still more difficult. Many years
ago Gegenbaur (1878) was struck by the correspondence of insect wings to
the tracheal gills of may-fly larvae, which are carried on the abdominal
segments somewhat as wings are on the thoracic segments. But Börner has
recently (1909) brought forward evidence that these abdominal gills
really correspond serially with legs. Moreover Gegenbaur's theory
suggests that the ancestral insects were aquatic, whereas the presence
of tubes for breathing atmospheric air in well-nigh all members of the
class, and the fact that aquatic adaptations, respiratory and otherwise,
in insect-larvae are secondary force the student to regard the ancestral
insects as terrestrial. It is indeed highly probable that insects had a
common origin with aquatic Crustacea, but all the evidence points to the
ancestors of insects having become breathers of atmospheric air before
they acquired wings. How the wings arose, what function their precursors
performed before they became capable of supporting flight, we can hardly
even guess.
Our study of the life-story of insects, therefore, while it has taught
us something of what is going on around us to-day, and has given us
hints of the course of a few threads of that long life-story which runs
through the ages, brings us face to face with the most instructive, if
humbling fact that 'there are many more things of which we are
ignorant.' The passage from creeping to flight, as the caterpillar
becomes transformed into the butterfly, was a mystery to those who first
observed it, and many of its aspects remain mysterious still. Perhaps
the most striking result of the study of insect transformation is the
appreciation of the divergent specialisation of larva and imago, and it
is a suggestive thought that of the two the larva has in many cases
diverged the more from the typical condition. The caterpillar crawling
over the leaf, or the fly-grub swimming through the water, may thus be
regarded as a creature preparing for a change to the true conditions of
its life. It is a strange irony that the preparation is often far longer
than the brief hours of achievement. But the light which research has
thrown on the nature of these wonderful life-stories, the demonstration
of the unseen presence and growth within the insect, during its time of
preparation among strange surroundings, of the organs required for
service in the coming life amid its native air, confirm surely the
intuition of the old-time students, who saw in these changes, so
familiar and yet so wonderful, a parable and a prophecy of the higher
nature of man.
OUTLINE CLASSIFICATION OF INSECTS
Class INSECTA or HEXAPODA.
Sub-class A, APTERYGOTA.
Order 1. _Thysanura_ (Bristle-tails).
2. _Collembola_ (Spring-tails).
Sub-class B, EXOPTERYGOTA.
Order 1. _Dermaptera_ (Earwigs).
2. _Orthoptera_ (Cockroaches, Grasshoppers, Crickets).
3. _Plecoptera_ (Stone-flies).
4. _Isoptera_ (Termites or 'White Ants').
5. _Corrodentia_
(_a_) _Copeognatha_ (Book-lice).
(_b_) _Mallophaga_ (Biting-lice).
6. _Ephemeroptera_ (May-flies).
7. _Odonata_ (Dragon-flies).
8. _Thysanoptera_ (Thrips).
9. _Hemiptera_
(_a_) _Heteroptera_ (Bugs, Pond-skaters)
(_b_) _Homoptera_ (Cicads, 'Greenfly,' Scales).
10. _Anoplura_ (Lice).
Sub-class C, ENDOPTERYGOTA.
Order 1. _Neuroptera_ (Alder-flies, Ant-lions, Lacewings).
2. _Coleoptera_ (Beetles).
3. _Mecaptera_ (Scorpion-flies).
4. _Trichoptera_ (Caddis-flies).
5. _Lepidoptera_ (Moths and Butterflies).
6. _Diptera_ (Two-winged flies)
(_a_) _Orthorrhapha_ (Crane-flies, Midges, Gnats)
(_b_) _Cyclorrhapha_ (Hover-flies, House-flies, Bot-flies, &c).
7. _Siphonaptera_ (Fleas).
8. _Hymenoptera_
(_a_) _Symphyta_ (Saw-flies)
(_b_) _Apocrita_ (Gall-flies, Ichneumon-flies, Wasps, Bees, Ants).
TABLE OF GEOLOGICAL SYSTEMS
These names, given by geologists to the various divisions of rocks, as
indicated by the fossils entombed in them, are arranged in 'descending'
order, the more recent formations above, the more ancient below, as
newer deposits necessarily lie over older beds.
CALNOZOIC OR TERTIARY GROUP.
Pleistocene.
Pliocene.
Miocene.
Eocene.
MESOZOIC OR SECONDARY GROUP.
Cretaceous.
Jurassic.
Triassic.
PALAEOZOIC OR PRIMARY GROUP.
Permian.
Carboniferous.
Devonian.
Silurian.
Cambrian.
BIBLIOGRAPHY
The following list of some books and papers, referred to in this little
volume or of especial service to the author in its preparation, is
needless to say very far from exhaustive. To save space, titles are
often abbreviated. Most of the works in the general list (A) contain
extensive lists of literature on insects and their transformations,
these should be consulted by the serious student.
A. GENERAL WORKS.
1909. C. Börner. Die Verwandlungen der Insekten. _Sitzb. d. Gesellsch.
naturforsch. Freunde, Berlin._
1869. F. Brauer. Betrachtung über die Verwandlung der Insekten.
_Verhandl. der K.K. zool.-bot. Gesellschaft in Wien._ XIX.
1899. G.H. Carpenter. Insects, their Structure and Life. London.
1859. C. Darwin. The Origin of Species. London.
1909. P. Deegener. Die Metamorphose der Insekten. Leipzig.
1906. J.W. Folsom. Entomology. London.
1878. C. Gegenbaur. Grundriss der Vergleichende Anatomie. Leipzig.
1906. A. Handlirsch. Die fossilen Insekten. Leipzig.
1904. L.F. Henneguy. Les Insectes. Paris.
1907. R. Heymons. Die verschiedenen Formen der Insectenmetamorphose.
_Ergebnisse der Zoologie._ I.
1899. A. Lameere. La raison d'être des Metamorphoses chez les Insectes.
_Ann. Soc. Entom. Bruxelles._ XLIII.
1874. J. Lubbock. The Origin and Metamorphoses of Insects. London.
1895. L.C. Miall. (_a_) The Transformations of Insects. _Nature._ LIII.
1895. ---- (_b_) The Natural History of Aquatic Insects. London.
1908. ---- Injurious and Useful Insects. 2nd edition. London.
1839. G. Newport. Insects. _Todd Cyclopaedia._ II. London.
1898. A.S. Packard. Text book of Entomology. New York.
1734-42. R.A.F. de Réaumur. Mémoires pour servir à l'Histoire naturelle
et à l'anatomie des Insectes. Paris.
1895-8. D. Sharp. The Cambridge Natural History, V, VI. London.
1899. ---- Some points in the Classification of Insects. IV. _Internat.
Zoolog. Congress._
1902. ---- Insects in _Encycl. Brit._ 10th Edition, XXIX. London.
1910. ---- and G.H. Carpenter. Hexapoda in _Encycl. Brit._ 11th
Edition. Cambridge.
1737. J. Swammerdam. Biblia Naturae. Leyden (incorporates works on
Insects published during the author's lifetime 1669-75).
1909. F.V. Theobald. Insect Pests of Fruit. Wye.
B. SPECIAL WORKS.
1881. H. Adler. Ueber den Generationswechsel den Eichen-Gallwespen.
_Zeitsch. f. wissensch. Zoologie._ XXXV.
1896. ---- and C.R. Straton. Alternating Generations. Oxford.
1902. J. Anglas. Nouvelles Observations sur les Métamorphoses Internes.
_Arch. d'Anat. Microscop._ IV.
1911. E.E. Austen. Handbook of the Tsetse-Flies. London (Brit. Museum).
1909. F. Balfour-Browne. Life-History of Agrionid Dragonfly. _Proc.
Zool. Soc. Lond._
1893, &c. C.G. Barrett. Lepidoptera of the British Islands. London.
1890. H. Beaurégard. Les Insectes Vésicants. Paris.
1909. C. Börner. Die Tracheenkiemen der Ephemeriden. _Zoolog. Anz._
xxxiii.
1863. F. Brauer. Monographie der Oestriden. Wien.
1894. C. Brongniart. Récherches pour servir à l'histoire des Insectes
fossiles des Temps Primaires. St Etienne.
1893. T.A. Chapman. Structure of Pupae of Heterocerous Lepidoptera.
_Trans. Entom. Soc. Lond._
1891. H. Dewitz. Das geschlossene Tracheensystem bei Insektenlarven.
_Zoolog. Anz._ xiii.
1857-8. J.H. Fabre. L'Hypermétamorphose et les Moeurs des Meloides.
_Ann. Sci. Nat._ (_Zool._), (4). VII. IX.
1869. M. Ganin. Die Entwicklungsgeschichte bei den Insekten. _Zeitsch.
f. wissensch. Zoolog._ xix.
1894. J. Gonin. La Métamorphose des Lepidoptères. _Bull. Soc. Vaud.
Sci. Nat._ xxx.
1870. O. Grimm. Die ungeschechtliche Fortpflanzung einer Chironomus.
_Mem. Acad. Impér. St Pétersbourg_ (7). xv.
1890. W. Hatchett-Jackson. Morphology of the Lepidoptera. _Trans. Linn.
Soc. (Zool.) Lond._ (2). v.
1896. R. Heymons. Flügelbildung bei der Larve von Tenebrio molitor.
_Sitzb. d, Gesellsch. Naturforsch. Freunde, Berlin._
1906. ---- Ueber die ersten Jugendformen von Machilis alternata. _Ib._
1908. W. Kahle. Die Paedogenesis der Cecidomyiden. _Zoologica._ IV.
1913. V.L. Kellogg. Distribution and Species-forming of Ectoparasites.
_Amer. Naturalist._ XLVII.
1887. A. Kowalevsky. Die nachembryonale Entwicklung der Musciden.
_Zeitsch. f. wissensch. Zool._ XLV.
1904. O.H. Latter. Natural History of Common Animals (chaps. III, IV,
V). Cambridge.
1890-95. B.T. Lowne. The Blowfly, 2 vols. London.
1863. J. Lubbock. Development of Chloeon. _Trans. Linn. Soc. Lond._
XXIII.
1762. P. Lyonet. Traité anatomique de la Chenille. Haag.
1669. M. Malpighi. De Bombyce. London.
1898. C.L. Marlatt. The periodical Cicada. _Entom. Bull._ 14, _U.S.
Dept. Agric._
1898. G.A.K. Marshall. Seasonal Dimorphism in Butterflies. _Ann. Mag.
Nat. Hist._ (7). II.
1900. L.C. Miall and A.B. Hammond. The Harlequin Fly. Oxford.
1901-3. R. Newstead. Coccidae of the British Isles. London.
1877. J.A. Palmén. Zur Morphologie des Tracheensystems. Leipzig.
1891. E.B. Poulton. External Morphology of the Lepidopterous Pupa.
_Trans. Linn. Soc. Zool._ (2). V.
1892. ---- Colour-relation between Lepidopterous Larvae &c. and their
surroundings. _Trans. Entom. Soc. Lond._
1880. C.V. Riley. Pupation of Butterflies. _Proc. Amer. Assoc._ XXVIII.
1902. E.D. Sanderson. Report of Entomologist. Delaware. U.S.A.
1885. E.O. Schmidt. Metamorphose und Anatomie des männlichen
Aspidiotus. _Archiv f. Naturgeschichte._ LI.
1885. S.H. Scudder. Insekten in Zittel's Paleontologie. II.
1907. A.J. Siltala. Die postembryonale Entwicklung der
Trichopteren-Larven. _Zoolog. Jahrb. Suppl._ IX.
1905. F. Silvestri. Metamorfosi e Costumi della Lebia scapularis.
_Redia._ II.
1900. J.B. Smith. The Apple Plant-louse. _New Jersey Agric. Exp.
Station Bull._ 143.
1888. J. Van Rees. Die innere Metamorphose von Musca. _Zoolog. Jahrb.
Anat._ III.
1911. K.W. Verhoeff. Ueber Felsenspringer, Machiloidea. _Zoolog. Anz._
XXXVIII.
1865. N. Wagner. Die viviparen Gallmückenlarven. _Zeitsch. f.
wissensch. Zoolog._ XV.
1901. E. Wasmann. Termitoxenia. _Zeitsch. f. wissensch. Zoolog._ LXX.
1864. A. Weismann. Die nachembryonale Entwicklung der Musciden.
_Zeitsch. f. wissensch. Zoolog._ XIV.
1865. ---- Die Metamorphose von Corethra. _Ib._ XVI.
1876. ---- Studien zur Descendenz-Theorie. Leipzig. (English
Translation by R. Meldola, London, 1882.)
INDEX
_Abraxas grossulariata_, 60, 83, 97-8
Adaptation of larvae, 57, 79, 114
Adephaga, 51
Adler, H., 94
Aeschnidae, 27, 29, 31
Agrionidae, 27, 28
_Agrotis segetum_, 98
Air-tubes, 2, 11, 23, 47, 70, 77, 87, 120
Alternation of generations, 17, 94
Ametabola, 11, 35
Anapterygota, 116
Anglas, J., 46
Ant-lions, 57
Ants, 64, 66
Aphidae, 17-20, 116
_Aphis pomi_, 18-19
Aphis-lion, 57
Apterygota, 41, 110
Aquatic insects, 23-34, 76-9, 120
_Araschnia levana_ and var. _prorsa_, 103
_Arctia caia_, 98
Arctiadae, 59
Arthropoda, 9
Austen, E.E., 91
Avebury, Lord, _see_ Lubbock, J.
Balfour-Browne, F., 28
Bark-beetles, 55
Barrett, C.G., 96, 99
Beaurégard, H., 56
Bees, 40, 46, 64, 83
Beetles, 40, 50-7, 80, 107, 112-3, 119
Bell Moths, 62
Bird-lice, 108
Birth, 18, 91
_Blatta orientalis_, 15
Blister-beetles, 56
Blowfly or Bluebottle, 43, 44, 46, 67, 71-3, 93, 114
Börner, C., 32, 120
Bot-flies, 73-4, 89, 91
Brain, 44
Brauer, F., 6, 52, 56, 67, 109
Bristle-tails, 11
Brongniart, C., 106
Butterflies, 1, 83, 95-6, 114
Cabbage-butterflies, 39, 41, 85, 100-1
Cabbage-fly, 73
Caddis-flies, 62-3, 86, 117
Cainozoic insects, 107
Calliphora, 43.
_See also_ Blowfly
Campodeiform larvae, 52, 56, 111
Carabidae, 52
Carboniferous insects, 107
_Carpocapsa pomonella_, 99-100
Carrion-beetles, 50
Caterpillar, 4, 36, 49, 58-62, 95-101, 109, 114
Cecidomyidae, 68-70, 90
Cerambycidae, 55
Cercopods, 12, 15
Chafers, 52
Chapman, T.A., 81, 84
Chironomus, 43, 77, 87, 91
Chloeon, 33
Chrysalis, 82.
_See also_ Pupa
Chrysomelidae, 53.
_See also_ Leaf-beetles
Chrysopa, 57
Cicads, 22, 93, 110
Classification, 122
Clearwing Moths, 62
Click-beetles, 52, 93
Clothes-moths, 62
Coccidae, 20, 110, 118
Coccinella, 113
Cockroaches, 11, 14, 15, 107, 115
Cocoons, 82
Codling Moth, 62, 99
Coleoptera, 50-6, 80, 112, 119
Collembola, 11
Complete transformation, 35, 107, 119.
_See also_ Endopterygota
Corethra, 43
Cossus, 38, 62, 82, 95
Crane-flies, 67, 70, 93, 117
Cremaster, 83
Crustacea, 7, 120
Culex, 43, 77, 86
Curculionidae, 55
Cuticle, 2, 9, 29, 37, 40, 50, 81, 87, 110
Cynipidae, 94.
_See also_ Gall-flies
Daddy-long-legs, 69-70
Darwin, C., 105
Deegener, P., 6, 114
Devonian insects, 107
Dewitz, H., 28
Digestive system, 10, 45-7
_Diplosis pyrivora_, 70
Diptera, 42, 64, 67-79, 81, 86-8, 91, 94, 107
Divergence between larva and imago, 110, 114, 121
Double-brooded Lepidoptera, 95, 100-4
Dragon-flies, 26-31, 107, 110
Drone-flies, 76
Duration of life, 34, 89, 92-3, 95
Dyticus, 51
Ecdysis, 10.
_See also_ Moult
Ectoderm, 9, 11, 47
Eggar Moths, 59, 89
Eggs, 6, 17-18, 26, 34, 65-7, 71, 90, 94-5, 97
Elateridae, 52
Endopterygota, 41, 49, 108, 112, 115-6
Ephemeroptera, 24.
_See also_ May-flies
Epidermis, 9, 40
Eristalis, 76
Eruciform larvae, 56, 58-70, 111
Evolution, 16, 103, 105-21
Exopterygota, 41, 108, 115-6, 118
Exoskeleton, 9
Fabre, J.H., 56
Fat-body, 47
Feeding-period, 27, 32, 36, 89, 111
Feelers, 1, 4, 42, 71
Fleas, 116
Fore-gut, 47
Free pupa, 80
Gall-flies, 64-6, 94, 115
Gall-midges, 68-70, 90
Ganin, M., 66
_Gastrophilus equi_, 73-4
Gegenbaur, C., 120
Geological history, 106-8, 123
Geometridae, 59
Gills, 24, 27, 32, 78, 87, 114, 120
Glossinia, 91
Glow-worm, 50, 113
Gnats, 43, 77, 86
Goat Moth, 38, 62, 82, 95
Gonin, J., 38, 41
Grasshoppers, 11, 14, 15
Grimm, O., 90
Ground-beetles, 52, 112
Growth, 9
Grub, 63-70.
_See also_ Caterpillar, Larva
Hairs, 59, 82, 98
Hammond, A.R., 43, 77, 87
Handlirsch, A., 106
Harvey, William, 7
Hatchett-Jackson, W., 83
Hawk Moths, 60
Heart, 45
Helodes, 50
Hemerobius, 57
Hemimetabola, 35
Hemiptera, 17, 110
Henneguy, L.F., 45, 48
Heymons, R., 6, 11, 119
Hibernation. _See_ Wintering stages
Hind-gut, 47
Hippoboscidae, 91
Histogenesis and Histolysis, 48
Holometabola, 35
House-fly, 67, 71, 73
Hover-flies, 74-6
Hymenoptera, 58, 64, 94, 107
Hypermetamorphosis, 56
_Hypoderma bovis_, 73-5
Hypodermis, 9
Ichneumon-flies, 64, 66, 82
Imaginal buds or discs, 34-48, 114, 117-8
Imago, 24, 34, 114
Instar, 13, 33, 56, 117-9
Jaws of imago and larva, 2, 4, 5, 32, 42, 89
Jurassic insects, 107
Kahle, W., 90
Kellogg, V.L., 108
Kowalevsky, A., 46
Labium, 2, 27
Lacewing-flies, 57, 107
Ladybirds, 113
Lameere, A., 111
Lampyris, 113
Larva, 4, 22, 26-7, 32, 49-79, 110-15
Larval reproduction, 90
Lasiocampidae, 59, 89
Latter, O.H., 28
Leaf-beetles, 53, 83, 92-3, 113
_Lebia scapularis_, 119
Lepidoptera, 1, 36, 38, 49, 58, 81, 95-104, 107
Libellulidae, 27
Lice, 116
Lipeurus, 108
Longhorn Beetles, 55
Looper caterpillars, 59, 61
Lowne, B.T., 42
Lubbock, J., 6, 32
Lymantriidae, 90
Lyonet, P., 38
Machilis, 11
Maggot, 44, 67, 71-6, 109, 114
Magpie Moth, 60, 82, 97-8
Mallophaga, 108
Mandibles, 4, 17, 26, 58, 67, 86
Mangel-fly, 73
Marlatt, C.L., 93
Marshall, G.A.K., 104
Maxillae, 2, 17, 37, 42
May-flies, 31-4, 107, 110, 117, 120
Meloidae, 56
Mesozoic insects, 107
Metabola, 35
Metamorphosis (in general), 6, 109;
(degrees of in insects) 8, 35, 109, 117-19
Miall, L.C., 6, 28, 33, 43, 77, 78, 87, 97, 113
Mosquito. _See_ Culex, Gnats
Moths, 1, 58-62, 84, 95-100, 117
Moult, 10, 32, 36, 41
_Musca domestica_, 71
Muscidae, 44
Muscles, 47
Nervous system, 44-5
Neuroptera, 57, 80, 112
Newport, G., 41, 44
Noctuidae, 60, 98
Nymph, 15, 28, 33
Oak-apples, 94
Obtect pupa, 81
Odonata, 24.
_See also_ Dragon-flies
_Oestrus ovis_, 91
Oil-beetles, 56, 112
_Orgyia antiqua_, 96-7
Orthoptera, 17, 35, 110
Owl Moths, 60, 98
Packard, A.S., 56, 118
Paedogenesis. _See_ Larval reproduction
Painted Lady Butterfly, 96
Palaeozoic insects, 107
Palmén, J.A., 25
Parasitic insects, 73-4, 108, 116
Parental care, 64-6
Parthenogenesis, 18
Partial transformation, 35, 37
Perla, 24
Permian insects, 107
Phagocytes, 48
Phyllodecta, 53, 113
Phyllotreta, 53
_Pieris brassicae_, 39, 41, 85, 100
_Pieris napi_ and var. _bryoniae_, 102-3
Platygaster, 66
Plecoptera, 24.
_See also_ Stone-flies
Pompilidae, 66-7
Poulton, E.B., 61, 82, 109
Precis, 104
Proctotrypidae, 66
Pro-legs, 4, 58-9, 84, 114
Pro-nymph, 118, 119
Protective coloration, 60-1
_Psylliodes chrysocephala_, 54
Ptinidae, 54
Pupa, 4, 37, 40, 79-88, 114, 117
Puparium, 88
Pupipara, 91
_Pyrameis cardui_, 96
Rat-tailed maggot, 76
Réaumur, R.A.F. de, 8, 28, 33, 41
Reproductive larvae, 90;
pupae, 91
Reproductive organs, 45
_Rhabdophaga heterobia_, 70
Riley, C.V., 83
Sanderson, E.D., 17
Sand-midges, 78
Sarcophaga, 91
Saw-flies, 58-9
Scale-insects, 20.
_See also_ Coccidae
Scarabaeidae, 52
Schmidt, E.O., 21
Scolytidae, 55
Scudder, S.H., 106
Seasonal changes, 89-104
Seasonal dimorphism, 102
Semi-pupa, 118
Sesiidae, 62
Sexual differences, 15, 20-1, 90
Sharp, D., 13, 36, 40, 115
Silk-spinning, 58, 62-3, 82
Silkworms, 82
Silpha, 50
Siltala, A.J., 63
Silvestri, F., 119
Simulium, 78, 87
Smith, J.B., 17
Sphegidae, 66-7
Sphingidae, 60
Spinneret, 58
Spiracles, 2, 23, 70, 72, 77, 86, 87
Spring-tails, 11
Stone-flies, 24, 107, 110
Sub-imago, 33, 117
Sucking insects, 17
Swammerdam, J., 33
Syrphus, 74-6
Tachininae, 73, 91
_Tenebrio molitor_, 119
Termitoxeniidae, 92
Theobald, F.V., 100
Thysanura, 11
Tiger Moths, 59, 82, 98
Timber-beetles, 54
Tineidae, 62
Tipulidae, 70
Tortoiseshell Butterfly, 45, 95
Tortricidae, 62
Tracheal system. _See_ Air-tubes, Spiracles
Transformation. _See_ Metamorphosis
Triassic insects, 107
Trichocera, 70
Trichoptera, 62-3, 76, 80, 86
Tsetse Flies, 91
Turnip-fly, 53, 92, 94
Turnip Moth, 98-9
Tussock Moths, 90, 97
_Vanessa urticae_, 45, 95
Van Rees, J., 42
Vapourer Moth, 96-7, 115
_Velia currens_, 116
Verhoeff, K.W., 11
Vermiculiform larvae, 67, 71-6, 111
Virgin stem-mothers, 18
Viviparous reproduction. _See_ Birth
Wagner, N., 90
Warble-fly, 73-4, 89, 108
Warning coloration, 60
Wasmann, E., 92
Wasps, 46, 64, 66-7, 83
Water-insects. _See_ Aquatic insects
Weevils, 55
Weismann, A., 38, 42, 102
White Butterflies, 41, 83, 85, 100-3
Willow-beetles, 53
Wingless insects, 15, 18, 20, 96, 115
Wing-rudiments, 13, 18, 20, 22, 24, 28, 33, 36-8, 40, 111, 115, 117-19
Wings, 1, 14, 115, 119-20
Winter broods, 102-3
Wintering stages, 93-101
Wireworms, 52, 93
Wood-wasps, 65
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