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.





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