THE COLOURATION OF INSECTS.
In the decoration of insects and birds, nature has exerted all her
power; and amongst the wealth of beauty here displayed we ought to find
crucial tests of the views herein advocated. It will be necessary,
therefore, to enter somewhat into detail, and we shall take butterflies
as our chief illustration, because in them we find the richest display
of colouring. The decoration of caterpillars will also be treated at
some length, partly because of their beauty, and partly because amongst
them sexual selection cannot possibly have had any influence.
Butterflies are so delicate in structure, so fragile in constitution, so
directly affected by changes of environment, that upon their wings we
have a record of the changes they have experienced, which gives to them
a value of the highest character in the study of biology. In them we can
study every variation that geographical distribution can effect; for
some species, like the Swallow-tail (_Papilio machaon_) and the Painted
Lady (_Cynthia cardui_), are almost universal, and others, like our now
extinct Large Copper (_Lycæna dispar_), are excessively local, being
confined to a very few square miles. From the arctic regions to the
tropics, from the mountain tops to the plains, on the arid deserts and
amidst luxuriant vegetation, butterflies are everywhere to be found.
Before entering into details, it will be as well to sketch some of the
broad features of butterfly decoration. In the first place they are all
day-fliers, and light having so strong an influence upon colour, there
is a marked difference in beauty between them and the night-flying
moths. A collection of butterflies viewed side by side with a collection
of moths brings out this fact more strongly than words can describe,
especially when the apparent exceptions are considered; for many moths
are as brightly coloured as butterflies. These will be found to belong
either to day-flying species, like the various Burnets (_Zygæna_), Tiger
Moths (_Arctia_), or evening flyers like the Hawk Moths (_Sphyngidæ_.)
The true night-flying, darkness-loving moths cannot in any way compare
with the insects that delight in sunshine. We see the same thing in
birds, for very few nocturnal species, so far as we are aware, are
brilliantly decorated.
Another salient feature is the difference that generally exists between
the upper and lower surfaces of the wings. As a rule, the upper surface
is the seat of the brightest colour. Most butterflies, perhaps all,
close their wings when at rest, and the upper wing is generally dropped
behind the under wing, so that only the tip is visible. The under
surface is very frequently so mottled and coloured as to resemble the
insect's natural surroundings, and so afford protection. It does not
follow that this protective colouring need be dull, and only when we
know the habit of the insect can we pronounce upon the value of such
colouring. The pretty Orange-tip has its under wings veined with green,
and is most conspicuous in a cabinet, but when at rest upon some
umbelliferous plant, with its orange tip hidden, these markings so
resemble the environment as to render the insect very inconspicuous. The
brilliant _Argynnis Lathonia_, with its underside adorned with plates of
metallic silver, is in the cabinet a most vivid and strongly-marked
species; but we have watched this insect alight among brown leaves, or
on brown stones, outside Florence, where it is very common, and find
that these very marks are a sure protection, for the insect at rest is
most difficult to see, even when it is marked down to its resting-place.
But some butterflies have parts of the under surface as gaily decorated
as the upper; and this not for protection. This may be seen to some
extent in our own species, for instance in the orange-tip of the
Orange-tip, and the red bar in the upper wing of the Red Admiral (_V.
atalanta_). If we watch these insects, the conviction that these are
true ornaments is soon forced upon us. The insect alights, perhaps
alarmed, closes its wings, and becomes practically invisible. With
returning confidence it will gradually open its wings and slowly vibrate
them, then close them again, and lift the upper wing to disclose the
colour. This it will do many times running, and the effect of the sudden
appearance and disappearance of the bright hues is as beautiful as it
is convincing. None can doubt the love of display exhibited in such
actions.
The delicacy of their organization renders butterflies peculiarly
susceptible to any change, and hence they exhibit strong tendencies to
variation, which make them most valuable studies. Not only do the
individuals vary, but the sexes are often differently coloured. Where
two broods occur in a season they are sometimes quite differently
decorated, and finally a species inhabiting widely different localities
may have local peculiarities.
We can thus study varieties of decoration in many ways, and we shall
treat of them as follows:--
1. _Simple Variation_, in which the different individuals of a
species vary in the same locality.
2. _Local Variation_, in which the species has marked peculiarities
in different localities.
3. _Sexual Dimorphism_, in which the sexes vary.
4. _Seasonal Dimorphism_, in which the successive broods differ.
[Illustration: Fig. 3. Diagram of Butterfly's Wing.
A. Upper Wing.
B. Lower Wing.
_a._ Costal Margin.
_b._ Hind Margin.
_c._ Inner "
_d._ Anal Angle.
_e._ Costa.
_f._ Costal nervure.
_g._ Sub-costal do.
_g_^{1-4}. Branches of do.
_h._ Median nervure.
_i._ Sub-median do.
_j._ Discoidal Cell.
_k._ Discoidal Veins.]
In order fully to understand the bearing of the following remarks it is
necessary to know something of the anatomy and nomenclature of
butterflies. Fig. 3 is an ideal butterfly. The wing margins are
described as the _Costal_, which is the upper strong edge of the wing,
the _Hind_ margin, forming the outside, and the _Inner_ margin, forming
the base. The nervures consist of four principal veins; the _Costal_, a
simple nervure under the costa, the _Sub-costal_, which runs parallel to
the costal and about halfway to the tip emits branches, generally four
in number; the _Median_ occupying the centre of the wing and sending off
branches, usually three in number, and the _Sub-median_ below which is
always simple. There are thus two simple nervures, one near the costal
the other near the inner margin, and between them are two others which
emit branches. Between these two latter is a wide plain space known as
the _discoidal cell_. Small veins called the _discoidal_ pass from the
hind margin towards the cell, and little transverse nervures, known as
sub-discoidal, often close the cell. By these nervures the wing is
mapped out into a series of spaces of which one, the discoidal cell, is
the most important.
The nervures have two functions, they support and strengthen the wing,
and being hollow serve to convey nutritive fluid and afterwards air to
the wing.
The wings are moved by powerful muscles attached to the base of the
wings close to the body and to the inside of the thorax, all the muscles
being necessarily internal. "There are two sets which depress the wings;
firstly a double dorsal muscle, running longitudinally upwards in the
meso-thorax;[28] and, secondly, the dorso-ventral muscles of the meso-
and meta-thorax,[29] which are attached to the articulations of the
wings above, and to the inside of the thorax beneath. Between these lie
the muscles which raise the wings and which run from the inner side of
the back of the thorax to the legs."[30] When we consider the immense
extent of wing as compared with the rest of the body, the small area of
attachment, and the great leverage that has to be worked in moving the
wings, it is clear that the area of articulation of the wing to the body
is one in which the most violent movement takes place. It is here that
the waste and repair of tissue must go on with greatest vigour, and we
should, on our theory, expect it to be the seat of strong emphasis.
Accordingly we commonly find it adorned with hairs, and in a vast number
of cases the general hue is darker than that of the rest of the wing,
and so far as we have been able to observe, never lighter than the body
of the wing. Even in the so-called whites (Pieris) this part of the wing
is dusky, and instances are numerous on Plate IV.
The scales, which give the colour to the wings, deserve more than a
passing notice. They are inserted by means of little stalks into
corresponding pits in the wing-membrane, and overlap like tiles on a
roof; occasionally the attachment is a ball and socket (_Morphinæ_), in
which case it is possible the insect has the power of erecting and
moving its scales. The shapes are very numerous, but as a rule they are
short. To this there is a remarkable exception on the wings of the males
of certain butterflies, consisting of elongated tufted prominences which
appear to be connected with sense-organs. They are probably
scent-glands, and thus we find, even in such minute parts as scales, a
difference of function emphasized by difference of ornamentation, here
showing itself in variety of forms; but, as we have said, ornamentation
in form is often closely allied to ornamentation in colours. In some
butterflies, indeed, these scales are aggregated into spots, as in
_Danais_, and have a different hue from the surrounding area.
The scales are not simple structures, but consist of two or more plates,
which are finely striated. The colouring matter consists of granules,
placed in rows between the striæ, and may exist upon the upper surface
of the upper membrane (epidermal), or the upper surface of the under or
middle plate (hypodermal), or the colour may be simple diffraction
colour, arising from the interference of the lightwaves by fine striæ.
Dr. Haagen, in the admirable paper before mentioned, has examined this
question thoroughly, and gives the results set forth in the following
table:--
_Epidermal Colours._
Metallic blues and greens }
Bronze }
Gold }
Silver } Persistent after death.
Black }
Brown }
Red (rarely) }
_Hypodermal Colours._
Blue }
Green }
Yellow }
Milk-white } Fading after death.
Orange and }
shades between }
Red }
The hypodermal colours are usually lighter than the epidermal, and are
sometimes changed by a voluntary act. Hypodermal and epidermal colours
are, of course, not peculiar to insects; and, as regards the former, it
is owing to their presence that the changing hues of fishes, like the
sole and plaice, and of the chameleon are due.
The great order Lepidoptera, including butterflies and moths, seems to
the non-scientific mind to be composed of members which are pretty much
alike, the differences being of slight importance; but this is not in
reality the case, for the lepidoptera might, with some accuracy, be
compared to the mammalia, with its two divisions of the placental and
non-placental animals. Comparing the butterflies (Rhopalocera) to the
placental mammals, we may look upon the different families as similar to
the orders of the mammalia. Were we as accustomed to notice the
differences of butterflies as we are to remark the various forms of
familiar animals, we should no longer consider them as slight, but
accord to them their true value. When in the mammalia we find animals
whose toes differ in number, like the three-toed rhinoceros and the
four-toed tapir, we admit the distinction to be great, even apart from
other outward forms. So, too, the seal and lion, though both belonging
to the carnivora, are readily recognized as distinct, but the seals may
easily be confounded by the casual observer with the manatees, which
belong to quite a different order.
Thus it is with the Lepidoptera, for from six-legged insects, whose pupæ
lie buried beneath the soil, like most moths, we pass to the highest
butterflies, whose fore-legs are atrophied, and whose pupæ hang
suspended in the open air; and this by easy intermediate stages. Surely,
if six-legged mammals were the rule, we should look upon four-legged
ones as very distinct; and this is the case with the butterflies. It is
necessary to make this clear at starting, in order that we may
appreciate to its full value the changes that have taken place in the
insects under study.
Butterflies (_Rhopalocera_) are grouped into four sub-families, as
under:--
1. _Nymphalidæ_, having the fore-legs rudimentary, and the pupæ
suspended from the base of the abdomen.
2. _Erycinidæ_, in which the males only have rudimentary fore-legs.
3. _Lycænidæ_, in which the fore-legs of the males are smaller than
those of the females, and terminate in a simple hook.
4. _Papilionidæ_, which have six perfect pairs of legs, and in
which the pupæ assume an upright posture, with a cincture round the
middle.
It may, at first sight, appear curious that the imperfect-legged
_Nymphalidæ_ should be placed at the head of the list, but this is based
upon sound reasoning. The larva consists of thirteen segments, and, in
passing to the mature stage, the second segment alone diminishes in
size, and it is to this segment that the first pair of legs is attached.
Looking now to the aerial habits of butterflies, we can understand how,
in the process of evolution towards perfect aerial structure, the legs,
used only for walking, would first become modified; and, naturally,
those attached to the segment which decreases with development would be
the first affected. When this is found to be combined with an almost
aerial position of the pupæ, we see at once how such insects approach
nearest to an ideal flying insect. It is a general law that suppression
of parts takes place as organisms become specialized. Thus, in the
mammalia, the greatest number of toes and teeth are found in the lowest
forms and in the oldest, simplest fossil species.
A butterfly is, indeed, little more than a beautiful flying machine; for
the expanse of wing, compared with the size of the body, is enormous.
[Illustration]
[28] The middle division of the thorax.
[29] Hinder division of thorax.
[30] Dallas in Cassell's Nat. Hist., vol. vi., p. 27.