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A BEE-HUNTER - THE PHILANTHUS AVIPORUS




To encounter among the Hymenoptera, those ardent lovers of flowers, a
species which goes a-hunting on its own account is, to say the least of
it, astonishing. That the larder of the larvae should be provisioned with
captured prey is natural enough; but that the provider, whose diet is
honey, should itself devour its captives is a fact both unexpected and
difficult to comprehend. We are surprised that a drinker of nectar
should become a drinker of blood. But our surprise abates if we consider
the matter closely. The double diet is more apparent than real; the
stomach which fills itself with the nectar of flowers does not gorge
itself with flesh. When she perforates the rump of her victim the
Odynerus does not touch the flesh, which is a diet absolutely contrary
to her tastes; she confines herself to drinking the defensive liquid
which the grub distils at the end of its intestine. For her this liquid
is doubtless a beverage of delicious flavour, with which she relieves
from time to time her staple diet of the honey distilled by flowers,
some highly spiced condiment, appetiser or aperient, or perhaps--who
knows?--a substitute for honey. Although the qualities of the liquid
escape me, I see at least that Odynerus cares nothing for the rest.
Once the pouch is emptied the larva is abandoned as useless offal, a
certain sign of non-carnivorous appetites. Under these conditions the
persecutor of Chrysomela can no longer be regarded as guilty of an
unnatural double dietary.

We may even wonder whether other species also are not apt to draw some
direct profit from the hunting imposed upon them by the needs of the
family. The procedure of Odynerus in opening the anal pouch is so far
removed from the usual that we should not anticipate many imitators; it
is a secondary detail, and impracticable with game of a different kind.
But there may well be a certain amount of variety in the means of direct
utilisation. Why, for example, when the victim which has just been
paralysed or rendered insensible by stinging contains in the stomach a
delicious meal, semi-liquid or liquid in consistency, should the hunter
scruple to rob the half-living body and force it to disgorge without
injuring the quality of its flesh? There may well be robbers of the
moribund, attracted not by their flesh but by the appetising contents of
their stomachs.

As a matter of fact there are such, and they are numerous. In the first
rank we may cite that hunter of the domestic bee, _Philanthus aviporus_
(Latreille). For a long time I suspected Philanthus of committing such
acts of brigandage for her own benefit, having many times surprised her
gluttonously licking the honey-smeared mouth of the bee; I suspected
that her hunting of the bee was not undertaken entirely for the benefit
of her larvae. The suspicion was worth experimental confirmation. At the
time I was interested in another question also: I wanted to study,
absolutely at leisure, the methods by which the various predatory
species dealt with their victims. In the case of Philanthus I made use
of the improvised cage already described; and Philanthus it was who
furnished me with my first data on the subject. She responded to my
hopes with such energy that I thought myself in possession of an
unequalled method of observation, by means of which I could witness
again and again, to satiety even, incidents of a kind so difficult to
surprise in a state of nature. Alas! the early days of my acquaintance
with Philanthus promised me more than the future had in store for me!
Not to anticipate, however, let us place under the bell-glass the hunter
and the game. I recommend the experiment to whomsoever would witness the
perfection with which the predatory Hymenoptera use their stings. The
result is not in doubt and the waiting is short; the moment the prey is
perceived in an attitude favourable to her designs, the bandit rushes at
it, and all is over. In detail, the tragedy develops as follows:

I place under a bell-glass a Philanthus and two or three domestic bees.
The prisoners climb the glass walls, on the more strongly lighted side;
they ascend, descend, and seek to escape; the polished, vertical surface
is for them quite easy to walk upon. They presently quiet down, and the
brigand begins to notice her surroundings. The antennae point forward,
seeking information; the hinder legs are drawn up with a slight
trembling, as of greed and rapacity, in the thighs; the head turns to
the right and the left, and follows the evolutions of the bees against
the glass. The posture of the scoundrelly insect is strikingly
expressive; one reads in it the brutal desires of a creature in ambush,
the cunning patience that postpones attack. The choice is made, and
Philanthus throws herself upon her victim.

Turn by turn tumbled and tumbling, the two insects roll over and over.
But the struggle soon quiets down, and the assassin commences to plunder
her prize. I have seen her adopt two methods. In the first, more usual
than the other, the bee is lying on the ground, upon its back, and
Philanthus, mouth to mouth and abdomen to abdomen, clasps it with her
six legs, while she seizes its neck in her mandibles. The abdomen is
then curved forward and gropes for a moment for the desired spot in the
upper part of the thorax, which it finally reaches. The sting plunges
into the victim, remains in the wound for a moment, and all is over.
Without loosing the victim, which is still tightly clasped, the murderer
restores her abdomen to the normal position and holds it pressed against
that of the bee.

By the second method Philanthus operates standing upright. Resting on
the hinder feet and the extremity of the folded wings, she rises proudly
to a vertical position, holding the bee facing her by her four anterior
claws. In order to get the bee into the proper position for the final
stroke, she swings the poor creature round and back again with the
careless roughness of a child dandling a doll. Her pose is magnificent,
solidly based upon her sustaining tripod, the two posterior thighs and
the end of the wings, she flexes the abdomen forwards and upwards, and,
as before, stings the bee in the upper part of the thorax. The
originality of her pose at the moment of striking surpasses anything I
have ever witnessed.

The love of knowledge in matters of natural history is not without its
cruelties. To make absolutely certain of the point attained by the
sting, and to inform myself completely concerning this horrible talent
for murder, I have provoked I dare not confess how many assassinations
in captivity. Without a single exception, the bee has always been stung
in the throat. In the preparations for the final blow the extremity of
the abdomen may of course touch here and there, at different points of
the thorax or abdomen, but it never remains there, nor is the sting
unsheathed, as may easily be seen. Once the struggle has commenced the
Philanthus is so absorbed in her operations that I can remove the glass
cover and follow every detail of the drama with my magnifying-glass.

The invariable situation of the wound being proved, I bend back the head
of the bee, so as to open the articulation. I see under what we may call
the chin of the bee a white spot, hardly a twenty-fifth of an inch
square, where the horny integuments are lacking, and the fine skin is
exposed uncovered. It is there, always there, in that tiny defect in the
bee's armour, that the sting is inserted. Why is this point attacked
rather than another? Is it the only point that is vulnerable? Stretch
open the articulation of the corselet to the rear of the first pair of
legs. There you will see an area of defenceless skin, fully as delicate
as that of the throat, but much more extensive. The horny armour of the
bee has no larger breach. If the Philanthus were guided solely by
considerations of vulnerability she would certainly strike there,
instead of insistently seeking the narrow breach in the throat. The
sting would not grope or hesitate, it would find its mark at the first
attempt. No; the poisoned thrust is not conditioned by mechanical
considerations; the murderer disdains the wide breach in the corselet
and prefers the lesser one beneath the chin, for purely logical reasons
which we will now attempt to elicit.

The moment the bee is stung I release it from the aggressor. I am struck
in the first place by the sudden inertia of the antennae and the various
members of the mouth; organs which continue to move for so long a time
in the victims of most predatory creatures. I see none of the
indications with which my previous studies of paralysed victims have
made me familiar: the antennae slowly waving, the mandibles opening and
closing, the palpae trembling for days, for weeks, even for months. The
thighs tremble for a minute or two at most; and the struggle is over.
Henceforth there is complete immobility. The significance of this sudden
inertia is forced upon me: the Philanthus has stabbed the cervical
ganglions. Hence the sudden immobility of all the organs of the head:
hence the real, not the apparent death of the bee. The Philanthus does
not paralyse merely, but kills.

This is one step gained. The murderer chooses the point below the chin
as the point of attack, in order to reach the principal centres of
innervation, the cephalic ganglions, and thus to abolish life at a
single blow. The vital centres being poisoned, immediate death must
follow. If the object of the Philanthus were merely to cause paralysis
she would plunge her sting into the defective corselet, as does the
Cerceris in attacking the weevil, whose armour is quite unlike the
bee's. Her aim is to kill outright, as we shall presently see; she wants
a corpse, not a paralytic. We must admit that her technique is
admirable; our human murderers could do no better.

Her posture of attack, which is very different to that of the
paralysers, is infallibly fatal to the victim. Whether she delivers the
attack in the erect position or prone, she holds the bee before her,
head to head and thorax to thorax. In this position it suffices to flex
the abdomen in order to reach the joint of the neck, and to plunge the
sting obliquely upwards into the head of the captive. If the bee were
seized in the inverse position, or if the sting were to go slightly
astray, the results would be totally different; the sting, penetrating
the bee in a downward direction, would poison the first thoracic
ganglion and provoke a partial paralysis only. What art, to destroy a
miserable bee! In what fencing-school did the slayer learn that terrible
upward thrust beneath the chin? And as she has learned it, how is it
that her victim, so learned in matters of architecture, so conversant
with the politics of Socialism, has so far learned nothing in her own
defence? As vigorous as the aggressor, she also carries a rapier, which
is even more formidable and more painful in its results--at all events,
when my finger is the victim! For centuries and centuries Philanthus has
stored her cellars with the corpses of bees, yet the innocent victim
submits, and the annual decimation of her race has not taught her how to
deliver herself from the scourge by a well-directed thrust. I am afraid
I shall never succeed in understanding how it is that the assailant has
acquired her genius for sudden murder while the assailed, better armed
and no less powerful, uses her dagger at random, and so far without
effect. If the one has learned something from the prolonged exercise of
the attack, then the other should also have learned something from the
prolonged exercise of defence, for attack and defence are of equal
significance in the struggle for life. Among the theorists of our day,
is there any so far-sighted as to be able to solve this enigma?

I will take this opportunity of presenting a second point which
embarrasses me; it is the carelessness--it is worse than that--the
imbecility of the bee in the presence of the Philanthus. One would
naturally suppose that the persecuted insect, gradually instructed by
family misfortune, would exhibit anxiety at the approach of the
ravisher, and would at least try to escape. But in my bell-glasses or
wire-gauze cages I see nothing of the kind. Once the first excitement
due to imprisonment has passed the bee takes next to no notice of its
terrible neighbour. I have seen it side by side with Philanthus on the
same flower; assassin and future victim were drinking from the same
goblet. I have seen it stupidly coming to inquire what the stranger
might be, as the latter crouched watching on the floor. When the
murderer springs it is usually upon some bee which passes before her,
and throws itself, so to speak, into her clutches; either thoughtlessly
or out of curiosity. There is no frantic terror, no sign of anxiety, no
tendency to escape. How is it that the experience of centuries, which is
said to teach so much to the lower creatures, has not taught the bee
even the beginning of apine wisdom: a deep-rooted horror of the
Philanthus? Does the bee count upon its sting? But the unhappy creature
is no fencer; it thrusts without method, at random. Nevertheless, let us
watch it at the final and fatal moment.

When the ravisher brings her sting into play the bee also uses its
sting, and with fury. I see the point thrusting now in this direction,
now in that; but in empty air, or grazing and slipping over the
convexity of the murderer's back, which is violently flexed. These blows
have no serious results. In the position assumed by the two as they
struggle the abdomen of the Philanthus is inside and that of the bee
outside; thus the sting of the latter has under its point only the
dorsal face of the enemy, which is convex and slippery, and almost
invulnerable, so well is it armoured. There is no breach there by which
the sting might possibly enter; and the operation takes place with the
certainty of a skilful surgeon using the lancet, despite the indignant
protests of the patient.

The fatal stroke once delivered, the murderer remains for some time on
the body of the victim, clasping it face to face, for reasons that we
must now consider. It may be that the position is perilous for
Philanthus. The posture of attack and self-protection is abandoned, and
the ventral area, more vulnerable than the back, is exposed to the sting
of the bee. Now the dead bee retains for some minutes the reflex use of
the sting, as I know to my cost: for removing the bee too soon from the
aggressor, and handling it carelessly, I have received a most effectual
sting. In her long embrace of the poisoned bee, how does Philanthus
avoid this sting, which does not willingly give up its life without
vengeance? Are there not sometimes unexpected accidents? Perhaps.

Here is a fact which encourages me in this belief. I had placed under
the bell-glass at the same time four bees and as many Eristales, in
order to judge of the entomological knowledge of Philanthus as
exemplified in the distinction of species. Reciprocal quarrels broke out
among the heterogeneous group. Suddenly, in the midst of the tumult,
the killer is killed. Who has struck the blow? Certainly not the
turbulent but pacific Eristales; it was one of the bees, which by chance
had thrust truly in the mellay. When and how? I do not know. This
accident is unique in my experience; but it throws a light upon the
question. The bee is capable of withstanding its adversary; it can, with
a thrust of its envenomed needle, kill the would-be killer. That it does
not defend itself more skilfully when it falls into the hands of its
enemy is due to ignorance of fencing, not to the weakness of the arm.
And here again arises, more insistently than before, the question I
asked but now: how is it that the Philanthus has learned for purposes of
attack what the bee has not learned for purposes of defence. To this
difficulty I see only one reply: the one knows without having learned
and the other does not know, being incapable of learning.

Let us now examine the motives which induce the Philanthus to kill its
bee instead of paralysing it. The murder once committed, it does not
release its victim for a moment, but holding it tightly clasped with its
six legs pressed against its body, it commences to ravage the corpse. I
see it with the utmost brutality rooting with its mandibles in the
articulation of the neck, and often also in the more ample articulation
of the corselet, behind the first pair of legs; perfectly aware of the
fine membrane in that part, although it does not take advantage of the
fact when employing its sting, although this vulnerable point is the
more accessible of the two breaches in the bee's armour. I see it
squeezing the bee's stomach, compressing it with its own abdomen,
crushing it as in a vice. The brutality of this manipulation is
striking; it shows that there is no more need of care and skill. The
bee is a corpse, and a little extra pushing and squeezing will not
deteriorate its quality as food, provided there is no effusion of blood;
and however rough the treatment, I have never been able to discover the
slightest wound.

These various manipulations, above all the compression of the throat,
lead to the desired result: the honey in the stomach of the bee ascends
to the mouth. I see the drops of honey welling out, lapped up by the
glutton as soon as they appear. The bandit greedily takes in its mouth
the extended and sugared tongue of the dead insect; then once more it
presses the neck and the thorax, and once more applies the pressure of
its abdomen to the honey-sac of the bee. The honey oozes forth and is
instantly licked up. This odious meal at the expense of the corpse is
taken in a truly sybaritic attitude: the Philanthus lies upon its side
with the bee between its legs. This atrocious meal lasts often half an
hour and longer. Finally the exhausted corpse is abandoned; regretfully,
it seems, for from time to time I have seen the ogre return to the feast
and repeat its manipulation of the body. After taking a turn round the
top of the bell-glass the robber of the dead returns to the victim,
squeezes it once more, and licks its mouth until the last trace of honey
has disappeared.

The frantic passion of the Philanthus for the honey of the bee is
betrayed in another fashion. When the first victim has been exhausted I
have introduced a second bee, which has been promptly stabbed under the
chin and squeezed as before in order to extract its honey. A third has
suffered the same fate without appeasing the bandit. I have offered a
fourth, a fifth; all are accepted. My notes record that a Philanthus
sacrificed six bees in succession before my eyes, and emptied them all
of honey in the approved manner. The killing came to an end not because
the glutton was satiated, but because my functions as provider were
becoming troublesome; the dry month of August leaves but few insects in
the flowerless garden. Six bees emptied of their honey--what a
gluttonous meal! Yet the famishing creature would doubtless have
welcomed a copious addition thereto had I had the means of furnishing
it!

We need not regret the failure of bees upon this occasion; for what I
have already written is sufficient testimony of the singular habits of
this murderer of bees. I am far from denying that the Philanthus has
honest methods of earning its living; I see it among the flowers, no
less assiduous than the rest of the Hymenoptera, peacefully drinking
from their cups of nectar. The male, indeed, being stingless, knows no
other means of supporting himself. The mothers, without neglecting the
flowers as a general thing, live by brigandage as well. It is said of
the Labba, that pirate of the seas, that it pounces upon sea-birds as
they rise from the waves with captured fish in their beaks. With a blow
of the beak delivered in the hollow of the stomach, the aggressor forces
the victim to drop its prey, and promptly catches it as it falls. The
victim at least escapes with nothing worse than a blow at the base of
the neck. The Philanthus, less scrupulous, falls upon the bee, stabs it
to death and makes it disgorge in order to nourish herself upon its
honey.

Nourish, I say, and I do not withdraw the expression. To support my
statement I have better reasons than those already presented. In the
cages in which various predatory Hymenoptera whose warlike habits I am
studying are confined, waiting until I have procured the desired
prey--not always an easy proceeding--I have planted a few heads of
flowers and a couple of thistle-heads sprinkled with drops of honey,
renewed at need. On these my captives feed. In the case of the
Philanthus the honeyed flowers, although welcomed, are not
indispensable. It is enough if from time to time I place in the cage a
few living bees. Half a dozen a day is about the proper allowance. With
no other diet than the honey extracted from their victims I keep my
specimens of Philanthus for a fortnight and three weeks.

So much is plain: in a state of freedom, when occasion offers, the
Philanthus must kill on her own account as she does in captivity. The
Odynerus asks nothing of the Chrysomela but a simple condiment, the
aromatic juice of the anal pouch; the Philanthus demands a full diet, or
at least a notable supplement thereto, in the form of the contents of
the stomach. What a hecatomb of bees must not a colony of these pirates
sacrifice for their personal consumption, to say nothing of their stores
of provisions! I recommend the Philanthus to the vengeance of apiarists.

For the moment we will not look further into the original causes of the
crime. Let us consider matters as we know them, with all their real or
apparent atrocity. In order to nourish herself the Philanthus levies
tribute upon the crop of the bee. This being granted, let us consider
the method of the aggressor more closely. She does not paralyse its
captives according to the customary rites of the predatory insects; she
kills them. Why? To the eyes of understanding the necessity of a sudden
death is as clear as day. Without eviscerating the bee, which would
result in the deterioration of its flesh considered as food for the
larvae; without having recourse to the bloody extirpation of the stomach,
the Philanthus intends to obtain its honey. By skilful manipulation, by
cunning massage, she must somehow make the bee disgorge. Suppose the bee
stung in the rear of the corselet and paralysed. It is deprived of
locomotion, but not of vitality. The digestive apparatus, in particular,
retains in full, or at least in part, its normal energies, as is proved
by the frequent dejections of paralysed victims so long as the intestine
is not emptied; a fact notably exemplified by the victims of the Sphex
family; helpless creatures which I have before now kept alive for forty
days with the aid of a little sugared water. Well! without therapeutic
means, without emetics or stomach-pumps, how is a stomach intact and in
good order to be persuaded to yield up its contents? That of the bee,
jealous of its treasure, will lend itself to such treatment less readily
than another. Paralysed, the creature is inert; but there are always
internal energies and organic resistances which will not yield to the
pressure of the manipulator. In vain would the Philanthus gnaw at the
throat and squeeze the flanks; the honey would not return to the mouth
as long as a trace of life kept the stomach closed.

Matters are different with a corpse. The springs relax; the muscles
yield; the resistance of the stomach ceases, and the vessels containing
the honey are emptied by the pressure of the thief. We see, therefore,
that the Philanthus is obliged to inflict a sudden death which
instantly destroys the contractile power of the organs. Where shall the
deadly blow be delivered? The slayer knows better than we, when she
pierces the victim beneath the chin. Through the narrow breach in the
throat the cerebral ganglions are reached and immediate death ensues.

The examination of these acts of brigandage is not sufficient in view of
my incorrigible habit of following every reply by another query, until
the granite wall of the unknowable rises before me. Although the
Philanthus is skilled in forcing the bee to disgorge, in emptying the
crop distended with honey, this diabolical skill cannot be merely an
alimentary resource, above all when in common with other insects she has
access to the refectory of the flowers. I cannot regard her talents as
inspired solely by the desire of a meal obtained by the labour of
emptying the stomach of another insect. Something must surely escape us
here: the real reason for emptying the stomach. Perhaps a respectable
reason is concealed by the horrors I have recorded. What is it?

Every one will understand the vagueness which fills the observer's mind
in respect of such a question as this. The reader has the right to be
doubtful. I will spare him my suspicions, my gropings for the truth, and
the checks encountered in the search, and give him the results of my
long inquiry. Everything has its appropriate and harmonious reason. I am
too fully persuaded of this to believe that the Philanthus commits her
profanation of corpses merely to satisfy her appetite. What does the
empty stomach mean? May it not--Yes!--But, after all, who knows? Well,
let us follow up the scent.

The first care of the mothers is the welfare of the family. So far all
we know of the Philanthus concerns her talent for murder. Let us
consider her as a mother. We have seen her hunt on her own account; let
us now watch her hunt for her offspring, for the race. Nothing is
simpler than to distinguish between the two kinds of hunting. When the
insect wants a few good mouthfuls of honey and nothing else, she
abandons the bee contemptuously when she has emptied its stomach. It is
so much valueless waste, which will shrivel where it lies and be
dissected by ants. If, on the other hand, she intends to place it in the
larder as a provision for her larvae, she clasps it with her two
intermediate legs, and, walking on the other four, drags it to and fro
along the edge of the bell-glass in search of an exit so that she may
fly off with her prey. Having recognised the circular wall as
impassable, she climbs its sides, now holding the bee in her mandibles
by the antennae, clinging as she climbs to the vertical polished surface
with all six feet. She gains the summit of the glass, stays for a little
while in the flask-like cavity of the terminal button or handle, returns
to the ground, and resumes her circuit of the glass and her climbing,
relinquishing the bee only after an obstinate attempt to escape with it.
The persistence with which the Philanthus retains her clasp upon the
encumbering burden shows plainly that the game would go straight to the
larder were the insect at liberty.

Those bees intended for the larvae are stung under the chin like the
others; they are true corpses; they are manipulated, squeezed, exhausted
of their honey, just as the others. There is no difference in the method
of capture nor in their after-treatment.

As captivity might possibly result in a few anomalies of action, I
decided to inquire how matters went forward in the open. In the
neighbourhood of some colonies of Philanthidae I lay in wait, watching
for perhaps a longer time than the question justified, as it was already
settled by what occurred in captivity. My scrupulous watching at various
times was rewarded. The majority of the hunters immediately entered
their nests, carrying the bees pressed against their bodies; some halted
on the neighbouring undergrowth; and these I saw treating the bee in the
usual manner, and lapping the honey from its mouth. After these
preparations the corpse was placed in the larder. All doubt was thus
destroyed: the bees provided for the larvae are previously carefully
emptied of their honey.

Since we are dealing with the subject, let us take the opportunity of
inquiring into the customs of the Philanthus in a state of freedom.
Making use of her victims when absolutely lifeless, so that they would
putrefy in the course of a few days, this hunter of bees cannot adopt
the customs of certain insects which paralyse their prey, and fill their
cellars before laying an egg. She must surely be obliged to follow the
method of the Bembex, whose larva receives, at intervals, the necessary
nourishment; the amount increasing as the larva grows. The facts confirm
this deduction. I spoke just now of the tediousness of my watching when
watching the colonies of the Philanthus. It was perhaps even more
tedious than when I was keeping an eye upon the Bembex. Before the
burrows of _Cerceris tuberculus_ and other devourers of the weevil, and
before that of the yellow-winged Sphex, the slayer of crickets, there
is plenty of distraction, owing to the busy movements of the community.
The mothers have scarcely entered the nest before they are off again,
returning quickly with fresh prey, only to set out once more. The going
and coming is almost continuous until the storehouse is full.

The burrows of the Philanthus know nothing of such animation, even in a
populous colony. In vain my vigils prolonged themselves into whole
mornings or afternoons, and only very rarely does the mother who has
entered with a bee set forth upon a second expedition. Two captures by
the same huntress is the most that I have seen in my long watches. Once
the family is provided with sufficient food for the moment the mother
postpones further hunting trips until hunting becomes necessary, and
busies herself with digging and burrowing in her underground dwelling.
Little cells are excavated, and I see the rubbish from them gradually
pushed up to the surface. With that exception there is no sign of
activity; it is as though the burrow were deserted.

To lay the nest bare is not easy. The burrow penetrates to a depth of
about three feet in a compact soil; sometimes in a vertical, sometimes
in a horizontal direction. The spade and pick, wielded by hands more
vigorous but less expert than my own, are indispensable; but the conduct
of the excavation is anything but satisfactory. At the extremity of the
long gallery--it seems as though the straw I use for sounding would
never reach the end--we finally discover the cells, egg-shaped cavities
with the longer axis horizontal. Their number and their mutual
disposition escape me.

Some already contain the cocoon--slender and translucid, like that of
the Cerceris, and, like it, recalling the shape of certain
homoeopathic phials, with oval bodies surmounted by a tapering neck.
By the extremity of the neck, which is blackened and hardened by the
dejecta of the larvae, the cocoon is fixed to the end of the cell without
any other support. It reminds one of a short club, planted by the end of
the handle, in a line with the horizontal axis of the cell. Other cells
contain the larva in a stage more or less advanced. The grub is eating
the last victim proffered; around it lie the remains of food already
consumed. Others, again, show me a bee, a single bee, still intact, and
having an egg deposited on the under-side of the thorax. This bee
represents the first instalment of rations; others will follow as the
grub matures. My expectations are thus confirmed; as with Bembex, slayer
of Diptera, so Philanthus, killer of bees, lays her egg upon the first
body stored, and completes, at intervals, the provisioning of the cells.

The problem of the dead bee is elucidated; there remains the other
problem, of incomparable interest--Why, before they are given over to
the larvae, are the bees robbed of their honey? I have said, and I
repeat, that the killing and emptying of the bee cannot be explained
solely by the gluttony of the Philanthus. To rob the worker of its booty
is nothing; such things are seen every day; but to slaughter it in order
to empty its stomach--no, gluttony cannot be the only motive. And as the
bees placed in the cells are squeezed dry no less than the others, the
idea occurs to me that as a beefsteak garnished with _confitures_ is not
to every one's taste, so the bee sweetened with honey may well be
distasteful or even harmful to the larvae of the Philanthus. What would
the grub do if, replete with blood and flesh, it were to find under its
mandibles the honey-bag of the bee?--if, gnawing at random, it were to
open the bees stomach and so drench its game with syrup? Would it
approve of the mixture? Would the little ogre pass without repugnance
from the gamey flavour of a corpse to the scent of flowers? To affirm or
deny is useless. We must see. Let us see.

I take the young larvae of the Philanthus, already well matured, but
instead of serving them with the provisions buried in their cells I
offer them game of my own catching--bees that have filled themselves
with nectar among the rosemary bushes. My bees, killed by crushing the
head, are thankfully accepted, and at first I see nothing to justify my
suspicions. Then my nurslings languish, show themselves disdainful of
their food, give a negligent bite here and there, and finally, one and
all, die beside their uncompleted meal. All my attempts miscarry; not
once do I succeed in rearing my larvae as far as the stage of spinning
the cocoon. Yet I am no novice in my duties as dry-nurse. How many
pupils have passed through my hands and have reached the final stage in
my old sardine-boxes as well as in their native burrows! I shall draw no
conclusions from this check, which my scruples may attribute to some
unknown cause. Perhaps the atmosphere of my cabinet and the dryness of
the sand serving them for a bed have been too much for my nurslings,
whose tender skins are used to the warm moisture of the subsoil. Let us
try another method.

To decide positively whether honey is or is not repugnant to the grubs
of the Philanthus was hardly practicable by the method just explained.
The first meals consisted of flesh, and after that nothing in
particular occurred. The honey is encountered later, when the bee is
largely consumed. If hesitation and repugnance were manifested at this
point they came too late to be conclusive; the sickness of the larvae
might be due to other causes, known or unknown. We must offer honey at
the very beginning, before artificial rearing has spoilt the grub's
appetite. To offer pure honey would, of course, be useless; no
carnivorous creature would touch it, even were it starving. I must
spread the honey on meat; that is, I must smear the dead bee with honey,
lightly varnishing it with a camel's-hair brush.

Under these conditions the problem is solved with the first few
mouthfuls. The grub, having bitten on the honeyed bee, draws back as
though disgusted; hesitates for a long time; then, urged by hunger,
begins again; tries first on one side, then on another; in the end it
refuses to touch the bee again. For a few days it pines upon its
rations, which are almost intact, then dies. As many as are subjected to
the same treatment perish in the same way.

Do they simply die of hunger in the presence of food which their
appetites reject, or are they poisoned by the small amount of honey
absorbed at the first bites? I cannot say; but, whether poisonous or
merely repugnant, the bee smeared with honey is always fatal to them; a
fact which explains more clearly than the unfavourable circumstances of
the former experiment my lack of success with the freshly killed bees.

This refusal to touch honey, whether poisonous or repugnant, is
connected with principles of alimentation too general to be a
gastronomic peculiarity of the Philanthus grub. Other carnivorous
larvae--at least in the series of the Hymenoptera--must share it. Let us
experiment. The method need not be changed. I exhume the larvae when in a
state of medium growth, to avoid the vicissitudes of extreme youth; I
collect the bodies of the grubs and insects which form their natural
diet and smear each body with honey, in which condition I return them to
the larvae. A distinction is apparent: all the larvae are not equally
suited to my experiment. Those larvae must be rejected which are
nourished upon one single corpulent insect, as is that of the Scolia.
The grub attacks its prey at a determined point, plunges its head and
neck into the body of the insect, skilfully divides the entrails in
order to keep the remains fresh until its meal is ended, and does not
emerge from the opening until all is consumed but the empty skin.

To interrupt the larva with the object of smearing the interior of its
prey with honey is doubly objectionable; I might extinguish the
lingering vitality which keeps putrefaction at bay in the victim, and I
might confuse the delicate art of the larva, which might not be able to
recover the lode at which it was working or to distinguish between those
parts which are lawfully and properly eaten and those which must not be
consumed until a later period. As I have shown in a previous volume, the
grub of the Scolia has taught me much in this respect. The only larvae
acceptable for this experiment are those which are fed on a number of
small insects, which are attacked without any special art, dismembered
at random, and quickly consumed. Among such larvae I have experimented
with those provided by chance--those of various Bembeces, fed on
Diptera; those of the Palaris, whose diet consists of a large variety of
Hymenoptera; those of the Tachytus, provided with young crickets; those
of the Odynerus, fed upon larvae of the Chrysomela; those of the
sand-dwelling Cerceris, endowed with a hecatomb of weevils. As will be
seen, both consumers and consumed offer plenty of variety. Well, in
every case their proper diet, seasoned with honey, is fatal. Whether
poisoned or disgusted, they all die in a few days.

A strange result! Honey, the nectar of the flowers, the sole diet of the
apiary under its two forms and the sole nourishment of the predatory
insect in its adult phase, is for the larva of the same insect an object
of insurmountable disgust, and probably a poison. The transfiguration of
the chrysalis surprises me less than this inversion of the appetite.
What change occurs in the stomach of the insect that the adult should
passionately seek that which the larva refuses under peril of death? It
is no question of organic debility unable to support a diet too
substantial, too hard, or too highly spiced. The grubs which consume the
larva of the Cetoniae, for example (the Rose-chafers), those which feed
upon the leathery cricket, and those whose diet is rich in nitrobenzine,
must assuredly have complacent gullets and adaptable stomachs. Yet these
robust eaters die of hunger or poison for no greater cause than a drop
of syrup, the lightest diet imaginable, adapted to the weakness of
extreme youth, and a delicacy to the adult! What a gulf of obscurity in
the stomach of a miserable worm!

These gastronomic experiments called for a counter-proof. The
carnivorous grub is killed by honey. Is the honey-fed grub, inversely,
killed by carnivorous diet? Here, again, we must make certain
exceptions, observe a certain choice, as in the previous experiments. It
would obviously be courting a flat refusal to offer a heap of young
crickets to the larvae of the Anthophorus and the Osmia, for example; the
honey-fed grub would not bite such food. It would be absolutely useless
to make such an experiment. We must find the equivalent of the bee
smeared with honey; that is, we must offer the larva its ordinary food
with a mixture of animal matter added. I shall experiment with albumen,
as provided by the egg of the hen; albumen being an isomer of fibrine,
which is the principal element of all flesh diet.

_Osmia tricornis_ will lend itself to my experiment better than any
other insect on account of its dry honey, or bee-bread, which is largely
formed of flowery pollen. I knead it with the albumen, graduating the
dose of the latter so that its weight largely exceeds that of the
bee-bread. Thus I obtain pastes of various degrees of consistency, but
all firm enough to support the larva without danger of immersion. With
too fluid a mixture there would be a danger of death by drowning.
Finally, on each cake of albuminous paste I install a larva of medium
growth.

This diet is not distasteful; far from it. The grubs attack it without
hesitation and devour it with every appearance of a normal appetite.
Matters could not go better if the food had not been modified according
to my recipes. All is eaten; even the portions which I feared contained
an excessive proportion of albumen. Moreover--a matter of still greater
importance--the larvae of the Osmia fed in this manner attain their
normal growth and spin their cocoons, from which adults issue in the
following year. Despite the albuminous diet the cycle of evolution
completes itself without mishap.

What are we to conclude from all this? I confess I am embarrassed. _Omne
vivum ex ovo_, says the physiologist. All animals are carnivorous in
their first beginnings; they are formed and nourished at the expense of
the egg, in which albumen predominates. The highest, the mammals, adhere
to this diet for a considerable time; they live by the maternal milk,
rich in casein, another isomer of albumen. The gramnivorous nestling is
fed first upon worms and grubs, which are best adapted to the delicacy
of its stomach; many newly born creatures among the lower orders, being
immediately left to their own devices, live on animal diet. In this way
the original method of alimentation is continued--the method which
builds flesh out of flesh and makes blood out of blood with no chemical
processes but those of simple reconstruction. In maturity, when the
stomach is more robust, a vegetable diet may be adopted, involving a
more complex chemistry, although the food itself is more easily
obtained. To milk succeeds fodder; to the worm, seeds and grain; to the
dead or paralysed insects of the natal burrow, the nectar of flowers.

Here is a partial explanation of the double system of the Hymenoptera
with their carnivorous larvae--the system of dead or paralysed insects
followed by honey. But here the point of interrogation, already
encountered elsewhere, erects itself once again. Why is the larva of
the Osmia, which thrives upon albumen, actually fed upon honey during
its early life? Why is a vegetable diet the rule in the hives of bees
from the very commencement, when the other members of the same series
live upon animal food?

If I were a "transformist" how I should delight in this question! Yes, I
should say: yes, by the fact of its germ every animal is originally
carnivorous. The insect in particular makes a beginning with albuminoid
materials. Many larvae adhere to the alimentation present in the egg, as
do many adult insects also. But the struggle to fill the belly, which is
actually the struggle for life, demands something better than the
precarious chances of the chase. Man, at first an eager hunter of game,
collected flocks and became a shepherd in order to profit by his
possessions in time of dearth. Further progress inspired him to till the
earth and sow; a method which assured him of a certain living. Evolution
from the defective to the mediocre, and from the mediocre to the
abundant, has led to the resources of agriculture.

The lower animals have preceded us on the way of progress. The ancestors
of the Philanthus, in the remote ages of the lacustrian tertiary
formations, lived by capturing prey in both phases--both as larvae and as
adults; they hunted for their own benefit as well as for the family.
They did not confine themselves to emptying the stomach of the bee, as
do their descendants to-day; they devoured the victim entire. From
beginning to end they remained carnivorous. Later there were fortunate
innovators, whose race supplanted the more conservative element, who
discovered an inexhaustible source of nourishment, to be obtained
without painful search or dangerous conflict: the saccharine exudation
of the flowers. The wasteful system of living upon prey, by no means
favourable to large populations, has been preserved for the feeble
larvae; but the vigorous adult has abandoned it for an easier and more
prosperous existence. Thus the Philanthus of our own days was gradually
developed; thus was formed the double system of nourishment practised by
the various predatory insects which we know.

The bee has done still better; from the moment of leaving the egg it
dispenses completely with chance-won aliments. It has invented honey,
the food of its larvae. Renouncing the chase for ever, and becoming
exclusively agricultural, this insect has acquired a degree of moral and
physical prosperity that the predatory species are far from sharing.
Hence the flourishing colonies of the Anthophorae, the Osmiae, the Eucerae,
the Halicti, and other makers of honey, while the hunters of prey work
in isolation; hence the societies in which the bee displays its
admirable talents, the supreme expression of instinct.

This is what I should say if I were a "transformist." All this is a
chain of highly logical deductions, and it hangs together with a certain
air of reality, such as we like to look for in a host of "transformist"
arguments which are put forward as irrefutable. Well, I make a present
of my deductive theory to whosoever desires it, and without the least
regret; I do not believe a single word of it, and I confess my profound
ignorance of the origin of the twofold system of diet.

One thing I do see more clearly after all my experiments and research:
the tactics of the Philanthus. As a witness of its ferocious feasting,
the true motive of which was unknown to me, I treated it to all the
unfavourable epithets I could think of; called it assassin, bandit,
pirate, robber of the dead. Ignorance is always abusive; the man who
does not know is full of violent affirmations and malign
interpretations. Undeceived by the facts, I hasten to apologise and
express my esteem for the Philanthus. In emptying the stomach of the bee
the mother is performing the most praiseworthy of all duties; she is
guarding her family against poison. If she sometimes kills on her own
account and abandons the body after exhausting it of honey, I dare not
call her action a crime. When the habit has once been formed of emptying
the bee's crop for the best of motives, the temptation is great to do so
with no other excuse than hunger. Moreover--who can say?--perhaps there
is always some afterthought that the larvae might profit by the
sacrifice. Although not carried into effect the intention excuses the
act.

I therefore withdraw my abusive epithets in order to express my
admiration of the creature's maternal logic. Honey would be harmful to
the grubs. How does the mother know that honey, in which she herself
delights, is noxious to her young? To this question our knowledge has no
reply. But honey, as we have seen, would endanger the lives of the
grubs. The bees must therefore be emptied of honey before they are fed
to them. The process must be effected without wounding the victim, for
the larva must receive the latter fresh and moist; and this would be
impracticable if the insect were paralysed on account of the natural
resistance of the organs. The bee must therefore be killed outright
instead of being paralysed, otherwise the honey could not be removed.
Instantaneous death can be assured only by a lesion of the primordial
centre of life. The sting must therefore pierce the cervical ganglions;
the centre of innervation upon which the rest of the organism is
dependent. This can only be reached in one way: through the neck. Here
it is that the sting will be inserted; and here it is inserted in a
breach in the armour no larger than a pin's head. Suppress a single link
of this closely knit chain, and the Philanthus reared upon the flesh of
bees becomes an impossibility.

That honey is fatal to larvae is a fact pregnant with consequences.
Various predatory insects feed their young with honey-makers. Such, to
my knowledge, are the _Philanthus coronatus_, Fabr., which stores its
burrows with the large Halictus; the _Philanthus raptor_, Lep., which
chases all the smaller Halictus indifferently, being itself a small
insect; the _Cerceris ornata_, Fabr., which also kills Halictus; and the
_Polaris flavipes_, Fabr., which by a strange eclecticism fills its
cells with specimens of most of the Hymenoptera which are not beyond its
powers. What do these four huntresses, and others of similar habits, do
with their victims when the crops of the latter are full of honey? They
must follow the example of the Philanthus or their offspring would
perish; they must squeeze and manipulate the dead bee until it yields up
its honey. Everything goes to prove as much; but for the actual
observation of what would be a notable proof of my theory I must trust
to the future.





Next: THE GREAT PEACOCK OR EMPEROR MOTH

Previous: THE SISYPHUS BEETLE - THE INSTINCT OF PATERNITY



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