INSECT COLOURING
_Phanaeus splendidulus_, the glittering, the resplendent: this is the
epithet selected by the official nomenclators to describe the
handsomest Dung-beetle of the pampas. The name is not at all
exaggerated. Combining the fire of gems with metallic lustre, the
insect, according to the incidence of the light, emits the green
reflections of the emerald or the gleam of ruddy copper. The
muck-raker would do honour to the jeweller's show-cases.
For the rest, our own Dung-beetles, though usually modest in their
attire, also have a leaning toward luxurious ornament. One Onthophagus
decorates his corselet with Florentine bronze; another wears garnets
on his wing-cases. Black above, the Mimic Geotrupes is the colour of
copper pyrites below; also black in all parts exposed to the light of
day, the Stercoraceous Geotrupes displays a ventral surface of a
glorious amethyst violet.
Many other series, of greatly varied habits, Carabi,[1] Cetoniae,
Buprestes, Chrysomelae,[2] rival and even surpass the magnificent
Dung-beetles in the matter of jewellery. At times we encounter
splendours which the imagination of a lapidary would not venture to
depict. The Azure Hoplia,[3] the inmate of the osier-beds and elders
by the banks of the mountain streams, is a wonderful blue, tenderer
and softer to the eye than the azure of the heavens. You could not
find an ornament to match it save on the throats of certain
Humming-birds and the wings of a few Butterflies in equatorial climes.
[Footnote 1: Cf. Chapter XIV. of the present volume.--_Translator's
Note_.]
[Footnote 2: Golden Apple-beetles.--_Translator's Note_.]
[Footnote 3: A genus of Cockchafer. Cf. _The Life of the Fly_: chap.
vii.--_Translator's Note_.]
To adorn itself like this, in what Golconda does the insect gather its
gems? In what diggings does it find its gold nuggets? What a pretty
problem is that of a Buprestis' wing-case! Here the chemistry of
colours ought to reap a delightful harvest; but the difficulties are
great, it seems, so much so that science cannot yet tell us the why
and the wherefore of the humblest costume. The answer will come in a
remote future, if indeed it ever comes completely, for life's
laboratory may well contain secrets denied to our retorts. For the
moment, I shall perhaps be contributing a grain of sand to the future
palace if I describe the little that I have seen.
My basic observation dates a long way back. I was at that time busy
with the Hunting Wasps, following their larval development from the
egg to the cocoon. Let us take an instance from my notes, which cover
nearly all the game-hunters of my district. I will choose the larva of
the Yellow-winged Sphex,[4] which, with its convenient size, will
furnish an easy object-lesson.
[Footnote 4: Cf. _The Hunting Wasps_: chap. iv.--_Translator's Note_.]
Under the transparent skin of the larva, which has been recently
hatched and is consuming its first Cricket, we soon perceive some fine
white spots, which rapidly increase in size and number and eventually
cover the whole body, except the first two or three segments. On
dissecting the grub, we find that these spots have to do with the
adipose layer, of which they form a considerable part, for, far from
being scattered only on the surface, they run through its whole
thickness and are present in such numbers that the forceps cannot
seize the least fragment of this tissue without picking up a few of
them.
Though perfectly visible without the help of a lens, these puzzling
spots call for the microscope if we wish to study them in detail. We
then find that the adipose tissue is made up of two kinds of vesicles:
some, bright yellow and transparent, are filled with oily drops; the
rest, opaque and starch-white, are distended with a very fine powder,
which spreads in a cloudy trail when the vesicle containing it is
broken on the object-slide. Intermingled without apparent order, the
two kinds of bags are of the same shape and the same size. The first
go to make up the nutritive reserves, the fatty tissue properly
so-called; the second form the white dots which we will study for a
moment.
An inspection under the microscope tells us that the contents of the
white cells are composed of very fine, opaque grains, insoluble in
water and of greater density. The use of chemical reagents on the
object-slide proves that nitric acid dissolves these grains, with
effervescence and without leaving the least residue, even when they
are still enclosed in their vesicles. On the other hand, the true
fatty cells suffer in no way when attacked by this acid; they merely
turn a little yellower.
Let us take advantage of this property to operate on a larger scale.
The adipose tissue taken from a number of larvae is treated with
nitric acid. The effervescence is as lively as if the reaction were
taking effect on a bit of chalk. When it has subsided, some yellow
clots are floating on the surface. These are easily separated. They
come from the fatty substance and the cellular membranes. There
remains a clear liquid containing the white granules in solution.
The riddle of these granules was being presented to me for the first
time; my predecessors had provided no physiological or anatomical data
to guide me; great therefore was my joy when, after a little fumbling,
I succeeded in hitting upon their characteristic feature.
The solution is evaporated in a small porcelain capsule, placed on the
hot embers. On the residue I pour a few drops of ammonia, or else
simply water. A glorious crimson colour at once makes its appearance.
The problem is solved: the colouring-matter which has just formed is
murexide; and consequently the powdery substance which filled the
cells was none other than uric acid, or more precisely ammonium urate.
A physiological fact of this importance can hardly stand alone.
Indeed, since this basic experiment I have discovered grains of uric
acid in the adipose tissue of the larvae of all the Hunting Wasps of
our parts, as well as in the Bees at the moment of the nymphosis. I
have observed them in many other insects, either in the larval or in
the perfect state; but in this respect there is none to equal the grub
of the game-hunting Wasp, which is all speckled with white. I think I
see the reason.
Let us consider two larvae which eat live prey: that of the Sphex and
that of the Hydrophilus.[5] Uric acid, the inevitable product of the
vital transformations, or at all events one of its analogues, must be
formed in both. But the Hydrophilus' larva shows no accumulation of it
in its adipose layer, whereas the Sphex' is full of it.
[Footnote 5: The Great Water-beetle.--_Translator's Note_.]
In the latter the duct through which the solid excretions pass is not
yet in working order; the digestive apparatus, tied at the lower end,
is not discharging an atom. The urinary products, being unable, for
want of an open outlet, to flow away as formed, accumulate in the
adipose tissue, which thus serves as a common store-house for the
residues of the present and the plastic material of the future organic
processes. Here something occurs analogous to what we see in the
higher animals after the removal of the kidneys; the urea at first
contained in the blood, in imperceptible quantities accumulates and
becomes manifest when the means by which it is eliminated disappear.
In the larva of the Hydrophilus, on the other hand, the excretions
enjoy a free outlet from the beginning; and the urinary products
escape as and when formed and are no longer deposited in the adipose
tissue. But during the intense labour of the metamorphosis, any
excretion becomes impossible; the uric acid must and does collect in
the adipose substance of the different larvae.
It would be out of place, despite its importance, to pursue the
problem of the uric residues any further. Our subject is coloration.
Let us return to it with the evidence supplied by the Sphex. Her
almost transparent larva has the neutral tint of fluid white of egg.
Under its fine translucid skin there is nothing coloured, save the
long digestive pouch, which is swollen a deep purple by the pulp of
the consumed Crickets. But against this indefinite, vitreous
background the opaque white uric cells stand out distinctly in their
myriads; and the effect of this stippling is a sketchy but by no means
inelegant costume. It is skimpy in the extreme, but at any rate it is
something.
With the urinary broth of which its intestine is unable to get rid,
the larva has discovered a means of making itself look a little smart.
The Anthidia have shown us how, in their cotton-wool wallets, they
manufacture a sort of jewellery with their ordure. The robe studded
with grains of alabaster is a no less ingenious invention.
To beautify themselves cheaply by using up their own refuse is a very
common method even among insects endowed with all that is wanted for
evacuating waste matter. While the larvae of the Hunting Wasps, unable
to do better, stipple themselves with uric acid, there are plenty of
industrious creatures that are able to make themselves a superb dress
by preserving their excretions in spite of their own open sewers. With
a view to self-embellishment, they collect and treasure up the dross
which others hasten to expel. They turn filth into finery.
One of these is the White-faced Decticus (_D. albifrons_, FAB.), the
biggest sabre-bearer of the Provencal fauna. A magnificent insect is
this Grasshopper, with a broad ivory face, a full, creamy-white belly
and long wings flecked with brown. In July, the season for the
wedding-dress, let us dissect him under water.
The adipose tissue, which is abundant and yellowish white, consists of
a lace of wide, irregular, criss-cross meshes. It is a tubular network
swollen with a powdery matter which condenses into minute chalk-white
spots, standing out very plainly against a transparent background.
When crushed in a drop of water, a fragment of this fabric yields a
milky cloud in which the microscope shows an infinite number of opaque
floating atoms, without revealing the smallest blob of oil, the sign
of fatty matter.
Here again we have ammonium urate. Treated with nitric acid, the
adipose tissue of the Decticus produces an effervescence similar to
that of chalk and yields enough murexide to redden a tumblerful of
water. A strange adipose body, this bundle of lace crammed with uric
acid without a trace of fatty matter! What would the Decticus do with
nutritive reserves, seeing that he is near his end, now that the
nuptial season has arrived? Delivered from the necessity of saving for
the future, he has only to spend in gaiety the few days left to him;
he has only to adorn himself for the supreme festival.
He therefore converts into a paint-factory what at first was a
warehouse for storing up foodstuffs; and with his chalk-like uric pulp
he lavishly daubs his belly, which turns a creamy white, and smears it
on his forehead, his face, his cheeks, until they assume the
appearance of old ivory. All those parts, in fact, which lie
immediately under the translucid skin are covered with a layer of
pigment which can be turned into murexide and is identical in nature
with the white powder of the adipose lace.
Biological chemistry can hardly offer a simpler and more striking
experiment than this analysis of the Decticus' finery. To those who
have not this curious Grasshopper handy, I recommend the Ephippiger of
the Vines, who is much more widely distributed. His ventral surface,
which also is of a creamy white, likewise owes its colour to a
plastering of uric acid. In the Grasshopper family many other species
of smaller size and requiring more delicate handling would give us the
same results in varying degrees.
White, slightly tinged with yellow, is all that the urinary palette of
the Locustidae shows us. A caterpillar, the Spurge Hawk-moth's, will
take us a little farther. Dappled red, black, white and yellow, its
livery is the most remarkable in our part of the country. Reaumur in
fact calls it _la Belle_. The flattering title is well-deserved. On
the black background of the larva, vermillion-red, chrome-yellow and
chalk-white figure side by side in circles, spots, freckles and
stripes, as clearly marked as the glaring patches of a harlequin's
dress.
Let us dissect the caterpillar and apply the lens to its mosaic. On
the inner surface of the skin, except in the portions coloured black,
we observe a pigmentary layer, a coating here red, there yellow or
white. We will cut a strip from this coat of many colours, after
depriving it of its muscular fibres, and subject it to the action of
nitric acid. The pigment, no matter what its hue, dissolves with
effervescence and afterwards yields murexide. Here again, then, it is
to uric acid, present, however, in small quantities in the adipose
tissue, that the caterpillar's rich livery is due.
The black parts are an exception. Unassailable by nitric acid, they
retain their sombre tint after treatment as before, whereas the
portions stripped of their pigment by the reagent become almost as
transparent as glass. The skin of the handsome caterpillar thus has
two sorts of coloured patches.
Those of an intense black may be likened to dyers' products: they are
completely impregnated with the colouring matter, which is part and
parcel of the molecular constitution and cannot be isolated by the
nitric solvent. The others, red, yellow or white, are actually
painted: on a translucid sheet is a wash of urinary pigment, which is
discharged by the minute ducts issuing from the adipose layer. When
the action of the nitric acid has ceased, the transparent circles of
the latter stand out against the black background of the former.
Yet one more example taken from a different order. As regards elegance
of costume, the Banded Epeira[6] is the most highly favoured of our
Spiders. On the upper surface of her corpulent belly alternate, in
transversal bands, bright black, a vivid yellow like that of yolk of
egg and a dazzling white like that of snow. The black and yellow also
show underneath, but arranged differently. The yellow, in particular,
forms two longitudinal ribbons, ending in orange-red on either side of
the spinnerets. A pale purple is faintly diffused over the sides.
[Footnote 6: Cf. _The Life of the Spider_: chaps. ii., vii., xi. and
xiii.--_Translator's Note_.]
Examined from the outside with the lens, the black parts reveal
nothing out of the common. The black is homogeneous and everywhere of
equal depth. On the other hand, in the coloured portions, we see
little polygonal, granular masses, forming a close-meshed network. By
cutting round the circumference of the abdomen with a pair of
scissors, the horny integument of the dorsal surface may readily be
removed in one piece, without any shreds of the organs which it
protected. This large strip of skin is transparent in the zones that
correspond with the white bands in the natural state; it is black or
yellow on the black or yellow bands. These last indeed owe their
colouring to a layer of pigment which the point of a paintbrush will
easily loosen and remove.
As for the white bands, their origin is this: once the skin has been
removed, the dorsal surface of the abdomen, whose graceful mosaic is
not in any way disturbed, reveals a layer of polygonal white spots,
distributed in belts, here densely and there less so. The denser belts
correspond with the white bands. It is their magnificent opaque white
granulations which, seen through the transparent skin, form the
snow-white stripes in the live Spider.
Treated with nitric acid on the microscopic slide, they do not
dissolve nor produce effervescence. Uric acid then is not present in
this case; and the substance must be guanine, an alkaloid known to be
the urinary product of the Spiders. The same is true of the yellow,
black, purple or orange pigment that forms a coating under the skin.
In short, by utilizing, in a different chemical combination, the waste
products of animal oxidization, the magnificent Spider rivals the
magnificent caterpillar; she beautifies herself with guanine as the
other does with its uric acid.
Let us abridge this dry subject; let us be content with these few
data, which could if necessary be corroborated by many others. What
does the little that we have learnt teach us? It tells us that the
materials rejected by the organism, guanine, uric acid and other dross
from life's refinery, play an important part in the coloration of the
insect.
Two cases are distinguishable, according as the colour is dyed or
simply painted. The skin, itself colourless and transparent, is in
places illumined by a coloured varnish, which can be removed by a
touch with a paintbrush. Here we have paint, the result of the urinary
compound laid on the inner surface of the covering, just as the
chromatic ingredients of our glass-painters are laid on our
stained-glass windows.
At other places the skin is coloured in its very substance; the
colouring-matter forms an integral part of it and can no longer be
swept away with a camel-hair brush. Here we have a dyed fabric,
represented in our windows by the panes of coloured glass which the
crucible decorates uniformly with this or that tint, by means of the
incorporated metallic oxides.
Whereas, in these two cases, there is a profound difference in the
distribution of the chromatic materials, is this true of their
chemical nature as well? The suggestion is hardly admissible. The
worker in stained glass dyes or paints with the same oxides. Life,
that incomparable artist, must even more readily obtain an infinite
variety of results by uniformity of method.
It shows us, on the back of the Spurge Caterpillar,[7] black spots
jumbled up with other spots, white, yellow or red. Paints and dyes lie
side by side. Is there on this side of the dividing line a paint-stuff
and on the other side a dye-stuff, absolutely different in character
from the first? While chemistry is not yet in a position to
demonstrate, with its reagents, the common origin of the two
substances, at least the most convincing analogies point to it.
[Footnote 7: The caterpillar of the Spurge Hawk-moth.--_Translator's
Note_.]
In this delicate problem of the insect's colouring, one single point
thus far comes within the domain of observed facts: the progressive
advance of chromatic evolution. The carbuncle of the Dung-Beetle of
the Pampas suggested the question. Let us then inquire of his near
neighbours, who will perhaps enable us to advance a step farther.
Newly stripped of his cast-off nymphal skin, the Sacred Beetle
possesses a strange costume, bearing no resemblance to the ebony black
which will be the portion of the mature insect. The head, legs and
thorax are a bright rusty red; the wing-cases and abdomen are white.
As a colour, the red is almost that of the Spurge Caterpillar, but it
is the result of a dye on which nitric acid has no effect as a
detector of urates. The same chromatic principle must certainly exist
in a more elaborate form and under a different molecular arrangement
in the skin of the abdomen and the wing-cases which will presently
replace white by red.
In two or three days the colourless becomes the coloured, a process
whose rapidity implies a fresh molecular structure rather than a
change of composition. The building-stone remains the same, but is
arranged in a different order; and the structure alters in appearance.
The Scarabaeus is now all red. The first brown stains show themselves
on the denticulations of the forehead and fore-legs, the sign of an
earlier maturity in the implements of labour, which are to acquire an
exceptional hardness. The smoky tinge spreads more or less all over
the insect, replaces the red, turns darker and finally becomes the
regulation black. In less than a week the colourless insect turns a
rusty red, next a sooty brown and then an ebony black. The process is
completed; the insect possesses its normal colouring.
Even so do the Copres, the Gymnopleuri,[8] the Onites, the Onthophagi
and many others behave; even so must the jewel of the pampas, the
Splendid Phanaeus set to work. With as much certainty as though I had
him before my eyes at the moment when he divests himself of his
nymphal swaddling-bands, I see him a dull red, rusty or crimson,
excepting on the wing-covers and the abdomen, which are at first
colourless and presently turn the same colour as the rest. In the
Sacred Beetle this initial red is followed by black; the Phanaeus
replaces it by the brilliance of copper and the reflections of the
emerald. Ebony, metal, the gem: have they the same origin here then?
Evidently.
[Footnote 8: Cf. _The Sacred Beetle and Others_: chap.
viii.--_Translator's Note_.]
The metallic lustre does not call for a change of nature; a mere
nothing is enough to produce it. Silver, when very finely subdivided
by the methods whereof chemistry knows the secret, becomes a dust as
poor to look at as soot. When pressed between two hard bodies, this
dirty powder, which might be dried mud, at once acquires the metallic
sheen and again becomes the silver which we know. A mere molecular
contact has wrought the miracle.
Dissolved in water, the murexide derived from uric acid is a
magnificent crimson. Solidified by crystallization, it rivals in
splendour the gold-green of the Cantharides. The widely-used fuschine
affords a well-known example of like properties.
Everything, then, appears to show that the same substance, derived
from urinary excretions, yields, according to the mode in which its
ultimate particles are grouped, the metallic red of the Phanaeus, as
well as the white, the dull red and the black of the Sacred Beetle. It
becomes black on the dorsal surface of the Stercoraceous Geotrupes and
the Mimic Geotrupes; and, with a quick change, it turns into amethyst
under the belly of the first and into copper pyrites under the belly
of the second. It covers the back of _Cetonia floricola_ with golden
bronze and the under surface with metallic purple. According to the
insect, according to the part of the body, it remains a dingy compound
or sparkles with reflections even more vivid and varied than those
possessed by the metals.
Light seems irrelevant to the development of these splendours; it
neither accelerates nor retards them. Since direct exposure to the
sun, owing to the excess of heat, is fatal to the delicate process of
the nymphosis, I shaded the solar rays with a screen of water
contained between slips of glass; and to the bright light thus
moderated in temperature I daily, throughout the period of chromatic
evolution, subjected a number of Sacred Beetles, Geotrupes and
Cetoniae. As standards of comparison I had witnesses of whom I kept
some in diffused light and others in complete darkness. My experiments
had no appreciable result. The development of the colours took place
in the sunlight and in the dark alike, neither more rapidly nor more
slowly and without difference in the tints.
This negative result was easy to foresee. The Buprestis emerging from
the depths of the trunk in which he has spent his larval life; the
Geotrupes and the Phanaeus leaving their natal burrows possess their
final adornments, which will not become richer in the rays of the sun,
at the time when they make their appearance in the open air. The
insect does not claim the assistance of the light for its colour
chemistry, not even the Cicada,[9] who bursts her larval scabbard and
changes from pale green to brown as easily in the darkness of my
apparatus as in the sunlight, in the usual manner.
[Footnote 9: Cf. _The Life of the Grasshopper_: chaps. i. to
v.--_Translator's Note_.]
The chromatics of the insect, having as its basis the urinary waste
products, might well be found in various animals of a higher order. We
know of at least one example. The pigment of a small American lizard
is converted into uric acid under the prolonged action of boiling
hydrochloric acid.[10] This cannot be an isolated instance; and there
is reason to believe that the reptilian class daubs its garments with
similar products.
[Footnote 10: A. B. Griffiths, Transactions of the Academie des
sciences, 26 November, 1894.--_Author's Note_.]
From the reptile to the bird is no great distance. Then the
Wood-pigeon's iridescent hues, the eyes on the Peacock's tail, the
Kingfisher's sea-blue, the Flamingo's carmine are more or less closely
connected with the urinary excretions? Why not? Nature, that sublime
economist, delights in these vast antitheses which upset all our
conceptions of the values of things. Of a pinch of common charcoal she
makes a diamond; of the same clay which the potter fashions into a
bowl for the Cat's supper she makes a ruby; of the filthy waste
products of the organism she makes the splendours of the insect and
the bird. The metallic marvels of the Buprestis and the Ground-beetle;
the amethyst, ruby, sapphire, emerald and topaz of the Humming-bird;
glories which would exhaust the language of the lapidary jeweller:
what are they in reality? Answer: a drop of urine.